Rapid recovery of tigers Panthera tigris in Parsa Wildlife Reserve, Nepal

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Rapid recovery of tigers Panthera tigris in Parsa Wildlife Reserve, Nepal
Rapid recovery of tigers Panthera tigris in Parsa
              Wildlife Reserve, Nepal
                   BABU RAM LAMICHHANE, CHIRANJIBI PRASAD POKHERAL, SHASHANK POUDEL
              DIPENDRA ADHIKARI, SAILENDRA RAJ GIRI, SANTOSH BHATTARAI, TEK RAJ BHATTA
                 ROB PICKLES, RAJAN AMIN, KRISHNA PRASAD ACHARYA, MAHESHWAR DHAKAL
                                    U B A R A J R E G M I , A S H O K K U M A R R A M and N A R E S H S U B E D I

              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.
              Resettlement of communities from the core area, reduced
              anthropogenic pressure, and improved security have made
                                                                                                          T     he tiger Panthera tigris, categorized as Endangered on
                                                                                                                the IUCN Red List (Goodrich et al., ), remains in
                                                                                                          % 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 baburaml@gmail.com, baburam@ntnc.org.np                                                  Within Nepal, tigers are restricted to five protected areas
              DIPENDRA ADHIKARI, SAILENDRA RAJ GIRI and TEK RAJ BHATTA Zoological Society of
                                                                                                          and surrounding forests in the Terai Arc Landscape in the
              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
              RAJAN AMIN Zoological Society of London, Regent’s Park, London, UK
                                                                                                          tiger recovery, the Chitwan–Parsa–Valmiki forest complex
                                                                                                          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 
              MAHESHWAR DHAKAL Ministry of Forests and Soil Conservation, Sighadurbar,
                                                                                                          (Wikramanayake et al., ). It also constitutes one of the
              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 . First published online  August .                                 km) (Karki et al., ) as a result of widespread hunting

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                                                                                                      Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
Recovery of tigers in Nepal                17

                                                                                                                                           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

           Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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. https://doi.org/10.1017/S0030605317000886
18         B. R. Lamichhane et al.

             (Fig. ). The local people practised subsistence agriculture,

                                                                                                               TABLE 1 Summary of survey effort for camera-trap surveys of tigers Panthera tigris in Parsa Wildlife Reserve (Fig. ) in , ,  and . Only a partial survey was conducted in ,

                                                                                                                                                                                                                                                                                                                                          Feb.–May 2016
             but activities such as livestock grazing, and collection of fod-
             der, fuelwood, timber and other forest products resulted in
             habitat degradation over a broad area (PWR, ). Conflict

                                                                                                                                                                                                                                                                                                                                                                                        3,681
                                                                                                                                                                                                                                                                                                                         Total

                                                                                                                                                                                                                                                                                                                                          (79)

                                                                                                                                                                                                                                                                                                                                                                                        627

                                                                                                                                                                                                                                                                                                                                                                                        158
             between local people and wildlife also resulted in retaliatory

                                                                                                                                                                                                                                                                                                                                          21
             killings (CNP, ). Both communities were resettled vol-
             untarily during –. In addition to the village reloca-

                                                                                                                                                                                                                                                                                                                                          Feb.–Mar. 2016
                                                                                                                                                                                                                                                                                                                         Extension area

                                                                                                                                                                                                                                                                                                                                                                                                                         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 comparison with other years.
             tions, Parsa’s management authorities began a programme
             of habitat enrichment in , maintaining nine artificial
             waterholes in the dry season and increasing the area of un-

                                                                                                                                                                                                                                                                                                                                          (27)

                                                                                                                                                                                                                                                                                                                                                                                        128
                                                                                                                                                                                                                                                                                                                                                                                        735
             gulate grazing by .  ha of grasslands.

