Dependence of the Leopard Panthera Pardus Fusca in Jaipur, India, on Domestic Animals

Dependence of the leopard Panthera pardus fusca in Jaipur, India, on domestic animals

S WAPNIL K UMBHOJKAR,REUVEN Y OSEF,JAKUB Z. KOSICKI P ATRYCJA K. KWIATKOWSKA and P IOTR T RYJANOWSKI

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 nowledge of prey selection and diet is important for prey base. However, in tropical countries where commu- Kunderstanding the life history strategies of carnivores nities are primarily agrarian, the high density of domestic and for planning the conservation of an apex predator  animals in human-dominated landscapes can be a potential (Miquelle et al., ). The density of carnivores is related prey source for large carnivores. We demonstrate almost to habitat quality, in particular to the availability of   complete dependence of the Vulnerable leopard Panthera prey (Fuller & Sievert, ; Carbone & Gittleman, ;   pardus fusca in the Jhalana Reserve Forest in Jaipur, north- Andheria et al., ; Karanth & Nichols, ). To enhance – west India on domestic animals as prey. We analysed  the cost benefit ratio of energetic intake, large carnivores leopard scats collected during the dry season of November may visit areas close to, or even inside, human settlements    –April . Domestic animals comprised the majority (Gehrt et al., ; Yirga et al., ; Athreya et al., ). of the leopards’ prey (.% frequency of occurrence): dogs Fear of carnivores, especially in the context of livestock dep- ’ Canis lupus familiaris (%), cats Felis catus (%), goats redation, can negatively affect people s well-being (Inskip &  Capra aegagrus hircus (%) and cattle Bos taurus (%). Zimmermann, ). It has been shown that the abundance Wild species, which occurred in the leopards’ diet at a and availability of wild and domestic prey is the predomin- relatively low frequency, were rodents, the hare Lepus ant factor that determines the potential carrying capacity of nigricollis, small Indian civet Viverricula indica, rhesus human-dominated landscapes for large carnivores (Boitani  macaque Macaca mulatta, northern plains grey langur & Powell, ). In human-dominated landscapes in Brazil,   Semnopithecus entellus and mongoose Herpestes edwardsii. Nepal and Kenya (Schaller, ; Seidensticker et al., ;  Diet is also a function of availability of potential prey, but no Mizutani, ) the biomass of potential domestic prey of data are available on the density of the leopard’s wild prey carnivores was higher than that of wild prey species. species in Jhalana Reserve Forest. Nevertheless, our results Discarded food waste and pet food may also contribute to  suggest that abundance of domestic prey around Jhalana the food resources of carnivores (Gehrt et al., ). This re- ’ Reserve Forest sustains the c.  known leopards. We con- duces carnivores fear of humans and, in consequence, the clude that these leopards, by preying on feral dogs in an density of carnivores in urban or semi-urban areas can be  urban environment, could be considered as suppliers of a higher than in the wild (Butler et al., ). service to the human population amongst whom they thrive, In India, the intrusion of large predators into urban although this potentially exposes the leopards to the canine areas is well documented. Wolves Canis lupus (Jhala &  distemper virus. Giles, ), Asiatic lions Panthera leo persica (Meena et al., ), striped hyaenas Hyaena hyaena (Singh et al., Keywords Diet, environmental service, human-dominated ) and tigers Panthera tigris (Karanth & Gopal, ) landscape, India, Jhalana, leopard, Panthera pardus fusca are known to attack livestock and persist in human- dominated landscapes. The leopard Panthera pardus fusca also lives successfully in the proximity of people (Athreya et al., ). Its broad diet includes amphibians, arthropods

