Determinants of Tiger Prey Base Distribution and Relative

Abundance in Bardia National Park and Adjoining Khata

Corridor

Naresh Khanal
Email: [email protected]
B.Sc. Forestry (2071-2075)
Institute of Forestry (IOF), Pokhara Campus

A Project Paper submitted for the partial fulfillment of Bachelor of Science in

Forestry Degree.

April 2019
 Determinants of Tiger Prey Base Distribution and Relative
Abundance in Bardia National Park and Adjoining Khata Corridor

Submitted To
Institute of Forestry (IOF)
Pokhara Campus, Pokhara

Principal Investigator

Naresh Khanal
Email: [email protected]
B.Sc. Forestry (2071-2075)
Institute of Forestry (IOF), Pokhara Campus

Research Advisor

Narayan Prasad Gautam

Asst. Professor,
Institute of Forestry (IOF), Pokhara Campus
Email: [email protected]

Co-Advisor
Santosh Rayamajhi, PhD
Professor, IOF Pokhara Campus
Email: [email protected]

A Project Paper submitted for the partial fulfillment of Bachelor of Science in

Forestry Degree.

April 2019
© Naresh Khanal

April 2019
E-mail:
[email protected]

Tribhuvan University
Institute of Forestry (IOF) Pokhara Campus, Pokhara Nepal
Website: www.iofpc.edu.np
Post Box No.: 43

Citation:
Khanal, N., 2019. Determinants of Tiger Prey Base Distribution and Abundance in Bardia
National Park and Adjoining Khata Corridor: A Project paper submitted for the partial
fulfillment for the degree in Bachelor of Science in Forestry, Tribhuvan University, Institute of
Forestry, Pokhara Campus, Pokhara, Nepal.
LETTER OF ACCEPTANCE

i
 DECLARATION

I, Naresh Khanal, hereby declare that this project paper entitled “Determinants of Tiger Prey
Base Distribution and Relative Abundance in Bardia National Park and Adjoining Khata
Corridor” is a presentation of my original study. Wherever contributions of others are involved,
every effort is made to indicate this clearly, with due reference to the literature, and
acknowledgement of collaborative research and discussions. This study was done under the
supervision of Mr. Narayan Prasad Gautam, Assistant Professor at the Tribhuvan University,
Institute of Forestry (IOF), Pokhara Campus. The results presented in this project paper have not
been submitted previously to this or any other university for a BSc or any other degree.

…………………..
Naresh Khanal
Batch: 2071-2075
Campus Roll no.: 18
TU Registration No.: 2-2-47-36-2014

ii
 ACKNOWLEDGEMENT

This project paper would not have been possible without the support of many people. My
foremost gratitude and deepest appreciation go to my advisor Assistant Professor Mr. Narayan
Prasad Gautam for his humble support, incredible guidance, advice and invaluable
encouragement throughout the study. I would like to extent my sincerest thanks to my respected
faculties of IOF Pokhara Campus Assistant Professors respectively Mrs. Nirjala Raut, Mr. Bijaya
Neupane, Mr. Bishnu Devkota, and Associate Professors Mr. Jeevnath Pandey and Dr. Arjun
Prasad Bastola for their constant guidance and continuous support during my study period. I
would like to thank Assistant Professor Dr. Thakur Silawal (Former for Campus Chief/IOF
Pokhara Campus) for his constructive comments and suggestions. I am thankful to the
Department of the National Park and Wildlife Conservation (DNPWC) for providing permission
to conduct this study. I am extremely thankful to Professor Dr. Santosh Rayamajhi and Pravin
Paudel for their technical guidance during my field work. My special thanks go to Mr. Hemraj
Acharya (Ecologist/DNPWC), Mr. Ambhika Prasad Khatiwada (NTNC Head Office), Mr.
Ashok Bhandari (Conservation Officer/BNP), Mr. Shyam Thapa (Incharge NTNC/BCP), Umesh
Paudel (NTNC/BCP), Rangers Mr. Saroj Khadka, Mr. Kush Raj Gole, Mr. Harsha Bahadur
Singh, Mr. Ram KC, Mr. Kabiraj Praja and game scouts Mr. Khum Bahadur Mahatara and Mr.
Kishor Mahat for their continuous support and suggestions throughout the field work. I thank
my senior brothers Mr. Navaraj Shahi and Mr. Sujan Khanal as well as my friends Mahesh
Prasad Tiwari, Milan Buda and Aavash Paudel for their help in GIS analysis and map
preparation. I would like to thank my room partner Madan Acharya and classmates Pravin,
Niranjan, Saroj, Prasun, Bhumi, Kushal, Kabiraj, Jawaf, Mausam, Amrit, Divas and Ashok for
their encouragement to conduct this work. I also thank Mr. Prakash Chaudhary for his overall
assistance during my field data collection. I also thank all my batch mates (Batch 071-075), my
seniors Mr. Rohit, Mr. Kabindra. Mr. Dipesh, Mr. Dipak, Mr. Keshav and juniors Apsana,
Saroon, Viddyaman, Kiran, Aashish, Sirish, Sabitri, Isha, Mrs. Sima khanal and entire CES
family who directly or indirectly supported me for this work. I am greatly indebted to the all the
IOF Family, Mr. Pradip Subedi and all other staffs for their great help. I owe a debt of gratitude
to all the authors of the literatures cited. Last but not the least, my deepest gratitude goes to my
family and my brother Gopal Khanal for their inspiration and support.

iii
 ABSTRACT
Background

Large carnivores are among the most important yet highly threatened group of species. Their
survival critically depends on prey populations and thus carnivore conservation efforts are often
focused on restoring depleted prey population. Conservation of endangered species like Panthera
tigris tigris and achieving the goal of doubling tiger population by 2022 requires recovery of
prey populations. Therefore, knowledge of prey population and their distributional response
along the human disturbance gradient is important in guiding tiger conservation efforts.

Objectives
The main objective of this study was to assess the impact of human disturbance (as indicated by
human encounter rate and livestock grazing intensity), habitat type (grassland; mixed forest;
riverine forest; agricultural land) and management regime (national park; buffer zone; outside
buffer zone) on relative abundance of key prey species of tiger in Bardia National Park, its
buffer zone and Khata Corridor area.

Methods and materials

I conducted sign surveys along 1-km transects in total of 126 transects in 32 grids of each 9 km2,
and recorded signs of each species for each 200 m segment of transects and evidence of human
presence, livestock grazing and habitat type. The total survey area was 288 km2. First, I
compared whether the sign encounter rate of each species differs according to the habitat type
and management regime. I then fit Generalized linear models to test the influence of four
predictor variables (human encounter rate, livestock grazing intensity, habitat type and
management regime) on Relative Abundance Index (RAI) or sign encounter rate per km of prey
species and checked the slope parameters or beta coefficient of predictor variables to infer their
influence/association with predictor variable.

Results
Chital was found to have highest mean relative abundance or mean encounter rate per km
(RAI=1.74, SD 1.96) whereas swamp deer had lowest encounter rate (RAI = 0.05, SD 0.31).
Overall species encounter rate showed mixed result for their distribution according to habitat
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type, some species preferring mixed forest (spotted deer 2.40 SER, wild boar 1.90 SER) while
some preferring grassland (hog deer 1.38 SER). Similarly, national park had highest encounter
rate of spotted deer, swamp deer, sambar, species whereas wild boar, blue bull and barking deer
species had highest encounter rate in outside buffer zone area. Generalized linear models showed
consistent negative influence of human encounter rate and livestock grazing influence on relative
abundance of all species expect wild boar and barking deer. Interestingly, both species had
positive association with human encounter rate but not with livestock grazing.

Discussions and recommendations

The result shows that prey species have differential response to human disturbance, habitat type
and management regime as indicated by their relative sign encounter rate/RAI. These results
suggest that some species are highly vulnerable to human disturbance (e.g., deer which had
negative association with human disturbances and other are to some extent resilient to human use
(e.g., wild boar and blue bull; as they had higher encounter rate outside the buffer zone).
Differential response to habitat type and management regime perhaps suggest that integrated
management of all habitat types and management regimes would be necessary to sustain the
collective prey base recovery of all species for tiger conservation. Most importantly, highest RAI
in mixed forest and grassland habitat indicates that appropriate mixture and distribution of
grassland and forest patches would be preferable from overall point of view of prey base
conservation. Moreover, the finding that some of the prey species occurring in buffer zone and
outside buffer zone area can be taken as an opportunity to include the local community in
conservations activities formulating appropriate plans.

