Rapid recovery of tigers Panthera tigris in Parsa Wildlife Reserve, Nepal
B ABU R AM L AMICHHANE,CHIRANJIBI P RASAD P OKHERAL,SHASHANK P OUDEL D IPENDRA A DHIKARI,SAILENDRA R AJ G IRI,SANTOSH B HATTARAI,TEK R AJ B HATTA R OB P ICKLES,RAJAN A MIN,KRISHNA P RASAD A CHARYA,MAHESHWAR D HAKAL U BA R AJ R EGMI,ASHOK K UMAR R AM and N ARESH S UBEDI
Abstract Information on density and abundance of globally Chitwan–Parsa complex should be managed as a single eco- threatened species such as tigers Panthera tigris is essential logical unit for conserving the Endangered tiger and other for effective conservation as well as to evaluate the success of wide-ranging species. conservation programmes. We monitored tigers in Parsa Keywords Camera trapping, conservation success, Nepal, Widlife Reserve, Nepal, using camera traps, in , Panthera tigris, Parsa Wildlife Reserve, source–sink, tiger and . Once believed to be a sink for tigers from adjacent population Chitwan National Park, Parsa now provides a new hope for tigers. Spatially explicit capture–recapture analysis over survey years revealed an increase in tiger density from . to . individuals per km from to . The tiger abundance was estimated to be seven (–), Introduction (–) and (–) in , and , respectively. he tiger Panthera tigris, categorized as Endangered on Resettlement of communities from the core area, reduced Tthe IUCN Red List (Goodrich et al., ), remains in anthropogenic pressure, and improved security have made % of its historical range (Joshi et al., ). The remaining Parsa Wildlife Reserve a suitable habitat for tigers. Tiger habitat is not occupied at optimum density because of abundance increased considerably within a km radius of poaching of tigers, hunting of their prey species and conflict the evacuated village sites, from two in to eight in with local communities (Goodrich, ; Walston et al., and in . Population turnover has remained ). With the aim to prevent extinction and double the moderate (, % per year), with persistence of individuals tiger population by , tiger range countries signed the in multiple years. Dispersing tigers from Chitwan’s source St Petersburg Declaration on Tiger Conservation in population accounted for a large portion (c. %) of the ti- (GTI, ), yet tiger populations continue to decline in gers detected in Parsa. Conservation efforts along with an- many countries (Goodrich et al., ). Studies show that nual monitoring should be continued in Parsa to sustain the the remaining habitat can support the global target of increase and monitor the persistence of tigers. The doubling the number of wild tigers to , by (Wikramanayake et al., ) if further degradation of habi- tat is prevented (Joshi et al., ) and core breeding source
BABU RAM LAMICHHANE* (Corresponding author), CHIRANJIBI PRASAD POKHERAL, sites are protected and embedded in larger conservation SHASHANK POUDEL,SANTOSH BHATTARAI and NARESH SUBEDI National Trust for landscapes (Walston et al., ). Nature Conservation, Khumaltar, POB 3712, Lalitpur, Nepal E-mail [email protected], [email protected] Within Nepal, tigers are restricted to five protected areas and surrounding forests in the Terai Arc Landscape in the DIPENDRA ADHIKARI,SAILENDRA RAJ GIRI and TEK RAJ BHATTA Zoological Society of London Nepal Office, Lazimpat, Kathmandu, Nepal south of the country, straddling the border with India
ROB PICKLES Panthera, New York, USA (Wikramanayake et al., ). Of global importance for tiger recovery, the Chitwan–Parsa–Valmiki forest complex RAJAN AMIN Zoological Society of London, Regent’s Park, London, UK of the Terai Arc Landscape was designated a Level I Tiger KRISHNA PRASAD ACHARYA Ministry of Forests and Soil Conservation, Department of Forests, Babarmahal, Kathmandu, Nepal Conservation Unit, a region of global priority, in (Wikramanayake et al., ). It also constitutes one of the MAHESHWAR DHAKAL Ministry of Forests and Soil Conservation, Sighadurbar, Kathmandu, Nepal global source sites of tigers (Walston et al., ). Despite
UBA RAJ REGMI and ASHOK KUMAR RAM Department of National Parks and this global recognition, Parsa had made little contribution to Wildlife Conservation, Kathmandu, Nepal global tiger recovery efforts until recently, with a density of *Also at: Institute of Cultural Anthropology and Development Sociology, only . tigers per km recorded in , in contrast to Faculty of Social and Behavioural Sciences, Leiden University, Leiden, neighbouring Chitwan’s . tigers per km (Dhakal Netherlands, and Evolutionary Ecology Group, Faculty of Sciences, University of Antwerp, Antwerp, Belgium et al., ). Prey density had also been low in Parsa (. Received March . Revision requested April . prey individuals per km ) compared to Chitwan (. per Accepted May . km ) (Karki et al., ) as a result of widespread hunting
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FIG. 1 Parsa Wildlife Reserve (with survey grid) and its buffer zone, and neighbouring Chitwan National Park, Nepal.
