Resource Partitioning Among Sympatric Carnivore Species Inhabiting Pir Lasura National Park (PLNP), AJ&K During 2014-2017
RESOURCE PARTITIONING AMONG SYMPATRIC
CARNIVORE SPECIES AT PIR LASURA NATIONAL PARK,
AZAD JAMMU & KASHMIR, PAKISTAN
FARAZ AKRIM 11-arid-3899
Department of Wildlife Management Faculty of Forestry, Range Management and Wildlife Pir Mehr Ali Shah Arid Agriculture University Rawalpindi, Pakistan 2018
RESOURCE PARTITIONING AMONG SYMPATRIC
CARNIVORE SPECIES AT PIR LASURA NATIONAL PARK,
AZAD JAMMU & KASHMIR, PAKISTAN
by
FARAZ AKRIM
(11-arid-3899)
A thesis submitted in partial fulfillment of
the requirements for the degree of
Doctor of Philosophy
in
Wildlife Management
Department of Wildlife Management Faculty of Forestry, Range Management and Wildlife Pir Mehr Ali Shah Arid Agriculture University Rawalpindi Pakistan 2018
IN THE NAME OF ALLAH
The Most Merciful
The Most Beneficent
The Most Knowing
“It is ALLAH who has made for
you the earth as a resting place and the sky as a canopy and has given you shape and made your shapes beautiful and has provided for you sustenance of things pure and good. Such is ALLAH
your Lord. So glory to ALLAH.
The Lord of the world”
GOLDEN SAYINGS OF THE HOLY PROPHET
HAZRAT MUHAMMAD
(Peace be upon him)
“Keep your thought well composed, and search
the fact of wisdom for it, otherwise mind
gets weary and the people get weary,
so ponder into the knowledge and
science thought fully and
search for new facts
and ideas”
I DEDICATE THIS EFFORT
TO
OUR BELOVED HOLY PROPHET HAZARAT MUHAMMAD (PEACE BE UPON HIM)
CONTENTS
Page
List of Tables xii
List of Figures xv
ACKNOWLEDGMENTS xxi
ABSTRACT xxiv
1 GENERAL INTRODUCTION 1
1.1 INTRODUCTION 1
1.1.1 Common Leopard 4
1.1.2 Asiatic/ Golden Jackal 5
1.1.3 Red Fox 7
1.1.4 Common/Asian Palm Civet 9
1.1.5 Small Indian Civet 10
1.1.6 Indian Grey Mongoose 12
1.1.7 Small Indian Mongoose 13
1.2 STUDY AREA 16
1.2.1 Study Objectives 17
1.3 RATIONALE OF THE STUDY 18
1.4 ORGANIZATION OF THE THESIS 18
2 DIVERSITY AND DISTRIBUTION OF CARNIVORES AT PIR 20
LASURA NATIONAL PARK, AZAD JAMMU AND KASHMIR,
PAKISTAN
2.1 INTRODUCTION 20
2.2 REVIEW OF LITERATURE 21
2.3 MATERIALS AND METHODS 24
2.3.1 Diversity of Carnivores 24
2.3.2 Distribution of Carnivore Species 25
2.3.3 Statistical Analysis 26
2.4 RESULTS 26
2.4.1 Diversity of Carnivore Species 26
2.4.2 Distribution of Carnivore Species 27
2.4.2.1 Distribution of common leopard 27
2.4.2.2 Distribution of Asiatic jackal 27
2.4.2.3 Distribution of Kashmir hill fox 31
2.4.2.4 Distribution of Asian palm civet 31
2.4.2.5 Distribution of small Indian civet 39
2.4.2.6 Distribution of Indian grey mongoose 39
2.4.2.7 Distribution of small Indian mongoose 44
2.5 DISCUSSION 44
2.6 CHAPTER SUMMARY 55
3 DIET COMPOSITION OF SYMPATRIC CARNIVORE SPECIES 56
AT PIR LASURA NATIONAL PARK
3.1 INTRODUCTION 56
3.2 REVIEW OF LITERATURE 57
3.3 MATERIALS AND METHODS 59
3.3.1 Molecular Identification of Carnivore Scats 59
3.3.2 Scat Analysis 63
3.3.2.1 Whole mount preparation 63
3.3.2.2 Scale replication 65
3.3.2.3 Identification of plant matter recovered 65
3.3.3 Statistical Analysis 65
3.4 RESULTS 67
3.4.1 Molecular Identification of Carnivore Scats 67
3.4.1.1 Accuracy and precision for molecular identification of 68 carnivore scats
3.4.2 Morphological Characteristics of Carnivore Scats 70
3.4.3 Diet Composition of Carnivore Species 70
3.4.3.1 Diet composition of common leopard 70
3.4.3.2 Diet of Asiatic jackal 79
3.4.3.3 Diet composition of Kashmir hill fox 84
3.4.3.4 Diet composition of Asian palm civet 88
3.4.3.5 Diet composition of small Indian civet 93
3.4.3.6 Diet composition of Indian grey mongoose 101
3.4.3.7 Diet composition of small mongoose 102
3.4.4 Diversity Index, Richness and Evenness of Prey 110
3.5 DISCUSSION 111
3.6 CHAPTER SUMMARY 127
4 ASSESMENT OF CARNIVORE PREY SPECIES ABUNDANCE 129
AT PIR LASURA NATIONAL PARK
4.1 INTRODUCTION 129
4.2 REVIEW OF LITERATURE 130
4.3 MATERIALS AND METHODS 131
4.4 RESULTS 132
4.5 DISCUSSION 135
4.6 CHAPTER SUMMARY 138
5 NICHE BREADTH AND NICHE OVERLAP AMONG SYMPATRIC 140
CARNIVORE SPECIES AT PIR LASURA NATIONAL PARK
5.1 INTRODUCTION 140
5.2 REVIEW OF LITERATURE 142
5.3 MATERIALS AND METHODS 143
5.3.1 Niche Breadth 143
5.3.2 Niche Overlap 144
5.4 RESULTS 145
5.4.1 Dietary Niche Breadth 145
5.4.2 Niche Overlap 148
5.5 DISCUSSION 148
5.6 CHAPTER SUMMARY 155
6 ASSESSMENT OF HUMAN-CARNIVORE CONFLICT AT PIR 157
LASUARA NATIONAL PARK
6.1 INTRODUCTION 157
6.2 REVIEW OF LITERATURE 159
6.3 MATERIALS AND METHODS 160
6.3.1 Methods 160
6.3.1.1 Questionnaire survey 162
6.3.1.2 Statistical analysis 165
6.4 RESULTS 167
6.4.1 Major Predator 167
6.4.2 Daily Attack Pattern 170
6.4.3 Monthly Attack Frequency 170
6.4.4 Seasonal and Annual Variation 170
6.4.5 Guarding, Herd Size and Penning 173
6.4.6 Local Perception About Conservation Status 173
6.4.7 Retaliatory Killing 174
6.4.8 Financial Losses 174
6.5 DISCUSSION 174
6.6 CHAPTER SUMMARY 179
7 GENERAL DISCUSSION 181
7.1 RECOMMENDATIONS 209
7.2 FUTURE PROSPECTIVE 210
SUMMARY 211
LITERATURE CITED 214
APPENDICES 246
List of Tables Table No. Page
2.1 Direct and indirect signs of carnivores in and around Pir Lasura 32
National Park, Azad Jammu and Kashmir, Pakistan
2. 2 Distribution of carnivore species in and around Pir Lasura National 33
Park, Azad Jammu and Kashmir, Pakistan, as shown by the
occurrence of any of the direct and indirect signs of the species. The
+ signs shows presence while – signs shows absence of the species
at particular site
3.1 Confusion matrix of scats of carnivore species identification using 71
morphological and molecular identification techniques
3.2 Error rates in species identification in the field corrected with 72
molecular identification of carnivore scats
3.3 Measurements of morphological characteristics of sympatric 73
carnivore scats (Mean ± SEM) sampled from in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan
3.4 Percent volume of prey items recovered from the scats of common 77
leopard
3.5 Percent frequency of occurrence of prey items in the scats of 77
common leopard (Panthera pardus) collected from the PLNP,
AJ&K
3.6 Percent volume of prey items recovered from the scats of Asiatic 85
jackal
3.7 Percent frequency of occurrence of prey items in diet of Asiatic 85
jackal
3.8 Percent volume of prey items recovered from the scats of Kashmir 89
hill fox
3.9 Percent frequency of occurrence of prey items in diet of Kashmir 89
hill fox
3.10 Percent volume of prey items recovered from the scats of Asian 94
Palm civet
3.11 Percent frequency of occurrence of prey items in diet of Asian palm 94
civet
3.12 Percent volume of prey items recovered from the scats of small 98
Indian civet
3.13 Percent frequency of occurrence of prey items in diet of small 98
Indian civet
3.14 Percent volume of prey items recovered from the scats of Indian 103
grey mongoose
3.15 Percent frequency of occurrence of prey items in diet of Indian grey 103
mongoose
3.16 Percent volume of prey items recovered from the scats of small 107
Indian mongoose
3.17 Percent frequency of occurrence of prey items in diet of small 107
Indian mongoose
3.18 Prey species diversity index, prey species richness and evenness in 113
diet of sympatric carnivore species in and around Pir Lasura
National Park, Azad Jammu and Kashmir, Pakistan
4.1 Abundance estimation of rodent prey species in the PLNP, AJ&K 133
4.2 Abundance estimation mammalian species in the PLNP, AJ&K 133
4.3 Abundance estimation of bird species in and around PLNP (20 133
transects, Transect 200m wide 500m long)
4.4 Abundance of amphibians and reptiles in the PLNP, AJ&K 136
4.5 Abundance estimation of invertebrates in the PLNP, AJ&K 136
4.6 Abundance of domestic prey species in district Kotli Azad 136
Kashmir. (Data source report DCR 2017.18-5-2017 data collected
from 52 units. Department of Livestock and dairy development
Kotli AJ&K. District area (1862 km sq.)
5.1 Niche breadth of sympatric carnivores occurring in and around Pir 146
Lasura National Park, Azad Jammu and Kashmir, Pakistan
L= Niche breadth; Lst= standardized niche breadth (Value 0-1)
5.2 Niche overlap between sympatric carnivore species occurring in 149
and around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
6.1 Livestock, dog, pet and poultry killed by various carnivore species 169
in and around Pir Lasura National Park (PLNP), Azad Jammu and
Kashmir during 2008-2015
6.2 Monthly variation in attacks of common leopard (Panthera pardus) 172
during 2008-2015 in and around Pir Lasura National Park, Azad
Jammu and Kashmir, Pakistan
List of Figures Fig. No. Page
1.1 Global distribution of common leopard. Distribution data retrieved 8
from IUCN database and map generated using Arc GIS software
version 10.1.
1.2 Global distribution of Asiatic jackal. Distribution data retrieved 8
from IUCN database and map generated using Arc GIS software
version 10.1
1.3 Global distribution of the red fox. Distribution data retrieved from 11
IUCN database and map generated using Arc GIS software version
10.1
1.4 Global distribution of Asian palm civet. Distribution data retrieved 11
from IUCN database and map generated using Arc GIS software
version 10.1
1.5 Global distribution of small Indian civet. Distribution data retrieved 14
from IUCN database and map generated using Arc GIS software
version 10.1
1.6 Global distribution of Indian grey mongoose. Distribution data 14
retrieved from IUCN database and map generated using Arc GIS
software version 10.1
1.7 Global distribution of small Indian mongoose. Distribution data 15
retrieved from IUCN database and map generated using Arc GIS
software version 10.1
2.1 Species accumulation curve showing diversity of carnivore species 28
in and around Pir Lasura National Park.
2.2 Distribution of common leopard (Panthera pardus) in and around Pir 29
Lasura National Park, Azad Jammu and Kashmir, Pakistan, as
indicated by various direct and indirect signs of the species
2.3 Kernel density estimation of common leopard (Panthera pardus) 30
signs in and around Pir Lasura National Park, AJ&K. Search radius
was 2000 m whereas, cell size used was 20 m. Red areas show sites
having high relative abundance of carnivore species signs
2.4 Map showing distribution of Asiatic jackal (Canis aureus) in and 35
around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan, as evident from various direct and indirect signs of the canid
species
2.5 Kernel density estimation of Asiatic jackal (Canis aureus) signs in 36
and around Pir Lasura National Park. Search radius was 2000 m
whereas cell size used was 20 m. Red areas show sites having high
relative abundance of carnivore species signs
2.6 Map showing distribution of Kashmir hill Fox (Vulpes vulpes 37
griffithi) in and around Pir Lasura National Park, Azad Jammu and
Kashmir, Pakistan, as indicated by various direct and indirect signs
of the species in the study area
2.7 Kernel density estimation of Kashmir hill fox (Vulpes vulpes 38
griffithi) signs in and around Pir Lasura National Park, AJ&K.
Search radius was 2000 m whereas, cell size used was 20 m. Red
areas show sites having high relative abundance of carnivore signs
2.8 Map showing distribution of Asian palm civet (Paradoxurus 40
hermaphroditus) in and around Pir Lasura National Park, Azad
Jammu and Kashmir, Pakistan, as indicated by various direct and
indirect signs of the species in the stud area
2.9 Kernel density estimation of Asian palm civet (Paradoxurus 41
hermaphroditus) signs in and around Pir Lasura National Park,
AJ&K. Search radius was 2000 m whereas, cell size used was 20
m. Red areas show sites having high relative abundance of
carnivore signs
2.10 Distribution of small Indian civet (Viverricula indica) in and around 42
Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan, as
indicated by its various direct and indirect signs in the study area
2.11 Kernel density estimation of small Indian civet (Viverricula indica) 43
signs in and around Pir Lasura National Park, AJ&K, Pakistan.
Search radius was 2000 m whereas, cell size used was 20 m. Red
areas show sites having high relative abundance of carnivore signs
2.12 Distribution of Indian grey mongoose (Herpestes edwardsii) in and 45
around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan, as is shown by occurrence of its various direct and indirect
signs in the study area
2.13 Kernel density estimation of Indian grey mongoose (Herpestes 46
edwardsii) signs in and around Pir Lasura National Park, AJ&K.
Search radius was 2000 m whereas, cell size used was 20 m. Red
areas show sites having high relative abundance of carnivore signs
2.14 Map showing distribution of small Indian mongoose (Herpestes 47
javanicus) in and around Pir Lasura National Park, Azad Jammu
and Kashmir, Pakistan, as indicated by occurrence of its various
direct and indirect signs in the study area
2.15 Kernel density estimation of small Indian mongoose (Herpestes 48
javanicus) signs in and around Pir Lasura National Park, AJ&K.
