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World Journal of Zoology 7 (2): 106-112, 2012 ISSN 1817-3098 © IDOSI Publications, 2012 DOI: 10.5829/idosi.wjz.2012.7.2.63139

Food Habits of Golden ( aureus) and Striped ( hyaena) in Sariska Reserve, Western

Pooja Chourasia Krishnendu Mondal, K. Sankar and Qamar Qureshi

Wildlife Institute of India, Dehradun, India . Abstract: Food habits of (Canis aureus) and (Hyaena hyaena) were investigated using scat analysis between November 2010 and June 2011 in , . Total 104 scats of golden jackal and 86 scats of striped hyena were collected and analyzed. The frequency of occurrence of each prey was estimated through bootstrapping using program SIMSTAT. Niche breadth of these two species was quantified using Levin’s measure. The diet overlap in the two species was assessed using Pianka’s index. Twelve food items were identified in golden jackal scats and nine in striped hyena scats. Vegetative matter contributed maximum (17.57%) in jackal’s diet followed by (15.77%), (10.81%), sambar (5.41%) and (4.05%). Nilgai and domestic contributed maximum (24.76% each) in the diet of striped hyena, followed by sambar (17.14%), chital (16.19%) and vegetative matter (10.48%). The estimated dietary overlap between striped hyena and golden jackal was 67%. Niche breadth for golden jackal was estimated as 0.69 and for striped hyena it was 0.57. The considerable overlap was attributed to mutual dependence on , which indicated high resource competition between the two species.

Key words: Canis aureus Dietary Niche Overlap Food Habits Hyaena hyaena Prey Size Scat

INTRODUCTION between predator and prey sizes. A confounding factor in predator-prey studies that include a wide size-range of The extent of and resource mammalian predators is the high variability in diet of partitioning determines the degree to which different smaller species, which can switch between insectivore, species can either coexist or competitively exclude each and . Canid and felid species above other [1, 2]. An important mode of resource partitioning is certain threshold weight class predate purely on the degree of dietary overlap between sympatric species vertebrates, while those below may be omnivorous or [3]. This overlap is influenced by each species’ physical predate on invertebrates and vertebrates. This implies that ability to obtain food [4, 5] and also by variation in the large (>21.5 kg) constitute a distinct functional spatial and temporal availability of food [6]. For distinct group from which predator-prey size relationships should carnivore guilds different mechanisms may be involved, emerge more clearly than from carnivore assemblages that such as different body sizes of predators [7] and prey are distributed across both sides of the body mass species [8, 9], activity pattern [10] and microhabitat use threshold [14]. The diet of the golden jackal and the [11]. However, whereas resources are limited, the strength striped hyena was studied through scat analysis in India of competition between sympatric species generally [15-18] and other countries [19-22]. Merve et al. [23] decreases with increased resource partitioning [12]. studied diet overlap between (Parahyaena Study by Dickman [13] showed that the smaller brunnea) and -backed jackal (Canis mesomelas) in predators in an insectivorous guild consumed Atherstone Nature Reserve, Pilanesberg National Park smaller prey, but switched over to larger prey when the and private (Mankwe Reserve) in South larger predators were removed. Consequently prey . They inferred that, despite the marked body size selection could depend more on competition among differences, brown hyena and black-backed jackal can be predator guild members than on any inherent relationship considered as either meso-carnivores or apex predators,