                                                                                                                                                                                                                                                                                                                                          21

                                                                                                                                                                                                                                                                                                                                                                                        31
                                                                                                                                                                                                                                                                                                                                          Nov. 2014–Jan. 2015 Mar.–May 2016
             Methods

             Camera trapping We set camera traps in a grid of  ×  km

                                                                                                                                                                                                                                                                                                                         Parsa

                                                                                                                                                                                                                                                                                                                                                                                        2,946
                                                                                                                                                                                                                                                                                                                  2016
             cells (DNPWC, ), deploying a pair of camera traps in

                                                                                                                                                                                                                                                                                                                                                              (79)

                                                                                                                                                                                                                                                                                                                                                                                        499

                                                                                                                                                                                                                                                                                                                                                                                        127
                                                                                                                                                                                                                                                                                                                                                              21
             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
             of  () to  days ( and ). The survey effort was
             ,–, camera-trap days (Table ). In ,  km of
             the extension area was covered with an additional  camera

                                                                                                                                                                                                                                                                                                                                                                                        3,549
                                                                                                                                                                                                                                                                                                                         2014

                                                                                                                                                                                                                                                                                                                                          (60)

                                                                                                                                                                                                                                                                                                                                                                                        499

                                                                                                                                                                                                                                                                                                                                                                                        130
             stations. The Reserve was divided into survey blocks, two in

                                                                                                                                                                                                                                                                                                                                          21
              and three in  and . The blocks were surveyed

                                                                                                                                                                                                                                                                                                                                          Apr.–May 2013
             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                                                                                                                                                                                                                                                    2

                                                                                                                                                                                                                                                                                                                                                                                        2,135
             identified. We positioned the camera traps  cm above
                                                                                                                                                                                                                                                                                                                         2013

                                                                                                                                                                                                                                                                                                                                          (59)

                                                                                                                                                                                                                                                                                                                                                                                        499

                                                                                                                                                                                                                                                                                                                                                                                        138
                                                                                                                                                                                                                                                                                                                                          15
             ground, perpendicular to, and – m either side of game
             trails, forest roads and riverbeds. Pairs of cameras were
                                                                                                                                                                                                                                                                                                                                          Dec. 2008–Mar. 2009

             placed at each sampling point to obtain images of both flanks
             of photographed individuals to assist in identification. We used
                                                                                                                                                                                                                                                                                                                         2009 (partial)

             Reconyx (Holmen, USA)  & , Bushnell (Overland Park,
             USA) Trophy Cam HD and Panthera (New York, USA) V &
                                                                                                               and information was obtained from a published report (Karki, ).

                                                                                                                                                                                                                                                                                                                         1

                                                                                                                                                                                                                                                                                                                                                                                        1,762

             V cameras. The cameras took three photographs per trigger
                                                                                                                                                                                                                                                                                                                                          (54)

                                                                                                                                                                                                                                                                                                                                                                                        353
                                                                                                                                                                                                                                                                                                                                          15

                                                                                                                                                                                                                                                                                                                                                                                        97

             with no delay, and used white flash to obtain colour images
             of tigers in low light to help with individual identification.
                                                                                                                                                                                                                                                                                                                                                                   Minimum no. of sampling occasions (1 day each) in a

             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
                                                                                                                                                                                                                                                                                                                                                                                                                         Parameters obtained from Karki ().

             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
                                                                                                                                                                                                                                                                                                                                          Sampling period (days)

                                                                                                                                                                                                                                                                                                                                                                   No. of camera stations
                                                                                                                                                                                                                                                                                                                                                                   Area surveyed (km2)

             maximum likelihood algorithm with function secr.fit was
                                                                                                                                                                                                                                                                                                                                                                   No. of trap nights

             used to fit the model. To determine the local importance
             of the former locations of villages for tigers, we
                                                                                                                                                                                                                                                                                                                         Parameters

             subsampled a buffer area of  km radius around the
                                                                                                                                                                                                                                                                                                                                                                    location

             village locations and calculated the number of tigers
             detected there.
                                                                                                                                                                                                                                                                                                                                                                                                                                                                                                 
                                                                                                                                                                                                                                                                                                                                                                                                                         

                                                                                          Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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. https://doi.org/10.1017/S0030605317000886
Recovery of tigers in Nepal                                                                             19

           Determining tiger dispersal and persistence We compared

                                                                                                       TABLE 2 Tiger capture rate, number of individual tigers captured, and population abundance and density estimates with % confidence intervals from camera-trap surveys in Parsa Wildlife

                                                                                                                                                                                                                                                                                                                                                              1.43 ± 0.33 (0.91–2.25)
           the individual tigers detected in  with camera-trap
           photographs of those recorded in ,  and  in