SWAPNIL KUMBHOJKAR Jhalana Wildlife Research Foundation, Pune, India and carrion, reducing dependence on water sources, which  REUVEN YOSEF (Corresponding author, orcid.org/0000-0003-4331-9866) Ben is obtained from the prey (Daniel, ; Kshettry et al., Gurion University of the Negev-Eilat Campus, P. O. Box 272, 88106 Eilat, Israel ). Compared to larger felids, the small body size of E-mail [email protected] the leopard reduces the territory required to sustain a popu- JAKUB Z. KOSICKI Department of Avian Biology & Ecology, Faculty of Biology, lation and allows it to survive and thrive in proximity to Adam Mickiewicz University, Poznań, Poland people (Daniel, ; Kshettry et al., ). ń PATRYCJA K. KWIATKOWSKA Institute of Zoology, Pozna University of Life To examine resource utilization by leopards in a rural, Sciences, Poznań, Poland semi-urban human-dominated landscape, we analysed the PIOTR TRYJANOWSKI Faculty of Environmental Sciences, Czech University of Life Sciences Prague, Prague, Czech Republic diet of leopards of the Jhalana Reserve Forest. Because the Received  June . Revision requested  July . Reserve is in close proximity to and surrounded by Accepted  September . First published online  October . human habitations, leopards have little fear of people and

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloaded fromdistribution, https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.202.126 the original work, on is 27 properly Sep 2021 cited. at 09:46:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2021, 55(5),. https://doi.org/10.1017/S0030605319001145 692–698 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001145 Leopard dependence on domestic animals 693

FIG. 1 Jhalana Reserve Forest in north-west India, and the surrounding city of Jaipur, with the locations of the leopard Panthera pardus fusca scats collected.

often feed opportunistically on easily accessible domestic felids (the leopard is the only large felid present in the prey (Athreya et al., , ). We thus hypothesized Reserve). Trained volunteers wearing gloves used forceps that the leopards’ diet comprises principally of domestic to collect the scats. A small portion of each scat was left, animals. Our objectives were to identify the leopards’ key so as to not disrupt the natural markings of the leopards prey species and examine their importance for human– (Schwarz & Fischer, ). The location of each scat was leopard coexistence and acceptance of the presence of leo- noted, with a GPS, at the time of collection. Scats were pards in the urban landscape (Kumbhojkar et al., ). stored in numbered polythene, zip-lock bags. Preliminary observations such as the presence of bones, claws, skin and other biological remains were noted if appropriate. Study area Highly degraded scats (n = ; .%) were excluded from  the analysis and only well-preserved scats (n = ) were ana- The  km Jhalana Reserve Forest lies on the south-east lysed. They were washed under running water and sundried. border of the city of Jaipur in north-west India (Fig. ). It The cuticular and medullar patterns of any hair remains were was designated on  November  in accordance with observed and photographed under a compound microscope. the provisions of Rajasthan Forest Act . The Reserve Scat analysis was based on Mukherjee et al. (), Mukherjee has a mean altitude of  m, with higher elevations in the &Mishra(), diet-related studies (Karanth & Sunquist, north in the form of low, flat-topped hills, and is character- ; Biswas & Sankar, ; Sankar & Johnsingh, ; ized by tropical dry deciduous forest. During the s Andheria et al., ; Khorozyan et al., ;Odden& the main valley was planted with the native species Acacia Wegge, ), and our own collections of prey remains. All tortilis and Acacia senegal. hair, hooves, claws, teeth, nails and bones were separated for further analysis. Prey species were identified based on com- Methods parison with reference slides of hair samples from domestic animals in the study area and from reference slides of hair The population of leopards in the Reserve has been estimated, samples from wild prey and from previous studies (Oli, using camera traps and recognition of individuals, to com- ;Tiwari,). prise  individuals (authors, unpubl. data). In a  survey To assess if the sample size was sufficient for accurate of the Reserve we found that leopards used trails and tourist diet analysis (Edgaonkar & Chellam, ;Kshettryetal., routes for defecation. We monitored the trails, collecting  ) we applied the rarefaction method, implemented scat samples in the dry season of November –April  in EstimateS . (Colwell, ). This method estimates (Kumbhojkar et al., ; Fig. ). the expected cumulative number of species, with % The identity of leopard scats was confirmed using their confidence intervals (the Mao Tau estimator; Colwell, occurrence in scrapes characteristic of those made by large ).

Oryx, 2021, 55(5), 692–698 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001145 Downloaded from https://www.cambridge.org/core. IP address: 170.106.202.126, on 27 Sep 2021 at 09:46:14, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319001145 694 S. Kumbhojkar et al.