Corresponding Author: K. Sankar, Wildlife Institute of India. P.O. Box 18, Chandrabani, Dehradun-248 001, (), India. Mob: +91 9412992784. 106 World J. Zool., 7 (2): 106-112, 2012 depending on the presence or absence of larger carnivores and such as tiger, , jungle carnivores [24]. As meso-predators, the potential food ( chaus), common ( availability increases as dietary items previously edwardsii), small Indian mongoose (H. auropunctatus), unattainable, are killed by apex predators and (H. smithi), common palm interspecific competition could decline concomitantly. ( hermaphroditus), The diet of black backed jackal comprises high proportion (Viverricula indica), ratel (Mellivora capensis) and prey of small and large , corroborating that both species like chital (Axis axis), sambar (Rusa unicolor), scavenge and hunt competently [25]. Brown , nilgai (Boselaphus tragocamelus), common langur on the other hand, are predominant [26]. (Semnopithecus entellus), wild pig (Sus scrofa), Thus, may be excluded from carcass sites by (Hystrix indica), rufous-tailed (Lepus nigricollis brown hyenas, eventually forcing them to hunt more ruficaudatus) and Indian (Pavo cristatus)[30]. frequently [27]. There are 32 villages within Sariska TR of which six are Both striped hyena (Hyaena hyaena) and golden located in the intensive study area of 160 km2 . A large jackal (Canis aureus) are found in semi-arid zone and number of brahmini cattle (Bos indicus), buffaloes tropical dry and moist of the country (Bubalus bubalis), (Capra hircus) and (Ovis and their diet ranges from wild ungulates to domestic aries) are kept by people living in villages. and rodents. Although numerous studies have investigated their diet separately, only a few studies [23] Scat Analysis: Estimation of food habits of golden jackal have investigated the overlap and resource partitioning, and striped hyena was done using scat analysis. Scats which may vary with the presence or absence of apex were collected on transects, trails and roads whenever predators like tiger ( ) and leopard encountered within the intensive study area. Each (Panthera pardus). In the present study, we assessed 1) collected scat sample was labeled with the name of the the relative contribution of different prey species to the species, date and location. The collected scats were sun diet of golden jackal and striped hyena and 2) the dietary dried and later broken down and washed under running overlap between these two sympatric species. In the water through a sieve. The scat contents were teased present study we estimated the food habits of striped apart and remain of different food items such as hair, hyena (Hyaena hyaena) and golden jackal (Canis aureus) feather, scales of , invertebrate remains and and compared the dietary overlap between them in Sariska vegetable matter (grass and fruit seeds) were separated. Tiger Reserve, Western India from November 2010 to June The frequency of occurrence of each food item was 2011. estimated through bootstrapping using program SIMSTAT [31]. This technique measures 95% confidence MATERIALS AND METHODS limits of the proportion of each food item in the diet, the limits of which range measure the random sampling errors Study Area: The study was conducted in Sariska Tiger [19]. Niche breadth of the two species was quantified Reserve (Sariska TR) (27°05'-27°33' N; 76°15'-76°35' E) using Levin’s measure [26] to measure the uniformity of between November 2010 and June 2011. Sariska TR is resources being utilized by each species. The overlap in situated in the Aravalli Hill Range in the semi-arid part of diet of the two species was assessed using Pianka’s [32] Western India [28]. The total area of the Tiger Reserve is index. 881 km22 , of which 273.8 km is notified National Park. The vegetation of Sariska corresponds to Northern tropical RESULTS dry deciduous and Northern tropical thorn forests [29]. Open areas are covered with scrub forests dominated by Altogether 104 scats of golden jackal and 86 scats of shrubs such as Zizyphus nummularia, Capparis sepiaria, striped hyena were collected and analyzed. Hair samples Capparis decidua, Adathoda vasica, from scats were identified from their histological structure and Acacia sp. The valleys are dominated by Zizyphus using microscope. In total 12 food items were identified in mauritiana mixed , gentle slopes are dominated by golden Jackal scats and nine in striped hyena scats. The Anogeissus pendula forest and the steep slopes are frequency of occurrence and percentage frequency of occupied by Boswellia serrata forest. Other than golden occurrence of each food item found in golden jackal and jackal and striped hyena, the Park supports various striped hyena scats are given in table 1.

107 World J. Zool., 7 (2): 106-112, 2012

Table 1: Frequency of occurrence and percentage frequency occurrence of each food item found in golden jackal and striped hyena scats in Sariska Tiger Reserve, Western India between November 2010 and June 2011 Golden jackal Striped hyena ------Food item Frequency of occurrence (±SD) Percentage frequency of occurrence Frequency of occurrence (±SD) Percentage frequency of occurrence Scales of reptiles 4±1.9 1.80 - - Vegetation 37.5±4.7 17.57 12.8±3.4 10.48 Fruit 19.1±3.8 9.01 - - 15.5±3.6 7.21 2.3±1.5 1.90 Insect 15.3±3.8 7.21 - - 34±4.5 15.77 1.2±1.1 0.95 Hare 8.9±2.9 4.05 1.2±1.1 0.95 Sambar 11.3±3.1 5.41 20.9±4.4 17.14 Chital 23.1±3.9 10.81 19.8±4.0 16.19 Nilgai 8.7±2.8 4.05 30.2±4.8 24.76 Cattle 32.9±4.5 15.32 30.2±4.8 24.76 3.7±1.9 1.80 3.5±2.1 2.86