Keywords: Tiger prey species, abundance, management regime, habitat type, Bardia National
Park

v
 ACRONYMS

AIC Akaike Information Criteria

AGL Agricultural Lands
BNP Bardia National Park
BZ Buffer Zone
CNP Chitwan National Park
DNPWC Department of National Park and Wildlife Conservation
FIG Figure
GIS Geographic Information System
GL Grasslands
GTRP Global Tiger Recovery Program
IUCN International Union for Conservation of Nature
KM Kilometer
KM2 Square Kilometer
MF Mixed Forests
NP National Park
OBZ outside Buffer Zone
NTNC National Trust for Nature Conservation
PA Protected Area
RF Riverine Forests
SE Standard Error
SPNP Sukhlaphanta National Park
TAL Terai Arc Landscape
TCL Tiger Conservation Landscape
TRC Tiger Range Countries
VDC Village Development Committee

vi
 Table of Contents

LETTER OF ACCEPTANCE ..................................................................................................... i

DECLARATION ....................................................................................................................... ii

ACKNOWLEDGEMENT ......................................................................................................... iii

ABSTRACT ............................................................................................................................. iv

ACRONYMS ............................................................................................................................ vi

CHAPTER 1: INTRODUCTION ................................................................................................1

1.1 Background .......................................................................................................................1

1.2 Rationale of the Study .......................................................................................................4

1.3 Research Objectives ..........................................................................................................5

1.3.1 General Objectives......................................................................................................5

1.3.2 Specific Objectives .....................................................................................................5

CHAPTER 2: LITERATURE REVIEW .....................................................................................6

CHAPTER 3: MATERIALS AND METHODS ..........................................................................9

3.1 Study Area ........................................................................................................................9

3.1.1 Bardia National Park ...................................................................................................9

3.1.2 Khata Corridor ............................................................................................................9

3.1.3 Climate ..................................................................................................................... 10

3.1.4 Vegetation ................................................................................................................ 10

3.1.5 Fauna ........................................................................................................................ 12

3.2 Data Collection................................................................................................................ 12

3.2.1 Sampling design ....................................................................................................... 12

3.2.2 Line transect surveys for distribution mapping .......................................................... 13

vii
 3.2.3 Relative abundance assessment ................................................................................. 14

3.2.4 Quantification of ecological and anthropogenic covariates ........................................ 14

3.3 Data Analysis .................................................................................................................. 15

3.3.1 Distribution mapping ................................................................................................ 15

3.3.2 Relative abundance along the gradient of human disturbance, and with respect to
habitat type and management regime. ................................................................................ 16

CHAPTER 4: RESULTS .......................................................................................................... 18

4.1 Distribution mapping ....................................................................................................... 18

4.2 Relative Abundance Index ............................................................................................... 20

4.2.2 Management Regime-wise RAI ................................................................................ 21

4.2.3 Habitat-wise RAI ...................................................................................................... 23

4.3 Species specific relative encounter rate according to habitat and management category ... 24

4.3.1 Spotted Deer ............................................................................................................. 24

4.3.2 Wild Boar ................................................................................................................. 25

4.2.3 Barking Deer ............................................................................................................ 26

4.2.4 Hog Deer .................................................................................................................. 28

4.2.5 Sambar ..................................................................................................................... 29

4.2.6 Blue Bull .................................................................................................................. 30

4.2.7 Swamp deer .............................................................................................................. 32

4.4 Species specific Generalized Linear Models (GLMs) ....................................................... 33

4.4.1 Spotted deer .............................................................................................................. 33

4.4.2 Wild Boar ................................................................................................................. 36

4.4.3 Blue Bull .................................................................................................................. 38

CHAPTER 5: DISCUSSIONS .................................................................................................. 41

5.1 Spotted Deer .................................................................................................................... 41

viii
 5.2 Wild Boar ........................................................................................................................ 42

5.3 Barking Deer ................................................................................................................... 42

5.4 Sambar ............................................................................................................................ 42

5.5 Hog Deer ......................................................................................................................... 43

5.6 Swamp Deer .................................................................................................................... 43

5.7 Blue Bull ......................................................................................................................... 43

CHAPTER 6: CONCLUSIONS AND RECOMMENDATIONS ............................................... 45

6.1 Conclusions ..................................................................................................................... 45

6.2 Recommendations ........................................................................................................... 45

REFERENCES ......................................................................................................................... 47

ANNEX 1: FIELD DATA COLLECTION FORM .................................................................... 52

ANNEX 2: PHOTO PLATES ................................................................................................... 53

List of Figures

Figure 1: Terai Arc Landscape showing various PAs of Nepal and India ................................... 10
Figure 2: Map showing study area and grids surveyed ............................................................... 11
Figure 3: Distribution map of barking deer Figure 4: Distribution map of spotted deer ..... 18
Figure 5: distribution map of blue bull Figure 6: distribution map of hog deer ..................... 19
Figure 7: distribution map of wild boar ..................................................................................... 19
Figure 8: Naïve occupancy of prey species ................................................................................ 19
Figure 9: Overall species encounter rate /RAI ........................................................................... 21
Figure 10: Overall species encounter rate according to the management category...................... 22
Figure 11: Overall species encounter rate according to the habitat type ..................................... 23
Figure 12: Relative encounter rate of spotted deer according to management regime ................. 24
Figure 13: Relative encounter rate of wild boar according to management regime ..................... 25
Figure 14: Relative encounter rate of barking deer according to management regime ................ 26
Figure 15: Relative encounter rate of hog deer according to habitat type ................................... 28
Figure 16: Relative encounter rate of hog deer according to management regime ...................... 28
Figure 17: Relative encounter rate of sambar according to habitat type ...................................... 29
Figure 18: Relative encounter rate of sambar according to management regime ........................ 30
Figure 19: Relative encounter rate of blue bull according to habitat type ................................... 31
Figure 20: Relative encounter rate of blue bull according to management regime ...................... 31
Figure 21: Relative encounter rate of swamp deer according to habitat type .............................. 32
Figure 22: Relative encounter rate of swamp deer according to management regime ................. 32

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Figure 23: Relationship between human encounter rate and relative abundance index of spotted
deer ........................................................................................................................................... 34
Figure 24: Relationship between livestock grazing intensity (encounter rate of livestock signs)
rate and relative abundance index of spotted deer ...................................................................... 35
Figure 25: Relationship between management regime (buffer zone=BZ, national park=CA, and
outside buffer zone (OBZ) and relative abundance index of spotted deer ................................... 35
Figure 26: Relationship between habitat type (MF, GL, MF & Fallow AG lands) and relative
abundance index of spotted deer ................................................................................................ 36
Figure 27: Relationship between human encounter rate and relative abundance index of wldboar
................................................................................................................................................. 37
Figure 28: Relationship between livestock grazing intensity (encounter rate of livestock signs)
rate and relative abundance index of wildboar ........................................................................... 37
Figure 29: Relationship between human encounter rate and relative abundance index of blue bull
................................................................................................................................................. 39
Figure 30: Relationship between livestock grazing intensity (encounter rate of livestock signs)
rate and relative abundance index of bluebull ............................................................................ 39

List of Tables
Table 1: Plausible ecological and anthropogenic covariates that are hypothesized to influence
prey species distribution. ........................................................................................................... 15
Table 2: Species sign encounter rate and associated standard deviation ..................................... 20
Table 3: RAI of Spotted deer ..................................................................................................... 25
Table 4: RAI of wild boar ......................................................................................................... 26
Table 5: RAI of barking deer ..................................................................................................... 27
Table 6: RAI of hog deer ........................................................................................................... 29
Table 7: RAI of sambar ............................................................................................................. 30
Table 8: RAI of blue bull .......................................................................................................... 31
Table 9: RAI of swamp deer...................................................................................................... 32

Table 10: Poisson regression models describing the relative abundance of spotted deer in BNP
(Fig. 2), ranked according to the Akaike Information Criterion adjusted for small sample size
(AICc). ...................................................................................................................................... 33

Table 11: Poisson regression models describing the relative abundance of wild boar in BNP (Fig.
2), ranked according to the Akaike information criterion adjusted for small sample size (AICc) 38

Table 12: Poisson regression models describing the relative abundance of blue bull in BNP (Fig.
2), ranked according to the Akaike information criterion adjusted for small sample size (AICc) 40

x
 CHAPTER 1: INTRODUCTION

1.1 Background

Large terrestrial carnivores are among the most charismatic and ecologically important group of
species. Globally, out of 245 species of terrestrial carnivores (order Carnivora), 31 are large
bodied (more than 15 kg) carnivore. Large mammalian carnivores (more than 15 kg, order
Carnivora) are characterized by their large body size and movement ranges, sparse population,
low density, high dietary requirements, and more importantly their charismatic nature in
capturing human emotions (Carbone et al., 1999; Carbone and Gittleman 2002; Ripple et al.,
2014). They play multiple direct and indirect role in structuring and maintaining ecosystem
balance through tropic cascades (Estes et al., 2011; Terborgh et al., 2001). The various researches
has found that these predators play leading role in regulating the ecosystem as they exert the top
down pressure on prey communities thereby reducing the herbivory of the plant communities
(Ripple and Beschta, 2012). For example, Canada et al., (2005) showed that wolves (Canis
lupus) reduced elk (Cervus canadensis) numbers which led to change in plant species
composition. Carnivores also have social and human benefits because they have become an
important aspects of the wildlife tourisms (Lindsey et al., 2007). Chitwan National Park, as an
example holds the high tourist value as people travel to see tiger in the wild (Thapa et al., 2017).
This has directly and indirectly helped grow the local as well as national economy of the country
(Karanth et al., 2012). Recently, Puma, a species found in North America has been found to act
as an ecosystem engineer (Barry et al., 2018) contributing in shaping of ecosystem as the large
portion of its abandoned prey is utilize by various invertebrates including beetles.