Search radius was 2000 m whereas, cell size used was 20 m. Red
areas show sites having high relative abundance of carnivore signs
3.1 Steps in molecular identification of carnivore scats 64
3.2 Process of carnivore scats analysis to investigate diet composition 64
of the carnivore species
3.3 Scats of sympatric carnivore species: A & B) Common leopard; C) 74
Asiatic jackal; D, E &F) Kashmir hill fox; G&H) Asian palm civet
3.4 Scats of sympatric carnivore species: A) Small Indian civet; B&C) 75
Indian grey mongoose; D) Small Indian mongoose
3.5 Variation in frequency of prey species in the diet of common 80
leopard in PLNP, AJ&K
3.6 A) Whole mount of Indian gerbil a) reference slide of Indian gerbil; 81
B) Whole mount of Turkistan rat, c) reference slide of Turkistan rat;
C) House mouse, c) reference slide of house mouse
3.7 A) Whole mount of house rat, b) reference slide of house rat; B) 82
Norway rat, b) reference slide of Norway rat; C) Desert hare, c)
reference slide of desert hare
3.8 A) Rhesus monkey a) reference rhesus monkey; B) Barking deer b) 83
reference barking deer; C) Goat, c) reference goat; D) wild boar, d)
reference wild boar
3.9 Variation in frequency of prey species in diet of Asiatic jackal 87
3.10 Variation in frequency of prey species in diet of Kashmir hill fox 91
3.11 Variation in frequency of prey species in diet of Asian palm civet 96
3.12 Variation in frequency of prey species in diet of small Indian civet 100
3.13 Variation in frequency of prey species in diet of Indian grey 105
mongoose
3.14 Variation in frequency of prey species in diet of Small Indian 109
mongoose
3.15 Prey species diversity index of dietary habits of sympatric carnivore 114
species in and around Pir Lasura National Park, Azad Jammu and
Kashmir, Pakistan
3.16 Prey species richness in diets of sympatric carnivore species in and 114
around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
3.17 Prey species evenness index of dietary habits of sympatric 115
carnivore species in and around Pir Lasura National Park, Azad
Jammu and Kashmir, Pakistan
5.1 Niche breadth of sympatric carnivore species occurring at Pir 147
Lasura National Park, Azad Jammu and Kashmir, Pakistan
5.2 Niche overlap between sympatric carnivore species at Pir Lasura 150
National Park, Azad Jammu and Kashmir, Pakistan
6.1 Locations of livestock depredation and retaliatory killing of 161
common leopard (Panthera pardus) during current study period in
and around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
6.2 Kernel density estimation for livestock depredation by common 168
leopard in and around Pir Lasura National Park. Search radius was
2000 m whereas, cell size used was 20m. Red areas shows areas
facing high livestock depredation by common leopard where
mitigation measures should be set up
6.3 Daily/hourly trend of common leopard (Panthera pardus) 171
depredation on livestock in and around PLNP, AJ&K, Pakistan
6.4 Monthly trend of common leopard (Panthera pardus) depredation 171
on livestock in and around PLNP, AJ&K
ACKNOWLEDGEMENTS
In the name of ALLAH, The Most Beneficent and The Most Merciful. All praises to
Almighty ALLAH who blessed me with the knowledge, courage and strength to work for the completion of my current thesis work in the field of Wildlife
Management, PMAS- Agriculture University, Rawalpindi. All and every respect is for His beloved and last Prophet, HAZRAT MUHAMMAD (PEACE BE UPON
HIM), who enabled us to recognize our Creator.
It is a pleasure to express my thanks to Prof. Dr. Maqsood Anwar, Dean Faculty of
Forestry, Range Management and Wildlife (FR&W) and Chairman Department of
Wildlife Management, for providing me all the facilities to complete this research work. I feel that words are inadequate to convey my sincere gratitude and deep sense of obligation and sincere thanks to my admirable supervisor Dr. Tariq Mahmood,
Assistant Professor, for his keen interest and inspiring guidance at every step of my work. His generous encouragement and gentle criticism has enabled me to produce this research work. His able guidance in this research work and attitude towards life has inspired me a lot. I am also highly obliged and thankful to the members of my supervisory committee Prof. Dr. Maqsood Anwar, and Dr. Muhammad Sajid
Nadeem, Assistant Professor, for their kind guidance and facilitation during my research work. I also wish to express my gratitude to my honorable teacher Dr.
Muhammad Rais, Assistant Professor, for his cooperation, guidance and moral support during my study period.
I am thankful to Higher Education Commission (HEC), Pakistan for providing me six-month scholarship (IRSIP) for University of Montana for molecular
identification of carnivore scats. I am also thankful to IDEA WILD (USA) for providing me equipment to conduct my research work.
I am also highly obliged and thankful to Professor L. Scott Mills for providing me facilities in University of Montana for molecular identification of carnivore scats. I am thankful to Tamara J. Max for training me to conduct my research work at
University of Montana. I am thankful to Professor Gordon Luikart for providing me lab space for DNA extractions and Tashi Dhendup for helping me in scat analysis,
Pedro Monterroso, Nathaniel Robinson (Panthera), Hugh Robinson (Panthera) for helping me in density mapping in GIS software.
There are many persons who helped me and supported me during my academic session. I cannot forget their sincere appreciation, continued encouragement and prayers for me. I also cannot forget the continued encouragement and prayers of my family members. I will extend my heartiest thanks to my wife Siddiqa Qasim for her cooperation.
Many special thanks to Shaista Andleeb, Farrukh Nazir, Shakeel Akram, Paras
Mehmood, Imad-ud-din Zangi, M. Shoaib Amjid and Danish Ameen for their cooperation and help during my research work. My extreme gratitude and sincere thanks are also due for my respected teachers, Dr. Siddique Awan, Riaz Aziz Minhas,
Shabbar Nazar, Abdul Razzaq, Abdul Rehman, Ishtiaq Choudhary, Gulzar Ahmed
Khan, Shapal Ahmed, Ikhlas Waheed for their kind guidance during my academic career.
Thanks, are also due for all lab staff of Department of Wildlife Management for their assistance in the laboratory for the current research work.
Last but not least, I acknowledge the support of my family members who always gave me hope for future and confidence in myself which has been a driving force throughout my life. They have helped me and prayed for my success and supported me morally, financially and spiritually achieving my aim in life.
I wish my well-wishers all the happiness and satisfaction both in this life and hereafter for their prayers.
Faraz Akrim
ABSTRACT
The current study aimed at investigating resource partitioning among sympatric carnivore species inhabiting Pir Lasura National Park (PLNP), AJ&K during 2014-2017. The main objectives included documenting the diversity and distribution of carnivores, investigation of the diet composition of carnivores, prey species availability, niche breadth and niche overlap among sympatric carnivores and the human- carnivore conflict at PLNP. We used direct (direct sightings, road killing) and indirect signs (scats) to document diversity and distribution of carnivore species. Scat analysis was used for diet composition study. Prey species abundance was estimated utilizing standardize method for different taxa. Niche breadth and niche overlap was computed using Levins and Pianka’s index while, human carnivore interaction was studied using questionnaire survey. The study recorded seven different species of carnivores in the Park including; common leopard
(Panthera pardus), Asiatic jackal (Canis aureus), Kashmir hill fox (Vulpes vulpes griffithi), Asian palm civet (Paradoxurus hermaphroditus), small Indian civet
(Viverricula indica), Indian grey mongoose (Herpestes edwardsii), and the small
Indian mongoose (Herpestes javanicus). Direct and indirect signs of common leopard, Asiatic jackal, Kashmir hill fox, Asian palm civet and small Indian mongoose were recorded in all 30 sampling sites having different elevational range.
Indian grey mongoose was found at 15 different sampling sites while signs of small
Indian civet were recorded at 7 sites.
The collected scats tested using molecular identification technique to confirm for the carnivore species. Accuracy for common leopard scats was found to be
95.8%, Kashmir hill fox 88.9%, Asiatic jackal 90.3%, Asian palm civet 74.3%, and
small Indian civet 92.4%. Morphological characteristics of scats showed that they overlap greatly among different species based on their diameter and length. Diet of common leopard comprised of 17 prey species (10 wild and 7 domestic). Frequency of occurrence of wild prey was 34.85 % of total leopard diet whereas domestic prey contributed 59.1%. Sixteen species of mammals, birds, insects and plants were identified from diet of Asiatic jackal. Among these species 10 were wild, 5 were domestic and 1 species of plant. Frequency of wild prey in diet of jackal was 18.48% and domestic was 59.78%. We recorded 21 species of mammals, birds, plants and insects in diet of fox. Among these 21 species 10 were wild, 5 domestic and 6 plant species. Frequency of occurrence of wild prey was 17.96%, domestic prey 50.9%, plants 27.54%. Analysis of 108 scats samples of Asian palm civet showed that 27 species of mammals, birds, invertebrates and plants were consumed. Among all recorded species 9 were wild, 3 domestic, and 15 plant species. Consumption of wild prey was 33.68% compared to 10.88% domestic prey species and 53.37% plants.
Analysis of 44 scats showed that 17 prey species occurred in diet of small Indian civet. Among them 8 were wild, 1 domestic, and 8 were plant species. Frequency of occurrence of wild prey in diet of small Indian civet was 30.65%, domestic prey
14.52%, and plants 51.61%. Analysis of 57 scat samples showed that 22 prey species occurred in diet of small Indian mongoose. Among them 15 were wild, 1 domestic, and 6 plant species. Diet of Indian grey mongoose consisted of (60%) wild species,
(19%) domestic prey species, and (14%) plant species. Analysis of 69 scat samples of small Indian mongoose showed that 17 prey species occurred in diet. Among them
10 were wild prey species, only 1 domestic prey species and 6 plant species.
Frequency of occurrence of wild prey was (59.68%), domestic prey (16.94%) and
plant matter (11.29%). The abundance of different vertebrate and invertebrate prey species was also estimated in the study area to confirm the availability of wild prey.
Among seven sympatric carnivores. Niche breadth niche breadth of Indian grey mongoose was widest 18 (0.72) followed by Asiatic jackal 14.2 (0.78), common leopard 13.88 (0.72), small Indian mongoose 12 (0.64), small Indian civet 10(0.52),
Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31). High niche overlap was found between Asian palm civet and small Indian civet (0.9), Indian grey mongoose and small Indian mongoose (0.89), Asiatic jackal and Kashmir hill fox
(0.81), while lowest niche overlap was recorded common leopard and small Indian civet (0.04). Data on livestock depredation by carnivores showed that 170 people lost 306 livestock, poultry, dogs, and pets to four different carnivore species; common leopard being the major predator killing (88.5%) livestock, with minor contributions from Asiatic jackal (5.2%), small Indian civet (3.2%) and Indian grey mongoose (2.9%). The depredated livestock by leopard mainly included goats
(82.2%) and dogs (9.2%). Common leopard was responsible for majority of the financial loss estimated approximately at US$ 80,000 and it negatively affected local people tolerance towards this top predator. The study concludes, niche of seven sympatric carnivore species overlaps in the study area with high overlap recorded for Asian palm civet and small Indian civet (0.9) and lowest was recorded for the leopard and small Indian civet (0.04).
Chapter 1
GENERAL INTRODUCTION
1.1 INTRODUCTION
Carnivores belong to order carnivora which is diverse order of eutherian
mammals. Carnivores are primarily meat eater, although few species are often
omnivorous such as mongooses, foxes and civets (Hinton and Dunn, 1967; Barun et
al., 2008; Mahmood et al., 2011). Carnivores range in size from the least weasels to
elephant seals. Carnivores are known to predate on variety of prey species. To
understand impact of a predator on dynamics of prey populations, the knowledge of
predator’s diet is vital (Oli, 1993). Diet composition of carnivores is usually
investigated using their fecal droppings (Shehzad et al., 2015). Faecal components
of carnivores may include remains of birds, bones, mammalian hairs, teeth, claws,
scales, parts of invertebrates, plant matter as well as mucus cells, and bacteria (Bang
and Dahlström, 1975; Bujne, 2000). Whereas, the size and quantity of carnivore scat
can be different based on age of carnivore, type of food eaten and absorption capacity
(Bang and Dahlström, 1975).
A variety of information can be obtained from scats and their contents, like animal identification (Seton, 1925; Camardella et al., 2000), activity centers of animal
(Walker, 1996), composition of diet (Chinchilla, 1997; Santos and Hartz, 1999;
Kauhala and Auniola, 2001), seasonal changes in diet (Aragona and Setz, 2001), prey species inventory (Camardella et al., 2000), role in seed dispersal (Fragoso and
Huffman, 2000; Williams et al., 2000), animal health condition, and entero-parasitosis dynamics (Page et al., 2001).
1
Species having similar ecological niches often divide their uses of resources to avoid competition, in habitats where they are sympatric (Schoener, 1986) and such a phenomenon is known as “resource partitioning” (Walter, 1991). Many factors play role in this phenomenon, like interspecific competition (Schoener, 1974), change in tolerance towards physical-chemical variables, environmental change, spatial and temporal change in availability of resources, predation (Ross, 1986) and intra-guild predation (Fedriani et al., 2000; Macdonald and Sillero-Zubiri, 2002 ). Interspecific competition is important force in structuring composition and resource composition of carnivore community (Delibes, 1983; Hersteinsson and Macdonald, 1992; White et al., 1994; Garcia and Virgos, 2007b). If competition does not occur in group of similar species, species may co-exist with population size lower than carrying capacity (suboptimal state) or to separate their resource utilization or to reduce competition and facilitate coexistence of these species (Lotka, 1932; Schoener, 1974;
Pianka, 1988). Extent of Resource partitioning depends upon difference in body size of the species involved (Schoener, 1974). The divergence in morphological traits is often associated with the divergence in ecological traits, for example, food (Brown and Wilson, 1956; Sih, 1993; Dayan and Simberloff, 1998; Dayan and Simberloff,
2005). Since larger animals consume larger prey and variety of eaten prey is greater as compared to the smaller animals, larger animals have broader trophic niche breadth as compared to the smaller (Schoener, 1974).
Association between body size and food resource partitioning has been previously reported in fishes (Ross, 1986; Schafer et al., 2002), invertebrates
(Woodward and Hildrew, 2002; Tall et al., 2006), shrews (Brannon, 2000) and carnivores (Martin, 1994; Jones, 1997; Dayan and Simberloff, 1998; Jones and
Barmuta, 2000; Garcia and Virgos, 2007a). Sympatric small carnivores consume rodents as their major food source in common and also took other secondary food items in size corresponding to their body size. Resource availability affects resource partitioning, generally, the degree of diet overlap among sympatric carnivore will increase with increase in abundance of common food and decrease with lean time
(Schoener, 1974; Schoener, 1982; Wiens, 1989; Smith, 1991) as the species will shift their food to different food items during the lean time. In addition, feeding strategy of different animals may also affect the results of resource partitioning; for example, opportunistic generalists may consume all of the remaining food resources. If these available food resources are similar among patches, as is the case usually during lean time, overlap in diet of these species will increase (Serafini and Lovari, 1993; Padial et al., 2002). But in a community, that consists of both generalist and specialist predators, diet overlap may increase or decrease between different species during lean time (Carvalho and Gomes, 2004).
Dietary overlap between carnivores can indicate level of interspecific competition. Manipulative studies can help to understand competition (Mac-Nally,
1983; Wiens, 1989), data on niche overlap is essential first step (Carrera et al., 2008;
Glen and Dickman, 2008). High overlap among carnivore species indicate level of intraspecific competition as well as strength of interference competition. Predators having similar diet often exhibit aggression, in order to eliminate competitor or predators may often encounter each other while searching for similar prey (Donadio and Buskirk, 2006; Polis et al., 1989). Carnivore living in premises of human occupied landscapes are reported to take up domestic livestock as prey (Sangay and
Vernes, 2008; Shehzad et al., 2015) and such cases usually result in antagonistic
human carnivore interaction.
Accurate data on distribution of a species is vital for its conservation and management. Recorded distribution of a species can be bias as it can reflect the distribution of recorders despite of species under study. Sometimes, it is only possible to survey for indirect signs of elusive species, like footprints, hairs and scats.
Through systematic surveys, correct data of indices can be collected (Gibbs, 2000).
Species distributed in areas where no significant research studies exists, are recorded by presence of their signs such as scat or hair (Busby et al., 2009).
1.1.1 Common Leopard
Common leopard (Panthera pardus) has widest distribution range of any felid species (Bailey, 1993; Nowell and Jackson, 1996). It is distributed from western and southern Africa to the Russian far east and Java (Stein and Hayssen, 2013)
(Figure 1.1). Such a large distribution range is result of high adaptability of the species, since it can survive diverse ecosystems ranging from boreal forest to tropical rainforests and arid savannah (Bertram, 1999). In Pakistan, Common Leopard is distributed in hilly forest areas of Punjab, Baluchistan, Sindh, Khyber-
Pakhtoonkhwa and Azad Jammu and Kashmir. It inhabits mountains of Waziristan,
Baluchistan and Sindh Kohistan in association with Acacia scrub forest (Roberts,
1997). In the north, it is found in the Murree Hills, Swat, Kohistan, Dir, Chitral,
Abbottabad, Kaghan valley, Gilgit, Margalla Hills National Park and Neelum Valley in AJ&K. Increasing human population and firearms has resulted in decline of its population in Pakistan (Roberts, 1997).