and habitat degradation through livestock grazing, leading (Fig. ). Initially established with an area of km, Parsa to Parsa being considered the sink to Chitwan’s source. was augmented by the allocation of a km buffer zone There are records of tigers dispersing from Chitwan being in , and in a further km was gazetted as core poisoned in retaliation by local people (Smith, ), or poa- area (hereafter referred to as the extension area; Fig. ; ched for their skin and bones. Dispersal of tigers from DNPWC, ). Parsa comprises highly porous alluvial sub- high-density areas to lower density areas is a common phe- strate and is dominated by the sal forested Churia Hills run- nomenon (Harihar et al., ; Harihar & Pandav, ). ning from east to west (into Chitwan) in the north of the Tigers are territorial, and adult males are known to have Reserve. The streams running off the Churia Hills permeate large territories encompassing – females (Sunquist, the porous sediment and flow underground, reappearing ). Subadults are known to disperse away from the south of the Reserve and restricting water availability in natal area at the age of – months to colonize relatively . % of Parsa throughout the dry months. As well as tigers, unoccupied areas (Smith, ). Parsa is home to other carnivores, including the leopard In Nepal committed to doubling the size of its tiger Panthera pardus, the dhole Cuon alpinus, the striped hyaena population by , with a goal of adults (GTI, ). Hyaena hyaena and the golden jackal Canis aureus. A wide Parsa is therefore a clear target for conservation interven- range of wild prey species of tigers are found there, including tions to meet the national goal. Here we examine the change the gaur Bos gaurus, the sambar Rusa unicolor, the nilgai in Parsa’s tiger population over a -year period since signifi- Boselaphus tragocamelus, the spotted deer Axis axis, the bark- cant conservation interventions began, and assess the fac- ing deer Muntiacus muntjak and the wild boar Sus scrofa tors responsible for the change. (Thapa et al., ). Historically, Parsa received less govern- mental and NGO support for tiger conservation compared to other protected areas in the Terai Arc Landscape. However, Study area since the Government of Nepal, with technical and fi- nancial support from NGOs, has undertaken significant ef- Parsa Wildlife Reserve is the easternmost protected area of forts to improve tiger conservation in Parsa (PWR, ). the Terai Arc Landscape. It is connected directly Two settlements, Rambhori Bhata ( ha, households) to Chitwan National Park along a km boundary and is and Ramauli Pratappur ( ha, households), had been , km from Valmiki Tiger Reserve’s easternmost edge located in Parsa prior to the establishment of the Reserve
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(Fig. ). The local people practised subsistence agriculture, ,
but activities such as livestock grazing, and collection of fod- der, fuelwood, timber and other forest products resulted in May 2016
habitat degradation over a broad area (PWR, ). Conflict – between local people and wildlife also resulted in retaliatory Feb. (79) killings (CNP, ). Both communities were resettled vol- untarily during –. In addition to the village reloca- tions, Parsa’s management authorities began a programme of habitat enrichment in , maintaining nine artificial Mar. 2016
waterholes in the dry season and increasing the area of un- –
gulate grazing by . ha of grasslands. Feb. (27) arison with other years.
