A cat among the dogs: leopard Panthera pardus diet in a human-dominated landscape in western Maharashtra, India
←
→
Page content transcription
If your browser does not render page correctly, please read the page content below
A cat among the dogs: leopard Panthera pardus diet
in a human-dominated landscape in western
Maharashtra, India
VIDYA ATHREYA, MORTEN ODDEN, JOHN D. C. LINNELL
J A G D I S H K R I S H N A S W A M Y and K . U L L A S K A R A N T H
Abstract The ecology and predator–prey dynamics of large Introduction
felids in the tropics have largely been studied in natural sys-
tems where wild ungulates constitute the majority of the
prey base. However, human-dominated landscapes can be S tudies have shown that the density of carnivores is re-
lated to the availability of prey biomass (Fuller &
Sievert, ; Carbone & Gittleman, ; Karanth et al.,
rich in potential prey for large carnivores because of the
high density of domestic animals, especially in tropical ; Khorozyan et al., ; Carbone et al., ). These
countries where pastoralism is an important livelihood analyses are based largely on studies of prey and predator
activity. We report the almost complete dependence of leo- species in natural or semi-natural ecosystems (Fuller &
pards Panthera pardus on domestic animals as prey in the Sievert, ; Carbone & Gittleman, ; Andheria et al.,
crop lands of Ahmednagar district, Maharashtra, India. ; Karanth & Nichols, ). However, more recent
From analysis of confirmed leopard scats, % of the leo- studies have found that large carnivores can also persist in
pard’s prey biomass consisted of domestic animals, with human-dominated areas (Yirga et al., ; Athreya et al.,
% consisting of domestic dogs Canis lupus familiaris ) by relying fully or partially on food resources asso-
alone. The only wild species that occurred in the leopard’s ciated with humans (Gehrt et al., ). The potential carry-
diet were rodents, small indian civet Viverricula indica, bon- ing capacity of human-dominated landscapes for large
net macaque Macaca radiata and other primates carnivores must therefore be investigated in terms of the
Semnopithecus spp., mongoose Herpestes spp., and birds. abundance and availability of domestic prey species as
Interviews conducted in households distributed random- well as wild prey (Boitani & Powell, ).
ly in the study area documented a high density of domestic The biomass of domestic animals in human-use land-
animals: adult cattle Bos taurus, calves, goats Capra aega- scapes can be higher than that of wild prey, as seen in
grus, dogs and cats Felis catus occurred at densities of , Brazil, Nepal and Kenya (Schaller, ; Seidensticker
, , and per km, respectively. Ivlev’s electivity et al., ; Mizutani, ). Anthropogenic food sources
index indicated that dogs and cats were over-represented such as garbage and pet food can also contribute to the
in the leopard’s diet, given the higher densities of goats diet of wild carnivores (Gehrt et al., ). These food re-
and cattle. The standing biomass of dogs and cats alone sources can be abundant, leading to densities of wild carni-
was sufficient to sustain the high density of carnivores at vores comparable to, or even greater than, their densities in
the study site. Our results show that the abundance of the wild. For example, densities of urban red foxes Vulpes
potential domestic prey biomass present in human-use vulpes were times higher (Bino et al., ), and of
areas supports a relatively high density of predators, black bear Ursus americanus three times higher
although this interaction could result in conflict with (Beckmann & Berger, ), in semi-urban areas than in
humans. natural habitats because of better foraging opportunities
from crops, garbage, livestock and artificial feeding.