                                                                                                                                                                                                                                                                                                                                                              20 ± 1.9 (19–26)
           Parsa. We also compared all tigers detected in Parsa with
           tigers recorded in Chitwan National Park in  and its
           buffer zone in  (Dhakal et al., ; CNP, ), and

                                                                                                                                                                                                                                                                                                          Total

                                                                                                                                                                                                                                                                                                                           3.80
                                                                                                                                                                                                                                                                                                                           140
           in Valmiki Tiger Reserve, India, in  (Maurya & Borah,

                                                                                                                                                                                                                                                                                                                           73

                                                                                                                                                                                                                                                                                                                                             19

                                                                                                                                                                                                                                                                                                                                             10
                                                                                                                                                                                                                                                                                                                                             9
           ; Chanchani et al., ).

                                                                                                                                                                                                                                                                                                          Extension area
           Results

                                                                                                                                                                                                                                                                                                                           3.67
                                                                                                                                                                                                                                                                                                                           11
                                                                                                                                                                                                                                                                                                                           27

                                                                                                                                                                                                                                                                                                                                            32

                                                                                                                                                                                                                                                                                                                                            22
                                                                                                                                                                                                                                                                                                                                            1
           Tiger density and abundance During – tiger

                                                                                                                                                                                                                                                                                                                                                              1.38 ± 0.34 (0.85–2.25)
           density in Parsa increased from . ± SE . to

                                                                                                                                                                                                                                                                                                                                                              17 ± 1.53 (17–20)
           . ± SE . tigers per  km (Table ). Five,  and 
           individual tigers were detected in ,  and ,
           respectively, with an additional two tigers detected in the
           extension area in . Tiger abundance increased

                                                                                                                                                                                                                                                                                                          Parsa
                                                                                                                                                                                                                                                                                                   2016

                                                                                                                                                                                                                                                                                                                           3.84
                                                                                                                                                                                                                                                                                                                           113
           gradually in Parsa during – (Fig. ).

                                                                                                                                                                                                                                                                                                                           62

                                                                                                                                                                                                                                                                                                                                             17
                                                                                                                                                                                                                                                                                                                                             8
                                                                                                                                                                                                                                                                                                                                             9

                                                                                                                                                                                                                                                                                                                                                              0.80 ± 0.25 (0.43–1.48)
           Evidence of dispersal from Chitwan Over the  sampling

                                                                                                                                                                                                                                                                                                                                                              11 ± 1.27 (10–16)
           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
           additional three individuals (two females and one male)                                                                                                                                                                                                                                        2014

                                                                                                                                                                                                                                                                                                                           2.17
                                                                                                                                                                                                                                                                                                                           45
                                                                                                                                                                                                                                                                                                                           77

                                                                                                                                                                                                                                                                                                                                             10
           were captured in  years ( and ). Ten new

                                                                                                                                                                                                                                                                                                                                             4
                                                                                                                                                                                                                                                                                                                                             6
           individuals were captured in . Of the  individuals

                                                                                                                                                                                                                                                                                                                                             0.78 ± 0.39 (0.31–1.98)
           recorded, nine were captured first in Chitwan (in  and
           ) and then dispersed to Parsa (in  and ). One

                                                                                                                                                                                                                                                                                                                                             7 ± 1.53 (6–13)
           tiger (F, Table ) was first captured in Parsa () and
           dispersed to Chitwan’s buffer zone () in the south
           (Someshwar Hills, contiguous to Valmiki). There was no
                                                                                                                                                                                                                                                                                                          2013

                                                                                                                                                                                                                                                                                                                           0.38
                                                                                                                                                                                                                                                                                                                           10

           evidence of tigers dispersing between Valmiki and Parsa.
                                                                                                                                                                                                                                                                                                                           9

                                                                                                                                                                                                                                                                                                                                             5
                                                                                                                                                                                                                                                                                                                                             2
                                                                                                                                                                                                                                                                                                                                             2
                                                                                                                                                                                                                                                                                                                                             1

                                                                                                                                                                                                                                                                                                                                                              0.61 ± 0.32 (0.23–1.62)

           Recolonization of tigers in evacuated village sites Tigers
           were confined to a narrow strip in the middle of the
                                                                                                                                                                                                                                                                                                                                                                                        A tigress was captured from both Parsa and the extension area.