Methods . Only a partial survey was conducted in May 2016
Camera trapping We set camera traps in a grid of × km – cells (DNPWC, ), deploying a pair of camera traps in and 2016 ParsaMar. (79) Extension area Total
each cell during the dry season in , and .The entire core area of Parsa was covered by – camera-trap ,
stations. Cameras were active in each station for a minimum , Jan. 2015
of ()to days ( and ). The survey effort was – ,–, camera-trap days (Table ). In , km of ) in the extension area was covered with an additional camera Fig.
stations. The Reserve was divided into survey blocks, two in 2014 Nov. 2014 (60) and three in and .Theblocksweresurveyed successively because of the limited availability of camera traps. Prior to deployment of cameras, potential sites where
tiger captures and camera safety could be maximized were May 2013 2 – identified. We positioned the camera traps cm above ground, perpendicular to, and – meithersideofgame Apr. (59) trails, forest roads and riverbeds. Pairs of cameras were in Parsa Wildlife Reserve ( placed at each sampling point to obtain images of both flanks of photographed individuals to assist in identification. We used Mar. 2009 Reconyx (Holmen, USA) & ,Bushnell(OverlandPark, –
USA) Trophy Cam HD and Panthera (New York, USA) V & (partial) 2013 ). 1 V cameras. The cameras took three photographs per trigger Panthera tigris (54) with no delay, and used white flash to obtain colour images 15 15 21 21 21 21 of tigers in low light to help with individual identification.
Estimating population abundance and density Tiger identification was conducted by three independent observers and cross-verified collectively by six or seven observers. We also used ExtractCompare (Conservation
Research Ltd, UK) to verify individual tigers identified ).
visually (Hiby et al., ). Tiger density and population size were estimated using spatially explicit capture– recapture models in the package secr (Efford, ) in R v. .. (R Development Core Team, ). The default ) 353 499 499 499 128 627 maximum likelihood algorithm with function secr.fit was 2 used to fit the model. To determine the local importance of the former locations of villages for tigers, we subsampled a buffer area of km radius around the 1 Summary of survey effort for camera-trap surveys of tigers village locations and calculated the number of tigers ABLE location Additional area in the south of the Reserve and some parts of the buffer zone in the east were also covered but we used data from the core area only, for comp Parameters obtained from Karki ( and information was obtained from a published report (Karki, Parameters 2009 T No. of trap nightsNo. of camera stations 97 1,762 2,135 138 3,549 130 2,946 127 735 31 3,681 158 Sampling period (days)Minimum no. of sampling occasions (1 day each) in a Dec. 2008 detected there. Area surveyed (km
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Determining tiger dispersal and persistence We compared
the individual tigers detected in with camera-trap 2.25) – photographs of those recorded in , and in 26) Parsa. We also compared all tigers detected in Parsa with –
tigers recorded in Chitwan National Park in and its 0.33 (0.91 1.9 (19 ±
buffer zone in (Dhakal et al., ; CNP, ), and ±
in Valmiki Tiger Reserve, India, in (Maurya & Borah, 19 10 ; Chanchani et al., ).
Results 2 2
Tiger density and abundance During – tiger
density in Parsa increased from . ± SE . to 2.25) 1.43 –
. ± SE . tigers per km (Table ). Five, and 20) 20 – individual tigers were detected in , and , respectively, with an additional two tigers detected in the 0.34 (0.85 1.53 (17 extension area in . Tiger abundance increased ± ±
gradually in Parsa during – (Fig. ). 2016 Parsa Extensionarea Total % confidence intervals from camera-trap surveys in Parsa Wildlife 1.48) 1.38
Evidence of dispersal from Chitwan Over the sampling – 16) 17
years a total of adult tigers ( females, males and – one of unknown sex) were detected (Table ). One male
and two females were detected in all years, and an 0.25 (0.43 1.27 (10 ±
additional three individuals (two females and one male) ± were captured in years ( and ). Ten new individuals were captured in . Of the individuals 1.98) 0.80
recorded, nine were captured first in Chitwan (in and – ) and then dispersed to Parsa (in and ). One 13) 11 tiger (F, Table ) was first captured in Parsa () and –
dispersed to Chitwan’s buffer zone () in the south 0.39 (0.31 ± (Someshwar Hills, contiguous to Valmiki). There was no 1.53 (6 ± 2013 2014 evidence of tigers dispersing between Valmiki and Parsa. 0.38 2.17 3.84 3.67 3.80 1.62) 0.78 Recolonization of tigers in evacuated village sites Tigers – were confined to a narrow strip in the middle of the Reserve during (Fig. a) but occupied most of the 0.32 (0.23 1 ). Reserve in the following years (Fig. b,c). The survey ± 0.47 7 ±
failed to detect a single tiger within the km buffer of
Ramauli Pratappur village, which was in the process of .