Leopard has broad diet as it can feed on broad spectrum of prey species and
it makes it successful predator among big cats whereas, its size results in ability to feed on a diverse prey species ranging from rodent to buffalo (Sankar and Johnsingh,
2002; Goyal and Chauhan, 2006; Qureshi and Advait, 2006; Andheria et al., 2007;
Arivazhagan et al., 2007; Ahmed and Khan, 2008; Chattha et al., 2015; Shehzad et al., 2015). Leopard is known to predate on domestic animals and dogs. In many parts of its range livestock has been reported as major part of its diet (Mukherjee and
Mishra, 2001) livestock has been reported to dominate wild prey in leopard’s diet
(Athreya et al., 2013; Shehzad et al., 2015). After goats and sheep, dogs have been reported as major prey of leopards (Shehzad et al., 2015; Athreya et al., 2016).
Common leopard is categorized as Vulnerable under IUCN red list of threatened species (Iucn, 2006). Anthropogenic activities are major threat to leopards. Habitat fragmentation, decline in prey species and antagonistic interaction with humans has resulted in reduced leopard population (Nowell and Jackson, 1996;
Ray et al., 2005; Hunter, 2013). The conversion of wild habitats into agricultural lands, livestock farms, and human settlements has resulted in population decline of leopards, with few exceptions (Athreya et al., 2013), such conversion usually results in depletion of natural prey species thereby population of predator declines in such areas.
1.1.2 Asiatic/ Golden Jackal
The jackal (Canis aureus) is distributed in North and north-east Africa, it is distributed from Senegal (Africa) to Egypt in the east, including Morocco, Algeria,
Libya in north to Nigeria, Chad and Tanzania in the south. Jackal is also distributed in Arabian Peninsula and have expanded into Europe having patchy distribution, and also distributed in Balkans, Hungary and south-western Ukraine. Jackal is vagrant in
Austria, Slovakia, Slovenia and north-eastern Italy (Krystufek et al., 1997).
Eastwards jackals are distributed in Turkey, Syria, Iraq, Iran, Central Asia, the entire
Indian subcontinent, then east and south to Sri Lanka, Myanmar, Thailand and parts of Indo-China (Jhala and Moehlman, 2008) (Figure 1.2).
The Asiatic jackal can survive in variety of habitats ranging from deserts to dense forests (Macdonald, 1984; Roberts, 1997), however, jackal prefers grasslands, scrubland, open areas and marshes up to 1,000 m in elevation (Anitei, 2008). In
Pakistan, jackal is distributed in plains of Sind, Punjab, Baluchistan and Khyber
Pakhtoonkhwa (KPK) provinces.
Although, jackal is not distributed at higher elevations in Himalayas, however, it has been recorded at lower elevations in Himalayan valleys such as Murree hills up to 2150 m or 7000 ft (Roberts, 1997). Asiatic jackal is an opportunistic feeder and it usually scavenge on garbage, human waste and animal carcasses. Jackal can hunt singly and in packs which are more successful in hunting. When single it can hunt smaller prey such as rodents, hares and birds by locating prey species by hearing. It can dig out gerbils (Tatera indica) from burrows and can hunt young, old, sick ungulates 4-5 times larger than its own size and body weight. Besides vertebrates,
Asiatic jackal has been reported to feed on invertebrates and plants matter. Jackals are omnivorous, and diet width of jackal is variable based on different seasons and habitat
(Wyman, 1967; Moehlman, 1983). They are generalist feeders and large quantities of vegetable matter and fruits are included in their diet, however, bulk of their food comprises of rodents and reptiles. They also supplement their diet with fruits and insects when available (Roberts, 1997).
Population of jackal is declining in entire distribution range, with exception of
protected areas. Traditional land use practices, are being replaced by industrialization and intense agricultural practices as a result wild as well as rural landscapes which were known to conducive to survival of jackals are rapidly urbanized. Jackal populations may persist for a while and try to adapt such a change in environment, but eventually disappear from such areas like other species of wildlife (Jhala and
Moehlman, 2008). The other known threats include local policies for extirpation of jackals and poisoning (for example, Israel and Morocco). Jackals are hunted as a game and eaten as food in Morocco (F. Cuzin pers. comm. 2007). There is little trade in jackal products such as skins and tails are occasionally sold (Jhala and Moehlman,
2008).
1.1.3 Red Fox
The red fox (Vulpes vulpes) is most widespread terrestrial carnivore species globally (Harris and Baker, 2001). It is distributed across the whole northern hemisphere from the Arctic circle to southern North America Europe, North Africa,
Asiatic steppes, Pakistan, Indian and Japan (Figure 1.3) (Gloor et al., 2001;
Wandeler et al., 2003; Soulsbury et al., 2010; Jenkins and Craig, 1992; Crooks et al., 2010).
The red fox is an omnivore (Flower, 1932; Macdonald, 1979; Osborn and
Helmy, 1980). It has broad diet which includes; invertebrates, small mammals, birds, fishes, fruits and carrion (Flower, 1932; Macdonald, 1979; Osborn and Helmy,
1980). Diet composition of the red fox depends on various factor including; habitat type, prey availability (Kolb and Hewson, 1979; Leckie et al., 1998; Sidorovich et al., 2006) and seasonal variation in food availability (Goszczyński, 1986;
Jędrzejewski and Jędrzejewska, 1992; Baltrunaite, 2001; Baltrunaite, 2002).
Figure 1.1 Global distribution of common leopard. Distribution data retrieved from
IUCN database and map generated using Arc GIS software version 10.1.
Figure 1.2 Global distribution of Asiatic jackal. Distribution data retrieved from
IUCN database and map generated using Arc GIS software version 10.1.
Anthropogenic activities like habitat degradation, habitat loss, fragmentation, exploitation, and direct and indirect persecution are major threats to the red fox. In
Mongolia the red foxes are near threatened according to local red list assessment
(Clark and Munkhbat, 2006) due to overhunting. In south Korea, habitat loss and poaching has resulted in extinction of this species (Yu et al., 2012).
1.1.4 Common/Asian Palm Civet
Common palm civet (Paradoxurus hermaphroditus) is widely distributed in south and south-east Asia from Afghanistan in the west to Hainan to the adjacent areas of Chinese coast in the east; it is distributed in south-east Asian islands (Figure
1.4) (Patou et al., 2010; Stevens et al., 2011; Veron et al., 2015). It has been introduced in Sulawesi and the Lesser Sundas, while the Philippine archipelago might have been colonized naturally but also might stem entirely from introductions
(Veron et al., 2015). Its distribution in China is restricted to Hainan, southern
Guangdong, south-western Guangxi, much of Yunnan and south-western Sichuan provinces (Wang and Fuller, 2003; Lau et al., 2010). It occurs on the islands of
Bawean (Indonesia), Con Son (Viet Nam), Koh Samui (Thailand), Koh Yao
(Thailand), and Telebon (Thailand) (Meiri, 2005), and on the Philippine islands of
Balabac, Busuanga, Camiguin, Culion, Leyte, Luzon, Marinduque, Mindanao,
Negros, Palawan, Sangasanga, Sibuyan (specimens) and Catanduanes, Biliran,
Maripipi and Panay (Heaney et al., 1998). It has been introduced in Japan, but this reflects confusion with masked palm civet Paguma larvata (S. Roy in litt. 2014). It occurs widely at sea-level; the highest distribution records globally are those at
2,400 m in North-east India (Choudhury, 2013) and in Afghanistan at 2,500 m
(Stevens et al., 2011).
Common palm civet is used as food in south-east Asia, and used for both, subsistence and trade to urban luxury restaurants. It is kept as a 'pet' and in captivity it is used for the production of coffee such as in Indonesia (Shepherd, 2012; Nijman et al., 2014).
1.1.5. Small Indian Civet
Small Indian civet (Viverricula indica) is categorized as least concern according to IUCN red list of threatened species (IUCN, 2006). Small Indian civet is distributed in Sri Lanka, Pakistan. India, Kashmir, Nepal, China, Bangladesh, mainland South-east Asia, and many islands of Indonesia (Sumatra, Java, Bali,
Bawean, Bintan, Kangean, Lombok, Panaitan, and Sumbawa; it should not be assumed to be native to those lying east of Bali) (Figure 1.5) (Roberts, 1997; Wang and Fuller, 2001; Wang and Xie, 2004; Than et al., 2008; Chen et al., 2009; Charoo et al., 2010; Lau et al., 2010; Jennings and Veron, 2011; Choudhury, 2013;
Mudappa, 2013; Chutipong et al., 2014).
Small Indian civet is used for wide range of purposes. It is harvested for skins in China (Lau et al., 2010). Its perineal gland secretions called “civet musk” are used in perfumes and medicines. These are kept captivity in India for such purposes and, in Thailand; these are harvested entirely from the wild (Balakrishnan and Sreedevi, 2007; Chutipong et al., 2014). In North-east India, southern China and northern south-east Asia it has been hunted using by using snaring, for domestic uses and for into urban and international wildlife trade. Small Indian civet is insulated from the heaviest pressures as such activities are restricted to closed evergreen forest, which is only a marginal habitat of this species. Civets are also persecuted by local communities as they cause damage to orchids (Jhala and Moehlman, 2008).
Figure 1.3 Global distribution of the red fox. Distribution data retrieved from IUCN
database and map generated using Arc GIS software version 10.1.
Figure 1.4 Global distribution of Asian palm civet. Distribution data retrieved from
IUCN database and map generated using Arc GIS software version 10.1.
1.1.6 Indian grey mongoose
Indian grey mongoose (Herpestes edwardsii) is mainly found in south Asia;
Pakistan, Afghanistan India, Nepal up to Ceylon (now Sri Lanka). The species also occupies coastal areas of Saudi Arabia and Iran (Ewer, 1973; Nowak, 2005;
Macdonald, 2006; Francis, 2008; Gilchrist et al., 2009). It has also been introduced to Japan and Peninsular Malaysia (Figure 1.6) (Francis, 2008; Gilchrist et al., 2009).
In Pakistan, grey mongoose is common in central and northern parts of Sindh particularly inhabiting the desert tracts of Tharparkar. It also occurs in some parts of
Punjab; Rawalpindi and the Salt Range. In Balochistan, grey mongoose is sparsely found in the southern parts. It also occurs in Peshawar, Kohat and Bannu districts in the province of Khyber Pakhtunkhwa (Roberts, 1997).
The grey mongoose is an opportunistic hunter, its common food items include mice, rats, lizards, snakes, beetles, ground birds and their eggs, and parts of plants; fruits, berries and roots. In India, it has been reported to feed on eggs and chicks of the red jungle fowl, peafowl, partridges, snakes, small mammals, and was also found searching food under stones on the beach side in Hawaii, (Santiapillai et al., 2000; Postanowicz, 2002). Grey mongoose has elongated skull and special teeth for hunting grasshoppers, scorpions, centipedes, frogs, crabs and fish. The protruded and pointed canines help to clamp up a snake's head and the molars with pointed cusps help in crushing insects (Whitfield, 1978).
Shekar (2003) reported that Indian grey mongoose is sold and also kept as a pet. In northern India it is captured for skin by using hook snares, these captured mongooses are sold in local markets in Nepal (Shekar, 2003). In India, all mongoose species are captured for wildlife trade (Van and Jayakumar, 2003). Their meat is
used as food in many tribes and the hair are used to make shaving brushes, paint brushes, and good luck charms (Hanfee and Hhmed, 2000).
1.1.7 Small Indian Mongoose
The small Indian mongoose (Herpestes javanicus) occurs naturally in the southern and south-eastern regions of Asia (Wozencraft, 2005), where it has a native range from Pakistan and northern India to southern China and the Malay Peninsula.
It is also found on Hainan Island and Java. In the west, it extends to southern Iran
(Corbet and Hill, 1992), south western Afghanistan (Hassinger, 1968), along the shore of Persian Gulf it extends up to Kuwait and Iraq (Figure 1.7).
In Pakistan, small Indian mongoose is widely distributed in Sind province; in
Tharparkar, Thatta and Dadu districts. In Punjab province, it occurs in Lahore,
Kasur, Sialkot, Gujranwala and Jhelum districts. It also occurs in the Salt Range and sparsely in Bahawalpur division. It is also found in the southern part of Baluchistan province but not yet been reported from Khyber Pakhtunkhwa (Roberts, 1997).
Mongooses are terrestrial, diurnal and solitary hunters. They use a quick trot, moving constantly, scanning the area for food (Santiapillai et al., 2000). The small
Indian mongoose is an important carnivore in biological niche. Small Indian mongoose is omnivore and its diet include; small mammals, birds, herpetofauna and invertebrates to plant materials. Some populations are insectivorous, others may consume fruits for a part of the year (Seaman and Randall, 1962). A few earlier studies conducted on food habits of the small Indian mongoose in Pakistan have focused this species in the central Punjab (Siddiqui et al., 2004) and the Pothwar
Plateau (Mahmood et al., 2011). In central Punjab region, the food composition of the species includes insects, birds feathers, bones, plant materials and hair, whereas,
Figure 1.5 Global distribution of small Indian civet. Distribution data retrieved from
IUCN database and map generated using Arc GIS software version 10.1.
Figure 1.6 Global distribution of Indian grey mongoose. Distribution data retrieved
from IUCN database and map generated using Arc GIS software version
10.1.
Figure 1.7 Global distribution of small Indian mongoose. Distribution data retrieved
from IUCN database and map generated using Arc GIS software version
10.1.
in the Pothwar Plateau, it feed on invertebrates such as insects, arthropods, small reptiles and vertebrate mammalian species (Mahmood et al., 2011).
This species is often captured and sold as pets (Shekar, 2003), and there is some commercial trade in India and Nepal. Mongooses are poached in India for their hairs, which are used for making paint and shaving brushes (Sahajpal et al., 2009).
1.2 STUDY AREA
The current study was carried out in and around Pir Lasura National Park
(PLNP) Tehsil Nakyal, District Kotli, Azad Jammu and Kashmir, Pakistan. The area is located in the south-eastern part of the State, close to the Line of Control between
o o o o 33 25.92 N to 33 29.31 N and 74 05.64 E to 74 03.02 E. The park encompasses
1580 ha area with elevation ranging between 1000 m – 2000 m above sea level (asl).
The lower elevations of the park consist of subtropical pine forest type vegetation; whereas; the tops/mountains comprised of sub-tropical dry evergreen forest pattern.
The climate of the study area is cold and humid forest. The average annual rainfall is 1500 mm.
The study area experiences four different seasons including summer (May-
July), autumn (August-October), winter (November-January), and spring (February-
April). Major wildlife species in the park include common leopard (Panthera pardus), rhesus monkey (Macaca mulatta), Asiatic jackal (Canis aureus), red fox
(Vulpes vulpes), small Indian mongoose (Herpestes javanicus), Indian grey mongoose (Herpestes edwardsii), barking deer (Muntiacus muntjak), Indian pangolin (Manis crassicaudata) and kaleej pheasant (Lophura leucomelanos).
However, there are no previous studies published reporting the population status of
carnivores or abundance of their prey species in the study area.
The park is surrounded by local communities especially on eastern side there is a whole Nakyal town and on western side there is Supply town having considerable human population. Also rest of surrounding and National Park itself have sparse human population. Local people keep a variety of animals including domestic cows, buffalos, goats, dogs, horses, poultry birds and rabbits. A reasonable majority of people are associated with professions of doing agriculture, government jobs, labor and shop keeping with average household income per month ranging from US$ 100-
200. Farmers, shopkeepers and labors usually keep livestock for milk and meat production and they depend on livestock for subsistence.
1.2.1 Study Objectives
The current study was designed to investigate ecological aspects of sympatric carnivore species at Pir Lasura national park, Azad Jammu and Kashmir, Pakistan, with hypothesizing that “Dietary niche overlap may occur among sympatric carnivore species inhabiting Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan” with following study objectives;
1- To document the diversity of carnivore species and their distribution in
Pir Lasura National Park.