Vegetative matter contributed maximum (17.57%) in jackal predating on goats in the study area but cattle golden jackal’s diet followed by rodents (15.77%), cattle by them was not recorded. The result of scat Bos indicus 15.32%), chital (10.81%), fruits (9.01%), analysis supports the typical diet generalist of golden (7.21%), sambar(5.41%), hare (4.05%), nilgai (4.05%), goat jackal, feeding on a variety of food source available to (1.80%) and scales (1.80%). No attempt was made them in the study area. to identify the rodent and reptile species in golden Out of 86 striped hyena scats, three scats had three jackal’s scat. Out of 104 golden jackal scats, one had six food items; 40 scats had two food items and 43 scats had food items, one had five food items, ten had four food single food items. Nilgai and cattle (Bos indicus) items, 23 had three food items, 34 had two food items and contributed maximum (24.76%) in the diet of striped hyena 35 had single food item. Zizyphus mauritiana seeds were followed by sambar (17.14%), chital (16.19%), vegetation the only fruit species found in golden jackal’s diet. (10.48%), goat (2.86%), unidentified bird (1.90%), hare Beetles (Class: Insecta), centipedes (Class: Chilopoda), (0.95%) and unidentified rodent (0.95%). The remains of millipedes (Class: Diplopoda) in golden jackal’s diet were larger ungulates such as sambar, nilgai and chital were identified on the basis of locomotory organs, i.e. shiny observed more in scat samples as compared to the elytra of beetles and round or oval shields of centipedes/ remains of goat, bird, rodent and hare. The above millipedes. Feathers of (Francolinus mentioned large ungulates are also the major food items pondicerianus) and peafowl (Pavo cristatus) were of tiger and leopard in the study area [33, 34]. This may identified in golden jackal’s diet. Narrow ventral scales lead to the inference that the occurrence of larger and translucent round scales were also identified to be of ungulates in hyena diet is perhaps the result of (Calotes sp.) and skinks (: Scincidae) but not scavenging. at species level. The estimated dietary overlap between striped hyena In some incidences (n=9) golden jackals were and golden jackal was 67% (Table 2). Niche breadth for found to prey upon chital fawns in the study area and this golden jackal was estimated as 0.69 and for striped hyena may be attributed to higher occurrence of chital in their it was 0.57 (Table 3). MacArthur and Levin’s [35] niche scats. Golden Jackal was never found to prey on sambar overlap was calculated as an overlap measure of species or nilgai (adult or fawn) in the study period, but they A on species B and vice-versa. The niche overlap of scavenge on sambar and nilgai kills made by tiger or golden jackal on striped hyena was 66% (0.66) i.e. out of leopard in the study area, which is reflected in lower total diet range of striped hyena, 66% of food items are occurrence in their scats. Golden Jackal were also found common with golden jackal and the niche overlap of scavenging on cattle carcasses at the dump sites outside striped hyena on golden jackal was 47% (0.47) i.e. out of villages in the study area, which is attributed to higher total diet range of golden Jackal, 47% of food items are occurrence of cattle in their diet. There were records of common with striped hyena.

108 World J. Zool., 7 (2): 106-112, 2012

Table 2: Dietary overlap between golden jackal and striped hyena as shown by scat analysis in Sariska Tiger Reserve, Western India between November 2010 and June 2011 Percentage frequency of Percentage frequency of

22 Food Item occurrence (striped hyena) (PiA) occurrence (golden jackal) (PiB) PiA*P iB (C) PiA (D) PiB (E) Vegetation 10.48 17.57 184.04 109.75 308.62 Bird 1.90 7.21 13.73 3.63 51.94 Rodent 0.95 15.77 15.02 0.91 248.56 Hare 0.95 4.05 3.86 0.91 16.44 Sambar 17.14 5.41 92.66 293.88 29.22 Chital 16.19 10.81 175.03 262.13 116.87 Nilgai 24.76 4.05 100.39 613.15 16.44 Cattle 24.76 15.32 379.24 613.15 234.56 Goat 2.86 1.80 5.15 8.16 3.25 Scales of reptiles 0.00 1.80 0.00 0.00 3.25 Fruit 0.00 9.01 0.00 0.00 81.16 969.11(F) 1905.67 (G) 1110.30 (H) 43.65 (I) 33.32 (J) 0.67 (O)

22 PiA-% of food item i in the diet of striped hyena; PiB-% of food item i in the diet of golden jackal; C = PiA * P iB; D = PiA ; E = PiB ; F= C; G = D and H = E; I = G0.5 ; J = H 0.5 , O = F/ (I*J).