relocation at the time. The following year four tigers were detected within the km buffer, two of which were redetected in the same area in (Fig. ). Within the and
km buffer surrounding the site of Rambhori Bhata ,
village, which was relocated in , two tigers were (95% CI) 0.61 2 detected in , four in and seven in (Fig. ). , SE (95% CI) 4 ± ) in Discussion SE per 100 km Fig. Our findings indicate a nearly threefold increase in tiger ± 2 Tiger capture rate, number of individual tigers captured, and population abundance and density estimates with numbers in Parsa within years. This is a unique scenario MaleFemaleUnknown 2 2 2 2 1 4 6 8 9 1 2 9 ABLE trap days) A tigress was captured from both Parsa and the extension area. Partial survey conducted; estimates are from Karki ( T Reserve ( Parameters 2009 No. of camera stations thatNo. captured of tigers independent detections ofTiger tigers capture rate (no. of detections per 100 9 10 45 77 62 113 11 27 73 140 Tiger abundance No. of individual tigers captured (Mt + 1) 4 5 10 17 3 of tiger recovery, beyond the potential of natural growth Density
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FIG. 2 Tiger abundance estimates for Parsa Wildlife Reserve (Fig. ), with % confidence interval, based on a spatially explicit capture– recapture model using the package secr in R (R Development Core Team, ). Published data (Karki et al., ) were used for the estimate. The timeline at the top of the figure highlights key events that strengthened the protection of Parsa.
TABLE 3 Persistence of individual tigers over years in Parsa Wildlife Reserve and neighbouring Chitwan National Park (Fig. ). Data from Chitwan in and data from Parsa in were not available for cross comparison. Blank cells indicate not surveyed.
Tiger ID 2013 2014 2015 2016 F01 Parsa Parsa Parsa F02 Parsa Parsa Parsa M01 Parsa & Chitwan Parsa Parsa M02 Parsa Not detected Not detected U01 Parsa Not detected Not detected F03 Chitwan Parsa Chitwan Parsa F04 Not detected Parsa Chitwan Not detected F05 Chitwan (Cub) Parsa Not detected F06 Not detected Parsa Parsa M03 Chitwan Parsa Chitwan M04 Not detected Parsa Parsa M05 Chitwan (Cub) Parsa Not detected F07 Not detected Not detected Chitwan Parsa F08 Not detected Not detected Parsa F09 Not detected Not detected Parsa F10 Not detected Not detected Parsa F11 Not detected Not detected Parsa (extension area only) F12 Not detected Not detected Parsa F13 Not detected Not detected Parsa (and extension area) M06 Not detected Not detected Chitwan Parsa M07 Not detected Not detected Parsa M08 Not detected Not detected Parsa M09 Not detected Not detected Parsa M10 Not detected Not detected Chitwan Parsa (extension area only) M11 Not detected Not detected Parsa
without immigration. In the camera-trap survey of only , . per km in ; Dhakal et al., ; Karki five tigers were detected (Dhakal et al., ). One of the vil- et al., ), with regular dispersal of subadults beyond lages (Ramauli Pratappur) was still in the core area of the the Park’s boundary. Of the tigers identified in Parsa during Reserve, where a tiger was killed in retaliation after it at- –, c. % had been detected previously in Chitwan, tacked livestock and people in (CNP, ; Fig. a). including two cubs detected in and subsequently de- Tigers recovered quickly, with a total of tigers detected tected in Parsa in as young adults (Table ). Habitat in Parsa, and two others in the extension area, in connectivity of continuous sal forest between Chitwan and (Fig. c,d). Parsa facilitated dispersal, particularly of young individuals, Chitwan’s source effect was instrumental in the recovery leading to a quick recovery. It has previously been suggested of Parsa’s tiger population. Tiger density in Chitwan has re- that the tigers in both protected areas comprise a single mained high and stable in recent years (. per km in population (Walston et al., ). This is supported by our