Unfortunately, despite this significant ecological and economic importance, large carnivores are
most threatened group of species (Ripple et al., 2014) in the contemporary world. The same
study by Ripple et al., (2014) has stated that although they have received considerable
conservation attention, funding and research efforts worldwide, they continue to decline in
numbers, distribution range (Wolf and Ripple, 2017), and their ecological functions due to
multiple threats (Cardillo et al., 2005), particularly human induced threats/human population
density (Cardillo et al., 2004), edge effect of human population growth along the protected area

1
boundaries (Balme et al., 2010; Joppa et al., 2009; Wittemyer et al., 2008; Woodroffe, 1998),
hunting and poaching for their furs and body parts to feed international trade (Alkemade et al.,
2017), habitat loss and fragmentation (Wikramanayake et al.1998; Miquelle et al., 1999;
Dinerstein et al., 2007), land use changes for agriculture and developmental activities such as
promoting roadside conflict (Function, 2009), prey depletion (Wolf and Ripple, 2016b), and
conservation conflicts (Treves and Karanth, 2003). According to the International Union for the
Conservation of Nature (IUCN) , about 61% of these large carnivores are classified as threatened
(vulnerable, endangered, or critically endangered) and face high risk of extinction (Ripple et al.,
2014). They have been wiped out from much of their historic distribution ranges (Wolf and
Ripple, 2017). Only 17 of the 31 species currently occupy on average only 47% (minimum <1%,
maximum 73%) of their historical ranges and 77% of these species continue to undergo decline
in their populations (Ripple et al., 2014).

While carnivores are most targeted group of species for conservation, they have never been
threatened at the level they are today (Ripple et al., 2014). On the one hand they continue to be
threatened by ever expanding human induced loss of their natural habitat, food resources,
poaching, disease, conflict over space and between conservation stakeholders (Di Minin et al.,
2016; Maxwell et al., 2016; Wolf and Ripple, 2016; Woodroffe and Ginsberg, 1998; Woodroffe,
2000). Whereas on the other hand, limited knowledge of their status and biology, despite some
carnivore being widely studied, have been key obstacles in informing appropriate measures to
conserve their habitats and populations. Particularly, research on prey depletion has received
comparatively lesser effort (Wolf and Ripple, 2016).

For the balanced survival of these large carnivores, they need abundant amount of prey (Carbone
and Gittleman, 2002; Carbone et al., 1999). Likewise, Carbone & Gittleman, 2002 has illustrated
the strong relationship between the mammalian prey and the abundance of carnivores. In the
same study, the author has found that irrespective of the species, approximately 10,000 kg of
prey is needed to support the overall survival of a carnivore of approximate 90 kg weight. There
is a chance of carnivore population decline and eventually being locally extinct if the prey is not
available in sufficient quantity (Wolf and Ripple, 2016). A recent comprehensive review of prey
depletion as threats to world’s large carnivore by the same study of Wolf and Ripple, (2016)
suggest that loss of prey base is a major and wide-ranging threat among large carnivore species

2
for their survival. Research work so far suggests that appropriate amount of prey availability to
key to survival of large carnivores and conservation management efforts including rewinding and
recovery of carnivore populations requires conservation of prey base.

For tigers, one of the most endangered species of large carnivore, which has suffered over 93%
of its historical range in last 100 years (Sanderson, 2015) and prey depletion has been key
conservation threats (Chapron et al., 2008; Karanth and Stith, 1999; Ramakrishnan et al., 1999).
Habitat loss, livestock grazing, resource extraction and subsistence hunting has largely
contributed to decline in wild ungulate prey species of tiger (Datta et al., 2008; Linkie et al.,
2018; Madhusudan, 2004). Other key threats to tigers include poaching (Sharma et al., 2014),
habitat loss and fragmentation (Wikramanayake et al., 1998) including human tiger conflict.
Over the last century global tiger population dropped from some 100,000 to less than 3000 in the
beginning of 21 century. Currently, it is distributed in about less than 5% of 1.5 million km 2
forested habitat (Walston et al., 2010) of 13 countries and population is estimated to be about
2154-3159 (Goodrich et al., 2015). Due to this precarious nature of tiger, the 13 tiger range
countries (TRCs) with support from international donor and conservation agencies (St.
Petersburg Tiger summit) in 2010 committed to a 12-year goal of doubling wild tiger numbers
by 2022. In order to achieve this ambitious goal, all the responsible authority must have a
commitment and must perform realistic actions. Implementing the targeted actions efficiently, it
is also needed systematic monitoring of the progress towards achieving the target (Harihar et al.,
2018). In order to achieve this goal, the Global Tiger Recovery Program (GTRP) is concentrating
the efforts and mobilizing the supports from various donor agencies, outlining the action plans to
strengthening policies building constitutional frameworks and securing financial commitments.
Apart from preventing habitat loss and fragmentation, ensuring continued conservation support,
maintaining connectivity, reducing poaching, ensuring sufficient prey populations is the key to
achieve the tiger population recovery targets. A recent study (Harihar et al., 2018) evaluating the
tiger population recovery potential of 18 sites suggests that recovery potential of the 15 of 18
sites is contingent on the initial recovery of prey populations. This study shows that in case of
Nepal, which holds approximately 235 tigers in five PAs, and which is one the way to double its
tiger population by 2022, increase in prey populations is crucial in meeting the tiger population
recovery targets. Increase in prey population and hence tiger population depends upon how well

3
the habitat is managed and how well the prey population is protected from other anthropogenic
threats. Designing appropriate habitat management plans or strategies to recover prey population
requires reliable scientific understanding of factors affecting prey population and density and
other management and ecological factors operating at different management scale, habitat type,
protected area and landscape scale.

1.2 Rationale of the Study

To achieve doubling the tiger population by 2022, strategies to recover wild tiger population
have been proposed by Waltson et al., (2010) and Wikramanayake et al., (2011), which basically
deal with maintaining the healthy habitat for tiger which constitute the enough ungulate
populations for their prey. While these strategies have been instrumental in guiding conservation
investments at appropriate locations, both local and regional scale conservation initiatives to date
however, are largely based on the limited understanding of distribution and preferences of the
prey population and tiger-prey-habitat relationships. Currently we are following the recovery
targets which are set by estimates of potential tiger densities assigned to biomes/ major habitat
types (Wikramanayake et al., 2011). However, they do not address the heterogeneity that
prevails in habitat features and management regimes within landscapes (Harihar et al., 2014).
The dynamic nature of forest ecosystem is not duly accounted in current tiger conservation
strategies.

Moreover, role of ungulates to determine the population size of carnivores is widely studied and
is considered to be one of the major factors for survival of predators (Carbone and Gittleman,
2002; Karanth et al., 2004). Similarly, Karanth et al., (2004) has stated that tiger density is a
function of prey densities. In order to recover the declined tiger populations and implementation
of effective conservation intervention in human dominated landscapes, we need scientific
understanding of the prey of tiger and the factors affecting the distribution of these prey species.
For on-the-ground implication of the effective and prioritize conservation intervention of tiger,
we require the sound understanding of the ecological and anthropogenic correlates of the
distribution of the prey species. While formulating the various strategies to protect tiger habitats
at different level i.e. protected area level and landscape level, in both the case, it is essential to
investigate the effect of different habitat and anthropogenic variables on ungulates occurring at

4
multiple scales. However, in the context of Nepal, we lack the adequate knowledge on the
ecological and anthropogenic factors responsible for the distribution of the ungulate species.
Though some of the studies outsides Nepal have provided some sorts of understanding about the
effects of prey abundance, human disturbances and habitat connectivity (e.g. Barber-Meyer et
al., 2018; Karanth et al., 2011), we still have knowledge deficit in understanding how the
different habitat type such as Sal forest, Mixed Sal forest, Riverine forest, Grassland area, etc.
and habitat quality including the human interventions influence the density and distribution of
these prey species. Because of this critical knowledge gap, we have remained unable to
effectively formulate and implement the evidence-based tiger conservation. Hence, I want to
study how the tiger prey species in a protected area and its surrounding buffer zone and
community forests are collectively influenced by different ecological and anthropogenic factors
including habitat types, livestock grazing and human disturbances.