           Reserve during  (Fig. a) but occupied most of the
                                                                                                                                                                                                                                                                                                                                                              4 ± 0.47

           Reserve in the following years (Fig. b,c). The  survey
                                                                                                                                                                                                                                                                                                                                                                                        Partial survey conducted; estimates are from Karki ().
                                                                                                                                                                                                                                                                                                          1
                                                                                                                                                                                                                                                                                                          2009

           failed to detect a single tiger within the  km buffer of
                                                                                                                                                                                                                                                                                                                                             4
                                                                                                                                                                                                                                                                                                                                             2
                                                                                                                                                                                                                                                                                                                                             2

           Ramauli Pratappur village, which was in the process of
                                                                                                       Reserve (Fig. ) in , ,  and .

                                                                                                                                                                                                                                                                                                                           Tiger capture rate (no. of detections per 100
                                                                                                                                                                                                                                                                                                                           No. of camera stations that captured tigers

           relocation at the time. The following year four tigers were
                                                                                                                                                                                                                                                                                                                           No. of individual tigers captured (Mt + 1)

           detected within the  km buffer, two of which were
                                                                                                                                                                                                                                                                                                                           No. of independent detections of tigers

           redetected in the same area in  (Fig. ). Within the
                                                                                                                                                                                                                                                                                                                           Density ± SE per 100 km2 (95% CI)

            km buffer surrounding the site of Rambhori Bhata
                                                                                                                                                                                                                                                                                                                           Tiger abundance ± SE (95% CI)

           village, which was relocated in , two tigers were
           detected in , four in  and seven in  (Fig. ).

           Discussion
                                                                                                                                                                                                                                                                                                                              Unknown
                                                                                                                                                                                                                                                                                                                            trap days)
                                                                                                                                                                                                                                                                                                          Parameters

                                                                                                                                                                                                                                                                                                                              Female

           Our findings indicate a nearly threefold increase in tiger
                                                                                                                                                                                                                                                                                                                              Male

           numbers in Parsa within  years. This is a unique scenario
           of tiger recovery, beyond the potential of natural growth
                                                                                                                                                                                                                                                                                                                                                                                                                
                                                                                                                                                                                                                                                                                                                                                                                        

           Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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20         B. R. Lamichhane et al.

                                                                                                                                             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

                                                                                          Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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Recovery of tigers in Nepal                21

                                                                                                 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

           Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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22         B. R. Lamichhane et al.

             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 –. Providing auxiliary support to Parsa’s security                            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.

                                                                                          Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
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. https://doi.org/10.1017/S0030605317000886
Recovery of tigers in Nepal                  23

