Status of leopard Panthera pardus and striped hyena Hyaena hyaena and their prey in Achanakmar Tiger Reserve, Central India

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1 The 2017; 4(4): ISSN JOZS 2017; 4(4): JOZS 2017 Received: Accepted: Status of leopard Panthera pardus and striped hyena Hyaena hyaena and their prey in Achanakmar Tiger Reserve, Central India Dibyendu Mandal 1,2, Krishnendu Basak 1,3, Rajendra Prasad Mishra 1, Rahul Kaul 1, Krishnendu Mondal 1,4 1 Wildlife Trust of India, F -13, Sector 8, Noida, U.P , India. 2 Wildlife Institute of India, Chandrabani, Dehradun , India. 3 Nova Nature Welfare Society, Raipur, Chhattisgarh , India 4 Zoological Survey of India, Kolkata, India. Corresponding Author: Mandal D Corresponding dibyendu86@gmail.com Abstract To understand the multi-predator ecosystem and their prey, the present study assessed status of leopard (Panthera pardus) and striped hyena (Hyaena hyaena) (co-predators of tiger) and their prey species in Achanakmar Tiger Reserve (ATR) between October 2010 and March 2011 to address issues concerning recovery of tiger population in the area. Camera trapping under markrecapture framework was followed to estimate the abundance of leopard and striped hyena and line transects based on distance sampling was used to estimate the abundance of prey species. The estimated leopard density in the study area was ± 2.98/ 100 km 2, whereas it was 4.54 ± 1.75/ 100 km 2 for striped hyena. The present study revealed that the combined wild ungulate prey base (32.62 ± 11.48/ km²) in the study area can support more tigers (19.57 ± 6.88/ 100 km²) than the existing population of the entire Tiger Reserve [12 (11-13)]. Livestock abundance was found to be high ± 11.10/ km 2 ), which may cause detrimental effect to the wild ungulates and their habitat and also enhance chances of negative human-wildlife interactions. Therefore, an effective protection and conservation strategy of habitat and prey and restocking tiger population in ATR is crucial for long-term survival of other carnivores and their prey. Keyword: Large carnivores, Prey abundance, Habitat management, Mark recapture, Distance sampling 1. Introduction The primary reasons behind the unprecedented decline in tiger population such as prey depletion, habitat loss and poaching are now a well-known fact [1]. Models based on tiger population dynamics have indicated that many of the fragmented tiger populations were unable to sustain poaching pressures despite high fecundity [2] and large prey base. In fact, two tiger populations in India such as Sariska Tiger Reserve, Rajasthan and Panna Tiger Reserve, Madhya Pradesh lost their tiger populations due to poaching despite having good prey base [1, 3]. With improvement in the efficiency of estimation techniques of tigers [4], more such areas are there in India being discovered which require huge conservation efforts to bring tigers back or revive tiger population. In areas, where tiger numbers deplete, it is usual for its co-predators to thrive [5]. As, in any multiple predator ecosystems, each predator has strong inter-relationship with the other [6-7] and understanding the intra-guild relationship of the co-predators is equally crucial for its conservation [8-9]. Removal of top predators from an eco-system may give way to a phenomenon coined as meso-predator release [10], which is often associated with expansion in the density and distribution in the middle rank predator and may lead to negative cascading effects on prey [11-12]. Evidence suggests that apex predators have a role in regulating the prey populations by suppressing populations of smaller predators (co-predators). Two of these co-predators, leopard (Panthera pardus) and striped hyena (Hyaena hyaena) are largely coexisting with tiger (Panthera tigris) in most of its distribution range in the Indian sub-continent [13]. Achanakmar Tiger Reserve (ATR) is one such reserve (total area 914 km 2 ), where both leopard and striped hyena coexist with very low density of tiger [13]. In fact, a single female tiger was photo-captured after an extensive effort in 2010 [13]. The estimated population is 1 with a density of 0.11/ 100 km² [13]. However, the location of ATR is strategic and significant for the long term prospect of central Indian tiger landscape which holds 35% of Indian tigers and around 25% of global tiger population. Achankmar is contiguous with Kanha and Bandhavgarh Tiger Reserve [13] through two tenuous links and important for maintaining meta-population dynamics in the landscape. In contrast, ATR bears high levels of anthropogenic disturbance [14] and subsistence level of poaching [13]. Page 34