 FIG. 2 Per cent of leopard scats collected in Jhalana Reserve FIG. 3 Cumulative number of prey species in leopard scats (solid   Forest (Fig. ) that contained various prey species/taxa (Table ). line), with % confidence interval (shaded area).

We calculated the relative biomass of prey species con- (Zar, ). All calculations were conducted in R .. sumed using the regression equation Y = .X + .Y, (R Core Team, ). where Y is the mass of prey consumed per scat and X is the mean mass of the prey (cf. Ramesh et al., ). The Results regression equation is derived based on feeding trials with captive mountain lions Puma concolor (Ackerman et al., The  scats had a mean of . ± SE . (.–.%CI)  ) to calculate relative biomass based on their relative prey species. Most of the scats (; %) were found along- proportions in scats. As mountain lions and leopards are side trails or jeep tracks. Of the , hairs selected from similar in size, this method has been used previously for scats,  (%) were unidentified. The scats contained a   leopards (Karanth & Sunquist, ; Andheria et al., ; total of  prey taxa:  individual species and rodents,   Khorozyan et al., ; Odden & Wegge, ; Wegge which were not identified to species level (Fig. ).   et al., ; Athreya et al., ). Seventy-nine per cent () of the scats contained only one The frequencies of occurrence of prey species (the per prey species; % contained two or more species. The cent of the total number of scats that contained a specific individual-based rarefaction curve shows that the increase prey item) were calculated but, as smaller prey species may in the cumulative number of species in scats was rapid for contribute more to scats than larger species (Karanth & the first c.  scats but began to level off thereafter (Fig. ).   Sunquist, ; Klare et al., ), this variable may poten- In the  leopard scats the majority (.%) of prey fre- tially be misleading. The relative biomass (D) was therefore quency of occurrence comprised domestic animals (domestic calculated using: dogs Canis lupus familiaris, %; domestic cats Felis catus, %; goats Capra aegagrus hircus, %; cattle Bos taurus, AY D = × 100 %) and wild prey comprised % of prey frequency of occur- AY rence; the difference was significant (G test; G = ., where A is the frequency of occurrence of the prey species. P , .). The mean biomass of domestic animal species The relative per cent of each prey species consumed (E)was detected in the scats was . and that of wild prey was . obtained using: kg; the difference was significant (t = .,P=.; Table ).

D Discussion E = X × 100 D In urban landscapes the relationship between large felids X and humans is complex, influenced by people’s fear and Because we found that the correlations between Y, D, and awe (Boomgaard, ; Loveridge et al., ). Aversion  E (R .–.) were all significant at P , ., we only occurs when the presence of a large felid results in damage tested differences in biomass (D) between domestic and to property or loss of human life (Treves et al., ), with wild prey species by t test with Cochran–Cox adjustment retaliatory killings resulting in significant felid mortality

TABLE 1 Mean weight, per cent frequency, mass consumed per scat, relative biomass consumed and relative per cent of prey, in decreasing order of frequency, for each of the  prey taxa recorded in the  scat samples of the leopard Panthera pardus fusca.

Taxon Weight1 (kg) Frequency (%) Y2 (kg) D2 (kg) E2 (%) Domestic dog Canis lupus familiaris 18.00 44.0 8.28 2.96 37.16 Domestic goat Capra aegagrus hircus 25.00 16.0 10.73 1.50 18.82 Domestic cattle (calf) Bos taurus 40.00 15.0 15.98 2.14 26.93 Domestic cat Felis catus 2.00 13.0 2.68 0.30 3.84 Indian grey mongoose Herpestes edwardsii 2.00 3.0 2.68 0.05 0.69 Rodents (Rodentia) 0.10 3.0 2.01 0.05 0.63 Nilgai Boselaphus tragocamelus 140.00 2.2 50.98 0.76 9.64 Hare Lepus nigricollis 2.10 1.5 2.71 0.02 0.35 Domestic pig Sus scrofa domesticus 21.00 1.5 9.33 0.09 1.20 Indian civet Viverricula indica 0.08 1.5 2.01 0.02 0.25 Langur Semnopithecus entellus 8.00 0.7 4.78 0.02 0.28 Rhesus macaque Macaca mulatta 7.00 0.7 4.43 0.02 0.25