Table 3: Dietary niche breadth between Golden Jackal and Striped Hyena as broader niches would be more closely linked to shown by scat analysis in Sariska Tiger Reserve November 2010 characteristics of favorable elements (cover or food) and to June 2011 the landscapes surrounding an element, because the Variable Golden jackal12Striped hyena combined effects of large size and a broad niche on the n 12 10 species would impose fewer constraints on its element B 8.604 6.157 and landscape use compared to smaller, more specialized 0.691 0.573 species [36]. In Sariska TR, golden jackals are found to be M MacArthur and Levins measure 0.663 0.475 confined in the valley areas where a range of smaller prey species as well as fruiting trees (Zizyphus sp.) are DISCUSSION available, while distribution of striped hyena was wider including valley and hilly areas [37]. In the In Sariska TR, the striped hyena occupies all the present study, broader dietary niche was observed for major types, compared to golden jackal that is golden jackal as compared to striped hyena, because mainly distributed in the open scrub land and valley golden jackal assumed to compensate their limited habitat. There are predictions that body size and distribution by utilizing broader spectrum of food. behavioral flexibility would influence the response of Many studies have explained dietary differences predators and this response would be evidenced in within the context of alternative feeding strategies (i.e. individual use of habitat elements and in the spatial generalist vs. specialist foraging) or intrinsic niche distribution of predators [36]. Specifically, larger species differentiation (i.e. disparity in habitat requirements or (i.e. striped hyena) would be more equitably distributed body size) [38-41]. However, not all species fall into such among all spatial habitat elements compared to smaller discrete categories [42]. Indeed, within a given species (i.e. golden jackal), because large size would community, the relative breadth of a carnivore’s diet (and confer an increased ability to traverse the habitat matrix by extension levels of interspecific dietary overlap) is best with minimal risk of predation. Similarly, species with a defined in relation to the feeding habits manifested by its broader niche would be more equitably distributed among tropical counterparts [43]. However, when prey spatial habitat elements because broader diet breadth abundance fluctuates dramatically over a period of time, would allow a species to use the majority of spatial even alleged specialist species may exhibit more generalist elements and exploit a wider range of resources. feeding patterns [44]. Moreover, a review of dietary Additionally, presence of larger species and species with breadth among herbivorous insects revealed that while