Keywords Conflict, diet, domestic dog, human-dominated
In India several carnivore species, such as wolves Canis
landscape, leopard, Panthera pardus lupus (Jhala & Giles, ), Asiatic lions Panthera leo persica
(Vijayan & Pati, ; Meena et al., ) and striped hyae-
VIDYA ATHREYA (Corresponding author) and K. ULLAS KARANTH Wildlife nas Hyaena hyaena (Shilpi et al., ; Singh et al., ),
Conservation Society–India, Centre for Wildlife Studies, 26-2, Aga Abbas Ali also occur in human-dominated landscapes and feed on
Road, Bangalore, Karnataka 560042, India. E-mail vidya.athreya@gmail.com
livestock. Tigers Panthera tigris are also known to attack
MORTEN ODDEN Faculty of Applied Ecology and Agricultural Sciences, Hedmark livestock in and around protected areas (Sekhar, ;
University College, Evenstad, 2480 Koppang, Norway
Karanth & Gopal, ; Woodroffe et al., ). Leopards
JOHN D. C. LINNELL Norwegian Institute for Nature Research, Trondheim,
Norway
Panthera pardus are adaptable, using a variety of habitats
and feeding on a range of wild and domestic prey
JAGDISH KRISHNASWAMY Ashoka Trust for Research in Ecology and the
Environment, Bangalore, India (Seidensticker et al., ; Daniel, ; Hunter, ),
Received July . Revision requested November . which enables them to live close to human settlements
Accepted January . First published online September . (Nasik Gazetteer, ; Daniel, ). A high density of
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106A cat among the dogs 157
FIG. 1 The Ahmednagar district of
western Maharashtra, India, where
leopard Panthera pardus scats were
collected during December –April
in a human-dominated
agricultural landscape. The rectangle on
the inset indicates the location of the
main figure in India.
domestic animals (Thornton et al., ; FAO, ) could Prionailurus rubiginosus (Athreya et al., ) in the area.
therefore constitute a stable and abundant prey base, facili- No wild ungulate species were recorded in this study, nor
tating the persistence of leopard populations in human- have any been reported by the Forest Department. There
dominated landscapes far from protected conservation are various occupational groups in the area, the dominant
areas (Athreya et al., ). one being settled farmers who own land and livestock.
Although widespread occurrence of leopards across Pastoral, migratory shepherds arrive annually in the dry sea-
India has been documented (Vijayan & Pati, ; Daniel, son, with herds of sheep and goats (each herder has .
; Athreya et al., ), few studies have assessed the leo- animals), to pasture on crop-residues in the fields.
pard’s diet and availability of prey in human-dominated
landscapes (Punjabi et al., ). To improve our under-
standing of resource usage by leopards in a rural, human-
dominated landscape, we analysed the diet of leopards, Methods
and estimated prey densities and biomass, in an agricultural
Government records indicate an overall livestock density of
landscape in western Maharashtra.
head of livestock per km across the district (Livestock
Census, ). This includes domestic and feral/semi-feral
Study area dogs Canis familiaris, pigs Sus scrofa and cats, which may
also constitute part of the leopard’s diet. To get a more de-
The study was conducted in an irrigated valley ( km), tailed overview of the prey available in the study area we in-
dominated by crop lands, around the town of Akole terviewed a random sample of households of resident
(human population c. ,) in the Ahmednagar district farmers to assess the number and species of domestic ani-
of western Maharashtra, India (Fig. ). The mean popula- mals owned. The density of houses was obtained by digitally
tion density reported for Ahmednagar district in mapping all residential houses, using imagery from
was people per km (Census of India, ). Google Earth v. ... (Google, Mountain View, USA),
Approximately % of the human population in the district and then ground-truthing a sample of homesteads to
is rural, with farming of millet, sugar cane and vegetables obtain the percentage of houses (as distinct from shops,
being the major sources of livelihood. Annual rainfall schools and other buildings). The interview data and esti-
is ,–, mm. The nearest protected area is the mated density of houses were used to estimate densities of
Kalsubai Harishchandragarh Wildlife Sanctuary ( domestic animals in the study area. Although the urban
km), km from the western edge of the study area. area of Akole has a large number of pigs, they are restricted
There are no patches of natural forest within the study area. to the town area and are not present in the wider crop-land
Camera-trapping surveys have recorded leopard, striped landscape. The Wadhari people, who claim ownership of
hyaena, golden jackal Canis aureus, Bengal fox Vulpes ben- the feral pigs, state that there are at least several hundred
galensis, jungle cat Felis chaus and rusty-spotted cat pigs in the town of Akole.
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106158 V. Athreya et al.
Initial scat surveys in the region indicated that leopards derived a regression equation to calculate the relative bio-
used trails such as foot paths, edges of fields, paved roads mass of different prey species consumed, based on their
and dry stream beds. These trails were identified using relative proportions in scats. Mountain lions are similar in
Google Earth and surveyed using three methods: () size to leopards and this method has been used previously
km of trails that were marked across the entire study area for leopards (Karanth & Sunquist, ; Andheria et al.,
were walked a total of three times each during December ; Khorozyan et al., ; Odden & Wegge, ;
–April , with a -week interval between each sam- Wegge et al., ).
pling session, () randomly selected -km grid cells were The regression equation is in the form:
overlaid on a Google Earth map and a mean of . ± SD .