1.3 Research Objectives

1.3.1 General Objectives

 To determine the tiger prey base distribution and relative abundance in Bardia National
Park and Adjoining Khata Corridor

1.3.2 Specific Objectives

 To assess the influence of different ecological and anthropogenic factors on distribution

of major prey species of tigers
 To compare the relative abundance of tiger prey species in with respect to habitat type
(mixed Sal forest, grassland, riverine forest and agriculture fallow lands) and
management regime ( national park, buffer zone, and outside buffer zone

5
 CHAPTER 2: LITERATURE REVIEW
Carnivores constitute basically the flesh-eating placental mammals of scrotiferan order. Broadly
we can classify the carnivores as Feliformia (cat like) and Caniformia (dog like). These carnivore
species including tiger is an important species from the ecological viewpoint. They play multiple
direct and indirect roles in structuring and maintaining ecosystem balance through tropic
cascades (Estes et al., 2011; Terborgh et al., 2001). The carnivores feed on prey species which in
turn creates indirect effect with large number of other species thus contribute in balancing the
ecosystem. For example, Canada et al., (2005) has found out that elk (Cervus canadensis)
numbers can be directly reduced or changed by the predator wolves (Canis lupus) which may
further lead to change in plant species composition. Prey-predator relationship naturally shapes
and controls the herbivore population in a terrestrial ecosystem and hence the predator is placed
at the top of ecosystem pyramid. The Royal Bengal Tiger (Panthera tigris tigris) is an important
species of the carnivore feliformia family. Not only from ecological viewpoint is the tiger
important but also it is important under the religious perspective. For an instance in Hindu
culture it is regarded as a vehicle of goddess Durga.
Taxonomic classification illustrates that there are 9 subspecies of the tiger out of which 3
subspecies (Bali tiger, Caspian tiger and Javan tiger) has already became extinct (IUCN, 2013).
Royal Bengal Tiger (Panthera tigris tigris), Asia’s largest predator’s home range once used to be
stretched from the Caspian sea to the Island of Bali in Indonesia including the entire Indian
Subcontinent (Dinerstein et al., 2007). Unfortunately, at the present, these giant predators range
merely a 7% of their former distribution range (Sanderson et al., 2010). About 3000 individuals
of the tiger are thought to be present in the wild; this is decreased by more than 95% in
comparison to last century (Dinerstein et al., 2007). Out of the total remaining individual of tiger,
about 70% of them inhabit 42 resource sites which occupy only 0.5% of their historical range
(Walston et al., 2010). Royal Bengal tiger has been included under Appendix I by CITES and
category Endangered by IUCN Red Data Book. Also, it has been protected by National Park and
Wildlife conservation Act 1973 in Nepal.

There is huge number of factors acting behind this miserable decline in the numbers of giant
predators. Many studies have shown that the major reason behind this includes habitat
degradation, prey depletion and poaching (Chapron et al., 2008; Karanth and Stith 1999;

6
Ramakrishnan et al., 1999). A study by Dinerstein and Wikramanayake, (1993) has revealed the
pitiful future for the tiger survival in Asia where the human population is increasing in geometric
ratio and for the life support of people; they are clearing the forest area, the natural habitat for
these predators. Nepal has too experienced the bitter cases of habitat depredation of wildlife as
the southern belt of Nepal was once used to be covered with wild forests and people started
clearing and making settlements once malaria was abolished from this range during 1950s and
1960s. Further these charismatic species are often persecuted by humans due to conflict over
livestock and shared prey (Wolf and Ripple 2016). As mentioned earlier, in the context of Nepal,
including all these threats, there are some other threats like developmental infrastructure inside
protected area such as highway where incidents like road kill frequently occurs and it is seriously
harming for the survival of tiger population. Although the above mentioned factors have
significant effect on the degrading the tiger population, very less attention has been given to prey
depletion in comparison to others (Wolf and Ripple, 2016).

Nepal as an active member of TRCs has been actively working for the 2022 vision. In order to
achieve the goal, Nepal has been highly committed and has been adopting the various strategies
such as maintaining the inviolate habitats that provide secure breeding, effective protection from
poaching, maintaining the abundant prey, direct involvement of the local people and making the
functional connection with source population (Canada et al., 2005; Harihar et al., 2009; Moe et
al., 2016; Steinmetz et al., 2014). Though in the meantime, Nepal is facing lots of difficulties and
challenges in tiger conservations. The various threats faced by tiger for its survival in the context
of Nepal include habitat degradation, prey depletion and poaching as mentioned earlier. Ahead
of many threats, the most important one is prey depletion and it is in immediate need for acting
upon (Chapron et al., 2008). There has been significant research to infer that prey depletion has
significant impact on the survival of tiger and many researchers have suggested that for effective
tiger conservation we must plan the ecology based conservation strategies so that the prey
species of these large predators could be conserved which automatically contribute in tiger
conservation (Canada et al., 2005; Ripple et al., 2014; Wikramanayake et al., 2011). Carbone and
Gittleman, (2002) has found out for the survival of 90 kg predators, it needs almost 10000 kg of
prey species irrespective of the species. Similarly, Karanth et al., (2004) has stated that tiger

7
density is a function of prey densities. But we lack systematic research on the prey species of
tiger such as their abundance, their preferred habitats, etc. specifically in the context of Nepal.

8
 CHAPTER 3: MATERIALS AND METHODS
3.1 Study Area

This research was conducted in Bardia National Park and adjoining Khata Corridor in Western
Terai Arc Landscape (TAL), Nepal.

3.1.1 Bardia National Park

Bardia National Park (IUCN category II) is located in the western lowland Terai-Bhabar tract on
the southwestern part of Nepal (N: 28.2630 to 28.6711; E: 80.1360 to 81.7645). The area
encompassed by BNP is 968 square kilometers. The park was established in 1976 with an area of
368 square kilometers as Royal Karnali Wildlife Reserve and extended to its current size in 1984
after relocating the settlements to the southern side of BNP. The park was established originally
to protect the representative ecosystems as well as to conserve the tiger and its prey species
(DNPWC, 2017). The park is bordered by East-west Highway, buffer zone forests and cultivated
lands in the south, crest of Siwalik range in the north, Ratna Highway in the east and Karnali
River Bank in the west. Karnali and Babai rivers drain through the park and also the park
includes by alluvial flood plain created by Karnali River in the west, and pristine ecosystem of
Babai valley in the northern side. Bardia is a part of Terai Arc Landscape, a transboundary tiger
conservation landscape in India and Nepal, identified as Level 1 core tiger conservation unit
(Wikramanayake et.al., 1998). The density of tigers in Bardia is 4.74/100 KM2 and prey density
is 77.5 animals/KM2 (DNPWC, 2018).

Tigers in BNP

 2009: 18
 2913: 44
 2018: 82-97 (DNPWC, 2018)

3.1.2 Khata Corridor

Khata corridor is about 9 km long stretch that connects the Bardia National Park with the
Katarniyaghat Wildlife Sanctuary in India. Geographically, the area is located between the N

9
28°27.342’ – E 81°12.591’ and N 28°22.19’ – E 81°13.605’ in the southwestern region of Nepal.
Tharu are indigenous communities living around BNP and adjoining Khata Corridor.

3.1.3 Climate

The climate in the study area is sub-tropical monsoonal type with three distinct seasons viz.
winter (November -February), Hot (March-June) and Monsoon (July-October). About 90% of
the rainfall occurs between mid-Julys to mid-September.

(Photo: WWF India)

Figure 1: Terai Arc Landscape showing various PAs of Nepal and India

3.1.4 Vegetation

BNP is known for its unique biota. It contains diverse ecosystems ranging from tall alluvial
floodplain grassland of early successional stage to climax stage sal forest established on uplands.

10
A vegetation study conducted by Dinerstein, (1979) classified into six major types. Jnawali and
Wegge, (1993) later modified into seven major vegetation types. These are:

 Sal Forest
 Khair-Sissoo Forest
 Moist Riverine Forest
 Mixed Hardwood Forest
 Wooded Grassland
 Phantas
 Tall Alluvial Floodplain Grassland

Figure 2: Map showing study area and grids surveyed

11
3.1.5 Fauna

The park supports exceptionally diverse wildlife populations. The park harbors a total of 56
species of mammals including five species of deer alone, 438 species of avifauna, 52 species of
herpetofauna and 121 species of fishes (DNPWC, 2016). Out of total mammal species that are
found in the park, 20 species are enlisted in the Appendices I of the CITES. Several species of
wildlife found in the park are listed as protected species according to National Park and Wildlife
Protection Act 1973. The protected mammals include tiger, one horned rhinoceros, asian wild
elephant, swamp deer, gangetic dolphins, striped hyena, four horned antelope and chinese
pangolin. Similarly, giant hornbill, black stork, sarus crane, bengal florican and lesser florican
are among protected avifauna in the park. Gharial crocodile and python are reptiles in the similar
category. In addition, large populations of spotted deer, hog deer, barking deer and wild boar
together with avian, herpetofauna and several invertebrates enrich the biological diversity.