 From Karanth & Sunquist (), Biswas & Sankar (), Sankar & Johnsingh () and Andheria et al. ().  Y, mass of prey consumed per scat; D, relative biomass; E, relative per cent of each prey species consumed (see text for further details).

the leopards. Hence, it is unsurprising that leopards move between the protected area and human habitat, despite the  m high barrier built around Jhalana to separate it from the city of Jaipur (Fig. ). There are no data available on the densities or numbers of the leopards’ wild prey species in Jhalana Reserve Forest. There are, however, data on the vaccination and sterilization of feral domestic dogs undertaken for the past  decades by the Help in Suffering Foundation, Jaipur (J. Reece, pers comm., ): a mean of c. , rabies inoculations were conducted annually over the past  years, to protect residents in the case of bites from feral dogs. Previous studies have shown that the availability of a large PLATE 1 Leopard Panthera pardus fusca preying on a domestic number of domestic animals in rural and urban areas facil- dog that strayed into the boundaries of Jhalana Reserve Forest. itates the survival of leopards near human habitats (Athreya Photo: Devam Shah. et al., ; Kshettry et al., ). Persistence of leopard populations in human-dominated habitats beyond protected (Inskip & Zimmermann, ). Studies have therefore areas is dependent on a stable and abundant domestic focused on so-called human–felid conflict (Inskip & prey base (Athreya et al., ). Zimmermann, ). Dependence of felids on domestic The Mao Tau diversity estimator indicated that – livestock has rarely been observed, however; domestic live- scat samples sufficed to give a representative sample of the stock usually contribute only a small proportion of the species included in the diet of the leopards (Fig. ). Biswas biomass consumed by felids (Athreya et al., ). In urban & Sankar () considered that a minimum of  scats landscapes the biomass of domestic animals is usually high needed to be analysed to examine the prey base of tigers. and usually exceeds that of wild prey in the neighbouring Scat analysis substantiated our hypothesis that the pre- forests (Schaller, ). So, if domestic animals lack anti- dominant prey of leopards in Jhalana Reserve Forest are do- predatory behaviour, they are more susceptible to attacks mestic animals, presumably from the neighbouring city and (Diamond, ). Jhalana Reserve Forest is surrounded by villages. Our results corroborate the results of Daniel () urban and rural areas with c.  million inhabitants (World and demonstrate that dogs are a major food resource for leop- Population Review, ). People living in the surrounding ards in the Reserve (Plate ). Scat analysis from the states of neighbourhoods and villages practice traditional livelihood Maharashtra and Jammu and Kashmir have also reported professions such as livestock farming. Goats and the calves of the importance of dogs as prey for leopards (Edgaonkar & cattle are available as leopard prey along with domestic dogs, Chellam, ; Shah et al., ). In West Bengal, however, cats and pigs. An estimated population of , domestic livestock (cattle and goats; %) predominated in the diet of dogs was documented in a  survey in Jaipur (Hiby leopards (Kshettry et al., ), whereas cattle and goats com- et al., ). This is a sizeable source of potential prey for prised % of the diet in Jhalana Reserve Forest. In Ayubia