Y = 1.98 + 0.35x
km of road/path within each cell was searched, on foot, for
scats, and () scats were collected opportunistically during where Y is the mass of prey consumed per scat and x is the
December –April . We used all three methods mean mass of the prey. The relative biomass (D) and the
because we lacked a priori information on where to locate relative numbers of each prey species consumed (E) were
leopard scats in a human-dominated landscape. The scats obtained using the equations
represent a sample from the dry season.
D = (A × Y)/S(A × Y) × 100
Two trained surveyors walked on either side of the trail
and collected all scats judged to be of carnivore origin, based E = (D/x)/S(D/x) × 100
on size, shape and ancillary evidence such as scrape signs where A is the frequency of occurrence of the prey item
and tracks. The scats were measured and stored in zip-lock in the scats. The prey biomass (Bprey = Dprey · Wprey;
bags, and the geographical coordinates of each location were Khorozyan et al., ) in the study area was calculated
recorded using a geographical positioning system. The scats using density estimates (Dprey) for the four most common
were later transferred to polythene bags and part of each scat prey species (domestic goats, dogs, calves and cats), based
was transferred into vials for storage in absolute alcohol for on interview data. The mean weight of the domestic animal
subsequent DNA analysis to identify leopard scats (Mondol species (Wprey) was estimated by a livestock veterinarian
et al., , ; Navya et al., unpubl. data). Visual identi- working in the region. Adult cattle were not included
fication of scats is not always reliable (Farrell et al., ), because compensation records indicate that only one cow
and therefore diet analysis was conducted only on scats con- was attacked in the study area in years, whereas calves
firmed to be of leopard origin, either using DNA methods or were preyed on in relatively higher numbers (Forest
having been collected from scrapes characteristic of those Department records).
made by large felids (as the leopard is the only large felid We used Ivlev’s () electivity index to assess prey
present in the study area). selection:
The scat analysis was carried out as described by r i − pi
Mukherjee et al. (), Mukherjee & Mishra () and E=
r i + pi
in other diet studies of large felids (Karanth & Sunquist,
; Andheria et al., ; Khorozyan et al., ; Odden where ri is the relative proportion of prey item i in the diet
& Wegge, ). The scats were washed and dried and the and pi is the relative proportion of prey item i in the envir-
prey species were identified from the presence of claws, onment. E is in the range −–+, where negative values indi-
hoofs or hair. For hair, we used a microscope to identify cate that the prey item is avoided or inaccessible and positive
the origin of randomly selected hair samples per scat. values indicate that it is selected for.
Prey species were identified based on comparison with ref-
erence slides of hair samples from domestic animals in the Results
study area and from reference slides of hair samples from
wild prey, from collections at the Centre for Wildlife Leopard scats, confirmed using DNA analysis, were found
Studies, Bangalore, and the Bombay Natural History throughout the sampled area, including close to houses,
Society, Mumbai. Scats that were highly degraded or had and on a variety of trail types, including dirt and paved
too few identifiable prey remains were not used in the roads. The mean distance from leopard scats to the nearest
analysis. house was m (range – m), and m (range
We calculated the frequencies of occurrence of the vari- – m) from roads. Two scats were found on the main
ous prey species (the percentage of the total number of scats street of Akole town.
that contained a specific prey item). However, this variable Of the scats collected were confirmed as leopard
can be misleading because smaller prey species contribute scats based on DNA analysis and were identified based
more to a scat than larger species (Karanth & Sunquist, on the presence of tracks or scrapes. Of these leopard
; Klare et al., ). Based on feeding trials on captive scats had usable remains for diet analysis. Thirty-eight
mountain lions Puma concolor, Ackerman et al. () scats could not be used because the remains were degraded
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106A cat among the dogs 159
TABLE 1 Prey species identified in the diet of leopards Panthera pardus in the Ahmednagar district of western Maharashtra, India (Fig. ),
from analysis of scats collected from a human-dominated landscape during December –April . Y is the correction factor from
Ackerman et al. ().