109 World J. Zool., 7 (2): 106-112, 2012 generalization characterized the diet of many insect presence of apex predators like tiger and leopard. This species, the dietary choices of individuals within suggests that niche partitioning in protected areas in the particular populations were often specialized (according presence of an results in a form of to the traditional framework described above), suggesting competitive exclusion of jackal at carcasses by hyena, that in many cases dietary breadth may be a local (i.e. resulting in a greater level of by the jackals, system-specific) phenomenon rather than an immutable connatural to the study in Africa [23]. species characteristic [42]. The considerable overlap Review of the studies showed marked interspecific (67%) characterizing the diets of striped hyena and golden difference in plant-matter consumption by golden jackal. jackal, may be attributed to mutual dependence on Whereas, plants did not play an important role in the ungulates, which intern indicates that resource feeding of golden jackals in [20], compared with competition between these two species may be high. the feeding of jackals in southern areas of [48]. In the However, striped hyena relied more on large ungulates, present study vegetative matter i.e. grass and Zizyphus whereas golden jackal preyed largely on small ungulates fruits comprised a higher proportion 26.58% in golden and rodents; these dietary specializations may have jackal’s diet. alleviated overall levels of competition between two Seasonal of small mammals was species. This pattern of exclusive use of a particular prey reported in golden jackal’s diet [49-51]. In the present items, despite the overall dietary similarity, characterizes study no attempt was made to quantify the seasonal diet the relationships between sympatric Canis of golden jackal. Study by Lanszki et al. [20] on feeding latrans and gray velox [45] and coyotes and habits and tropical niche overlap between golden jackal red foxes Vulpes vulpes [46]. (Canis aureus) and red (Vulpes vulpes) in the Depending on food availability, golden jackals may Pannonia eco-region, Hungary revealed that functional be solitary hunters, co-operating in pairs or hunting as response of the golden jackal to a limited but favored groups (mainly when the parents teach the offspring to food item was more rapid than that of the . hunt). The differences in large mammal and small mammal The golden jackal shifted from small prey to other food proportions between the diets resulted in a significant items earlier when the availability of small mammals difference between golden jackal and striped hyena diets. declined and also returned to rodent hunting earlier than However, at 67%, the niche overlap between the species did the red fox. Long term ecological studies including was still high. Result of the present study is comparable seasonal food habits on golden jackal and striped hyena with the studies by Nowell and Jackson [46] and Walton are much desired across different landscapes in the and Joly [25] on black-backed jackal (Canis mesomelas) in country. Africa. Studies by Owens and Owens [26, 27] on brown hyenas showed that they are predominantly scavengers ACKNOWLEDGEMENTS and black-backed jackals are generally excluded from carcass sites by the much larger brown hyenas, forcing We thank Forest Department for them to hunt more frequently. The similar case may likely facilitation of this work in Sariska, as a part of ‘Ecology of happen in Sariska TR where striped hyenas occur in high Leopard’ research project conducted by the Wildlife density i.e. 24.5 individual/ 100 sq. km [47]. Study by Institute of India. We thank Director and Dean, WII, for Merwe et al. [23] on assessment of diet overlap between their encouragement and support provided for the study. brown hyena and black-backed jackal suggested that in We thank our field assistants Jairam, Omprakash and black backed jackal’s diet, more large mammals were Ramesh for their assistance in field. consumed outside protected areas whereas more small mammals were consumed inside protected areas in South REFERENCES Africa’s North West Province. This observation supports the fact that golden jackal are being excluded from 1. Pianka, E.R., 1973. The structure of scavenging sites and forced to actively hunt when communities. Annual Review of Ecology and competing with striped hyena in the present study. The , 4: 53-74. low frequency of smaller prey species in the diet of the 2. Carvalho, J.C. and P. Gomes, 2004. Feeding resource striped hyena may reflect sufficient carrion resources, as partitioning among four sympatric carnivores in the most samples originated in the study area where the Peneda-Gereˆ s National Park (). J. of Zool. hyenas are relegated to meso-carnivore status by the (Lond.), 263: 275-283.