2- To investigate food habits of different carnivore species in the park.
3- To assess abundance of carnivore prey species in the study area.
4- To study dietary niche breadth and niche overlap among the sympatric
carnivore species of the study area.
5- To investigate human-carnivore conflict in the park.
1.3 RATIONALE OF THE STUDY
The Pir Lasura National Park in AJ&K is located in North-eastern locale of
Pakistan which is known for rich fauna and flora. Despite of having rich biodiversity,
Pir Lasura National Park has not been focused for wildlife studies in the past. Data on diversity, distribution, diet composition of sympatric carnivores, seasonal variation in their diet, availability of prey species to carnivores, niche breadth, niche overlap and human carnivore interaction is very important from conservation and management point of view. However, such data was lacking from Pir Lasura national park and current study was undertaken with the aim to understand carnivore ecology to fill the gap by generating required scientific information, prerequisite for conservation and management of carnivores and their prey species in the study area.
1.4 ORGANIZATION OF THE THESIS
This study was designed to explore and document diversity, distribution, diet composition, seasonal variation in diet, seasonal variation in availability of prey species, niche breadth, niche overlap, and human carnivore interaction in Pir Lasura
National Park Azad Jammu and Kashmir, which is located in the North-eastern
Himalayan region of Pakistan. To meet the study objectives the research work and thesis has been divided into different chapters corresponding to the five objectives described above.
The chapter 1 describes the general introduction of the research study explaining the distribution, general diet and economic importance of seven sympatric carnivore species recorded during current study. In chapter 2, diversity and distribution of sympatric carnivore species in the study area is explained. The chapter
3, documents the errors involved in observer identification of carnivore scats (based on morphological characteristics) vs molecular identification, variation in morphological characteristics of carnivore scats, diet composition of 7 sympatric carnivore species and seasonal variation in their diet. The chapter 4, investigates the seasonal variation in abundance of prey species of carnivores occurring in the study area. The chapter 5 focuses the niche breadth and niche overlap among sympatric carnivore species and chapter 6 investigates human-carnivore interaction in the study area with major emphasis on common leopard. In chapter 7 overall results of the study are discussed.
Chapter 2
DIVERSITY AND DISTRIBUTION OF CARNIVORES AT PIR
LASURA NATIONAL PARK, AZAD JAMMU AND KASHMIR,
PAKISTAN
2.1 INTRODUCTION
The order carnivora is one of the largest group of species among mammals
(Gittleman et al., 2001). Species belonging to order carnivora are primarily meat eater and are often omnivorous (Barun et al., 2008). Carnivores play important role in determining the structure of the communities (Terborgh et al., 1999; Gittleman and Gompper, 2005). There is great variation in size of species belonging to order carnivora which range in size from the least weasel to the southern elephant seal.
Accurate knowledge of species distribution is very important for conservation, protection, and management of wildlife. Sometimes, it is only possible to survey for indirect signs of the elusive species, such as hairs, scats, or footprints but it depends upon correct identification of indices. Through systematic surveys, correct data of indices can be collected (Gibbs, 2000). Species distributed in areas where no significant research studies exist, are recorded by presence of their signs such as scat or hair (Busby et al., 2009).
Many species belonging to order carnivora are the most threatened species in the world since they have specific requirements and they need conservation efforts for their survival. Many species of carnivores need larger areas to sustain viable populations and such species can be argued to be keystone species. Carnivores show
20
extreme variation in many characteristics which include; size, reproductive rates, habitat use pattern, home range and social structure (Gittleman et al., 2001). A protected area can have many species of carnivores which may range from 250 kg
Panthera leo to 300g dwarf mongoose. Such variability among the order carnivora makes it priority for conservation as conserving these carnivore species will effectively preserve a larger degree of biodiversity (Gittleman et al., 2001).
Carnivores, especially big cats can be argued to be the indicator of ecosystem health as their presence indicate the prey species and in turn, prey species depend on productive vegetation. Explicit information of diversity and distribution of carnivore species in a protracted area is prerequisite for conservation and management purposes. The Pir Lasura National Park, Azad Jammu and Kashmir part of Pakistan is such an area which lacks baseline ecological information on diversity and distribution pattern of carnivore species. Therefore, the current study was designed to investigate diversity and distribution of carnivore species in the study area in order to improve our understanding of species composition and their distribution pattern in and around the protected area.
2.2 REVIEW OF LITERATURE
Accurate data of species distribution is vital for conservation, and management of wildlife. Sometimes, it is only possible to survey for indirect signs of elusive species, like scats, hairs, and footprints. Through systematic surveys, correct data of indices can be collected (Gibbs, 2000). Species distributed in areas where no significant research studies exists, are recorded by presence of their signs such as scats or hairs (Busby et al., 2009).
Distribution range of common leopard (Panthera pardus), one of the top predators in an ecosystem, is widest as compared to other felid species (Bailey, 1993;
Nowell and Jackson, 1996). Its distribution has been recorded in Africa, Russia to
Java (Stein and Hayssen, 2013). Large distribution of common leopard is result of its adaptability as it can survive in a diversity of habitats (Bertram, 1999). In
Pakistan, common leopard is distributed in the provinces of Punjab, Baluchistan,
Sindh, Khyber-Pakhtoonkhwa, and Azad Jammu and Kashmir (AJ&K). It is found throughout Waziristan, Baluchistan and Sindh Kohistan (Roberts, 1997). In the northern mountainous region, it is found in the Murree Hills, Swat, Kohistan, Dir,
Chitral, Abbottabad, Kaghan valley, Gilgit, Margalla Hills National Park and
Neelum valley in AJ&K. Increased human settlements and firearms have resulted in decreased distribution range of leopards in Pakistan (Roberts, 1997).
The Asiatic jackal (Canis aureus) is distributed in Africa to Egypt to
Morocco, Algeria, and Libya, Nigeria, Chad and Tanzania. It is also reported in
Arabian Peninsula to Europe, having patchy distribution. It is also found in Balkans,
Hungary and Ukraine. It also occurs as a vagrant in Austria, Slovakia, Slovenia and
Italy (Krystufek et al., 1997). In the east, the canid species is found in Turkey, Syria,
Iraq, Iran, Central Asia, the entire Indian subcontinent, then east and south to Sri
Lanka, Myanmar, Thailand and parts of Indo-China (Jhala and Moehlman, 2008). In
Pakistan, it occurs throughout the plains of province Sind, Punjab, Balochistan and
Khyber Pakhtoonkhwa (KPK). Although it does not penetrate higher elevations, but it does occur in most of the broader Himalayan valleys such as Murree hills up to
2150 m or 7000 feet (Roberts, 1997).
The red fox (Vulpes vulpes) is most widespread terrestrial carnivore species
globally (Harris and Baker, 2001). It is distributed across the whole northern hemisphere from the Arctic circle to southern North America, Europe, North Africa,
Asiatic steppes, Pakistan, India and Japan (Gloor et al., 2001; Wandeler et al., 2003;
Soulsbury et al., 2010; Jenkins and Craig, 1992; Crooks et al., 2010).
Distribution of Asian/common palm civet (Paradoxurus hermaphroditus) has been recorded in South and South-east Asia from Afghanistan in the west to
Hainan and in the east Chinese coast. It is also found in South-east Asian islands
(Patou et al., 2010; Stevens et al., 2011; Veron et al., 2015). It was introduced in
Sulawesi and the Lesser Sundas, while it might have been colonized naturally in the
Philippine (Veron et al., 2015). Its distribution in China is restricted, to
Hainan, Guangdong, Guangxi, Yunnan and Sichuan provinces (Wang and Fuller,
2003; Lau et al., 2010). It occurs on the small islands of Bawean in Indonesia, Con
Son in Vietnam, Koh Samui and Telebon in Thailand (Meiri, 2005), and on the
Philippine islands of Balabac, Busuanga, Camiguin, Culion, Leyte, Luzon,
Marinduque, Mindanao, Negros, Palawan, Sangasanga, Sibuyan (specimens) and
Catanduanes, Biliran, Maripipi and Panay (Heaney et al., 1998). It has been recorded at 2,400 m in North-east India (Choudhury, 2013) and in Afghanistan at 2,500 m
(Stevens et al., 2011).
Small Indian civet (Viverricula indica) has been recorded in Pakistan, India,
Sri Lanka, Nepal, China, Bangladesh, South-east Asia and many islands of Indonesia
(Sumatra, Java, Bali, Bawean, Bintan, Kangean, Lombok, Panaitan, and Sumbawa)
(Roberts, 1997; Wang and Fuller, 2001; Wang and Xie, 2004; Than et al., 2008;
Chen et al., 2009; Charoo et al., 2010; Lau et al., 2010; Jennings and Veron, 2011;
Choudhury, 2013; Mudappa, 2013; Chutipong et al., 2014).
Indian grey mongoose (Herpestes edwardsii), a small carnivore, is mainly found in south Asia; Pakistan, Afghanistan, India, Nepal up to Ceylon (now Sri
Lanka). The species also occupies coastal areas of Saudi Arabia and Iran (Ewer,
1973; Nowak, 2005; Macdonald, 2006; Francis, 2008; Gilchrist et al., 2009). It has also been introduced to Japan and Peninsular Malaysia (Francis, 2008; Gilchrist et al., 2009). In Pakistan, grey mongoose is common in central and northern parts of
Sindh province particularly inhabiting the desert tracts of Tharparkar. It also occurs in some parts of the Punjab; Rawalpindi and the Salt Range. In Balochistan, grey mongoose is sparsely found in the southern parts. It also occurs in Peshawar, Kohat and Bannu districts in the province of Khyber Pakhtunkhwa (Roberts, 1997).
The small Indian mongoose (Herpestes javanicus), another small carnivore, has been recorded in Asia (South and south-eastern part) (Wozencraft, 2005), where it has a native range from Pakistan to northern India, southern part of China and the
Malay Peninsula. It has been reported from Hainan Island, Java and southern Iran
(Corbet and Hill, 1992), Afghanistan (Hassinger, 1968), Kuwait and Iraq (Harrison,
1968). In Pakistan, small Indian mongoose is distributed in Sind, Punjab and
Baluchistan. In Punjab, it is reported from Lahore, Kasur, Sialkot, Gujranwala and
Jhelum districts. It also occurs in the Salt Range and sparsely in Bahawalpur division.
It has not yet been reported from Khyber Pakhtunkhwa (Roberts, 1997).
2.3 MATERIALS AND METHODS
2.3.1 Diversity of Carnivores
For recording diversity of carnivore species in the study area, we used both direct and indirect methods which included; direct observation of carnivore species
by surveying study area using established transects in different study sites (N=30) in and around the National Park area. All transects were surveyed during day and night times. Moreover, we also recorded all opportunistically encountered carnivore species while moving on car or on foot. The road killed carnivore species were also recorded. Indirect methods included; collection of carnivore scats and their morphological identification (n = 551), followed by refinement of morphological identification using a subsample of 149 scats from total of 551 scats for species identification using Sanger Sequencing. Morphological identification of scats was corrected using molecular identification data and 473 scats were confirmed to be originating from wild carnivore species while rest of sample were either from non- target species or remained unidentified. Furthermore, field staff of the Department of Wildlife and Fisheries Azad Jammu and Kashmir, Pakistan and local communities were approached for data collection regarding presence or absence of the carnivore species in the study area.
2.3.2 Distribution of Carnivore Species
Extensive field surveys were conducted to document distribution of carnivore species in and around Pir Lasura National Park during the study period from 2014 to
2017. Carnivore species occurring in the park and their distribution were studied by recording direct (direct sightings and road killed carnivore species) and indirect signs like scats (morphological followed by molecular identification) of all carnivore species in the study area at 30 study sites (Wemmer et al., 1996). Data were also collected from local community living in and around the park and field staff of department of Fisheries and Wildlife AJ&K, who were also approached for data
collection. Data on site, geographic location, elevation, date and species identification for each scat were recorded. The sites having signs of carnivore species were marked as positive for their presence and vice versa. The data were processed in Quantum GIS (Version 2.2.3) and Arc GIS (Version 10.1) to produce distribution maps.
2.3.3 Statistical Analysis
Estimate S software package was used to analyzed data related to diversity of carnivore species occurring in and around Pir Lasura National Park, AJ&K. The data related to the species detected in each survey were subjected to analysis in
Estimate S software (version 9.1.0) and AEC, Chao 1 and Chao 2 estimators were used for computing carnivore species diversity.
To estimate abundance of carnivore species using their signs, we used a kernel density analysis. Kernel density analysis expresses sign density of each carnivore species (including direct and indirect signs) per kilometer square of study area. Mapping kernel density allowed us to identify areas having high sign density of carnivore species. To estimate a density value, we used bandwidth of 2000m as a search radius to calculate sign abundance of carnivores per square kilometer and cell size was 20m. Estimated density values were then classified using Jenks methods
(Jenks and Caspall, 1971).
2.4 RESULTS
2.4.1 Diversity of Carnivore Species
Seven species of carnivores were recorded in and around the Pir Lasura
National Park, Azad Jammu and Kashmir, Pakistan which included; Common leopard (Panthera pardus), Asiatic jackal (Canis aureus), Kashmir hill fox (Vulpes
Vulpes griffithi), Asian palm civet (Paradoxurus hermaphroditus), small Indian civet
(Viverricula indica), Indian grey mongoose (Herpestes edwardsii) and small Indian mongoose (Herpestes javanicus) (Table 2.1; Table 2.2). The AEC, Chao 1 and Chao
2 estimators were used to compute carnivore species diversity in and around Pir
Lasura National Park and each estimator estimated 7 carnivore species in the study area (Figure 2.1).
2.4.2 Distribution of Carnivore Species
2.4.2.1 Distribution of common leopard
Among all surveyed sites (N=30) distribution of the common leopard was recorded as positive at all sites surveyed. Altitudinal range of distribution of the cat species was between 757 m- 1891 m. Scats of common leopard (n = 39) were found and collected from 7 different sites which included; Kothian (N=4; 10.2%), Palani
(N=1; 2.5%), Panagali (N=8; 20.5%), Pir Kana (N=8; 20.5%), Pothi sairi (N=3;
7.7%), Sairi (N=12; 30.7%) and Supply (N=3; 7.7%). Livestock depredation by common leopard was reported from 22 different sites of the study area while common leopard was directly field observed at one sampling site named “Sairi” and its sightings were reported from 6 other sites in the study area. A high sign density of common leopard was recorded at Pir Kana and Sairi sampling sites (Table 2.1;
Table 2.2; Fig. 2.2; Fig. 2.3).
2.4.2.2 Distribution of Asiatic jackal
Asiatic jackal was recorded at all 30 sites surveyed in the study area.
S Mean (runs) ACE Mean Chao 1 Mean Chao 2 Mean 7.6 7.4 7.2 7 6.8 6.6
6.4 No. of of No. species 6.2 6 5.8 5.6 1 2 3 4 5 6 7 8 9 101112131415161718192021222324 Months / Surveys
Figure 2.1 Species accumulation curve showing diversity of carnivore species in and
around Pir Lasura National Park.
Figure 2.2 Distribution of common leopard (Panthera pardus) in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan, as indicated by
various direct and indirect signs of the species
Figure 2.3 Kernel density estimation of common leopard (Panthera pardus) signs in
and around Pir Lasura National Park, AJ&K. Search radius was 2000 m
whereas, cell size used was 20 m. Red areas show sites having high relative
abundance of carnivore species signs.
Altitudinal range of its distribution was 721 m – 1690 m asl. Scats of the jackal (n =
64) were found and collected from 7 different sites which were Kothian (n = 8;
12.5%), Panagali (n = 6; 9.3%), Pir kana (n = 13; 20.3%), PLNP (n = 3; 4.7%), Pothi sairi (n = 15; 9.3%), Sairi (n = 15; 23.4%) and Supply (n = 13;20.3%).