3.2 Data Collection

3.2.1 Sampling design

The objective of this study was three folds:

i) Map the distribution of major prey species of tiger

ii) Assess influence of different ecological and anthropogenic factors on relative
abundance of prey species of tiger, and
iii) Compare the relative abundance of major tiger prey species in relation to habitat type
(e.g., forest versus grassland) and management regime of habitat (protected area versus
buffer zone versus outside protected area).
In order to achieve these objectives, I needed to sample different habitat types in different
management regime. And also, since tiger prey species are medium ranging species as their daily
movement range can be roughly 5-6 square kilometer based on their home range estimates; I
needed to design sampling strategy to encompass their movement range and the spatial nature of
their distribution. Generally, wild ungulate home range varies between 2 km2 to 3.5 km2

12
(Shankar, 1994; Moe and Wegge 1994), and therefore my choice of 9 km2 cell is valid to make
inference on distribution of tiger prey species.
Importantly, I had to quantify two response variables, a) distribution and b) relative abundance.
One of best sampling strategy to quantify these variables and answer my objective is to divide
the whole study area into geographic grid cells of adequate spatial coverage, select representative
grid cells for all habitat types and management regimes and then conduct appropriate surveys to
quantify the response variables. Therefore, I first divided my whole study area into 3 strata and
selected the strata containing the core area, buffer zone and corridor area. I then, overlaid 3 km *
3 km geographic square grid cells into the land-cover matrix of the study area (both inside and
outside BNP using QGIS tools (Fig. 2). This yielded 288 number of grid cells. I then purposively
selected 32 grids to ensure representation of different habitat types and management regimes.
The sampling intensity became 18%. Out of 32 grids, 10 grids were inside the national park area,
12 in buffer zone area and 10 grid cells outside the buffer zone and national park area. Now these
grid cells represented my sampling units. Now the objective of sampling within these sampling
units is to first quantify response variables (i. distribution/ ii. relative abundance) and predictor
variables (elevation, forest cover, habitat type, management regime, distance to park boundary,
intensity of livestock grazing/human disturbance etc.) and use statistical modeling to see the
association between response variable and predictor variables.

3.2.2 Line transect surveys for distribution mapping

To quantify the distribution status of prey species within each sampling units, I needed to
quantify somehow whether species of interest in present/absent within each sampling unit. One
way to obtain this data is to do line transect surveys in each grid cell, do either temporal or
spatial replication in these transects and record detection (1) and non-detection (0) data for each
species of interest in these transects. To reduce any sort of systematic and personal bias in
transect placement, I first generated one random point within each grid cells (two random points
in adjacent grid cell being at least 500 m apart) and then started transect surveys for this random
point. For each grid, at least 4 transects of each 1 km were surveyed. Any direct (sighting) and
indirect (signs) evidence of specie presence were recorded by looking along each transects and
associated habitat variables (e.g. Habitat type, number of people, evidence of human disturbance

13
such as livestock grazing, logging and lopping etc.) were recorded. Each transects were
considered as spatial replicates and to ensure independence among replicates each transects were
at least about 250 m apart.

3.2.3 Relative abundance assessment

As I also wanted to quantify and compare the relative abundance of major prey species in
different habitat and management category, I quantified the relative abundance at transect scale.
To do this, I sub-divided each 1 km transect into five-200 m segments and evidence of species
presence (either direct/indirect) was recorded as either 0 or 1 for those segments. It is possible to
have more than 1 signs per segments but these signs may not be independent. So, I recorded 1 for
any number of species signs and 0 for non-detection of species occurrence for each 200 m
segment. This means I could have maximum of 5 signs for each one transects, 20 for 4 transects
within grid. This means the relative abundance within each grid can be maximum 20 signs. This
means the response variable for each sampling grid could range from 0 to 20.

3.2.4 Quantification of ecological and anthropogenic covariates

As I was also interested in assessing the influence of ecological and anthropogenic factors on
distribution of prey species of tiger, I quantified all the variables (Table 1) at grid cell level. For
example, human disturbance was quantified using field level data as encounter rate of evidence
of human presence. For this, I recorded presence or absence of livestock grazing, number of
people encountered for each 200 m segments and was later summarized at both transect level and
the grid cell level. I also recorded presence or absence of livestock grazing, number of people
encountered for each 200 m segment, habitat type such as Riverine, Low density mixed forest,
Grassland, Wooded land etc. and availability of water resources etc.

14
Table 1: Plausible ecological and anthropogenic covariates that are hypothesized to
influence prey species distribution.
Variable Measurement Expected influence on distribution
Habitat type Categorical (Sal forest, mixed sal Different habitat type would have
forest, grassland, riverine forest) different effect on ungulate species
distribution
Human disturbance Sum of the proportion of 200 m Negative effect on occupancy and
long segments of 1 km spatial detectability by affecting ungulate
replicates within a cell containing abundance due to direct competition with
signs of human disturbance livestock for forage and indirectly through
including, timber logging, vehicle other, correlated human disturbance.
etc.
Management regime National Park area (CA), Buffer I expected different prey species to have
zone area (BZ), and Outside buffer different responses to habitat management
zone area ( OBZ) regime as they have different type of
habitat requirements. But as national park
area is most protected I expected it would
have highest encounter rate of all species.
Livestock grazing Sum of the proportion of 200 m Negative effect on relative abundance of
long segments of 1 km spatial ungulate due to direct competition with
replicates within a cell containing livestock for forage and associated human
signs of livestock encounter such as impact etc.
hoof marks, dung, etc.

3.3 Data Analysis

3.3.1 Distribution mapping

First, I prepared the detection non-detection data/history (1010) for each grid cells and for seven
prey species. This detection history tells us that we had four transects (spatial replicates) in
particular grid cell, and species of interest (e.g., spotted deer, was detected in first and third
transect but no in second and fourth transect). This also means that this particular grid cell had

15
four occasions for the detection history. I mapped distribution based on naïve occupancy. The
grid cells where I encountered species sign was given code and the species where I didn’t
encounter species was given code 0 and based on this detection non-detection information I
updated the attribute table of the GIS shape file of the grid cells and prepared distribution map
with different color codes.

3.3.2 Relative abundance along the gradient of human disturbance, and with respect to
habitat type and management regime.

I wanted to assess how the relative abundance index of species changes along the gradient of
human disturbance. And I also wanted to assess whether relative abundance vary with respect to
habitat type and management regime. I tested the influence of four key variables: human
encounter rate, livestock grazing encounter rate, Habitat type (grassland, riverine forest, mixed
forest and agricultural land) and management regime (national park, buffer zone and outside
buffer zone). I had 126 data points, that means for 126 transects (1km each). For each transect, I
had total number of indirect signs detected (for every 200 m segment) that is relative abundance
as a response variable. I therefore ran a couple of Poisson Regression or Generalized Linear
Models (GLMs) with Poisson error distribution and log link function since the response variable
was count (number of signs encountered in each transect) to test the relative influence of these
variables on response variable.

I defined six a priori set of models, including the null model. I ranked the models using the
Akaike Information Criterion adjusted for small samples (AICc) (Burnham, K. P., & Anderson,
2002). I examined the beta-coefficient (slope) of covariates to test the significance of their
effect on response variable (number of signs per transect). Confidence intervals that included
zero indicated no significant effect of the predictor variables on response variables. All analyses
were carried out in R 3.1.2 (R Core Team, 2016).

16
3.3.4 Report Writing

The final report was summed up in Microsoft word document 2010 using appropriate charts and
diagrams. The citation for the various research articles were included following APA format in
Mendeley v 1.19.3.

17
 CHAPTER 4: RESULTS

4.1 Distribution mapping

I surveyed a total of 32 grid cells/sampling units/ sites covering an area of c. 288 km2. The
overall survey effort was 126 km (126 times, 1-km long transects). Based on the field data of
detection and non-detection of prey species, distribution mapping was done and also naïve
occupancy for each species was calculated and it is shown in the figures 3,4,5,6, 7 & 8. Spotted
deer was found in 28 grid cells (fig 4), so the naïve occupancy was 0.81. Similarly, the signs of
sambar deer were found in 9 grids with naïve occupancy 0.28. Similarly wild boar, hog deer,
barking deer, blue bull and swamp deer were detected in 22 (fig 7), 3 (fig 6), 7 (fig 3), 11 (fig 5)
and 2 grids with naïve occupancy 0.69, 0.09, 0.23, 0.34 and 0.06 respectively. The results
revealed that spotted deer was found to be most abundant species whereas swamp deer was least
abundant across my study area.

Figure 3: Distribution map of barking deer Figure 4: Distribution map of spotted deer

18
Figure 5: distribution map of blue bull Figure 6: distribution map of hog deer

Figure 7: distribution map of wild boar

0.80 0.75
0.69
0.70
0.60
0.50
0.40 0.34
0.28 Naïve Occupancy
0.30 0.22
0.20
0.09 0.06
0.10
0.00
Spotted Wild boar Hog Deer Barking Blue Bull Sambar Swamp
Deer Deer Deer

Figure 8: Naïve occupancy of prey species

19
4.2 Relative Abundance Index

The Relative Abundance Index (RAI) or encounter rate per km was highest for Chital (1.73
signs/km) and was lowest for swamp deer (0.05 signs/km), Barking deer (0.30 signs/km), hog
deer (0.24 signs/km), sambar (0.22 signs/km) and blue bull (0.21 signs/km) (Fig. 9 & Table: 2).
Note that sign of each species seen in each 200 m segments was considered as 1 sign and in 1 km
transect (5 segments) there can be maximum 5 signs as per my criteria to maintain independence
in sign.