110 World J. Zool., 7 (2): 106-112, 2012

3. Hayward, M.W. and G.I.H. Kerley, 2008. Prey 16. Mukherjee, S., S.P. Goyal, A.J.T. Johnsingh and preferences and dietary overlap amongst Africa’s M.R.P.L. Pitman. 2004. The importance of rodents in large predators. South African Journal of Wildlife the diet of (Felis chaus), (Caracal Research, 38: 93-108. caracal) and Golden Jackal (Canis aureus) in Sariska 4. Radloff, F.G.T. and J.T. Du Toit, 2004. Tiger Reserve, Rajasthan, India. Journal of Zoology, Large predators and their prey in a southern 262: 405-411. African : a predator’s size determines its 17. Aiyadurai, A. and Y.V. Jhala, 2006. Foraging and prey size range. Journal of Ecology, habitat use by Golden Jackal (Canis aureus) in the 73: 410-423. Bhal Region, , India. J. Bombay Nat. Hist. 5. Owen-Smith, N. and M.G.L. Mills, 2008. Shifting prey Soc., 103(1): 5-12. selection generates contrasting herbivore dynamics 18. Singh, P., 2008. and feeding within a large-mammal predator-prey web. Ecology, ecology of striped hyena (Hyaena hyaena) in relation 89: 1120-1133. to land-use patterns in an arid region of Rajasthan. 6. Azevedo, F.C.C., V. Lester, W. Gorsuch, S. Lariviere, M.Sc. dissertation submitted to Manipal University, A.J. Wirsing and D.L. Murray, 2006. Dietary NCBS, Banglore, pp: 56. breadth and overlap among five sympatric 19. Reynolds, J.C. and H.S. Aebischer, 1991. Comparison prairie carnivores. Journal of Zoology, , and quantification of carnivore diet by faecal 269: 127-135. analysis: a critique, with recommendations, based 7. Carvalho, J.C. and P. Gomes, 2004. Feeding resource on a study of Fox (Vulpes vulpes). Mammal Review, partitioning among four sympatric carnivores in 21: 97-122. the Peneda-Gereˆ s National Park (Portugal). J. of 20. Lanszki, J., M. Heltai and L. Szabo, 2006. Feeding habits and trophic niche overlap between sympatric Zool. (Lond.), 263: 275-283. golden jackal (Canis aureus) and red fox (Vulpes 8. Karanth, K.U. and M.E. Sunquist, 2000. Behavioral vulpes) in the Pannonian ecoregion (Hungary). Can. correlates of predation by tiger (Panthera tigris), J. Zool., 84: 1647-1656. leopard (Panthera pardus) and (Cuon alpinus) 21. Giannatos, G., Y. Marinos, P. Maragou and in Nagarahole, India. J. Zool. (Lond.), 250: 255-265. G. Catsadorakis, 2005. The status of the golden jackal 9. Juarez, K.M. and J. Marinho-Filho, 2002. Diet, habitat (Canis aureus L.) in Greece. Belg. J. Zool., 134: 37-42. use and home ranges of sympatric canids in central 22. Yom-Tov, Y., S. Ashkenazi and O. Viner, 1995. Cattle Brazil. J. Mammal., 83: 925-933. predation by the golden jackal Canis aureus in the 10. Loveridge, A.J. and D.W. MacDonald, 2003. Niche , . Biol. Conserv, 73: 19-22. separation in sympatric jackals (Canis mesomelas 23. Merwe, I.V.D., C.J. Tambling, M. Thorn, D.M. Scott, and Canis adustus). J. Zool. (Lond.), 259: 143-153. R.W. Yarnell, M. Green, E.Z. Cameron and P.W. 11. Johnson, W.E. and W.L. Franklin, 1994. Spatial Bateman, 2009. An assessment of diet overlap of resource partitioning by sympatric grey fox two mesocarnivores in the North West Province, (Dusicyon griseus) and fox (Dusicyon . African Zoology, 44(2): 288-291. culpaeus) in southern Chile. Can. J. Zool., 24. Ritchie, E.G. and C.N. Johnson, 2009. Predator 72: 1788-1793. interactions, mesopredator release and biodiversity 12. Pacala, S.W. and J. Roughgarden, 1985. Population conservation. Ecology Letters, 12: 982-998. experiments with the Anolis lizards of St. Maarten 25. Walton, L.R. and D.O. Joly, 2003. Canis mesomelas. and St. Eustatius. Ecology, 66: 129-141. Mammalian Species, 715: 1-9. 13. Dickman, C.R., 1988. Body size, prey size and 26. Owens, M.J. and D.D. Owens, 1978b. Feeding community structure in insectivorous mammals. ecology and its influence on social organization in Ecology, 69: 569-580. brown hyenas (Hyanea brunnea, Thunberg) of the 14. Ray, J.C. and M.E. Sunquist, 2001. Trophic relations central . East African Wildlife Journal, in a community of African carnivores. 16: 113-135. Oecologia, 127: 395-408. 27. Owens, M.J. and D.D. Owens, 1978a. Feeding 15. Sankar, K., 1988. Some observations on food habits ecology and its influence on social organization of jackals (Canis aureus) in Keolaeo National Park, in brown hyena (Hyaena brunnea, Thunberg) of the Bharatpur, as shown by scat analysis. J. Bombay. central Kalahari Desert. East African Wildlife Journal, Nat. Hist. Soc., 85: 185-186. 16: 113-135.