Frequency of
occurrence in Relative biomass Relative no. of
Prey species scats (%) Mean mass (kg) Y (kg per scat) in leopard diet (%) individuals consumed
Primate spp. 0.8 10.0 2.3 0.7 0.3
Small Indian civet 2.3 2.0 2.1 1.9 3.6
Viverricula indica
Mongoose Herpestes spp. 3.1 2.0 2.1 2.5 4.8
Pig Sus scrofa 3.1 20.0 2.7 3.2 0.6
Bird spp. 3.9 1.0 2.0 3.0 11.8
Sheep Ovis aries 5.4 30.0 3.0 6.4 0.8
Rodent spp. 6.9 0.5 2.0 5.4 42.3
Goat Capra aegagrus 10.0 25.0 2.9 11.2 1.7
Cow Bos taurus 10.8 40.0 3.4 14.2 1.4
Cat Felis catus 15.4 2.0 2.1 12.3 24.1
Dog Canis lupus 38.5 18.0 2.6 39.2 8.5
and, although they contained hair and bone, they could
not be identified visually or under the microscope. Of
intact leopard scats the mean diameter at the thickest
section was . ± SD . mm (range .–. mm). The
scats contained a total of prey items, comprising
prey species (Table ). Fifty-six percent of the scats con-
tained one prey species, % two species, % three species,
% four species and only one scat contained five species.
Domestic animals (pig, sheep, cat, dog, goat, cow) consti-
tuted % of the prey biomass consumed by leopards. The
wild prey present in the scats were civet Viverricula indica,
rodent, primate, bird and mongoose Herpestes spp.
(Table ). In the case of rodents a mean mass of g
(Table ) was considered because of the presence of the
bandicoot rat Bandicota spp. in the crop lands. Dogs FIG. 2 Ivlev’s index (Ivlev, ) for goat Capra aegagrus, calf Bos
were the most significant constituent of the leopard’s taurus, cat Felis catus and dog Canis lupus familiaris, based on
diet, accounting for % of the biomass consumed. scat analysis of leopard prey in Ahmednagar district of western
Maharashtra. The index is based on the frequency of prey species
Ivlev’s index indicates that despite the higher biomass of
in scats relative to the availability of these species in the area.
goats and calves available, dogs and cats were preyed Species with positive index values are more selected for/more
upon to a greater extent than expected, which could be available than species with negative values.
attributable to preference or greater accessibility (Fig. ;
Table ).
depredation of livestock is a widespread occurrence
Discussion (Linnell et al., ), domestic livestock usually constitute
only a small part of the diet of large felids and complete
The relationship between large felids and humans is com- dependency on domestic species has rarely been observed.
plex and the spectrum of interactions ranges from fascin- The biomass of livestock can be high in human-dominated
ation to fear (Boomgaard, ; Loveridge et al., ). areas, exceeding that of wild prey in surrounding forest areas
Large felids are often portrayed as flagship species for con- in Brazil, Nepal and Kenya (Schaller, ; Seidensticker
servation (Treves & Karanth, ) but conflict occurs at the et al., ; Mizutani, ). Domestic animals are easier
local level, where the presence of a large carnivore can result to attack because they lack anti-predatory behaviour, unlike
in damage to property and loss of human life (Treves et al., their wild counterparts (Diamond, ). However, avail-
). Retaliatory killings are a significant cause of mortal- ability does not always indicate accessibility as livestock
ity of large felids, and studies have focused on human–felid may be guarded by day and enclosed in predator-proof en-
conflict (Inskip & Zimmermann, ). Although closures at night.
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106160 V. Athreya et al.
TABLE 2 Density (km−) of domestic animals in the study area, from interviews of a random sample of households (n = ) in the town of
Akole in the Ahmednagar district of western Maharashtra (Fig. ). The interviews were conducted during September –September
.