Table 2: Species sign encounter rate and associated standard deviation

Species RAI (mean encounter rate/km) Standard deviation

Spotted deer 1.74 1.96
Wild boar 1.03 1.67
Barking deer 0.30 0.86
Hog deer 0.24 0.96
Sambar 0.22 0.65
Blue bull 0.21 0.58
Swamp deer 0.05 0.31

20
 Prey species
Figure 9: Overall species encounter rate /RAI

4.2.2 Management Regime-wise RAI

Sambar deer, swamp deer and hog deer were almost exclusively found inside national park.
Spotted deer was abundant across all habitat type but higher encounter rate was found inside
national park. Importantly, although blue bull, wild boar and barking deer also found inside
national park and buffer zone, they were found with higher encounter rate outside the buffer zone
in community forests lands. Apart from their ecological behaviours, another possible reason for
this is in order to get rid of predators pressures, these ungulates seems to stay away from tiger
dominated landscapes i.e. core area (Kumar et al., 2008).
The graph showed below (fig 10) represents the management regime wise encounter rate of prey
species in the study area. Here Y axis represent the percentage total encounter of each species,
For example, out of all encounters of swamp deer, all of the sign encounters were inside national
park.

21
 100% 0.00 0.00 0.00
0.73
90%

80%
1.47
70% 0.53 0.43

60%

50% 3.98 0.75 0.70 0.18

40%
0.78
0.13
30%
0.20
20%
0.30 0.98
10%
0.68 0.11
0% 0.00 0.00 0.00
Barking Spotted Wild boar Blue bull Hog deer Sambar Swamp deer
deer deer
Species
Buffer zone National park Outside BZ

Figure 10: Overall species encounter rate according to the management category

The graph shows that hog deer, sambar and swamp deer were exclusively detected inside the
core area of BNP. Being specific, the various signs of hog deer and swamp deer were recorded
around Karnali Flood Plain area with abundant grasses whereas sambar was recorded mostly
around mixed sal forest in core area (also refer to fig 11). The Relative Encounter Rate of hog
deer, sambar and swamp deer in core area of BNP is 0.75, 0.70 and 0.18 respectively (fig 10).
Similarly, the RAI for barking deer, wild boar and blue bull is relatively higher in outside buffer
zone which accounts for 0.53, 1.47 and 0.43 respectively. The RAI of the barking deer, wild boar
and blue bull in buffer zone is 0.30, 0.98 and 0.11 whereas in core area is 0.13, 0.78 and 0.20
respectively. Spotted deer is distributed across the entire management regime though most of the
detection was in core area. The RAI for spotted deer in CA, BZ and OBZ is 3.98, 0.68 and 0.73

22
respectively. This result shows that, spotted deer doesn’t have that much selective habitat pattern
as comparing to other species.

4.2.3 Habitat-wise RAI

Sign encounter rate of spotted deer, wild boar, blue bull, sambar and barking deer was
consistently higher in mixed sal forest whereas swamp and hog deer had higher encounter rate in
grassland. Encounter rate was consistently lower for all species in agricultural lands (Fig. 11).
The graph shown below represents the habitat wise encounter rate of prey species in the study
area. Y axis represents the species and X axis indicates the encounter rate per km.

Spotted deer had encounter of 2.3signs /km in mixed sal forests, 2.1 signs /km in grassland,
1.4signs/km in riverine forests, and 0.1 signs /km in agricultural lands. Similarly, barking deer
has RAI 0.60 in mixed sal forest, 0.09 in grasslands and 0.05 in riverine forests. It was not
detected in agriculture fallow lands.

2.36

2.13

1.90
Sign Enounter rate per km

1.45
1.30

0.60
0.48
0.36 0.34 0.36
0.26 0.30 0.26
0.09 0.05 0.04 0.04 0.05
0.00 0.00 0.00 0.00
0.00 0.00 0.00 0.00 0.02
0.00

Barking deer Spotted deer Wild boar Blue bull Hog deer Sambar Swamp deer
Agriculture lands Grasslands Mixed sal forests Riverine forests

Figure 11: Overall species encounter rate according to the habitat type

23
Similarly, wild boar has highest encounter rate in mixed forest with RAI 1.90 followed by
grassland 0.48, riverine forest 0.36 and agriculture lands 0.04. Blue bull and sambar has highest
detection in mixed forest types. In case of hog deer (1.30) and swamp deer (0.26), almost entire
of their detection were in grasslands.

4.3 Species specific relative encounter rate according to habitat and

management category

4.3.1 Spotted Deer

Spotted deer had higher encounter rate inside

national park in comparison to other
management regime (Fig. 12) however its
encounter rate is more or less similar in
mixed sal forest and grassland but lower in
riverine forest and lowest in agricultural and
fallow lands (fig. 13). The RAI of spotted
deer in CA is 3.98 whereas BZ is 0.68. Its
RAI in OBZ is 0.73. Similarly, it has highest
encounter in mixed sal forest (2.36) followed
by grassland (2.13), riverine forest (1.45) and
least in agriculture and fallow lands (0.04). Figure 12: Relative encounter rate of spotted
deer according to management regime

24
 Figure 13: Relative encounter rate of spotted deer according to habitat type

Table 3: RAI of Spotted deer

Categories RAI (sign encounter rate per km)

BZ 0.68
Management Regime CA 3.98
OBZ 0.73
AGL 0.04
GL 2.13
Habitat Types
MF 2.36
RF 1.45

4.3.2 Wild Boar

Wild boar had higher encounter rate outside buffer zone
(1.47) followed by buffer zone (0.98) and national park
(0.78) (fig. 14, table.4), whereas in terms of habitat type,
RAI was higher in mixed forest (1.90) followed by
grasslands (0.48), riverine forests (0.36) and fallow lands
(0.04) (fig 15, table 4)

Figure 13: Relative encounter rate of

wild boar according to management
regime

25
 Figure 15: Relative encounter rate of wild boar according to habitat type

Table 4: RAI of wild boar

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.98
CA 0.78
OBZ 1.47
Habitat Types AGL 0.04
GL 0.48
MF 1.90
RF 0.36

4.2.3 Barking Deer

Figure 14: Relative encounter rate of barking deer

according to management regime
26
Barking deer had higher encounter rate outside the buffer zone area (0.53) in comparison to other
management regime; buffer zone (0.30) and core area (0.13), (Fig. 16 and table 5). However its
encounter rate is lowest in national park. Similarly the result shows its highest encounter rate in
mixed forest (0.60) followed by grassland (0.09) and riverine forest (0.05), zero in agriculture
fallow lands (fig. 17, table 5).

Figure 17: Relative encounter rate of barking deer according to habitat type

Table 5: RAI of barking deer

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.30
CA 0.13
OBZ 0.53
Habitat Types AGL 0.00
GL 0.09
MF 0.60
RF 0.05

27
4.2.4 Hog Deer

Hog deer was exclusively encountered

inside national park and no encounter
in other management regimes. Its RAI
in core area is 0.75. Moreover, it was
only encountered in grassland patches
in Karnali flood plain area of national
park (fig 18 & 19).

Figure 15: Relative encounter rate of hog deer according to

habitat type

In grassland habitat, its RAI is 1.30. The major reason behind the non-detection of hog deer
outside the core area may be due to absence of the flood plain area that supports various habitat
requirements of this species (Mondol et al., 2009).

Figure 16: Relative encounter rate of hog deer according to management regime

28
Table 6: RAI of hog deer

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.00
CA 0.75
OBZ 0.00
Habitat Types AGL 0.00
GL 1.30
MF 0.00
RF 0.00

4.2.5 Sambar

My study has found that sambar is typically

found in mixed forests and grasslands inside
core area of national park and its sign
encounter in buffer zone and outside buffer
zone is almost zero ( Fig. 20 & 21 and table
7). The RAI for sambar deer in CA is highest
(0.70) whereas it was not detected in buffer
zone and outside buffer zone. While talking
about habitat types, mixed forest (0.36) in
national park is more preferred by sambar than Figure 17: Relative encounter rate of
grasslands (0.30) of national park. It was not detected in sambar according to habitat type

fallow lands and riverine forests.

29
 Figure 18: Relative encounter rate of sambar
according to management regime

Table 7: RAI of sambar

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.00
CA 0.70
OBZ 0.00
Habitat Types AGL 0.00
GL 0.30
MF 0.36
RF 0.00

4.2.6 Blue Bull

Blue bull had higher encounter rate outside buffer zone area in comparison to other management
regime. The RAI of blue bull in OBZ is 0.43 which is followed by core area 0.20 and buffer zone
0.11. However its encounter rate is more or less similar in mixed sal forest (0.34) and grassland
(0.26) but lower in riverine forest (0.05) and zero in agricultural and fallow lands (fig. 22 and
table 8).

30
 Figure 19: Relative encounter rate of blue bull
according to habitat type

Figure 20: Relative encounter rate of blue bull according to management regime

Table 8: RAI of blue bull

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.11
CA 0.20
OBZ 0.43
Habitat Types AGL 0.00
GL 0.26
MF 0.34
RF 0.05

31
4.2.7 Swamp deer

The encounter rate of swamp deer was inside national park (0.18) in comparison to other
management regime (The buffer zone and outside buffer zone had zero encounter). Similarly, it
is mostly found in grassland (0.26) but very little in mixed forest as well (Fig 24 & 25).