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results, which show two-way movement of three individuals between the protected areas, and one individual holding ter- ritories that straddle the Park boundary. We therefore rec- ommend managing the two areas as a single entity. Although dispersal of tigers from Chitwan to Parsa is not a new phenomenon, this is the first recording of recoloniza- tion of vacant habitat in a sink habitat by substantial num- bers of individuals from a neighbouring source in Nepal. In addition to increasing tiger density in Parsa as a result of the influx from Chitwan, population turnover has remained moderate, and persistence of individuals high. When considered together with the Chitwan dataset, % of tigers identified in persisted into and were re- detected, and % of the cohort was redetected in (Table ). Tigers are not simply passing through Parsa but are holding territories and breeding. Two females (F &F) and one male (M) were detected in all survey years in Parsa. Pugmarks of at least two cubs were also dis- covered in and within km of the evacuated site of Rambhori Bhata village, indicating that, in addition to im- migration, reproduction is now also contributing to the in- crease in Parsa’s tiger population. The proximate reason for the recovery of tigers in Parsa was predictable, given the high density of tigers in neigh- bouring Chitwan and the vacation of territory in Parsa fol- lowing the alleviation of pressures that had historically constrained the increase of tiger numbers there. The recol- onization of Parsa can be attributed to two major interven- tions: resettlement of villages from the core area, and security enhancement in the core area and buffer zone.
Resettlement of villages from the core area
Our camera-trap results indicate two things. Firstly, tigers started using the vacant areas immediately after evacuation of the villages. Secondly, there is greater intensity of usage by tigers of the land in the locations of the former villages than across the rest of the Reserve, a phenomenon which would have been impossible to achieve had the villages remained. Given the limited availability of water in Parsa, the location of the settlements close to the Reserve’s two principal water sources and natural grasslands created competition for this re- source between people and wildlife. Conflict with crop-raiding elephants Elephas maximus and cattle-raiding tigers was com- mon (CNP, ). The removal of households and their de- mands on the forest’s natural resources substantially reduced local habitat degradation and disturbance in tiger habitat in Parsa’s core, and reduced poaching opportunities through re- duced access to the Reserve, as has been recorded extensively in India (Karanth & Madhusudan, ;Karanth,). The conversion of c. ha of farmland and settlement FIG. 3 Camera-trap locations and tiger detection polygons in Parsa Wildlife Reserve, Nepal (Fig. ) in (a) , (b) and into productive grasslands for ungulate grazing, and the in- (c) . creased access of wildlife to water are not insignificant. Prey
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density increased from . ungulates per km in (Karki protection was strengthened there (Wegge et al., ). In et al., ) to ungulates per km in (Dhakal et al., India’s Rajaji National Park in the Western Terai the tiger ). In the greater one-horned rhinoceros Rhinoceros population increased rapidly, with a high immigration rate unicornis also returned to northern Parsa, grazing on the following the relocation of Gujjars (the local tribal people) site of the evacuated villages (DNPWC, ). and thousands of their livestock (Harihar et al., ). Rambhori Bhata village was resettled km south of the With the increase in the tiger population, conflict be- Parsa boundary in and Ramauli Pratappur was re- tween tigers and communities on the edges of Parsa is likely settled c. km north of Parsa in and (PWR, to rise, as has been reported from Chitwan (Gurung et al., ). Resettlement of communities from within protected ). Preventive and mitigation measures therefore need areas frequently attracts criticism (Schmidt-Soltau & to be initiated. Although the core area of the Reserve is Brockington, ) but studies from Chitwan have shown free of settlements, pressure from grazing, fuelwood and that people being resettled were positive about the experi- timber extraction continues to encroach from the buffer ence (McLean & Straede, ) and benefited socio- zone in the north and from the communities south of