Buffalo Cow (adult) Calf Goat Sheep Dog Cat Chicken
Mean no. per household (range) 0.11 (0–1) 2.7 (0–18) 0.9 (0–8) 2.9 (0–40) 0.06 (0–1) 0.40 (0–5) 1 (0–6) 1.8 (0–50)
Density (km−2) 7 162 54 174 4 24 61 109
Biomass (kg per km2) 2,450 56,700 2,440 4,350 120 432 108 109
Our results show that a large predator such as a leopard estimates from a camera-trap study (Athreya et al., ).
can attain relatively high densities in a rural landscape The reason for the relatively low density of leopards despite
(Athreya et al., ) while subsisting almost entirely on a the high biomass of prey is probably related to the low avail-
diet of domestic animals. Despite the density of goats ability of domestic stock because they are protected by farm-
being seven times that of domestic dogs, goats constituted ers (Athreya et al., unpubl. data) and the fact that large stock
only % of the prey biomass of leopards, whereas dogs con- are not predated by leopards. Most cattle in the study area are
stituted %. This is probably because goats are less access- hybrid varieties that are larger than the indigenous breeds.
ible than dogs, being actively herded by day and enclosed in The selection of domestic dogs and cats as prey means
sheds at night. The results show that dogs are an important that the economic impact of predation by the leopard on
food resource for leopards and they occur at high density in valuable livestock is lower than expected. Thus human–
the study area. The proclivity of leopards towards killing and leopard conflict is largely driven by people’s fear of leopards
eating dogs has been noted in anecdotal, historical literature foraging in the proximity of their houses, and the sentimen-
(Daniel, ). Two studies carried out within protected tal value of dogs as pets.
areas in the states of Maharashtra and Jammu & Kashmir Our findings are similar to those of studies on rural and
have reported the importance of dogs as prey for leopards urban carnivores in western Europe and North America,
(Edgaonkar & Chellam, ; Shah et al., ). where wild carnivores reside in modified human-dominated
Domestic dogs are ubiquitous in the Indian landscape landscapes and are totally dependent on anthropogenic
and density estimates from an adjoining human-use land- sources of food (Gehrt et al., ). Our work contributes
scape in Maharashtra range from (farmland) to to a growing awareness of the potential conservation value
km− (village area; Punjabi et al., ; Hughes & of private lands and non-protected areas in the tropics
Macdonald, ). Domestic cats also appear to be an import- (Negrões et al., ).
ant component of the leopard’s diet in our study area, contrib-
uting % of the biomass consumed. Based on interviews, the
densities of goats and cows in the study area were and Acknowledgements
km−, respectively. Thus, rural landscapes in India can
be prey-rich areas for wild carnivores because of the import- We thank Ashok Ghule, Abhijit Kulkarni, Avinash Kulkarni
ance of animal husbandry in the livelihoods of rural people. and Kiran Rahalkar for taking part in collection of scats,
In natural ecosystems predator density is correlated with Uma Ramakrishnan, R. Navya and Chandrima Home for
prey biomass (Carbone & Gittleman, ; Karanth et al., carrying out the DNA analysis and standardization of the
; Marker & Dickman, ; Beckmann & Lackey, scat DNA procedures, the Maharashtra Forest Department
; Khorozyan et al., ; Bino et al., ). If we consider for their support, and the Centre for Wildlife Studies and
the density of the four common prey species (goats, calves, the Asian Nature Conservation Foundation, Bangalore, for
cats and dogs) identified in the scat analysis, the potential institutional support.
prey biomass for leopards in the study area is , kg
per km (Table ). The regression equation of Carbone
& Gittleman () estimates that , kg of prey is re- References
quired per kg of predator, irrespective of predator size.
A C K E R M A N , B.B., L I N D Z E Y , F.G. & H E M K E R , T.P. () Cougar food
Based on this equation the total amount of prey biomass habits in southern Utah. The Journal of Wildlife Management, ,
(including goats and cattle) in our study area could, in the- –.
ory, support . times the number of leopards that are pre- A N D H E R I A , A.P., K A R A N T H , K.U. & K U M A R , N.S. () Diet and prey
sent. The biomass of the owned dogs and cats alone profiles of three sympatric large carnivores in Bandipur Tiger
constitutes c. , kg per km, which would be pre- Reserve, India. Journal of Zoology, , –.
A T H R E Y A , V., O D D E N , M., L I N N E L L , J.D.C., K R I S H N A S WA M Y , J. &
dicted to support a kg predator at a density of . individ- K A R A N T H , K.U. () Big cats in our backyards: persistence of large
uals per km, which is close to the combined density of carnivores in a human dominated landscape in India. PLoS ONE,
leopards and striped hyaenas in the study area, based on (), e.