2 The core area of Achanakmar TR has extensive human interference which includes 25 villages (with 1774 families), frequently visited by the outsiders and cattle camps inside the Tiger Reserve boundary [14]. The Tiger Reserve also has a large domestic livestock population of approximately 9000 including buffalo (Bubalus bubalis), cow (Bos indicus) and goat (Capra hircus). In such a disturbed area, where the tiger population has already gone down, the present study assessed the status of leopard and striped hyena (co-predators) and their prey in Achanakmar Tiger Reserve between October 2010 and March 2011 to address issues concerning recovery of tigers in the area. 2. Material and methods 2.1 Study area Achanakmar Tiger Reserve or ATR (Fig. 1), located in the state of Chhattisgarh in Central India covers an area of 914 Km 2 with Km 2 of core zone and Km 2 area are buffer zone. It is situated between 81⁰30 E, 22⁰39 N and 82⁰30 E, 22⁰19 N at the Eastern part of Maikal hills of Satpuda Ranges and North West of Bilaspur and newly constituted Mungeli Districts of Chhattisgarh. Climate here is tropical with distinct winter (November-February), summer (April-mid June) and warm rainy seasons (July- September). Temperature ranges varies from monthly maximum temperature C to monthly minimum temperature C in the study area. The average annual rainfall is 1300 mm of which 82% occurs during rainy season [15]. Study area is mostly hilly and undulating. The forest is tropical moist deciduous type [15]. Valleys are dominated mostly with sal (Shorea robusta) but in many areas gregarious association with different tree species like Diospyros melanoxylum, Terminalia tomentosa, Adina cordifolia, Pterocarpus marsupium, Anogeissus latifolia, Tectona grandis, Madhuca indica were observed. Lower and higher slopes have bamboo (Dendrocalamus strictus) with miscellaneous tree species. Fig 1: Achanakmar Tiger Reserve in Chhattisgarh covers an area of 914 Km2 with Km2 of core zone and Km2 area are buffer zone. Apart from tiger, leopard and hyena, the other carnivores present in ATR are dhole (Cuon alpinus) and golden jackal (Canis aureus), jungle cat (Felis chaus) and Indian fox (Vulpes bengalensis). Sloth bear (Melursus ursinus) is the only bear species found in this tiger reserve. Chital (Axis axis), sambar (Rusa unicolor), gaur (Bos gaurus), nilgai (Boselaphus tragocamelus), wild pig (Sus scrofa), barking deer (Muntiacus vaginalis) and chowsingha (Tetraceros quadricornis) are the wild ungulate species found in Page 35

3 the study area. The common langur (Semnopithecus entellus) and rhesus macaque (Macaca mulatta) represents the primate fauna of the area. The Indian porcupine (Hystrix indica), black-naped hare (Lepus nigricollis) and Indian giant squirrel (Ratufa indica) are also found in ATR. Domestic livestock such as domestic buffaloes, cows and goats are present inside the Tiger Reserve. 2.2 Estimating abundance of sympatric carnivores The abundance of co-predators of tiger (here leopard and striped hyena) was estimated using camera traps under mark-recapture framework. An initial survey was carried out in the study area in more than 300 km 2 in the month of November - December 2010 to record indirect signs, like scats, pugmark and rake marks of leopard and striped hyena. Based on the observed encounter rates of indirect signs, potential camera trap locations were identified and recorded for the carnivore species. The intensive study area (~200 km 2 ) was divided into 40 grids (5 km 2 ) and of these, three grids were found unsuitable for camera trapping due to high anthropogenic disturbances (Fig. 1). A total of 37 pairs of camera traps were deployed in 37 grids, approximately 2 km (average inter-trap distance was 1689 m) apart to maximize the capture probability for all the carnivore species [16] for 46 consecutive occasions (one day) resulting into 1702 trap nights. A combination of distinguishing characters such as position and shape of rosettes on flanks, limbs and forequarter and pelage markings on hind limbs, forelimbs and stripes on flanks were used to identify individual leopards and striped hyena respectively from the photographs obtained from camera traps [4, 17-20]. Following the identification, unique identification numbers were given to individual striped hyenas and leopards and capture histories (X matrix) were developed [21]. Estimation of population size using closed capture models require the population under investigation to be both demographically and geographically closed. We used the closure test (Z) to test the null hypothesis of population closure [22]. The density (D) of striped hyena and leopard in the study area was estimated using both MMDM/2 and robust Maximum Likelihood Spatially Explicit Capture Recapture or MLSECR using Density 4.4 software [23]. 