National Park, Pakistan, the leopard similarly subsisted main- to the human population amongst whom they thrive, al- ly on domestic animals, with goats predominating (.%) the though this potentially exposes the leopards to the canine frequency of occurrence (Shehzad et al., ). distemper virus. The differences between our results and other studies are a consequence of the high number of dogs and cats in our Acknowledgements We thank Sudharshan Sharma, Deputy Conservator of Forests, for his guidance, the Rajasthan Forest study area. The total dog and cat populations in the area are Department for permission to conduct research on the leopards of     estimated to be c. , and , individuals, respectively Jhalana and for collecting scat and biological samples; Pratiksha (J. Reece, pers. comm., ); these species are thus widely Vedpathak, Benazir Bagwan, Aseem Shendye and Abhay Shendye available. Cattle and goats are relatively less abundant (%) from Swasti Agro and Bio-Products Pvt. Ltd. for technical support because these animals are herded during the day and pro- and assistance with scat analysis; Satish Pande, Satish Karmalkar, Pramod Deshpande, Sanjay Khatavkar and Rajkumar Pawar of Ela tected in enclosures at night. Similarly, in Sanjay Gandhi Foundation for provision of equipment and laboratory facilities for National Park, Mumbai, leopards have adapted to the de- scat analysis; Rahul Lonkar; Bablu Gurjar, Abhishek Gurjar, Dayal pleted wild ungulate prey base by feeding on dogs found Gurjar, Shubham Saini, Varad Bansod and Hrishikesh Wagh for in the surrounding areas and by scavenging on buffalo car- help with field surveys, scat collection and washing of the scats; G.V. casses (Edgaonkar & Chellam, ). Reddy, Head of Forest Office, Rajasthan Forest Department arranged In spite of the higher frequency of goats in the leopard training at Sanjay Gandhi National Park, Mumbai, for scat analysis; Vidhya Athreya, Nikit Surve and his team of volunteers for training scats we analysed, the relative biomass of cattle was higher, in the field; and Abhinav Mehta for drafting Fig. 1. a result of the greater weight of cattle. A similar finding was made in the study of leopard diet in Sri Lanka (Mukenhirn Author contributions Study design, fieldwork: SK, RY; data ana- & Eisenberg, ). In our study the leopards were probably lyses, writing: all authors. feeding on discarded carcasses of calves as the Rajasthan Conflicts of interest None. Forest Department does not have any records of claims for compensation for predation of cattle. Ethical standards This research do not involve handling of animals The fact that leopards prey on the most easily available and otherwise abided by the Oryx guidelines on ethical standards. species (the most numerous), has already been demonstrated SK and RY worked under research permit 171 of the Rajasthan Forest (Schaller, ; Seidensticker, ;Kshettryetal.). With Department. a diet dominated by domestic dogs and cats rather than by domestic livestock, the potential economic impact of the References leopards of Jhalana Reserve Forest is lower compared to areas  where predation of livestock is high and there is a consequent ACKERMAN, B.B., LINDZEY, F.G. & HEMKER,T.P.( ) Cougar food habits in southern Utah. Wildlife Magazine, , –. lack of tolerance for such damage (cf. Margulies & Karanth, ANDHERIA, A.P., KARANTH, K.U. & KUMAR, N.S. () Diet and ). In addition a large proportion of the rural population prey profiles of three sympatric large carnivores in Bandipur surrounding Jhalana Reserve Forest are Jains, who practise Tiger Reserve, India. Journal of Zoology, , –. the conservation of biodiversity, and especially of large felids ATHREYA, V.R., THAKUR, S.S., CHAUDHURI,S.&BELSARE, A.V.  (Athreya et al., ); this facilitates human–leopard coexis- ( ) Leopards in human-dominated areas: a spillover from  sustained translocations into nearby forests. Journal of Bombay tence (Kumbhojkar et al., ). Natural History Society, , –. Domestic dogs are a public health problem in India and ATHREYA, V.R., THAKUR, S.S., CHAUDHURI,S.&BELSARE, A.V. elsewhere (Reece & Chawla, ) and occasionally attack () A Study of the Man–Leopard Conflict in the Junnar Forest other small domestic animals (Srinivasan, ). For people, Division, Pune District, Maharashtra. Submitted to the Office of the dog bites can result in infection, disfigurement, incapacity, Chief Wildlife Warden, Maharashtra State Forest Department, and the Wildlife Protection Society of India, New Delhi, India. post-traumatic stress, and even death, and bites from in-  .  projectwaghoba.in/docs/junnar_conflict_report_athreya_et_ _ fected dogs account for % of rabies cases (Seligsohn, condensed.pdf [accessed January ]. ; Taylor et al., ). Only % of the total dog population ATHREYA, V., ODDEN, M., LINNELL, J.D.C., KRISHNASWAMY,J.& in the study area are vaccinated (J. Reece, pers. comm., ; KARANTH, K.U. () Big cats in our backyards: persistence of large Reece & Chawla ). Dogs could also transfer infectious carnivores in a human dominated landscape in India. PLOS ONE,   diseases to leopards. Canine distemper virus is the second ,e . 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