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106A cat among the dogs 161
B E C K M A N N , J.P. & B E R G E R , J. () Rapid ecological and behavioural K A R A N T H , K.U. & N I C H O L S , J.D. () Non-invasive survey
changes in carnivores: the responses of black bears (Ursus methods for assessing tiger populations. In Tigers of the World: The
americanus) to altered food. Journal of Zoology, , –. Science, Politics and Conservation of Panthera tigris (eds R.L. Tilson
B E C K M A N N , J.P. & L AC K E Y , C.W. () Carnivores, urban & P.J. Nyhus), pp. –. Elsevier, New York, USA.
landscapes, and longitudinal studies: a case history of black bears. K A R A N T H , K.U., N I C H O L S , J.D., K U M A R , N.S., L I N K , W.A. & H I N E S ,
Human–Wildlife Conflicts, , –. J.E. () Tigers and their prey: predicting carnivore densities
B I N O , G., D O L E V , A., Y O S H A , D., G U T E R , A., K I N G , R., S A LT Z , D. & from prey abundance. Proceedings of the National Academy of
K A R K , S. () Abrupt spatial and numerical responses of Sciences of the United States of America, , –.
overabundant foxes to a reduction in anthropogenic resources. K A R A N T H , K.U. & S U N Q U I S T , M.E. () Prey selection by tiger,
Journal of Applied Ecology, , –. leopard and dhole in tropical forests. Journal of Animal Ecology, ,
B O I TA N I , L. & P O W E L L , R.A. (eds) () Carnivore Ecology and –.
Conservation: A Handbook of Techniques. Oxford University Press, K H O R O Z Y A N , I.G., M A L K H A S Y A N , A.G. & A B R A M OV , A.V. ()
New York, USA. Presence–absence surveys of prey and their use in predicting
B O O M G A A R D , P. () Frontiers of Fear: Tigers and People in leopard (Panthera pardus) densities: a case study from Armenia.
the Malay World, –. Yale University Press, New Haven, Integrative Zoology, , –.
USA. K L A R E , U., K A M L E R , J.F. & M AC D O N A L D , D.W. () A comparison
C A R B O N E , C. & G I T T L E M A N , J.L. () A common rule for the and critique of different scat-analysis methods for determining
scaling of carnivore density. Science, , –. carnivore diet. Mammal Review, , –.
C A R B O N E , C., P E T T O R E L L I , N. & S T E P H E N S , P.A. () The bigger L I N N E L L , J., O D D E N , J. & M E R T E N S , A. () Mitigation methods for
they come, the harder they fall: body size and prey abundance conflicts associated with carnivore depredation on livestock. In
influence predator–prey ratios. Biology Letters, , –. Carnivore Ecology and Conservation: A Handbook of Techniques
C E N S U S O F I N D I A () Http://censusindia.gov.in/ [accessed May (eds L. Boitani & R.A. Powell), pp. –. Oxford University Press,
]. Oxford, UK.
D A N I E L , J.C. () The Leopard in India: A Natural History. Natraj L I V E S T O C K C E N S U S () Directorate of Economics and Statistics,
Publishers, Dehradun, India. Planning Department, Government of Maharashtra, Mumbai,
D I A M O N D , J. () Evolution, consequences and future of plant and India.
animal domestication. Nature, , –. L O V E R I D G E , A.J., W A N G , S.W., F R A N K , L.G. & S E I D E N S T I C K E R , J.
D I C K M A N , A.J. () Complexities of conflict: the importance of () People and wild felids: conservation of cats and
considering social factors for effectively resolving human–wildlife management of conflicts. In The Biology and Conservation of Wild
conflict. Animal Conservation, , –. Felids (eds D.W. Macdonald & A.J. Loveridge), pp. –. Oxford
E D G AO N K A R , A. & C H E L L A M , R. () Food habit of the leopard, University Press, Oxford, UK.