2.3 Estimating abundance of prey species Prey abundance was estimated through distance sampling methods using line transects [24]. A total of 29 line transects [24] of 2 km each were laid randomly in 29 different forest beats within the effective trapping area selected for camera trapping (Fig. 1). Each line transect was walked three times in early morning time (6:00 am to 8:30 am) in the study period. Line transects were laid using see-through compass and measuring tapes. Start and end locations of the line transects were recorded using GPS. Based on food habit study in Indian sub-continent by various authors [7, 25-29], spotted deer, barking deer, common langur, rhesus macaque, cattle, wild pig, peafowl, red jungle fowl, hare and giant squirrel were considered as potential prey species of leopard and striped hyena. The distances of the encountered prey species from the line transect were measured using laser range finder and the animal bearings were measured by see-through compass [1]. Overall prey density was estimated using DISTANCE 5 program [24, 30]. Minimum Akaike Information Criteria was used to select best fit models [24]. Density estimates for respective prey species were estimated using the stratification option in DISTANCE 5, keeping the global detection probability constant for all species. 3. Results 3.1 Abundance of leopard and striped hyena In total, 19 striped hyena photographs and 42 leopard photographs were obtained from the intensive camera trapping, from which we identified 10 individual hyenas and 23 individual leopards. Out of the total 23 leopard individuals, four male (17.4%) and 15 female leopards (65.2%) were identified. Sex of four individual leopards could not be identified from the photographs. Both, striped hyena and leopard populations were demographically and geographically closed (hyena: Z = 0.413, P = 0.66 and leopard: Z = , P = 0.274) during the sampling period. Although, model M(o) (null) was identified as the most appropriate model for both the species in our study, we used M(h) model as heterogeneity is observed in any biological population. The model selection scores for striped hyena were as follows: M(o) = 1, M(h) = 0.92, M(b) = 0.31, M(bh) = 0.52, M(t) = 0.00, M(th) = 0.24, M(tb) = 0.35 and M (tbh) = The model selection score for leopard were as follows: M(o) = 1, M(h) = 1, M(b) = 0.51, M(bh) = 0.75, M(t) = 0.00, M (th) = 0.40, M (tb) = 0.52 and M (tbh) = The estimated population (N) of striped hyena was 12 (± SE 2.3) and leopard was 30 (± SE 4.2) whereas estimated MMDM/2 density was 3.84 (± SE 0.94)/ 100 km 2 for striped hyena and (± SE 1.89)/ 100 km 2 for leopard and MLSECR density was 4.54 (± SE 1.75)/ 100 Km 2 for striped hyena and (± SE 2.98)/ 100 km2 for leopard (Table 1). Page 36

4 Table 1: Density estimates of striped hyena and leopard in Achanakmar Tiger Reserve, Chhattisgarh, Central India (December 2010 to March 2011). Species Model Population (± SE) MCP (Km 2 ) Methods for calculating ETA Width (Km) ETA (Km 2 ) Density/ 100 Km 2 (± SE) Striped Hyena M(O) 12 ± MMDM/2 ML Dens ± ± 1.75 Leopard M(O) 30 ± MMDM/2 ML Dens ± ± 2.98 (MCP= Minimum Convex Polygon, ETA= Effective Trapping Area, D= Density estimate, MMDM= Mean Maximum Distance Moved, ML Dens= Maximum Likelihood Density, SE= Standard Error) 3.2 Abundance of prey species A total of 219 observations were made on the transect surveys (Table 2). Although, giant squirrel and black-naped hare were spotted on the line transect, we did not calculate their density because of low sample size. Among all the prey species, the estimated density of common langur was highest, followed by domestic cattle (cow and buffalo), rhesus macaque, wild pig, spotted deer, gaur, peafowl, red jungle fowl and barking deer (Table 2). Of the groups encountered during line transect, 71.88% were small-bodied animals (common langur, rhesus macaque and peafowl, <20 Kg; Density (D) = ± individuals/ km 2, Biomass (BM) = ± kg/ km 2 ), 20.71% were medium-sized animals (spotted deer, wild pig and barking deer, kg; D= ± 7.79 individuals/ km 2, BM: ± kg/ km 2 ) and 7.40% were large-bodied animals (Gaur, >50 kg; D = 8.59±3.38 individuals/ km 2, BM: ± kg/ km 2 ) (Table 2). The overall or pooled estimated density of all prey species including domestic livestock was 140.8/ km 2 and total prey biomass was kg/ km² (Table 2). Combined wild ungulate prey density was ± animals/ km². Livestock density of the