Figure 21: Relative encounter rate of swamp deer according to habitat type

Figure 22: Relative encounter rate of swamp deer according to management regime

Table 9: RAI of swamp deer

Categories RAI (sign encounter rate per km)

Management Regime BZ 0.00
CA 0.18
OBZ 0.00
Habitat Types AGL 0.00

32
 GL 0.26
MF 0.00
RF 0.00

4.4 Species specific Generalized Linear Models (GLMs)

4.4.1 Spotted deer

The model selection results showed that human encounter rate and livestock encounter rate as
proxy of human disturbance consistently had negative influence on relative abundance of spotted
deer (Fig. 26 and Fig. 27). The beta-coefficient for human encounter rate was (βhumanER (SE0.03)
= -0.31 (p value= <0.05; 95 % CI=-0.37- (-0.24)) and for livestock grazing encounter rate it was
(βlivestockER (SE 0.05) = -0.41 (p value <0.005; 95 % CI= -0.54- (-0.31). Since the 95 %
confidence interval of the beta-coefficients don’t overlap, the effects of these variables are
significant. Habitat type also had influence on relative abundance of spotted deer. Highest
relative abundance was mixed sal forest (MF) and grassland (GL) and lowest in agricultural land
(Fig 28). In terms of management category, agricultural land had negative influence (β Agriculture
(SE) = -3.13 (0.99) whereas other habitat categories had positive influence but magnitude differ
(βmixed sal forest (SE) = 3.99 (1.00); βgrassland(SE) = 3.89 (1.00); βgrassland (SE) = 3.50 (1.01)). The
figure 26 shows that the relative abundance of spotted deer decreases with increase in human
disturbance along the human disturbance gradient. The figure 27 shows that the relative
abundance of spotted deer decreases with increase in livestock grazing intensity along the human
disturbance gradient. The maximum likelihood based model selection showed management
regime as best supporting model the data with 100 % Akaike Weight (table 10)

Table 10: Poisson regression models describing the relative abundance of spotted deer in
BNP (Fig. 2), ranked according to the Akaike Information Criterion adjusted for
small sample size (AICc).

Model AICc Delta weight Df logLik

Mgmt. regime 377.58 0.00 1.00 3 -185.69

HumanER + LivestcokER 408.60 31.02 0.00 3 -201.20

33
HumanER 435.25 57.67 0.00 2 -215.58

Habitat type 448.10 70.52 0.00 4 -219.89

LivestockER 453.41 75.83 0.00 2 -224.65

Intercept only 527.02 149.44 0.00 1 -262.50

Figure 23: Relationship between human encounter rate and relative abundance index of
spotted deer

34
Figure 24: Relationship between livestock grazing intensity (encounter rate of livestock
signs) rate and relative abundance index of spotted deer

Similarly, the effect of management regime is significant with higher RAI in CA, and higher
RAI in MF. The figure below shows that the median relative abundance of spotted deer is
highest inside the national park and lowest in outside the buffer zone area. The figure above
shows that the median relative abundance of spotted is highest in mixed forest and grassland.

Figure 25: Relationship between management regime (buffer zone=BZ, national park=CA,
and outside buffer zone (OBZ) and relative abundance index of spotted deer

35
 Figure 26: Relationship between habitat type (MF, GL, MF & Fallow AG lands) and relative
abundance index of spotted deer

4.4.2 Wild Boar

The model selection results showed that human encounter rate and livestock encounter rate as
proxy of human disturbance consistently had significant influence on relative abundance of wild
boar (Fig. 30 and Fig.31, table.11). The beta-coefficient for human encounter rate was (βhumanER
(SE) = 0.11 (0.04, positive p value 0.02 significant; 95 % CI: 0.02-0.22). This suggests that the
human encounter and encounter rate of wild boar has positive relation i.e. the wild boar
encounter rate increases along with increase in human disturbances. And for livestock grazing
encounter rate it was (βlivestockER (SE 0.08) = -0.47 (The 95 % CI is -0.6520962 -0.3214686, p
value less than 0.05). This does not overlap zero suggesting significant negative effect. Since the
95 % confidence interval of the beta-coefficients don’t overlap, the effects of these variables are
significant. The encounter rate of wild boar increased with the outside buffer zone factor of the
categorical variable-management regime. The slope parameter of this factor was (βoustide
BZ(SE0.20), p value 0.04). Other factor inside NP were negative but not significant (β NP (), -
0.24(SE 0.22, p value 0.29) and (β BZ(SE0.13), -0.02 (p value 0.89).

36
 Figure 27: Relationship between human encounter rate and relative abundance index of
wldboar

Figure 28: Relationship between livestock grazing intensity (encounter rate of livestock
signs) rate and relative abundance index of wildboar

Habitat type emerged as the best model in model section results based on AIC criteria (table 4).
It had influence on relative abundance of wild boar. Interestingly, although agricultural land had

37
negative influence (βAgriculture (SE) = -3.31 (1.00), other habitat types had significant positive
influence on RAI ((βmixed sal forest (SE) = 3.77 (1.00); βgrassland(SE) = 2.39 (1.00); βriverine (SE) =
1.12 (1.06))) suggesting its most generalist nature of response.

Table 11: Poisson regression models describing the relative abundance of wild boar in BNP
(Fig. 2), ranked according to the Akaike information criterion adjusted for small
sample size (AICc)

Model AICc delta weight Df logLik

habitat type 346.50 0.00 1.00 4 -169.08

HumanER + LivestcokER 362.46 15.96 0.00 3 -178.13

LivestockER 383.27 36.77 0.00 2 -189.58

Mgmt.regime 430.57 84.07 0.00 3 -212.19

HumanER 430.76 84.26 0.00 2 -213.33

Intercept only 434.20 87.70 0.00 1 -216.08

4.4.3 Blue Bull

The model selection results showed that human encounter rate and livestock encounter rate as
proxy of human disturbance did not have significant influence on relative abundance of blue bull
(fig. 32 and fig.33, table.12). Here in 95% Confidence Interval of the beta coefficient of the
variable humanER is: lower CI= βhumanER– 1.96 * SE (0.09) = -0.2374076 and upper 95 % CI =
βhumanER+ 1.96 * SE (0.09) = 0.1333121. This means 95 % CI of the beta coefficient ranges from
-0.2374076 ( negative value) to 0.1333121 (positive value), which means it overlaps zero,
suggesting we are not sure if its actual effect is negative or positive- so the effect is statistically
undiscernible. Here effects of both human disturbance factors on blue bull are not significant
(better word- weak and undiscernible effect size (beta estimate/coefficient of predictor variable).
The model selection results show that habitat type model had best support for data (table 12). But
habitat factors individually were not significant at 5% level of significance.

38
 Figure 29: Relationship between human encounter rate and relative abundance index of
blue bull

Figure 30: Relationship between livestock grazing intensity (encounter rate of livestock signs)
rate and relative abundance index of bluebull

39
Table 12: Poisson regression models describing the relative abundance of blue bull in BNP
(Fig. 2), ranked according to the Akaike information criterion adjusted for small
sample size (AICc)

Model AICc Delta Weight Df logLik

habitat type 140.75 0.00 0.96 4 -66.21

Mgmt.regime 148.02 7.27 0.03 3 -70.91

LivestockER 150.45 9.70 0.01 2 -73.18

HumanER + LivestcokER 152.54 11.79 0.00 3 -73.17

Intercept only 152.74 11.99 0.00 1 -75.35

HumanER 154.41 13.67 0.00 2 -75.16

Unfortunately for rest of the species i.e. sambar, swamp deer, barking deer and hog deer, the
relative influence of various anthropogenic factors couldn’t be assessed as they had very less
encounters.

40
 CHAPTER 5: DISCUSSIONS

Understanding ecological and anthropogenic determinants of a species is a key to prioritizing

conservation efforts. In this study, I examined the relative influence of (management category,
habitat type as well as human disturbance factor) on relative abundance of various tiger prey
species in Bardia national park and adjoining Khata corridor. The findings of this study show
that prey species have differential response to human disturbance, habitat type and management
regime as indicated by their relative sign encounter rate/RAI. These results suggest that some
species are highly vulnerable to human disturbance (e.g., deer which had negative association
with human disturbances and other are to some extent resilient to human use (e.g., wild boar and
blue bull; as they had higher encounter rate outside the buffer zone).

5.1 Spotted Deer

Spotted deer was found to have highest detection rate among the studied species (table 2).
Management regime appeared as most important predictor variable in influencing relative
abundance of or sign encounter rate of spotted deer as indicated by its 100% Akaike weight
(table. 10). Sign encounter rate was significantly lower in buffer zone and outside buffer zone in
comparison to the national park area (fig. 12) suggesting that national park is critical for its
conservation. Although spotted deer are described as a habitat generalist (Schaller, 1967) and
prefer to live in open grassland intermixed with forest for shelter, my study showed that
encounter rate of spotted deer was higher in mixed forest type in comparison to other habitat type
(Fig. 13). This means that mixed habitat type with appropriate interspersion of grassland and sal
forest patches is important for spotted deer. A similar study from Suklaphanta National Park has
shown that spotted deer had highest occupancy in grid cells those had highest proportion of
forest area (Lamichane, 2018).