Asiatic jackal was directly sighted at 11 study sites, while four road killings of the species were observed at four sites (Table 2.2 Sightings of the jackal were also reported at 19 sites in the study area. A high abundance of the jackal signs was recorded at Pir Kana, Sairi, and Katera, sampling sites (Table 2.1; Table 2.2; Fig.
2.4; Fig. 2.5).
2.4.2.3 Distribution of Kashmir hill fox
Direct and indirect signs of Kashmir hill fox (Vulpes Vulpes griffithi) were recorded from all 30 sites surveyed in the study area. Altitudinal range of the fox in and around Pir Lasura national park was recorded between 815m-1646 m asl. Scats of the fox (n = 92) were collected from 6 different study sites including; Kothian (n
= 5; 5.4%), Panagali (n = 14; 15.2%), Pir Kana (n = 16; 17.4%), Pothisairi (n = 8;
8.7%), Sairi (n = 10; 10.9%), and Supply (n = 39; 43.4%).
The fox was directly sighted at 23 different sites. Road killed foxes were recorded at two locations, whereas sightings of the fox were reported from 5 different sites. A high density of the fox signs was recorded at Supply sampling site (Table
2.1; Table 2.2; Fig. 2.6; Fig. 2.7).
2.4.2.4 Distribution of Asian palm civet
Asian palm civet was found distributed at all sampling sites (n=30) surveyed, with an altitudinal range of occurrence between 896 m – 1922m asl. Scats of Asian
Table 2.1 Direct and indirect signs of carnivores in and around Pir Lasura National
Park, Azad Jammu and Kashmir, Pakistan
Species Direct Scats Road Reported Total sightings killed sightings Common leopard 1 39 0 6 46 Asiatic jackal 19 64 4 21 108 Kashmir hill fox 28 92 2 5 127 Asian palm civet 6 108 1 21 136 Small Indian civet 0 44 2 2 48 Indian grey mongoose 18 57 0 0 75 Small Indian mongoose 31 69 4 11 115
Table 2. 2 Distribution of carnivore species in and around Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan, as shown by the
occurrence of any of the direct and indirect signs of the species. The + signs shows presence while – signs shows absence of the
species at particular site.
Asian Small Small Common Asiatic Red Grey Site name Elevation palm Indian Indian Geographical coordinates leopard Jackal Fox mongoose (m) civet civet mongoose 1 Sarda 33° 31.147'N, 73° 55.213'E 788 + + + + - + + 2 Chitibakri 33° 29.844'N, 73° 57.255'E 944 + + + + - - + 3 Shakyali 33° 28.453'N, 73° 57.168'E 740 + + + + - + + 4 Kothian 33° 29.583'N, 73° 58.067'E 1088 + + + + - - + 5 Phagwarmorah 33° 29.298'N, 73° 58.328'E 1111 + + + + - - + 6 Panagali 33° 29.249'N, 73° 59.105'E 1164 + + + + - - + 7 Qamrooti 33° 29.973'N, 74° 1.927'E 953 + + + + - - + 8 Supply 33° 28.957'N, 74° 1.810'E 1219 + + + + - + + 9 Pir kana 33° 28.578'N, 74° 4.055'E 1626 + + + + - - + 10 Nakyal 33° 29.177'N, 74° 6.224'E 1325 + + + + + + + 11 Sairi 33° 27.878'N, 74° 4.759'E 1802 + + + + + - + 12 GDC Nakyal 33° 29.166'N, 74° 5.305'E 1369 + + + + + + + 13 Pothi Sairi 33° 28.993'N, 74° 4.974'E 1523 + + + + + + + 14 Majhan 33° 27.200'N, 74° 7.371'E 1110 + + + + + + + 15 Karela 33° 25.803'N, 74° 6.659'E 1310 + + + + + + +
16 Katera 33° 28.318'N,74° 5.222'E 1368 + + + + + + + 17 Mathrani 33° 23.459'N, 74° 9.164'E 1141 + + + + - - + 18 Mendhatar 33° 25.992'N, 74° 3.937'E 998 + + + + - - + 19 Barmoch 33° 27.140'N, 74° 3.953'E 1292 + + + + - + + 20 Klinjar 33° 29.552'N, 74° 7.857'E 1699 + + + + - - + 21 Palani 33° 28.041'N, 74° 9.480'E 1412 + + + + - - + 22 Datote 33° 29.128'N, 74° 9.781'E 1582 + + + + - + + 23 Tarkundi 33° 25.851'N, 74° 9.793'E 1673 + + + + - - + 24 Jandroot 33° 32.004'N,74° 3.099'E 1285 + + + + - - + 25 Nerghal 33° 25.188'N, 74° 9.294'E 1409 + + + + - + + 26 Sairi Methrani 33° 22.978'N, 74° 9.462'E 1064 + + + + - + + 27 Kallar Galla 33° 24.433'N, 74° 8.902'E 1251 + + + + - + + 28 Banala 33° 30.578'N, 74° 5.436'E 1056 + + + + - + + 29 Pir Lasura 33° 28.982'N, 74° 3.804'E 1377 + + + + - - + 30 Gala 33° 29.438'N, 73° 58.316'E 1119 + + + + - - +
30 30 30 30 7 15 30
Figure 2.4 Map showing distribution of Asiatic jackal (Canis aureus) in and around
Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan, as evident
from various direct and indirect signs of the canid species.
Figure 2.5 Kernel density estimation of Asiatic jackal (Canis aureus) signs in and
around Pir Lasura National Park. Search radius was 2000 m whereas cell size
used was 20 m. Red areas show sites having high relative abundance of
carnivore species signs.
Figure 2.6 Map showing distribution of Kashmir hill fox (Vulpes vulpes griffithi) in
and around Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan,
as indicated by various direct and indirect signs of the species in the study
area.
Figure 2.7 Kernel density estimation of Kashmir hill fox (Vulpes vulpes griffithi)
signs in and around Pir Lasura National Park, AJ&K. Search radius was 2000
m whereas, cell size used was 20 m. Red areas show sites having high relative
abundance of carnivore signs.
palm civet were found and collected from six different sites which included; Kothian
(n = 8; 7.4%), Panagali (n = 34; 31.5%), Pir Lasura (PLNP) (n = 6; 5.5%), Pir Kana
(n = 17; 15.7%), Sairi (n=30; 27.8%), and Supply (n=13; 12%). Asian palm civet was directly sighted at 6 different locations, only one road killed Asian palm civet was found at Kothina site, the sightings of the species were reported from 21 sites.
While a high density of Asian palm civet signs was recorded at Sairi and Panagali sampling sites (Table 2.1; Table 2.2; Fig. 2.8; Fig. 2.9).
2.4.2.5 Distribution of small Indian civet
Distribution of small Indian civet was recorded at seven different sampling sites, with an altitudinal range between 1171m-1857m asl. Scats of small Indian civet were encountered at 4 different sites including; GDC Nakyal (n = 11; 25%), Katera
(n = 3; 6.8%), Pothi Sairi (n = 11; 25%) and Sairi (n = 19; 43.2%). Two road killed civets were found from GDC Nakyal and Nakyal sites, respectively. Only two filed sightings of the species were reported, one from Karela and the other from Majhan site. A high density of small Indian civet signs was recorded at GDC Nakyal and
Pothi Sairi sampling sites in the study area (Table 2.1; Table 2.2; Fig. 2.10; Fig.
2.11).
2.4.2.6 Distribution of Indian grey mongoose
Distribution of Indian grey mongoose was recorded at fifteen sampling sites with an elevation ranging from 699m – 1559 m a.s.l. Scats of the Indian grey mongoose (n = 57) were collected from five different study sites which included;
GDC Nakyal (n = 8; 14%), Katera (n = 14; 24.5%), Nakyal (n = 19; 33.3%), Pothi sairi (n = 5; 8.8%), and Supply (n = 11; 19.3%), direct field sightings of the species
Figure 2.8 Map showing distribution of Asian palm civet (Paradoxurus
hermaphroditus) in and around Pir Lasura National Park, Azad Jammu and
Kashmir, Pakistan, as indicated by various direct and indirect signs of the
species in the stud area.
Figure 2.9 Kernel density estimation of Asian palm civet (Paradoxurus
hermaphroditus) signs in and around Pir Lasura National Park, AJ&K.
Search radius was 2000 m whereas, cell size used was 20 m. Red areas show
sites having high relative abundance of carnivore signs.
Figure 2.10 Distribution of small Indian civet (Viverricula indica) in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan, as indicated by
its various direct and indirect signs in the study area.
Figure 2.11 Kernel density estimation of small Indian civet (Viverricula indica) signs
in and around Pir Lasura National Park, AJ&K, Pakistan. Search radius was
2000 m whereas, cell size used was 20 m. Red areas show sites having high
relative abundance of carnivore signs.
were recorded at 14 different sites. While a high sign density of Indian grey mongoose was found at Nakyal sampling site (Table 2.1; Table 2. 2; Fig. 2.12; Fig.
2.13).
2.4.2.7 Distribution of small Indian mongoose (Herpestes javanicus)
The small Indian mongoose was found distributed at 30 different sites in the study area, having an elevation range between 691 m- 1624 m asl. Scats of small
Indian mongoose were found and collected from 7 different sampling sites which included; Katera (n = 6; 8.7%), Kothian (n = 10; 14.5%), Panagali (n = 8; 11.6%),
Pir kana (n = 17; 24.6%), Pothi sairi (n = 4; 5.8%), Sairi (n = 6; 8.7%) and Supply
(n =18; 26%).
Small Indian mongoose was directly field sighted at 18 sites, road killed individuals of the small Indian mongoose were encountered at 4 sites, while indirect sightings of the mongoose species were reported at 11 different sites. A high density of small Indian mongoose signs was estimated at Pir Kana, Sairi, Katera and Supply sampling sites (Table 2.1; Table 2.2; Fig. 2.14; Fig. 2.15).
2.5 DISCUSSION
Zoogeography relates with distribution of animals on earth, and distribution is the phenomenon of dispersal of animals or plants over a geographical space.
Species are distributed on earth in different patterns. There are three basic patterns of animal distribution which includes; uniform distribution, random distribution and clumped distribution. The distribution of a species can be affected by many biotic and abiotic factors which include; food availability, vegetation characteristics, competition, power of animal dispersions, geographic barriers, climate and weather,
Figure 2.12 Distribution of Indian grey mongoose (Herpestes edwardsii) in and
around Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan, as is
shown by occurrence of its various direct and indirect signs in the study area.
Figure 2.13 Kernel density estimation of Indian grey mongoose (Herpestes
edwardsii) signs in and around Pir Lasura National Park, AJ&K. Search
radius was 2000 m whereas, cell size used was 20 m. Red areas show sites
having high relative abundance of carnivore signs.
Figure 2.14 Map showing distribution of small Indian mongoose (Herpestes
javanicus) in and around Pir Lasura National Park, Azad Jammu and
Kashmir, Pakistan, as indicated by occurrence of its various direct and
indirect signs in the study area.
Figure 2.15 Kernel density estimation of small Indian mongoose (Herpestes
javanicus) signs in and around Pir Lasura National Park, AJ&K. Search
radius was 2000 m whereas, cell size used was 20 m. Red areas show sites
having high relative abundance of carnivore signs.
and habitat quality. These factors and many others can determine distribution of a species and can limit distribution in a country or in a region (Sclater and Sclater,
1899; Roberts, 1997).
During current study, we recorded common leopard in all (100%) surveyed sites and direct or indirect signs of common leopard were recorded between 757 m-
1891 m. The Common leopard was found evenly distributed in the study area. This could be due to the fact that it can survive and adapt to variety of habitats. No previous published studies are available that report on the distribution of common leopard in the study area. Roberts (1997), however, described that four sub-species of Panthera pardus are known to occur in Pakistan including; firstly, Panthera pardus saxicolor (Pocock, 1927) found in Baluchistan (also found in Persia), secondly, Panthera pardus sindica (Pocock, 1930) occurring in Kirthar, Sindh, thirdly, Panthera pardus fusca found in whole of India and fourth sub species
Panthera pardus millardi (Pocock, 1930) is known to occur in state of Kashmir.
Siddiqui (1961) stated that all above four sub-species occur in Pakistan although at present it might be hard to distinguish individual specimens into sub-species while population from Sindh and Balochistan are so small as compared to northern
Himalayan population. Throughout its range, there is considerable variation in the pattern and density of rosettes or spots on the body of leopards since leopards in Swat and Hazara districts have longer and more luxuriant pelage during winter season.
Roberts (1997) described the distribution of leopards in Pakistan as it is confined to the forest of Himalayan region up to tree line or at lower altitudes in valleys which are more arid hilly regions in the north. It also occurs in hilly areas associated with
Acacia modesta and Acacia senegal scrub forests of Waziristan, Balochistan and
Sindh Kohistan. It was once inhabitant of the “Salt Range” and still survives in Kala
Chitta hills but is not found in human settlement areas, cultivated lands, riverain tracts for many decades. But during the current study, signs of leopard have been recorded near human habitations. This might be because of the reason that scarcity of wild prey in forests is now resulting in leopards’ interaction with human populations and it is known to predate on their livestock. It is also reported in Kirthar hills, Kalat and Mekran, Ziarat, Murree hills, Margallah Hills, Chitral, and the Chilas district of Gilgit-Baltistan.
The second sympatric carnivore, the Asiatic jackal was found distributed in all (100%) surveyed sites during the current study period. No previously published on distribution of jackal is available for comparison from the study area, however, there are few reports from the other parts of the country. Roberts (1997) described
Asiatic jackal as very adaptable animal and readily found in mountainous regions, forest plantations and riverine tracks. It is well adapted to dry, open habitat and avoids dense forests. During the current study, direct and indirect signs of the jackal were recorded near human settlements, cultivated lands, and in open and close forest areas. Roberts (1997) had reported Asiatic jackal occurring at 2150m elevation in
Hazara district, Shogran, and Murree hills and reported that it did not penetrate high mountain region. During the current study, we have recorded distribution of the jackal at elevation ranging from 721 m to 1690 m asl. However, signs of the jackal were not recorded at top of the mountains at elevation around 2000 m.
The red fox is among the least studied carnivores in Pakistan. During current study, we recorded direct and indirect signs of Kashmir hill fox (a sub-species of the
red fox that occurs in AJ&K) from all (100%) surveyed sites in and around Pir Lasura
National Park. Altitudinal range of its distribution was 815m-1646 m a. s. l. in the study area. No previous published studies are available for comparison form study area; however, few studies do report distribution of the species in Pakistan.
According to Roberts (1997) the species avoids dense forests and it can be found in open areas in the country. In Indus plain, it is reported to prefer extensive uncultivated tracts with sand dunes. During the present study, signs of the fox species were recorded in all habitat types surveyed which included; near human dwellings, cultivated lands, open forests as well as close forests. Roberts (1997) also, had reported that it occurs throughout the mountainous regions of Baluchistan, Khyber
Pakhtunkhwa and Himalayas, both in valleys and hilly mountainous regions. We also recorded signs of the fox in valleys as well as in the hilly mountains regions.
We recorded direct as well as indirect signs of Asian palm civet in all (100%) study sites. Initially, during surveys we found no information regarding distribution of this species in the study area, but then local people helped us identify scats of palm civet and based on their identification they confirmed that these are scats of yellow throated martin. However, after molecular identification of scats, it was confirmed to be originating from Asian palm civet and later we also found road killed specimens of this species in the study area as well. No yellow throated martin was found in the study area based on both the direct signs as well as indirect signs. Altitudinal range of Asian palm civet was recorded between 896 m – 1922m asl. Asian palm civet looks to be a wide spread species and its scats were encountered easily near human habitations on boundary walls of settlements, stones and hills. Scats of this species were usually encountered at rides of the mountains. The species was recorded at all
the sampling sites surveyed and often encountered by local people especially at night raiding fruits. Distribution of Asian palm civet/ common palm civet in Pakistan was previously described by Roberts (1997). According to Roberts, it occurs in extreme north-western boundary in Pakistan and is believed to have very local and restricted distribution in Pakistan. It has been recorded in Rawalpindi, Lahore, Sargodha,
Multan, Sahiwal, Bahawalpur, and Dera Ghazi Khan districts (Ahmad and Ghalib,
1979). It occurred in sub-Himalayan tracts, Mansehra in district Hazara at an elevation of 2670m and Rawalpindi district. It was also trapped from Margalla hills at Chak Jabri.