sample area was ± 11.10/ km 2. Comparing with other studies in Tropical Deciduous Forests, it can be well said that Achanakmar supports a good prey base across all body size which is crucial for maintaining viable populations of different carnivores [1]. Table 2: Density estimates of prey species in Achanakmar Tiger Reserve, Chattisgarh, Central India (December 2010 to March 2011). Prey Species Number of Sighting Group Size Density Group Size SE D SE Gaur Spotted deer Wild pig Barking deer Common langur Rhesus macaque Peafowl Livestock (cattle and buffalo) Discussion Since, ungulates contribute majorly to carnivore diet [7, 31], this factor primarily influences distribution and abundance of large and medium sized carnivores. Wild ungulate prey density (32.5/ km 2 ) in the present study area was considerably lower than the Page 37

5 density estimated in Rajaji National Park (90.8/ km 2 [32] ) and Sariska Tiger Reserve (103.4/ km 2 [33] ) which sustained similar predators such as leopard [18, 33], striped hyena [17, 19] and tiger [3, 32]. Perhaps, the wild ungulate density was supplemented by a fairly high density (25.4/ km 2 ) of livestock in the study area to sustain the carnivore population in the study area. However, other tiger reserves in the neighbor-hood like Kanha, Pench and Bandhavgarh sustain a higher wild ungulate prey density than Achanakmar TR [29]. The reason for this could be attributed to the fact that chital and gaur, being ruminant grazers like cattle and buffalo, were probably more affected by the high abundance of livestock (Table 2) in the study area. In Bandipur National Park, Madhusudan [34] documented spatial exclusion and recovery of ruminant grazers (chital and gaur) following a 49% decline of livestock populations, indicating that livestock did cause resource limitation on wild herbivores. Harihar et al. [32] also documented recovery of chital population in terms of increased reproductive success following the resettlement of 193 Gujjar families from within Chilla Range of Rajaji National Park. As the tiger density is a function of wild prey density, applying formula given by Karanth et al. [35], we estimated that current combined wild ungulate prey base (32.62 ± 11.48/ km²) in the intensive study area can support up to 6.52 ± 2.29 tigers/ 100 km² corresponding to a population of ± 6.88 tigers across the sampled area only (300 km 2 approximately), which is more than the current tiger population of the entire Achanakmar landscape [12 ± 1 tigers in 855 km 2 ]. Photographic capture-recapture sampling methodology has been used widely to estimate the density of tigers in many protected areas throughout India [16, 35], similar density estimates of leopard and striped hyena are available from very few Tiger Reserves such as Rajaji National Park in the Himalayan foothills [18, 19] and semi-arid Sariska Tiger Reserve [17, 20]. Density estimates of striped hyena using similar methods are available only from Kumbhalgarh WLS and Esrana Forest Range [36] and Sariska Tiger Reserve [17]. The density estimate for leopard in the present study is lower than the density estimated from the Rajaji National Park, however, considerably higher (Table 3) than density estimate from the Sariska Tiger Reserve [20]. Both the above mentioned areas have low tiger populations, similar to ATR [13]. Striped hyena density estimate in the present study is comparatively similar to the estimates obtained from Rajaji National Park, Kumbhalgrah WLS and Esrana Forest Range, however, comparatively lower than the density estimated from Sariska Tiger Reserve which is the highest striped hyena density estimate (Table 3) available in India and Africa [17, 36-38]. Achanakmar TR appears to be behaving typically of areas which have suppressed populations of apex predators [39]. In low productivity systems, apex predators may become extinct even in absence of human influence and meso-predator is then regulated by the limited supply of food rather than by predation [40]. However, ATR cannot be called a low productivity system as it was found from the historical records that this area did support reasonable tiger densities in the recent past and its depletion is a relatively recent phenomenon. Declines, therefore, appear to have occurred due to immense anthropogenic issues, in the form of either poaching or retaliatory killings. This is also to some extent corroborated by the fact that several cases (n=32) of livestock depredation were attributed to tiger killing during Knowing that the tiger density is low in the study area, this appears to be a high number and could attract retaliatory killing. Thus, it