Panthera pardus, in the Sanjay Gandhi National Park, Maharashtra, M A R K E R , L.L. & D I C K M A N , A.J. () Factors affecting leopard
India. Mammalia, , –. (Panthera pardus) spatial ecology, with particular reference to
FAO (F O O D A N D A G R I C U LT U R E O R G A N I Z A T I O N O F T H E U N I T E D Namibian farmlands. South African Journal of Wildlife Research, ,
N AT I O N S ) () Livestock Sector Brief: India. http://www.fao.org/ –.
ag/againfo/resources/en/publications/sector_briefs/lsb_IND.pdf M E E N A , V., J H A L A , Y.V., C H E L L A M , R. & P AT H A K , B. ()
[accessed January ]. Implications of diet composition of Asiatic lions for their
F A R R E L L , L.E., R O M A N , J. & S U N Q U I S T , M.E. () Dietary conservation. Journal of Zoology, , –.
separation of sympatric carnivores identified by molecular analysis M I Z U TA N I , F. () Biomass density of wild and domestic herbivores
of scats. Molecular Ecology, , –. and carrying capacity on a working ranch in Laikipia District,
F U L L E R , T.K. & S I E V E R T , P.R. () Carnivore demography and the Kenya. African Journal of Ecology, , –.
consequences of changes in prey availability. In Carnivore M O N D O L , S., R A M E S H , N., A T H R E YA , V., S U N A G A R , K., S E LVA R A J ,
Conservation (eds J.L. Gittleman, S.M. Funk, D.W. Macdonald & V.M. & R A M A K R I S H N A N , U. () A panel of microsatellites to
R.K. Wayne), pp. –. Cambridge University Press, individually identify leopards, and its application to leopard
Cambridge, UK. monitoring in human-dominated landscapes. BMC Genetics,
G E H R T , S.D., R I L E Y , S.P.D. & C Y P H E R , B.L. (eds) () Urban , –.
Carnivores: Ecology, Conflict and Conservation. The Johns Hopkins M O N D O L , S., T H AT T E , P., Y A D AV , P. & R A M A K R I S H N A N , U. () A
University Press, Baltimore, USA. set of mini STRs for population genetic analyses of tigers (Panthera
H U G H E S , J. & M AC D O N A L D , D.W. () A review of the interactions tigris) with cross-species amplification for seven other Felidae.
between free-roaming domestic dogs and wildlife. Biological Conservation Genetics Resources, , –.
Conservation, , –. M U K H E R J E E , S., G O Y A L , S.P. & C H E L L A M , R. () Standardisation of
H U N T E R , L. () Carnivores of the World (Princeton Field Guides). scat analysis techniques for leopard (Panthera pardus) in Gir
Princeton University Press, Princeton, USA. National Park, Western India. Mammalia, , –.
I N S K I P , C. & Z I M M E R M A N N , A. () Human–felid conflict: a review M U K H E R J E E , S. & M I S H R A , C. () Predation by leopard Panthera
of patterns and priorities worldwide. Oryx, , –. pardus in Majhatal Harsang Wildlife Sanctuary, W. Himalayas.
I V L E V , V.S. () Experimental Ecology of the Feeding of Fishes. Yale Journal of the Bombay Natural History Society, , –.
University Press, New Haven, USA. N A S I K G A Z E T T E E R () Https://cultural.maharashtra.gov.in/english/
J H A L A , Y.V. & G I L E S , R.H. () The status and conservation of the wolf gazetteer/Nasik/%General/%WildAnimalsAndBirds.
in Gujarat and Rajasthan, India. Conservation Biology, , –. htm [accessed June ].
K A R A N T H , K.U. & G O P A L , R. () An ecology-based policy N E G R Õ E S , N., R E V I L L A , E., F O N S E C A , C., S O A R E S , A.M.V.M., J Á C O M O ,
framework for human–tiger coexistence in India. In People and A.T.A. & S I LV E I R A , L. () Private forest reserves can aid in
Wildlife: Conflict Or Co-existence? (eds R. Woodroffe, S. Thirgood & preserving the community of medium and large-sized vertebrates in
A. Rabinowitz), pp. –. Cambridge University Press, the Amazon arc of deforestation. Biodiversity Conservation, , –
Cambridge, UK. .
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106162 V. Athreya et al.
O D D E N , M. & W E G G E , P. () Kill rates and food consumption V I J AYA N , S. & P AT I , B.P. () Impact of changing cropping patterns
of leopards in Bardia National Park, Nepal. Acta Theriologica, , on man–animal conflicts around Gir Protected Area with specific
–. reference to Talala Sub-District, Gujarat, India. Population and
P U N J A B I , G.A., A T H R E Y A , V. & L I N N E L L , J.D.C. () Using natural Environment, , –.
marks to estimate free-ranging dog Canis familiaris abundance in a W E G G E , P., O D D E N , M., P O K H A R E L , C.P. & S T O R A A S , T. ()
MARK–RESIGHT framework in suburban Mumbai, India. Tropical Predator–prey relationships and responses of ungulates and their
Conservation Science, , –. predators to the establishment of protected areas: a case study of
S C H A L L E R , G.B. () Mammals and their biomass on a Brazilian tigers, leopards and their prey in Bardia National Park, Nepal.
ranch. Arquivos de Zoologia, , –. Biological Conservation, , –.