Distribution of small Indian civet was recorded at seven sampling sites (23%) in the study area during surveys, at an altitudinal range of 1171m-1857m asl. Since no previous studies have focused this species in Pir Lasura national park so far, therefore, current study results cannot be compared. However, according to Roberts
(1997) small Indian civet is best adapted species among all civets in sub-continent to terrestrial hunting and semidesert landscapes. Published literature reports that the small Indian civet occurs in variety of habitats in Indus basin such as, riverine jungles and sand-dune deserts. It is also found in irrigated forest plantations and avoids high human settlements and cultivated areas as well as mountainous regions. In Pakistan, it is decidedly uncommon and erratic in its distribution. In the current study, we recorded signs of small Indian civet from few sampling sites with no single direct sighting except recovery of road killed specimens. Signs of this species were also less common in the study area while its scats were found in dense or open areas having high understory vegetative cover, and areas having sparse human settlements surrounded by dense forests.
Distribution of Indian grey mongoose was recorded at fifteen different sampling sites (50%) in the study area, with an elevational range of 699m – 1559 m a.s.l. According to Roberts (1997), the grey mongoose is better adapted to arid conditions, it occurs in the Salt Range and is plentiful around Rawalpindi district.
The grey mongoose is reported to prefer more arid tracts and so the evidences of its high occurrence in natural vegetated area and in medium human activity areas, cultivated lands and avoidance to live near human dwellings is consistent with previous published literature such as by Roberts (1997); Santiapillai et al. (2000);
Francis (2008); Hussain et al. (2017). Roberts (1997) had reported that grey mongoose does not penetrate into the Murree foothills, however, in the current study, we recorded its distribution at 1559m elevation in Azad Kashmir which is almost the same elevation as that of Murree hills. Hussain et al. (2017) reported that in Potohar region mongooses are distributed at elevational range of 203m to 874m elevation.
Both Roberts (1997) and Hussain et al. (2017) reported that Indian grey mongoose avoids human settlement areas with high disturbance. However, in the current study, we recorded distribution of Indian grey mongoose in areas having human settlements surrounded by wild areas or cultivated land. Indian grey mongooses were found distributed in nullas having undisturbed semi-wild habitat near dense human habituations. They were often sighted in or near human settlements predating on domestic chickens. During present study, local community reported, and the author/researcher also sighted that Indian grey mongooses enter in their homes through sewerage pipelines. However, in highly populated areas having no undisturbed habitats around, we did not record direct or indirect signs of the Indian grey mongoose.
Distribution of small Indian mongoose was recorded at all 30 (100%) sampling sites surveyed in the current study. It was found distributed at elevational range of 691 m- 1624 m asl. Earlier on, Roberts (1997) had reported small Indian mongoose as one of the commonest small carnivore in southern Sind and North-
Eastern Punjab, having its distribution around Jhelum, Gujranwala and the Salt
Range. The species is well adapted to live in rocky areas with stunted thorn scrub typical of the Salt range. During the present study, this species was recorded in all study sites even in dense human populations. Distribution of the small Indian mongoose within or near the high human habitation and medium human activity areas or in cultivated lands had also been reported earlier (Roberts, 1997; Mahmood et al., 2011; Hussain et al., 2017), where they recorded the occurrence of the small
Indian mongoose near human habitations, near poultry farms and in cultivated lands.
Roberts (1997) recorded that the small Indian mongoose is well adapted to live in the outskirts of villages and towns.
This study has reported high sign density of carnivore species outside of protected area. The sites having high sign density outside the protected area should be included in Pir Lasura national park. This study provides baseline data on diversity and distribution of carnivore species and it will help in conservation and management of carnivores in the study area. The results of current study showed that seven species of carnivores occur at Pir Lasura National Park, and distribution range of seven species overlaps in the study area.
It is interesting to compare that some sampling sites support higher numbers of carnivore species than other sites. For example, Nakyal sampling site, GDC
nakyal, Pothi Sairi, Majhan, Karela, and Katera, support maximum numbers of carnivores occurring (n = 7). This could be due to the fact that these sites are rich in resources, and have high carrying capacity, and so these sites harbor large numbers of carnivores in comparison to some other sites which are poor in resources and low in carrying capacity, so they can support only fewer species.
2.6 CHAPTER SUMMARY
The current study aimed at documenting the diversity and distribution of carnivores at Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan. The study recorded seven different species of carnivores in the Park including; common leopard (Panthera pardus), Asiatic jackal (Canis aureus), Kashmir hill fox (Vulpes vulpes griffithi), Asian palm civet (Paradoxurus hermaphroditus), small Indian civet
(Vivercula indica), Indian grey mongoose (Herpestes edwardsii), and the small
Indian mongoose (Herpestes javanicus). Direct and indirect signs of common leopard, Asiatic jackal, Kashmir hill fox, Asian palm civet and small Indian mongoose were recorded in all 30 sampling sites having different elevational range.
Indian grey mongoose was found at 15 different sampling sites while signs of small
Indian civet were recorded at 7 sites. The current study concludes that seven species of carnivores occur at Pir Lasura National Park, and their distribution range overlaps.
Chapter 3
DIET COMPOSITION OF SYMPATRIC CARNIVORE
SPECIES AT PIR LASURA NATIONAL PARK
3.1 INTRODUCTION
Mammalian scats have been commonly used in biological studies to estimate population size (Kohn et al., 1999; Webbon et al., 2004), distribution patterns or species richness (Dalén et al., 2004), as they are abundant and easily found (Sanz et al., 2007). In many cases, it is assumed that scats are correctly identified, but it is difficult using scat morphology alone (Davison et al., 2002; Prugh and Ritland,
2005). It becomes more difficult when sympatric species have similar body features, behavior and feeding habits, and so the visual identification of scats becomes error prone (Ruiz-González et al., 2008).
Knowledge of a predator’s diet is vital to understand its ecology and to predicting its effect on the dynamics of prey populations (Oli, 1993). The diets of red foxes and dogs have been studied in Australia (Triggs et al., 1984; Brown and
Triggs, 1990; Lunney et al., 1990; Glen, 2005) and both are opportunistic predators, and diet comprised of mammals. Although, the diets of both species can vary in different habitats and seasons (Mitchell and Banks, 2005).
Faecal components of carnivores can comprise of feathers, bones, hairs, teeth, claws, scales, arthropod chitin, plant matter, mucus cells, and bacteria (Bang and Dahlström, 1975; Bujne, 2000). Whereas, the quantity and size of carnivore scats can be different based on age of individuals, prey species consumed and absorption capacity (Bang and Dahlström, 1975). Scats can be used for animal identification
56
(Seton, 1925; Camardella et al., 2000), activity centers of animal (Walker, 1996), composition of diet (Chinchilla, 1997; Santos and Hartz, 1999; Kauhala and Auniola,
2001), seasonal changes in diet (Aragona and Setz, 2001; Basuony et al., 2005;
Baker et al., 2006), prey species inventory (Camardella et al., 2000), role in seed dispersal (Fragoso and Huffman, 2000; Williams et al., 2000), animal health condition, and entero-parasitosis dynamics (Page et al., 2001).
Knowledge of diet composition of carnivores is important from conservation point of view. The Pir Lasura National Park in AJ&K is known for important carnivore species including common leopard, Asiatic jackal, civets, red fox and mongooses. Despite of widespread occurrence of carnivores across Azad Jammu and
Kashmir, Pakistan, only a few studies have documented morphological characteristics of carnivore scats, errors in morphological identification, diet composition and seasonal variation in diet of different carnivore species. To improve our understanding of vital ecological parameters we, in the current study, investigated morphological characteristics of carnivore scats, errors in morphological identification, diet composition and seasonal variation in diet of different carnivore species in human-dominated landscape in and around Pir Lasura
National Park, Azad Jammu and Kashmir, Pakistan.
3.2 REVIEW OF LITERATURE
Kabir (2011) investigated diet composition of common leopard at Machiara
National Park AJ&K. Twenty-three prey species were identified, including seven species of large mammals, eleven meso mammals and five small mammals. Leopard preyed mainly on domestic livestock (40.43%), meso mammals (29.75%), small
mammals (18.97%), and wild ungulates (5.13%).
The diet composition of Asiatic jackal (Canis aureus) in the Margalla Hills
National Park, Islamabad revealed, that diet of jackal comprised of 27 dietary items.
The diet of jackal comprised of rodents, mongooses, and domestic hen, wild boar and livestock. Grasses, and fruits were also consumed. Prey species richness was high in summer and least in autumn. Diet diversity was highest during summer season and it was low during autumn season. The prey species evenness (E) was high in winter and low in summer season (Mahmood et al., 2013). A similar study by
Nadeem et al. (2012) on Asiatic Jackal from Potohar Plateau, Pakistan showed that diet of jackal comprised of rodents, livestock, fruits grains and grasses. Consumption of mammalian bio mass was higher.
Shabbir et al. (2013) investigated diet composition and overlap of Wolf and
Jackal in Pakistan. Wolf and jackal consumed 14 and 13 prey species respectively.
Both predators had 11 species common in their diet. Dominant species in diet of wolf were sheep, palm civet and golden marmot. The diet of jackal was dominated by palm civet, golden marmot, cape hare and wood mouse. There was high nich overlap among two carnivores. Which showed that both carnivore species area competing for resources.
Food habits of small Indian mongoose were investigated by Mahmood et al.
(2011) in Potohar region. The major part of diet comprised of mammals followed by insects and plants. Siddiqui et al., (2004) investigated food habits of small Indian mongoose in Faisalabad, Pakistan. Diet comprised of mainly insects followed by birds, plants and mammals.
3.3 MATERIALS AND METHODS
Diet composition of different carnivore species occurring in the study area were investigated by analysis of their scat samples. We conducted surveys to collect scats of carnivore in Summer (May-July), Autumn (August-October), Winter
(November-January) and Spring (February-April) seasons during 2014-2016 using area searches technique. Three people participated in survey and only one (author) was responsible for identification of carnivore scats. When any scat was encountered, the field identification was determined based on its morphology including diameter, length, shape, color, odor, physical appearance such as characteristics contents (hairs, bones and plant material) (Seton, 1925; Jackson and
Hunter, 1995). Additional criteria included nature of scat deposit site, and presence of tracks or signs of activity of the species under study. The diameter at widest point, length, disjoint segments and weight of each scats sample was measured and samples were preserved in 95% ethanol for molecular identification and further analysis.
When scats consisted of disjoint segments total length was determined by summing up the length of each segment. All those scats which lacked typical structure and shape, for which measurement was not possible were excluded from the analysis.
3.3.1 Molecular Identification of Carnivore Scats
Results of the scat analysis of carnivores identified on morphological basis in the field may convey incorrect information about the carnivore species if not identified on molecular basis. Therefore, the collected scats of all carnivores in the study area were processed for molecular identification. This part or component of the current study was carried out at Non-invasive & Environmental DNA Lab
(NIEL) Conservation Genomics group (CGG) dedicated to DNA extractions in
University of Montana, Missoula, USA (Figure 3.1), while rest of the procedure for molecular identification of the carnivore species from scats was conducted at
Genomic core Laboratory, University of Montana, Missoula, USA, availing IRSIP
(International Research Support Initiative Program) program of the Higher
Education Commission (HEC) Islamabad, Pakistan.
Out of 551 scats of carnivores collected during field surveys, 149 were subjected to molecular identification and their results were later on used to correct morphologically identified scat samples, and hence 473 scats were confirmed to be originating from target species. We did not include scats of Indian grey mongoose and small Indian mongoose for molecular identification since these scats were different from other carnivores and all collected from outside of their respective burrows. We extracted fecal DNA in Non-invasive & Environmental DNA Lab
(NIEL) Conservation Genomics group (CGG) dedicated to DNA extractions in
University of Montana, Missoula, USA. We used QIAamp DNA Stool Mini Kits
(Qiagen, INC., Valencia, CA) for extraction of DNA from scats. We used negative control to keep track of cross contamination during extraction (Beja-Pereira et al.,
2009). The total volume of DNA extracts from each scat sample were 100 µL.
Initially, we used five different primers to validate them for identification of known carnivore species such as wolf, bobcat and martin. These primers included
12S/V5 100bp (Riaz et al., 2011), ATP6 172bp (Chaves et al., 2012), CytB (Chaves et al., 2012), CytB 146 bp (Farrell et al., 2000) and CytB 400 bp. Among these primers, 3 were selected based on their success to identify carnivore species which included 12S/V5_100bp, CytB_400bp, and CytB_Farrel. Then we tested these three primers for our 10 scats samples and one negative control to check for possible cross
contamination and one positive control which was known DNA of Wolf. The primer pair having highest success rate for species identification was selected for the study.
The PCR for all scats samples was carried out in a total volume of 50 µL.
The recipe of our master mix (MM) per sample was 20.375 µL H2O, 5 µL buffer (7
µL MgCl2, 0.375 µL BSA, 2 µL dNTP, 2.5 µL 12S/V5 primer F, 2.5 µL 12S/V5 primer R, 0.25 µL Taq polymerase and 10 µL DNA as extract template for each scat sample. The PCR conditions were denaturation at 95 °C for 5 min., then 40 cycles of PCR starting at 95 °C for 1 min., then annealing at 55 °C for 1 min. and elongation at 72 °C for 1:30 min. Then a final elongation at 72 °C for 5 min. at the end and 4
°C for infinity till product was removed from PCR. All PCRs were conducted on
Eppendrof vapo. protect Master cycler® pro and all reactions included a negative and positive control.
To clean PCR product, we used ExoSAP. We centrifuged ExoSAP at 500 rpm (revolution per minute) and placed it on ice, then transferred 5 µL of PCR product into a new plate and added 2 µL of ExoSAP in each sample, again centrifuged the plate and incubated in thermocycler. After cleaning PCR product, sequencing of all samples was carried out.
For each sample reaction, we used 0 2.25 µL 10 mM MgCl2, 3.0 µL Big Dye buffer (5X), 5.85 µL molecular grade H2O for each sample, 0.5 µL Big Dye (2X)
(should be added at end). We had 147 samples, so I multiplied each of these with
147 and then final volume of each reagent was mixed in falcon tube. For cycle sequencing we prepared plate by add 2.5 µL of PCR clean product in new plate (for positive samples 1 µL product was used). Then 1.1 µL primer was added to each
sample (1 either forward or reverse, not both). Then 11.5 µL master mix (MM) was added to each sample. Finally, 1 µL PGEM vector and its primer 1 µL m13 was added, also 11.5 µL MM was added in positive sample as well.
Then sample plate was centrifuged for 30 sec. at 4680 rpm and it was placed in thermocycler under following conditions; 96 °C for 5 min then 35 cycles at 96 °C for 10 sec, 50°C for 5 sec and 60°C for 2 min. At the end temperature was 4C for infinity till samples were removed from thermocycler.
For Cleaning of Sequenced product, we used Omega Bio Tek seq DTR Kit
(beads). Firstly, we brought bead mix to room temperature (30 min before use), centrifuged Sanger PCR plate. Then we added 11 µL bead mix to each sample, pipette up and down a couple of times to mix beads with DNA. Then added 50 µL
85% EtOH to each sample, pipette mix 15 times or until sample was homogenous.