111 World J. Zool., 7 (2): 106-112, 2012

28. Rodgers, W.A. and H.S. Panwar, 1988. Planning a 40. Brown, J.H., 1975. Geographical ecology of desert Wildlife Network in India. Vol. I, rodents. In Ecology and of communitie, Wildlife Institute of India, Dehradun, pp: 341. pp: 315-341. 29. Champion, H.G. and S.K. Seth, 1968. A revised 41. Tokeshi, M., 1999. Species coexistence. Ecological survey of forest types of India. Manager of and evolutionary perspectives. UK: Blackwell Publications, , New Delhi, Science. pp: 404. 42. Fox, L.R. and P.A. Morrow, 1981. Specialization: 30. Sankar, K., Q. Qureshi, K. Mondal, D. Worah, species property or local phenomenon? Science, T. Shrivastava, S. Gupta and S. Basu, 2009. Ecological 211: 887-893. studies in Sariska Tiger Reserve, Final report 43. Azevedo, F.C.C., V. Lester, W. Gorsuch, S. Lariviere, submitted to National Authority, A.J. Wirsing and D.L. Murray, 2006. Dietary breadth Gov. of India, New Delhi. Wildlife Institute of India, and overlap among five sympatric prairie carnivores. Dehradun, pp: 145. Journal of Zoology, London, 269: 127-135. 31. Efron, B. and R. Tibshirani, 1986. Bootstrap methods 44. O’Donoghue, M., S. Boutin, C.J. Krebs, G. Zutla, D.L. for standard errors, confidence intervals and other Murray and E.J. Hofer, 1998. Functional responses measures of statistical accuracy. Statistical Science, of coyotes and to the snowshoe hare cycle. 1: 54-77. Ecology, 79: 1209-1222. 32. Pianka, E.R., 1981. Competition and niche theory. 45. Kitchen, A.M., E.M. Gese and E.R. Schauster, Theoretical ecology: principles and applications 1999. Resource partitioning between coyotes (R.M. May, eds). Blackwell Scientific Publications, and swift foxes: space, time and diet. Can. J. Oxford, United Kingdom, pp: 167-196. Zool., 77: 1645-1656. 33. Mondal, K., S. Gupta, Q. Qureshi and K. Sankar, 2011. 46. Nowell, K. and P. Jackson, 1996. Wild : Status Prey Selection and food habit of leopard (Panthera Survey and Conservation Action Plan. IUCN/SSC Cat pardus fusca) in Sariska tiger reserve, Rajasthan. Specialist Group, Gland, and Cambridge, Mammalia, 75: 201-205. UK. IUCN. Gland, Switzerland. 34. Sankar, K., Q. Qureshi, P. Nigam, P.K. Malik, 47. Gupta, S., K. Mondal, K. Sankar and Q. Qureshi, P.R. Sinha, R.N. Mehrotra, R. Gopal, S. Bhattacharjee, 2010. Estimation of Striped hyena (Hyaena hyaena) K. Mondal and S. Gupta, 2010. Monitoring of population using in Sariska Tiger reintroduced in Sariska Tiger Reserve, Western Reserve, Rajasthan, India. J. Bombay. Nat. His. Soc., India: preliminary findings on , prey 106(3): 284-288. selection and food habits. Journal of Tropical 48. Balasubramaniam, P. and P.V. Bole, 1993. Seed Conservation Science, 3(3): 301-318. dispersal by mammals at Point Calimere wildlife 35. MacArthur, R. and R. Levins, 1967. The limiting sanctuary. . J. Bombay Nat. Hist. Soc., similarity, convergence and divergence of coexisting 90: 33-44. species. American Naturalist, 101: 377-385. 49. Taryannikov, V.I., 1974. Feeding of Canis aureus 36. Gehring, T.M. and R.K. Swihart, 2003. Body size, aureus in the Syrdarja Basin. Zool. Zh., 53: 1539-1547. niche breadth and ecologically scaled responses to [In Russian with English summary]. : mammalian predators in an 50. Ishunin, G.I., 1980. The boar, jackal, red fox and agricultural landscape. Biological Conservation, in the Aydarsk solonceks inundated by the 109: 283-295. waters of the Syr Daria. Bull. Soc. Nat. Biol. 37. Gupta, S., 2011. Ecology of medium and small sized Ser., 85: 43-51. [In Russian with English summary]. carnivore in Sariska tiger reserve, Rajasthan, India. 51. Mukherjee, S., S.P. Goyal, A.J.T. Johnsingh and Ph.D Thesis Submitted to the Saurashtra University. M.R.P.L. Pitman, 2004. The importance of rodents in Rajkot. Gujarat, pp: 156. the diet of Jungle Cat (Felis chaus), Caracal (Caracal 38. Rosenzweig, M.L., 1966. Community structure in caracal) and Golden Jackal (Canis aureus) in Sariska sympatric . J. Mammal., 47: 602-612. Tiger Reserve, Rajasthan, India. Journal of Zoology, 39. Schoner, T.W., 1974. Some methods for calculating 262: 405-411. competition coefficients from resource-utilization spectra. Am. Nat., 108: 332-340.

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