S E I D E N S T I C K E R , J., S U N Q U I S T , M. & M C D O U G A L , C. () Leopards W O O D R O F F E , R., T H I R G O O D , S. & R A B I N O W I T Z , A. () The impact
living at the edge of the Royal Chitwan National Park, Nepal. In of human–wildlife conflict on natural systems. In People and
Conservation in Developing Countries: Problems and Prospects Wildlife: Conflict or Co-Existence? (eds R. Woodroffe, S. Thirgood &
(eds J.C. Daniel & J.S. Serrao), pp. –. Oxford University Press, A. Rabinowitz), pp. –. Cambridge University Press, Cambridge,
Bombay, India. UK.
S E K H A R , N.U. () Crop and livestock depredation caused by wild Y I R G A , G., D E I O N G H , H.H., L E I R S , H., G E B R I H I W OT , K., D E C K E R S , J.
animals in protected areas: the case of Sariska Tiger Reserve, & B A U E R , H. () Adaptability of large carnivores to changing
Rajasthan, India. Environmental Conservation, , –. anthropogenic food sources: diet change of spotted hyena (Crocuta
S H A H , G.M., J A N , U., B H A T , B.A., A H M A D , F. & A H M A D , J. () crocuta) during Christian fasting period in northern Ethiopia.
Food habits of the leopard Panthera pardus in Dachigam National Journal of Animal Ecology, , –.
Park, Kashmir, India. Journal of Threatened Taxa, , –.
S H I L P I , G., K R I S H N E N D U , M., S H A N K A R , K. & Q A M A R , Q. ()
Estimation of striped hyaena Hyaena hyaena population using
camera traps in Sariska Tiger Reserve, Rajasthan, India. Journal of Biographical sketches
the Bombay Natural History Society, , –.
S I N G H , P., G O P A L A S WA M Y , A.M. & K A R A N T H , K.U. () Factors V I D Y A AT H R E Y A ’s research has focused on understanding the ecol-
influencing densities of striped hyenas (Hyaena hyaena) in arid ogy of large carnivores when they reside in human-use landscapes,
regions of India. Journal of Mammalogy, , –. and translating her findings into informed management actions.
T H O R N T O N , P.K., K R U S K A , R.L., H E N N I N G E R , N., K R I S T J A N S O N , P.M., M O R T E N O D D E N has worked on large carnivores in Nepal and on
R E I D , R.S., A T I E N O , F. et al. () Mapping Poverty and Livestock human–leopard interactions in India. He is interested in predator–
in the Developing World. Http://www.ilri.org/InfoServ/Webpub/ prey interactions and in space use of large carnivores. J O H N
fulldocs/InvestAnim/Book/media/PDF_chapters/B_Front.pdf L I N N E L L conducts multi-disciplinary research on the relationships
[accessed January ]. between wildlife and humans, with a focus on large carnivores and her-
T R E V E S , A. & K A R A N T H , K.U. () Human–carnivore conflict and bivores. He works in Europe, South America, India and South-east
perspectives on carnivore management worldwide. Conservation Asia. J A G D I S H K R I S H N A S W A M Y is interested in ecohydrology, land-
Biology, , –. scape ecology and applications of Bayesian statistics in ecology and
T R E V E S , A., W A L L AC E , R.B., N A U G H T O N -T R E V E S , L. & M O R A L E S , A. environmental science. U L L A S K A R A N T H studies the ecology of
() Co-managing human–wildlife conflicts: a review. Human large carnivores, focusing on population modelling and estimation,
Dimensions of Wildlife, , –. and applies the results to practical conservation efforts.
Oryx, 2016, 50(1), 156–162 © 2014 Fauna & Flora International doi:10.1017/S0030605314000106
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 Jan 2022 at 01:30:15, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms.
https://doi.org/10.1017/S0030605314000106You can also read