We placed sample on magnetic rack and waited until solution was clear. Supernatant was pipette out and discarded. Then added 180 µL of 85% EtOH to each sample and removed and discarded EtOH. Again, added 180 µL of 85% EtOH to each sample, and then removed and discarded EtOH. Then placed a temporary seal on the PCR plate and spun it briefly in centrifuge up to 500 rpm. Then placed the plate back on magnetic rack. Using a 10 µL pipette tip removed excess EtOH and waited for one minute so that beads may get dry. Then removed plate from magnetic rack and added
50 µL of ddH2O to each sample and mix it with pipette (usually we add 30 µL H2O but for my samples we added 50 µL because we were getting high peaks in 3130.
Also, we should add 40 µL normally when PCR amplicon size is less than 200bp).
Waited for no less than 2 minutes, then placed plate back on magnetic rack, and waited until solution became clear. Then added supernatant to 3130 genetic analyzer
compatible semi-skirted plate (AB-1100) and added septa to the plate and spun that in centrifuge to ensure that there were no bubbles. Placed the plate in 3130 retainer, and it was made sure not to snap or jostle the plate in as this will cause the sample to splash up or bubbles to form. Then we added the white retaining lid, again it was made sure that it was secure. Then we run 3130 as per the 3130-run protocol. The
DNA sequences of all samples were blasted on NCBI blast.
3.3.2 Scat Analysis
Based on molecular identification performed, the collected scat samples were correctly assigned to each of the carnivore species, for diet analysis. The scat samples were sun dried. After drying scat samples morphological characteristics of scats such as length, breadth and weight were recorded (Figure 3.2).
For disintegration scats samples were soaked in warm water and scats were washed under tap water in a sieve to remove dust and mucus and to segregated different prey items such as hairs, bones, insects, bird feathers and plant parts
(Mahmood et al., 2013). These prey parts were dried and divided into different groups such as plant based diet, and animal based diet. The weight of each dietary item such including hairs, bones, feathers, insects and plant parts were recorded using electronic weighing balance in order to compute percent volume.
3.3.2.1 Whole mount preparation
We used hairs for identification of mammalian prey species. For this purpose, slides of the hairs of prey species were prepared. Hairs were washed in carbon tetrachloride 15-20 minutes). Long hairs were cut into small pieces and jumbled up hairs were separated. For whole mount preparation, we used transparent nail polish.
Scat collection
NCBI blast for species ID Preservation
3130 genetic analyzer to read DNA sequence DNA extraction
Sequencing
Gel electrophoreses
DNA amplification Figure 3.1 Steps in molecular identification of carnivore scats
Scat collection
Prey spp. identification whole mount and scale replication Reference material collection
Frequency & volume of prey items Segregation
Figure 3.2 Process of scat analysis to investigate diet composition of the carnivore
species.
Prey species of carnivores were identified using medullary pattern and cuticle cast pattern of the hairs recovered from scat samples as described by Moore et al., (1974).
Prepared slides were then compared with reference hair slides (n = >60 species) for identification. Similarly, bones, feathers of birds, invertebrates such as insects (with help of Entomologist) and plant matter including seeds were identified. The hairs were identified using Light microscope, at 10x, 40x and 100x magnification.
3.3.2.2 Scale replication
Cuticular scale patterns of mammalian hair were identified by slightly modifying procedure of Lavoie (1971). Two to three drops of transparent nail polish were placed and spread evenly on glass slide. A small hair was placed in vertical position along axis of slide so as one end of hair projected out of slide. After the nail polish was dry the end of hair projecting out was plucked with single attempt using forceps to get cast of hair on nail polish. The cast of hair prepared was exact duplicate of scales of the hair and was studied under microscope against reference for identification.
3.3.2.3 Identification of plant matter recovered
Plant matter recovered form scats of carnivore species mainly comprised of seeds and fruit parts. Recovered seeds and fruit remains were compared with reference material collected from field and were identified. Seeds were also sow for species identification.
3.3.3 Statistical Analysis
We computed misclassification rate based on field identification for each
carnivore species. We used the confusion matrices for each species to calculate True positive, False positives, true negative, false negative, accuracy, misidentification rate, false positive rate, specificity, precision and prevalence for each predator species. If a sample was identified in field as a leopard scat and confirmed to be a leopard with molecular identification the result was a true positive. If sample was identified in the field as a leopard scat and molecular identification determine it to be from dog or jackal or any other species, the outcome was false positive. But if a sample was identified as another species in field but identified as a leopard scat using molecular identification the outcome was false negative or if a scat was identified in field as another species (not leopard) and molecular identification determines that it is not a leopard scat then outcome was true negative.
Accuracy was calculated as the sum of true positives and true negatives divided by the sum of all possible outcomes (true positive, true negative, false positive, false negative). Thus, accuracy decreases with high false-positive or false- negative rates and increases with high true-positive and true negative rates.
True-positive rate was calculated as; true positive/ (true positive + false negative). True positive rate is a measure of how often did we identify a leopard scat correctly?
True-negative rate was determined as; true negative/ (true negative + false
positive). It is measure of how often did we correctly predict a scat was not a leopard scat? False-positive rate was computed as; false positive/ (true negative + false positive). It gave us measure of how often did we incorrectly identify scat from another species as leopard? False-negative rate was determined as; false negative/
(true positive + false negative). It is measure of how many times did we incorrectly
predict a scat was not a leopard scat?
The prey species diversity index (H'), prey richness (S) and prey evenness
(E) indices were calculated for each carnivore species during different seasons. Prey
Species Richness (S) was the total number of prey species consumed by each predator during each season. Diversity Index (H') was calculated by using the following formula:
H'= -Σ [pi × ln pi]
Where pi is prey index,
The Evenness Index (E) was calculated by using the formula:
E = H'/ln of S
Where, S represents the prey species richness and H' represents diversity index.
All data was analyzed using SPSS (version 23) software (SPSS Inc., Chicago,
USA) and Excel statistics.
We compared total frequency of dietary items consumed by each carnivore species for statistical differences. To compare seasonal variation in diet composition of each carnivore species we used general linear model (GLM). Similarly, we compared seasonal variation in consumption of wild prey species, domestic prey species, and plant matter. We repeated GLM for variation in consumption of each dietary item consumed by each carnivore species. All analysis was conducted in
(SPSS version 23).
3.4 RESULTS
3.4.1 Molecular Identification of Carnivore Scats
Molecular identification of carnivore scats was carried out in the University of Montana, USA. Out of 149 scats, we successfully extracted DNA from 144 scats with success rate of 96.6 %. Based on field identification of carnivore scats we had predicted that out of 149 scats only 11 (7.6%) scats were from common leopard, 38
(26.4%) from the red fox, 29 (20.1) from Asiatic jackal, 37 (25.7%) from yellow throated marten, 14 (9.7%) from Asian palm civet and 15(10.4%) from small Indian civet.
However, molecular identification process showed and confirmed that 11
(7.6%) scats were from Canis lupus Spp., 9 (6.24%) scats were from common leopard, 3 (2%) scats were from domestic dog, 40 (27.8 %) scats were from the red fox, 1 (0.7 %) scats were from porcupine, 21 (14.6%) scats were from Asiatic jackal,
45 (31.25%) scats were from Asian palm civet, 2 (1.4%) were from rhesus monkey and 12 (8.3%) scats were from small Indian civet (Table 3.1).
3.4.1.1 Accuracy and precision for molecular identification of carnivore scats
Accuracy for common leopard scats were 95.8%. Which showed that we correctly identified leopard scats. True positive rate was high (77.8%) and false negative rate was low (22.22%) which shows that morphological identification of leopard scats that was carried in the field was accurate one. Low false positive rate
(3%) and high true negative rate (97%) suggested the scats of leopard were not misclassified with other species in the field (Table 3.2).
For Kashmir hill fox scats, field identification accuracy was found to be
88.9% which suggested that scats of the fox species were correctly identified in the field. High true positive rate (77.5%) and low false negative rate (22.50%) also
showed correct identification of fox scats in the field. Low false positive rate (6.7%) and high true negative rate (93.26%) indicated that scats samples from other carnivore species were misclassified as those from fox species (Table 3.2).
Accuracy rate for field identification of Asiatic jackal scats was 90.3%. High accuracy showed that we often correctly identified the scats of Asiatic jackal in the field. High true positive rate (85.7%) and low false negative (14.28%) rate indicated that scats of Asiatic jackal in the field were correctly identified. Whereas, low false positive rate (8.9%) and high true negative rate (91.05%) suggested that scats of other predators were not misclassified as those from the Asiatic jackal (Table 3.2).
Field identification accuracy for yellow throated marten was (74.3%). True positive rate was (0%) which showed that scats of yellow throated marten were identified incorrectly. Similarly, false negative rate was 0% which showed that scats of yellow throated marten were always identified incorrectly, and yellow throated marten did not occur in study area (Table 3.2).
Field accuracy rate of Asian palm civet scats was 74.3%. Low true positive rate (24.4%) and high false negative rate (75.55%) suggested that scats of Asian palm civet were not correctly identified in the field. However, low false positive rate (3%) and high true negative rate (96.97%) showed that scats of other carnivore species were also not misclassified as those from Asian palm civet. The molecular identification confirmed that scats of Asian palm civet were always classified as those of yellow throated marten in the field (Table 3.2).
Accuracy rate for identification of small Indian civet scats was found to be
92.4%. High accuracy rate for small Indian civet showed that we often correctly
identified scats of small Indian civet in the field. True positive rate was 66.7% and false negative rate was 33.33% which showed that scats of this species were often misidentified. Low false positive rate (5.3%) and high true negative rate (94.69%) showed that scats of other carnivore species were misclassified as those from small
Indian civet (Table 3.2).
3.4.2 Morphological Characteristics of Carnivore Scats
The scats of seven different carnivore species collected from the field were measured for their morphological characteristics in the laboratory (Table 3.3). The scats of common leopard were found heaviest in weight (32.73 ± 2.3 g), followed by those of Asiatic Jackal (16.58 ± 1.0 g) while those of small Indian mongoose were found lightest (2.26 ± 0.05 g) in weight. Similarly scats of common leopard were found having maximum length (11.36 ± 0.57 cm) and diameter (2.63 ± 0.05 cm) whereas those of small Indian mongoose had minimum length (4.78 ± 0.1 cm) and diameter (0.69 ± 0.01 cm) among all seven carnivore scats (Table 3.3; Fig. 3.3 and
3.4).
3.4.3 Diet Composition of Carnivore Species
Diet of seven carnivore species was studied by analyzing a total of 473 scats samples collected from the field. Among those, were 39 from common leopard,
Asiatic jackal 64, Kashmir hill fox 92, Asian palm civet 108, small Indian civet 44,
Indian grey mongoose 57, and small Indian mongoose 69 scat samples.
3.4.3.1 Diet composition of common leopard
Among 39 collected scats of common leopard, 13 samples were collected during summer season, 11 during autumn season, 6 during winter season, and 9
Table 3.1 Confusion matrix of scats of carnivore species identification using morphological and molecular identification techniques.
Field identification
Panthera Vulpes Canis Martes Paradoxurus. Viverricula pardus vulpes aureus flavicola hermaphroditus indica Total
Genetic identification Canis lupus spp. 2 1 7 1 0 0 11 Common leopard (Panthera pardus) 7 0 2 0 0 0 9 Dog (Canis lupus familiaris) 2 0 0 0 1 0 3 Kashmir hill fox (Vulpes Vulpes griffithi) 0 31 0 2 0 7 40 Porcupine (Hystrix indica) 0 0 0 0 1 0 1 Asiatic Jackal (Canis aureus) 0 2 18 0 1 0 21 Asian palm civet (P. hermaphroditus) 0 0 0 34 11 0 45 Rhesus monkey (M. mulatta) 0 0 2 0 0 0 2 Small Indian civet (Vivercula indica) 0 4 0 0 0 8 12 Totals 11 38 29 37 14 15 144
Table 3.2 Error rates in species identification in the field corrected with molecular identification of carnivore scats.
Panthera Vulpes vulpes Canis Martes Paradoxurus Vivercula pardus griffithi aureus flavicola hermaphroditus indica
True positives 7 31 18 0 11 8 False positives 4 7 11 37 3 7 True Negatives 131 97 112 107 96 125 False negatives 2 9 3 0 34 4 Accuracy 95.8% 88.9% 90.3% 74.3% 74.3% 92.4% Misidentification rate 4.2% 11.1% 9.7% 25.7% 25.7% 7.6% True positive rate 77.8% 77.5% 85.7% 0% 24.4% 66.7% False positive rate 3.0% 6.7% 8.9% 25.7% 3.0% 5.3% True negative rate 97% 93.26% 91.05% 74.30% 96.97% 94.69% False negative rate 22.22% 22.50% 14.28% 0% 75.55% 33.33%
Table 3.3 Measurements of morphological characteristics of sympatric carnivore scats (Mean ± SEM) sampled from in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan
Carnivore Species n Diameter (cm) Length (cm) Disjoint segments Weight (g)
Common Leopard (Panthera pardus) 39 2.63 ± 0.05 11.36 ± 0.57 4.05 ± 0.24 32.73 ± 2.3
Asiatic Jackal (Canis aureus) 64 2.11 ± 0.03 8 ± 0.33 1.59 ± 0.08 16.58 ± 1
Kashmir hill fox (Vulpes vulpes griffithi) 92 1.29 ± .02 5.87 ± 0.25 1.23 ± 0.05 6.21 ± 0.39
Asian Palm Civet (Paradoxurus hermaphroditus) 108 1.4 ± 0.043 5.54 ± 0.2 1.11 ± 0.035 6.26 ± 0.38
Small Indian Civet (Viverricula indica) 44 1.16 ± 0.02 5.18 ± 0.36 1.16 ± 0.05 8.72 ± 0.42
Indian Grey Mongoose (Herpestes edwardsii) 57 1.13 ± 0.02 6.53 ± 0.19 1.07 ± 0.03 3.2 ± 0.11
Small Indian Mongoose (Herpestes javanicus) 69 0.69 ± 0.01 4.78 ± 0.1 1.05 ± 0.02 2.26 ± 0.05
Figure 3.3 Scats of sympatric carnivore species: A & B) Common leopard; C)
Asiatic jackal; D, E &F) Kashmir hill fox; G&H) Asian palm civet
Figure 3.4 Scats of sympatric carnivore species: A) Small Indian civet; B&C)
Indian grey mongoose; D) Small Indian mongoose
during spring season. Analysis of common leopard scats (Table 3.4) revealed that average percent volume (%V) of prey components recovered, consisted of hairs
(70.25 ± 1.9), feathers (1.43 ± 0.37), bones (4.19 ± 1.15), plants (1.09 ± 0.51), anthropogenic materials (0.41 ± 0.18) and sand/clay/unidentified (22.62 ± 2.23).
In general, diet of common leopard comprised of 17 prey species including mammals (n=14 species) and birds (03 species). Among 17 prey species, 10 species included wild prey (mammals 8; birds 2) and 7 species were domestic (6 mammals;
1 bird) animals. Frequency of occurrence of wild prey was 34.85 % of total leopard diet whereas domestic prey contributed 59.1% of leopard diet. The anthropogenic items contributed 6.06%.
Among wild prey species, frequency of occurrence of Rhesus monkey was highest (10.61%) while among domestic prey, frequency of occurrence of goat was found highest (28.79%) Comparison of dietary items using GLM showed that consumption of different diet items significantly differed F=9.26, df=17, p=0.000.
The model explained 74.5.3% (R squared =0.745) of variation in diet of leopard
(Appendix I) (Table 3.5) (Figure 3.5, 3.6, 3.7, 3.8).
To document seasonal variation in diet of common leopard, we analyzed 13 scats of leopard during summer season, 11 during autumn, 6 during winter, and 9 during spring season. Frequency of occurrence of wild prey was high during spring season (40%) and low during winter season (27.27%). Frequency of occurrence of domestic prey species was high during winter season (72.73%) whereas, consumption of domestic prey was low during Autumn season (52.63%). GLM showed that there is no statistical significant difference in seasonal diet of common leopard F=0.904, df=3, p=0.443.