           Leiden University, Netherlands supported the first author                                           G O O D R I C H , J.M., L Y N A M , A., M I Q U E L L E , D., W I B I S O N O , H.,
           during manuscript preparation and Open Access publica-                                                  K AWA N I S H I , K., P AT TA N AV I B O O L , A. et al. () Panthera tigris. In
                                                                                                                   The IUCN Red List of Threatened Species : e.TA.
           tion. The camera-trap surveys would not have been possible
                                                                                                                   Http://dx.doi.org/./IUCN.UK.-.RLTS.TA.
           without the tireless effort of field staff, including officers,                                         en [accessed  June ].
           rangers and game scouts of Parsa Wildlife Reserve, wildlife                                         GTI (G LO B A L T I G E R I N I T I AT I V E ) () 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.
                                                                                                               GTI (G LO B A L T I G E R I N I T I AT I V E ) () Global Tiger Recovery
           Kathmandu University. We acknowledge the contribution
                                                                                                                   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                                                  G U R U N G , B., S M I T H , J.L.D., M C D O U G A L , C., K A R K I , J.B. & B A R LO W ,
           for their valuable comments on the article.                                                             A. () Factors associated with human-killing tigers in Chitwan
                                                                                                                   National Park, Nepal. Biological Conservation, , –.
                                                                                                               H A R I H A R , A. & P A N D AV , 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,                                            H A R I H A R , A., P A N D AV , B. & G O YA L , S.P. () Responses of tiger
                                                                                                                   (Panthera tigris) and their prey to removal of anthropogenic
           SB and URR implemented the fieldwork and collected data.
                                                                                                                   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.                                                                               H I B Y , L., L O V E L L , P., P AT I L , N., K U M A R , N.S., G O P A L A S WA M Y , A.M.
                                                                                                                   & K A R A N T H , K.U. () A tiger cannot change its stripes: using a
                                                                                                                   three-dimensional model to match images of living tigers and tiger
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               Department of National Parks and Wildlife Conservation,                                             ten years later. Ambio, , –.
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               R package version ... Https://CRAN.R-project.org/package=                                       Adhabhar, Bara, Nepal.
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             S C H M I D T -S O LT A U , K. & B R O C K I N G T O N , D. () Protected areas                           conservation strategy to double the wild tiger population.
                 and resettlement: what scope for voluntary relocation? World                                             Conservation Letters, , –.
                 Development, , –.                                                                            W I K R A M A N AYA K E , E., M C K N I G H T , M., D I N E R S T E I N , E., J O S H I , A.,
             S M I T H , J.L.D. () The role of dispersal in structuring the Chitwan                                   G U R U N G , B. & S M I T H , D. () Designing a conservation
                 tiger population. Behaviour, , –.                                                               landscape for tigers in human‐dominated environments.
             S U N Q U I S T , M.E. () The Social Organization of Tigers (Panthera                                    Conservation Biology, , –.
                 tigris) in Royal Chitawan National Park. Smithsonian Institution
                 Press, Washington, DC, USA.
             T H A P A , K., S H R E S T H A , R., K A R K I , J., T H A P A , G.J., S U B E D I , N.,
                 P R A D H A N , 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/.                                     BA B U RA M LA M I C H H A N E is interested in the application of geograph-
                 //.                                                                                      ical information systems (GIS) for wildlife management, and human–
             W A L S T O N , J., R O B I N S O N , J.G., B E N N E T T , E.L., B R E I T E N M O S E R , U., D A        wildlife interactions. CH I R A N J I B I PR A S A D PO K H E R A L is interested in
                 F O N S E C A , G.A.B., G O O D R I C H , J. et al. () Bringing the tiger back                     large carnivore biology and interspecies interactions. SH A S H A N K
                 from the brink—the six percent solution. PLoS Biology, (),                                           PO U D E L is interested in conservation communication. DI P E N D R A
                 e.                                                                                              AD H I K A R I is interested in small mammals. SA I L E N D R A RA J GI R I is in-
             W E G G E , P., O D D E N , M., P O K H A R E L , C.P. & S T O R A A S , T. ()                         terested in applications of GIS and remote sensing in wildlife conserva-
                 Predator–prey relationships and responses of ungulates and their                                       tion. SA N T O S H BH A T T A R A I is interested in herpetology. TE K RA J
                 predators to the establishment of protected areas: a case study of                                     BH A T T A is involved in development and implementation of conserva-
                 tigers, leopards and their prey in Bardia National Park, Nepal.                                        tion projects. RO B PI C K L E S specializes in tiger monitoring using camera
                 Biological Conservation, , –.                                                                 traps. RA J A N AM I N specializes in the conservation of Asian and African
             W I K R A M A N A YA K E , E., D I N E R S T E I N , E., R O B I N S O N , J.G.,                           grassland and forest ecosystems. KR I S H N A PR A S A D AC H A R Y A is inter-
                 K A R A N T H , U., R A B I N O W I T Z , A., O L S O N , D. et al. () An                          ested in resolving human–wildlife conflicts. MA H E S H W A R DH A K A L is
                 ecology‐based method for defining priorities for large mammal                                          involved in policy analysis of protected areas. UB A RA J RE G M I has
                 conservation: the tiger as case study. Conservation Biology, ,                                       served as manager of Parsa Wildlife Reserve and other protected areas
                 –.                                                                                               in Nepal. AS H O K KU M A R RA M is involved in geospatial modelling of
             W I K R A M A N A YA K E , E., D I N E R S T E I N , E., S E I D E N S T I C K E R , J., L U M P K I N ,   wild elephants in Central Nepal. NA R E S H SU B E D I is interested in the
                 S., P A N D AV , B., S H R E S T H A , M. et al. () A landscape-based                              ecology of rhinoceroses and invasive species.

                                                                                                                Oryx, 2018, 52(1), 16–24 © 2017 Fauna & Flora International doi:10.1017/S0030605317000886
Downloaded from https://www.cambridge.org/core. IP address: 46.4.80.155, on 10 May 2021 at 18:24:58, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms
. https://doi.org/10.1017/S0030605317000886
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