is imperative that tiger numbers be augmented in the Tiger Reserve. This may involve pro-active measures like re-introductions/ re-stocking as the numbers are too low to build the populations by itself and the connectivity with other areas may not be viable. Table 3: Estimated leopard and striped hyena densities in different protected areas of India Study Site Leopard Density/ 100 Km 2 Author Striped hyena density/ 100 Km 2 Present Study Author Sariska Tiger Reserve 7.1 Mondal et al. 2012b 15.1 Gupta et al Rajaji National Park 14.9 Harihar et al Harihar 2009 Satpura National Park 7-10 Edgaonkar 2008 [42] - Mudumalai Tiger Reserve 14.9 Kalle et al [43] - Kumbhalgarh WLS Singh 2010 Esrana Forest Range Singh 2010 Pench Tiger Reserve, Madhya Pradesh 10.6±3.2 Majumder Page 38

6 As livestock causes detrimental effect on the wild ungulate habitat as observed by Shukla and Khare (1998) [41], in Central India, their movements need to be restricted for the protection of habitat for wild ungulates. Also, believing the fact that tigers require pristine habitat for rearing cubs [29], relocation of villages at least from the core areas is also essential to create a disturbance free breeding habitat for tiger in ATR. Although, the present study was short in nature due to logistical constraints and lacks spatial coverage of the entire Reserve, however, the present study indicates that the existing prey base can sustain good numbers of tigers in the Reserve which is contiguous with other forests in the important tiger landscape of central India. 5. Acknowledgements We sincerely thank National Tiger Conservation Authority (NTCA) and Wildlife Institute of India for providing us financial support and opportunity to carry out the field research work. We also thank Chhattisgarh Forest Department for granting us necessary permission and cooperation during the field work. In Wildlife Trust of India, we would like to thank Mr. Vivek Menon for his overall support, Dr Anil Singh, Mr Milind Pariwakam, Ms. Smita Bodhankar and Mr. Moiz Ahmad for their assistance during the study period. Special thanks Dr. Aniruddha Majumder for helping in preparation of the manuscript. 6. References 1. Sankar K, Qureshi Q, Mondal K, Worah D, Srivastava T, Gupta S, Basu S. Ecological studies in Sariska Tiger Reserve, Final report submitted to National Tiger Conservation Authority, Govt of India, New Delhi. Wildlife Institute of India, Dehradun. 2009, pp Chapron G, Miquelle DG, Lambert A, Goodrich JM, Legendre S, Clobert J. The impact on tigers of poaching versus prey depletion. Journal of Applied Ecology 2008; 45(6): Sankar K, Qureshi Q, Nigam P, Malik PK, Sinha PR, Mehrotra RN et al. Monitoring of reintroduced tigers in Sariska Tiger Reserve, Western India: Preliminary findings on home range, prey selection and food habits. Tropical Conservation Science 2010; 3(3): Karanth KU. Estimation Tiger population through camera trap data using Capture recapture model. Biological Conservation. 1995; 71: Ramakrishnan U, Coss RG, Pelkey NW. Tiger decline caused by the reduction of large ungulate prey: supporting evidence from a comparative study of leopard diets in two reserves in southern India. Biological Conservation 1999; 89: Curio E. The ethology of predation. Springer-Verlag, New York, 1976; x Johnsingh AJT. Large mammalian prey-predator in Bandipur. Journal of Bombay Natural History Society 1983; 80(1): Terborgh J, Lopez L, Nuñez VP, Rao M, Shahabuddin G, Orijuela G et al. Ecological meltdown in predator-free forest fragments. Science. 2001; 294: Mondal, K, Gupta, S, Bhattacharjee, S, Qureshi, Q, Sankar, K. Response of leopard to reintroduced tigers in Sariska Tiger Reserve, Western India. International Journal of Biodiversity and Conservation 2012a; 4(5): Soulé ME, Bolger DT, Alberts AC, Sauvajot RS, Wright J, Sorice M, Hill S. Reconstructed dynamics of rapid extinctions of chaparral-requiring birds in urban habitat islands. Conservation Biology 1988; 2: Berger KM, Gese EM, Berger J. Indirect effects and traditional trophic cascades: A test involving wolves, coyotes, and pronghorn. Ecology. 2008; 89(3): Brashares JS, Prugh LR, Stoner CJ, Epps CW. Ecological and conservation implications of mesopredator release. In Terborgh J, Estes JA, eds. Trophic cascades: predators, prey, and the changing dynamics of nature. Island Press, Washington, 2010; Jhala YV, Qureshi Q, Gopal R, Sinha PR. (Eds.). Status of tigers, copredators and prey in India, National Tiger Conservation Authority, Govt. of India, New Delhi and Wildlife Institute of India, Dehra Dun. 2011, pp302. Page 39

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