41
5.2 Wild Boar

Wild boar was second species after spotted deer in terms of sign encounter rate in my study
area. The habitat types appeared to be the most important predictor variable in influencing
relative abundance or sign encounter rate of wild boar as indicated by its 100% Akaike weight
(table. 11). The sign encounter rate of wild boar in mixed forest was far more in comparison to
other habitat category (fig 14). The major reason for this includes its preferences with the
human disturbance factors (Bhattarai and Kindlmann, 2013). Similarly, it was found that it
mostly lives outside the core area i.e. around buffer zone and outside buffer zone. The possible
reason for this could be in order to get rid of predators pressures, the wild boars seems to stay
away from tiger dominated landscapes (Kumar et al., 2008).

5.3 Barking Deer

Barking deer was third mostly encountered species of deer family after spotted deer and sambar
in terms of sign encounter in my study area (fig 8, table 2). Barking deer had higher encounter
rate outside the buffer zone area in comparison to other management regime. It was mostly
observed in the community forests with abundant mixed sal forests outside the buffer zone.
However its encounter rate was lowest in national park. The findings are similar to that of
Bhattarai and Kindlmann, (2013) who has found that barking deer were mostly detected in mixed
forest followed by grasslands and riverine forests. The encounter rate for barking deer was
almost zero in agriculture and fallow lands. Similarly, Pokhrel et al., (2015) have also found the
strong response of barking deer to riverine and sal forest with the availability of water;
unfortunately, I have not included water availability as predictor variable in my study.

5.4 Sambar

Out of the total grids surveyed, the sambar sign were recorded in 9 grids. The entire grids were
of core area. It was fourth most detected species in my study area in terms of its sign encounter
rate. It was found that sambar is typically found in mixed forests and grasslands inside core area
of national park and its sign encounter in buffer zone and outside buffer zone is almost zero.
Moreover, their sign encounter rates were slightly higher in mixed forest than grassland of BNP

42
(fig 20). The findings of this study that mixed forest of national park is more preferred by sambar
than grasslands of national park is similar with the findings of (Bhattarai and Kindlmann, 2013).

5.5 Hog Deer

Hog deer is in second position in least detected species in terms of its sign encounter. It was
exclusively encountered in grassland patches of Karnali flood plain in my study area. It is similar
to the previous findings that have revealed an almost exclusive utilization of grassland by hog
deer both as feeding site and shelter from predators (Dhungel and O’gara, 1991; Dinerstein,
1987; Mishra, 1982).

5.6 Swamp Deer

The least encountered species in my study area was swamp deer. It was exclusively encountered
in two grids of core area of BNP. These two grids were in Karnali Flood Plain area consisting of
abundant grasslands. Moreover, I have noticed that the burned patches were mostly preferred by
swamp deer which is similar to the findings of Lamichane, (2018) in Sukhlaphanta National
Park. Swamp deer mostly preferred grassland and I got the result similar to Bhatta, (2004); Khan
et al., (2004); Martin, (1975); Pokharel, (2008); Qureshi et al., (1995); Schaff, (1978); Schaller
(1967); Singh, (1984) and who had reported that basically swamp deer is a grazer who largely
fed on grasses. The finding that swamp deer tries to avoid the mixed forest area is similar to the
findings of Gyawali, (2005) who reported the negligence of forest by swamp deer due to the
absence of preferred food plants and water.

5.7 Blue Bull

Blue bull was found to have third highest naïve occupancy after spotted deer and wild boar (fig
8) in my study area. Both habitat type and management regime appeared to be an important
predictor variable in influencing relative abundance or sign encounter rate of blue bull as
indicated by its 100% Akaike weight (table. 12). It was observed that blue bull preferred to stay
around buffer zone and outside the buffer zone near to human dominant areas instead of core

43
area. The area with mixed forest type had higher encounter of blue bull followed by grasslands.
However, the model selection results showed that human encounter rate and livestock encounter
rate as proxy of human disturbance did not have significant influence on relative abundance of
blue bull (fig. 32 and fig.33, table.12). The undiscernible effect size of beta estimate/coefficient
of predictor variable made difficult to predict the exact response of human disturbance and
livestock grazing intensity in the relative abundance of blue bull. This might mean this species is
a bit resilient to human disturbance or may be its habitat inside PAs is not great because there is
more competition for forage for conspecifics or predator pressure from tiger etc. However, these
finding contradicts with the findings of Aryal et al., (2016) and Bisht, (2015) who had found out
that mixed forest area with human disturbance had the strong positive influence in the occupancy
of blue bull in Rupendehi district and Sukhlaphanta National Park respectively.

Prey populations are chief components in shaping the tiger’s survival in an area. In case of my
study area, I came to know that the prey species are widely distributed in various habitat types
such as mixed sal forest, riverine forest, grasslands, etc. across various management regimes
i.e. core area, buffer zone and outside buffer zone. Aryal et al., (2016) has stated that the
current PAs are insufficient to bear the doubling of tiger population goal by 2022 and hence
their habitat must be extended outside the current PAs. In support with their argument, I found
that the major habitat components of tiger, i.e. availability are good in buffer zone and outside
buffer zone around BNP. So, this could be a topic of discussion that in case we require
expanding the current tiger habitat to BZ and OBZ, the prey requirement is positive and other
requirements needs to be assessed and comprehensive study must be carried out.

44
 CHAPTER 6: CONCLUSIONS AND RECOMMENDATIONS

6.1 Conclusions

Overall, the results of this study show that tiger prey species have differential response to human
disturbance, habitat type and management regime as indicated by their relative sign encounter
rate/RAI. These results suggest there are some species which are highly vulnerable to human
disturbance and other are to some extent resilient to human disturbances. For an instance, it was
found that deer had negative association with human disturbances whereas wild boar and blue
bull had higher encounter rate outside the buffer zone in human dominated landscapes. These
results also show the generalists and specialist behaviour of prey species of tiger. Differential
response to habitat type and management regime perhaps suggest that integrated management of
all habitat types and management regimes would be necessary to sustain the collective prey base
recovery of all species for tiger conservation. Most importantly, highest RAI in mixed forest and
grassland habitat indicates that appropriate mixture and distribution of grassland and forest
patches would be preferable from overall point of view of prey base conservation. However
while doing so species specific requirement and ecology of species must be taken into
consideration. Moreover the finding that some of the prey species preferring to distribute in
buffer zone and outside buffer zone area can be taken an opportunity to indulge include the local
community in conservations activities formulating appropriate plans. These results can be useful
to protected area managers in formulating appropriate habitat management and conservation
intervention for prey species conservation.

6.2 Recommendations

The recommendations can be summarized as:

 The policy makers should make the policy which must be based on integrated
management of all habitat types, because the tiger prey species are widely distributed in
various management regimes and habitat types and in order to sustain the collective
prey base recovery of all species for tiger conservation, we need integrated approach.

45
 Further, appropriate mixture and distribution of grassland and forest patches would be
preferable for overall conservation of BNP and Khata corridor. Because, the grids cells
containing various habitat types has higher RAI of prey species.
 And the prey species are distributed in buffer zone and outside buffer zone area, the
management officials must take this as an opportunity to include local community in
conservations activities.
 Further, the concerned authority must have due concentration of the human disturbances
factors while formulating the specific recovery plan of prey species as they have
significant influence in the relative abundance of prey species.
 Prey base monitoring as well as further study should be done in periodic basis for the
effective conservation and management intervention as the various ecological and
anthropogenic factors which has great influence on the prey species frequently
observed in the study area and they could be dynamic.

46
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 ANNEX 1: FIELD DATA COLLECTION FORM

Aff3sf] cfxf/f k|hftL अनुगमन फारम २०७५

सर्वेक्षकको नाम ब्लकको नाम शुरु गरेको स्थान

अन्तिम स्थान
शुरुको न्िन्िएस लोके शन अन्तिमको न्िन्िएस लोके शन

शुरु समय अत्य समय

ट्यासेगमे न्ट जजजिएस उचा जचन्ह जचन्ह जचन्ह जचन्हको आहारा मानजि बासस्थान कैजफएि
न्जे नं. लोकेशन ई छ/छै को भएको उमे र प्रजाजि य प्रकार
क्टMeter न प्रकार जजमन (छ छै न) गजिजि
जाि र जि
नं संख्या
.
0
0-200
201-400
401-600
601-800
801-
1000

न्ितहको प्रकारः िग माकक स, स्राि, स््याट, सेतट माकसक, ्ल माकसक

न्ितह भएको िन्मनः खोला ककनार, मातछे मात्र न्हड्ने बाटो, clUg /]vf,
न्ितहको उमेरः धेरै नया, नया ,िुरानो, धेरै िुरानो
बासस्थानको प्रकार SF - Sal forest, MF - Mixed Forest, RF - Riverine Forest, TG - Tall Grassland, SG - Short
Grassland, W-Wetland, S – Streamed.
Form Source: Tiger and Prey Species Monitoring Protocol by DNPWC

52
 ANNEX 2: PHOTO PLATES

Sambar pellet Spotted deer pellet

Swamp deer pellet Hog deer pellet

Blue bull pellet Tiger pug mark

53
54