the economically in their new location (Dhakal et al., ). – km strip of forest in the south of the Reserve that is man- Households from the two villages within Parsa petitioned aged as a collaborative forest. This strip, which is not part of for relocation because of the problems they were facing re- the designated buffer zone, is used by wildlife as a refuge, but garding conflict with wildlife and limited access to markets, they face the threat of poaching and persecution from the health and education. All households were granted land al- communities in the south. Eight of the tigers recorded lotments in areas outside the Reserve boundary and received in Parsa were camera-trapped on the southern boundary financial support from the Government of Nepal for reloca- of the Reserve in and we do not know if they went fur- tion and house construction (PWR, ). ther south into the collaborative forest. Including the collab- orative forest in future surveys will provide valuable Security enhancement in the core area and buffer zone information about Parsa’s tigers. The ongoing work of the Buffer Zone Programme is an essential counterpart to the Until security in Parsa was controlled by a company of the strengthened security measures within Parsa, reducing Nepal Army comprising c. men in seven guard posts, the demands of local people on the Reserve’s resources equating to one man per . km.In the company was through alternative livelihoods, technical innovation and upgraded to full battalion strength and shared with Chitwan, improved governance of communal resources such as the increasing the standing force to c. men, or approximately community forest. This integrated conservation effort of one man per km in Parsa. Two additional guard posts were strong security coupled with community support should constructed near the northern boundary and three more in be continued to sustain the recovery of Parsa’s tiger the extension area (Dhudhaura, Sahajnath & Ratanpuri) dur- population. ing –.ProvidingauxiliarysupporttoParsa’ssecurity Parsa presents a striking example of tiger population re- force, community-based anti-poaching units were formed covery and progress towards achieving Nepal’s national goal in ,comprising. local youths from communities of doubling tiger numbers by . Following the manage- around Parsa, who serve voluntarily in controlling illegal graz- ment interventions undertaken by the Reserve authority and ing, hunting and forest resource extraction, and provide infor- conservation partners in recent years, Parsa has put in place mation on poachers and smugglers to the Reserve authority. the foundation to facilitate tiger recolonization and popula- The increase in security personnel and expansion of for- tion recovery, and illustrates the rapidity with which tiger est guard posts in Parsa from seven in to nine in recovery can occur given the appropriate conditions of increased protection of tigers and their prey from poaching. controlled poaching, inviolate space and connectivity to a Strategically, the placement of the additional guard posts source population. was important, particularly two posts in the northern part of the Reserve, where there had been no patrols previously. The guard posts near the two evacuated village sites secured Acknowledgements the new grasslands that evolved there. With a year-round water source, these grasslands are ideal for both ungulates We thank the Department of National Parks and Wildlife and tigers, and would have been at risk from poaching in Conservation of the Government of Nepal for giving per- the absence of security. The presence of anti-poaching pa- mission to conduct this research, and the Parsa Wildlife trols in these areas deterred poaching attempts and facili- Reserve Office and the National Trust for Nature tated the persistence of recolonizing tigers. Conservation for their support in implementing the Recovery of tigers has been reported from other parts of camera-trap survey. We acknowledge the funding received Nepal and India (e.g. Panwar, ). Tiger and prey popula- from ZSL Nepal, Panthera, the USAID Haiyo Ban tions recovered in Bardia National Park, Nepal, after Program, U.S. Fish & Wildlife Service and WWF Nepal.
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Leiden University, Netherlands supported the first author GOODRICH, J.M., LYNAM, A., MIQUELLE, D., WIBISONO, H., during manuscript preparation and Open Access publica- KAWANISHI, K., PATTANAVIBOOL, A. et al. () Panthera tigris. In tion. The camera-trap surveys would not have been possible The IUCN Red List of Threatened Species : e.TA. Http://dx.doi.org/./IUCN.UK.-.RLTS.TA. without the tireless effort of field staff, including officers, en [accessed June ]. rangers and game scouts of Parsa Wildlife Reserve, wildlife GTI (GLOBAL TIGER INITIATIVE)() Global Tiger Recovery technicians of the National Trust for Nature Conservation, Program. Global Tiger Initiative Secretariat, The World Bank, and student volunteers from Tribhuvan University and Washington, DC, USA. Kathmandu University. We acknowledge the contribution GTI (GLOBAL TIGER INITIATIVE)() Global Tiger Recovery Program: Implementation Plan –. Global Tiger Initiative of Mr Ashish Bista in data analysis of surveys and Secretariat, The World Bank, Washington, DC, USA. thank Abhishek Harihar and two anonymous reviewers GURUNG, B., SMITH, J.L.D., MCDOUGAL, C., KARKI, J.B. & BARLOW, for their valuable comments on the article. A. () Factors associated with human-killing tigers in Chitwan National Park, Nepal. Biological Conservation, , –. HARIHAR, A. & PANDAV, B. () Influence of connectivity, wild prey Author contributions and disturbance on occupancy of tigers in the human-dominated western Terai Arc Landscape. PLoS ONE, (), e. BRL, NS, CPP, RP and RA designed the study. SP, DA, SRG, HARIHAR, A., PANDAV, B. & GOYAL, S.P. () Responses of tiger SB and URR implemented the fieldwork and collected data. (Panthera tigris) and their prey to removal of anthropogenic influences in Rajaji National Park, India. European Journal of BRL, RP and RA analysed the data. All authors wrote and Wildlife Research, , –. reviewed the article. 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SCHMIDT-SOLTAU, K. & BROCKINGTON, D. () Protected areas conservation strategy to double the wild tiger population. and resettlement: what scope for voluntary relocation? World Conservation Letters, , –. Development, , –. WIKRAMANAYAKE, E., MCKNIGHT, M., DINERSTEIN, E., JOSHI, A., SMITH, J.L.D. () The role of dispersal in structuring the Chitwan GURUNG, B. & SMITH, D. () Designing a conservation tiger population. Behaviour, , –. landscape for tigers in human‐dominated environments. SUNQUIST, M.E. () The Social Organization of Tigers (Panthera Conservation Biology, , –. tigris) in Royal Chitawan National Park. Smithsonian Institution Press, Washington, DC, USA. THAPA, K., SHRESTHA, R., KARKI, J., THAPA, G.J., SUBEDI, N., PRADHAN, N.M.B. et al. () Leopard Panthera pardus fusca Biographical sketches density in the seasonally dry, subtropical forest in the Bhabhar of Terai Arc, Nepal. Advances in Ecology, , http://dx.doi.org/. BABU RAM LAMICHHANE is interested in the application of geograph- //. ical information systems (GIS) for wildlife management, and human– WALSTON, J., ROBINSON, J.G., BENNETT, E.L., BREITENMOSER, U., DA wildlife interactions. CHIRANJIBI PRASAD POKHERAL is interested in FONSECA, G.A.B., GOODRICH, J. et al. () Bringing the tiger back large carnivore biology and interspecies interactions. SHASHANK from the brink—the six percent solution. PLoS Biology, (), POUDEL is interested in conservation communication. DIPENDRA e. ADHIKARI is interested in small mammals. SAILENDRA RAJ GIRI is in- WEGGE, P., ODDEN, M., POKHAREL, C.P. & STORAAS,T.() terested in applications of GIS and remote sensing in wildlife conserva- Predator–prey relationships and responses of ungulates and their tion. SANTOSH BHATTARAI is interested in herpetology. TEK RAJ predators to the establishment of protected areas: a case study of BHATTA is involved in development and implementation of conserva- tigers, leopards and their prey in Bardia National Park, Nepal. tion projects. ROB PICKLES specializes in tiger monitoring using camera Biological Conservation, , –. traps. RAJAN AMIN specializes in the conservation of Asian and African WIKRAMANAYAKE, E., DINERSTEIN, E., ROBINSON, J.G., grassland and forest ecosystems. KRISHNA PRASAD ACHARYA is inter- KARANTH, U., RABINOWITZ, A., OLSON, D. et al. () An ested in resolving human–wildlife conflicts. MAHESHWAR DHAKAL is ecology‐based method for defining priorities for large mammal involved in policy analysis of protected areas. UBA RAJ REGMI has conservation: the tiger as case study. Conservation Biology, , served as manager of Parsa Wildlife Reserve and other protected areas –. in Nepal. ASHOK KUMAR RAM is involved in geospatial modelling of WIKRAMANAYAKE, E., DINERSTEIN, E., SEIDENSTICKER, J., LUMPKIN, wild elephants in Central Nepal. NARESH SUBEDI is interested in the S., PANDAV, B., SHRESTHA, M. et al. () A landscape-based ecology of rhinoceroses and invasive species.
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