Table 3.4 Percent volume of prey items recovered from the scats of common leopard
Sr. Prey Summer Autumn Winter Spring Total Mean no. species/items (n=13) (n=11) (n=6) (n=9) (n=39) ± SE recovered 1 Hairs 83.46 75.59 69.41 69.48 281.03 70.25±1.9 2 Feathers 2.39 1.61 1.1 0.62 5.72 1.43±0.37 3 Bones 4.11 2.59 2.59 7.47 16.76 4.19±1.15 5 Plants 0.69 1.24 0 2.43 4.36 1.09±0.51 7 Anthropogenic 0.28 0.84 0 0.51 1.63 0.41±0.18 8 Sand/clay/ 25.98 18.13 26.9 19.49 90.5 22.62±2.23 unidentified 100 100 100 100 400
Table 3.5 Percent frequency of occurrence of prey items in the scats of common
leopard (Panthera pardus) collected from the PLNP, AJ&K.
Summer Autumn Winter Spring Prey species Total (n=13) (n=11) (n=6) (n=9) Wild prey Barking deer (Muntiacus 4.76 5.26 0 0 3.03 muntjac) Kashmir hill fox (Vulpes 4.76 0 0 0 1.52 Vulpes griffithi) Rhesus monkey (Macaca 4.76 10.53 9.09 20 10.61 mulatta) Indian Gerbil (Tetra 4.76 0 0 0 1.52 indica) Wild boar (Sus scrofa) 0 5.26 9.09 0 3.03 Indian crested porcupine 0 5.26 9.09 0 3.03 (Hystrix indica) Asian palm civet (Paradoxurus 4.76 0 0 6.67 3.03 hermaphroditus) Desert hare (Lepus 0 0 0 6.67 1.52 nigricollis dayanus) Kalij pheasant (Lophura 4.76 5.26 0 6.67 4.55 leucomelanos) Indian/common peafowl 4.76 5.26 0 0 3.03 (Pavo cristatus) Total wild prey 33.33 36.84 27.27 40 34.85
Domestic prey Goat (Capra hircus) 28.57 31.58 27.27 26.67 28.79 Dog (Canis lupus 19.05 10.53 18.18 13.33 15.15 familiaris) Sheep (Ovis aries) 4.76 0 0 0 1.52 Cow (Bos taurus) 4.76 0 18.18 6.67 6.06 Buffalo (Bubalus 4.76 10.53 0 0 4.55 bubalis) Horse (Equus ferus 0 0 0 6.67 1.52 caballus) Poultry (Gallus gallus 0 0 9.09 0 1.52 domesticus) Total domestic prey 61.9 52.63 72.73 53.33 59.1 Anthropogenic 4.76 10.53 0 6.67 6.06
3.4.3.2 Diet of Asiatic jackal
Mean percent volume of hairs recovered from scats of Asiatic jackal during all seasons was found to be 20.34±3.14, birds 34.44±3.95, bones 33.8±7.54, insects
0.04±0.04, plants 0.2±0.14, grits 0.16±0.07, anthropogenic 0.34±0.11, and sand/clay/ unidentified 10.7±2.75. During summer season percent volume of bones was high 46.52 %, followed by birds 29.10%, hairs 13.02 and lowest was of grits
0.04%. During autumn season percent volume of bones was high 47.02% followed by birds 28.87%, hairs 19.85% and lowest was of grits 0.26%. Percent volume of birds was high during winter season 34.04%, followed by hairs 28.37%, bones
22.85% and lowest was of grits 0.03%. During spring season, highest percent volume was shared by birds 45.75%, followed by hairs 20.12% and then bones 18.75% and low was of grits 0.31% (Table 3.6).
Sixteen species of mammals, birds, insects and plants were identified from diet of Asiatic jackal. Among these species 10 were wild, 5 were domestic and 1 species of plant. Frequency of wild prey in diet of jackal was 18.48% and domestic was 59.78%. Among wild prey frequency of occurrence of rhesus monkey was high
5.43% followed by Norway rat 3.26%. Among domestic species consumption of poultry was high 27.17% followed by goat 20.65%. Consumption of plant matter was (3.26%) (Table 3.7) (Figure 3.6, 3.7, 3.8, 3.9). General linear model fitted explained 82.6% of variation in prey item consumption of Asiatic jackal. The consumption of dietary items varied significantly F=15.02, df=17, p=0.000
(Appendix II).
Among 64 collected scats of Asiatic jackal 17 were collected during summer
35
30
25
20
15
10
5 Frequency of prey species species prey of Frequency
0
Goat Indian/commonpeafowl Dog Rhesus monkey Cow Anthropogenic pheasantKalij Buffalo Barking deer boarWild Indian crested porcupine palm Asian civet Kashmirfoxhill Indian Gerbil Desert hare Sheep Horse Poultry
Figure 3.5 Variation in frequency of prey species in the diet of common leopard in
PLNP, AJ&K.
Figure 3.6 A) Whole mount of Indian gerbil a) reference slide of Indian gerbil; B)
Whole mount of Turkistan rat, c) reference slide of Turkistan rat; C) House
mouse, c) reference slide of house mouse
Figure 3.7 A) Whole mount of house rat, b) reference slide of house rat; B) Norway
rat, b) reference slide of Norway rat; C) Desert hare, c) reference slide of
desert hare
Figure 3.8 A) Rhesus monkey a) reference rhesus monkey; B) Barking deer b)
reference barking deer; C) Goat, c) reference goat; D) wild boar, d) reference
wild boar
season, 23 were collected during autumn season, 14 were collected during winter season and 10 were collected during spring season. Consumption of wild prey species was higher during Autumn season (21.88%) and low during spring season
(12.5%). Similarly, consumption of domestic prey was high during winter season
(66.67%) and low during autumn and spring season (56.25% each). GLM showed that there is no statistically significant difference in seasonal diet consumption of
Asiatic jackal during the current study period F=0.946, df=3, p=0.423. Similarly, consumption of wild prey species did not differ significantly during four seasons
F=1.082, df=3, p=0.371. Consumption of domestic prey species did not differ during different seasons F=0.346, df=3, p=0.792.
3.4.3.3 Diet composition of Kashmir hill fox
Scats of red fox were segregated into different categories and percent volume of each category was computed. The percent volume of birds was high 53.34±9.35 followed by plants 22.23±9.7, hairs 16.4±3.73 and lowest was anthropogenic material 0.21±0.16. During summer season percent volume of plants was high
42.41% followed by birds 38.67%, hairs 6.71%, bones 6.69%, and lowest were grits
0.17%. During autumn season percent volume of birds was high 37.30% followed by plants 34.49%, hairs 21.1% and lowest were insects 0.07%. During winter season, highest percent volume was contributed by birds 75.97%, followed by hairs 14.5%, bones 5.56% and lowest were plants and grits 1.23 each. During spring season percent volume of birds was high 61.43% followed by hairs 23.3%, plants 10.78% and lowest were grits 0.08% (Table 3.8).
To document diet composition of Kashmir hill fox 92 scats samples were
Table 3.6 Percent volume of prey items recovered from the scats of Asiatic jackal
Prey Summer Autumn Winter Spring Total Mean ± SE species/items (n=17) (n=23) (n=14) (n=10) (n=64) Hairs 13.02 19.85 28.37 20.12 81.36 20.34±3.14 Feathers 29.10 28.87 34.04 45.75 137.76 34.44±3.95 Bones 46.52 47.02 22.85 18.75 135.14 33.8±7.54 Insects 0 0 0.14 0 0.14 0.04±0.04 Plants 0.18 0.6 0 0 0.78 0.2±0.14 Grits 0.04 0.26 0.04 0.31 0.65 0.16±0.07 Anthropogenic 0.54 0.46 0.03 0.33 1.36 0.34±0.11 Sand/clay 10.6 2.94 14.53 14.74 42.81 10.7±2.75 100 100 100 100 400
Table 3.7 Percent frequency of occurrence of prey items in diet of Asiatic jackal
Summer Autumn Winter Spring Prey Species Total (n=17) (n=23) (n=14) (n=10) Wild prey Rhesus monkey (Macaca 4.35 9.38 0 6.25 5.43 mulatta) Wild boar (Sus scrofa) 4.35 3.13 0 0 2.17 Desert hare (Lepus 0 3.13 0 0 1.09 nigricollis dayanus) Indian Gerbil (Tetra indica) 4.35 0 0 0 1.09 Roof or house rat (Rattus 0 3.13 0 0 1.09 rattus) House mouse (Mus 0 0 4.76 0 1.09 musculus) Norway rat (Rattus 0 0 9.52 6.25 3.26 norvegicus) Kalij pheasant (Lophura 0 3.13 0 0 1.09 leucomelanos) Spotted dove (Streptopelia 4.35 0 0 0 1.09 chinensis) Insects (Orthoptera) 0 0 4.76 0 1.09 Grasshopper Total wild prey 17.39 21.88 19.05 12.5 18.48 Domestic prey Goat (Capra hircus) 21.74 21.88 23.81 12.5 20.65 Sheep (Ovis aries) 4.35 0 4.76 6.25 3.26 Cow (Bos taurus) 8.7 9.38 9.52 0 7.61 Buffalo (Bubalus bubalis) 4.35 0 0 0 1.09
Poultry (Gallus gallus 21.74 25 28.57 37.5 27.17 domesticus) Total domestic prey 60.87 56.25 66.67 56.25 59.78 Jaro grass (Thameda 4.35 6.25 0 0 3.26 anathera) Grits 4.35 3.13 4.76 6.25 4.35 Anthropogenic 13.04 12.5 9.52 25 14.13
30
25
20
15
Frequency of prey species prey of Frequency 10
5
0
Figure 3.9 Variation in frequency of prey species in diet of Asiatic jackal.
analyzed. We recorded 21 species of mammals, birds, plants and insects in diet of red fox. Among these 21 species 10 were wild, 5 domestic and 6 plant species.
Frequency of occurrence of wild prey was 17.96%, domestic prey 50.9%, plants
27.54%. Among wild prey species frequency of occurrence of wild boar was high
2.99% followed by house mouse 2.4%. Among domestic prey frequency of occurrence of poultry was high 29.34% followed by goat 13.77%. (Table 3.9) (Figure
3.6, 3.7, 3.8, 3.10). General linear model explained 73.9% variation in consumption of different dietary items by Kashmir hill fox (R squared =0.739). The consumption of different dietary items varied significantly in diet of Kashmir hill fox (F=8.86, df=22, p=0.000) (Appendix III).
To document seasonal variation in diet composition of Kashmir hill fox we analyzed 27 scats in summer, 31 in autumn, 19 in winter and 15 in spring season.
GLM showed that diet of fox did not differ across different seasons F=1.14, df=3, p=0.337. Consumption of wild prey species did not differ across all four seasons
F=1.025, df=3, p=0.395. Consumption of domestic prey species did not among all four seasons F=0.201, df=3, p=0.894.
Consumption of wild prey species was high during spring season (25%) and low during summer season (9.43%). Consumption of domestic prey species was high during winter season 66.67% and low during autumn season (40.35%). Consumption of plant species was high during summer season (35.85%) and low during winter season (9.09%).
3.4.3.4 Diet composition of Asian palm civet
Diet of Asian palm civet mainly comprised of plant matter by its volume.
Table 3.8 Percent volume of prey items recovered from the scats of Kashmir hill fox
Prey Sr. Summer Autumn Winter Spring Total Mean ± SE species/items no. (n=27) (n=31) (n=19) (n=15) (n=92) recovered 1 Hairs 6.71 21.1 14.5 23.3 65.61 16.4±3.73 2 Feathers 38.67 37.30 75.97 61.43 213.37 53.34±9.35 3 Bones 6.69 1.2 5.56 4.25 17.7 4.43±1.18 4 Insects 1.35 0.07 0 0.16 1.58 0.4±0.32 5 Plants 42.41 34.49 1.23 10.78 88.91 22.23±9.7 6 Grits 0.17 0.07 1.23 0 1.47 0.37±0.29 7 Anthropogenic 0.66 0.17 0 0 0.83 0.21±0.16 Sand/clay/ 8 3.34 5.6 1.51 0.08 10.53 2.63±1.19 unidentified 100 100 100 100 400
Table 3.9 Percent frequency of occurrence of prey items in diet of Kashmir hill fox
Prey species/items Summer Autumn Winter Spring Total recovered (n=27) (n=31) (n=19) (n=15) Wild prey Desert hare (Lepus nigricollis dayanus) 3.77 0 0 0 1.2 Wild boar (Sus scrofa) 0 5.26 3.03 4.17 2.99 Rhesus monkey (Macaca mulatta) 0 3.51 0 0 1.2 Turkistan rat (Rattus pyctoris) 0 1.75 3.03 0 1.2 Indian Gerbil (Tetra indica) 0 0 6.06 0 1.2 House mouse (Mus musculus) 0 0 6.06 8.33 2.4 Roof or house rat (Rattus rattus) 0 0 0 4.17 0.6 Red-vented bulbul (Pycnonotus cafer) 0 1.75 6.06 0 1.8 Spotted dove (Streptopelia chinensis) 1.89 1.75 0 0 1.2 Insects Hymanoptera (ants bees) 3.77 5.26 0 8.33 4.2 Total wild prey 9.43 19.3 24.24 25 17.96 Domestic prey Goat (Capra hircus) 13.21 14.04 15.15 12.5 13.77
Sheep (Ovis aries) 3.77 7.018 9.09 0 5.39 Cow (Bos taurus) 3.77 0 0 4.17 1.8 Buffalo (Bubalus bubalis) 1.89 0 0 0 0.6 Poultry (Gallus gallus domesticus) 26.42 19.3 42.42 41.67 29.34 Total domestic prey 49.06 40.35 66.67 58.33 50.9 Plant species Wild Himalayan pear (Dhandali) (Pyrus pashia) 22.64 21.05 0 0 14.37 Jaru grass (Themeda anathera) 5.66 3.51 3.03 4.17 4.19 Kov (Olea ferruginea) 5.66 7.02 6.06 0 5.39 Wheat (Triticum aestivum) 0 1.75 0 0 0.6 Dhania (Coriandrum sativum) 0 1.75 0 0 0.6 Choti berry (Ziziphus oxyphylla) 1.89 0 0 12.5 2.4 Total plant species 35.85 35.09 9.09 16.67 27.54 Grits 3.77 1.75 0 0 1.8 Anthropogenic 1.89 3.51 0 0 1.8
35
30
25
20
15
10 Frequency Frequency prey of species
5
0
Hymanoptera Wildboar Poultry HimalayanpearWild Goat Sheep Kov grass Jaru House mouse Choti berry Red-vented bulbul Cow Grits Anthropogenic Desert hare Rhesus monkey Turkistanrat Indian Gerbil Spotteddove orRoof rathouse Buffalo Wheat Corainder
Figure 3.10 Variation in frequency of prey species in diet of Kashmir hill fox
Percent volume of plant species was 85.67±2.32 of all 108 scats analyzed for diet composition study, followed by birds 5.84, Sand/clay/ unidentified 3.99±1.93, hairs 3.05±1.27 and lowest was anthropogenic material 0.01±0.01. During summer season percent volume of plants was high 83.52%, followed by birds 7.78%,
Sand/clay/ unidentified 7% and hairs 1.34%, bones 0.26%, snail 0.08%, grits 0.025% and lowest were insects 0.006%. During autumn season percent volume of plant matter was 81.1 %, followed by Sand/clay/ unidentified 7.67%, birds 7.35%, bones
3%, snails 0.46%, hairs 0.40% and lowest were insect 0.015%. During winter season percent volume of plants was 91.93% followed by hairs 5.15%, insects 1.31 %,
Sand/clay/ unidentified 0.82%, snails 0.33%, birds 0.23%, grits 0.17% and lowest was anthropogenic material 0.06%. During spring season percent volume of plant matter was 86.16% followed by birds 8.03%, hairs 5.32% and lowest was Sand/clay/ unidentified 0.49%.
Analysis of 108 scats samples of Asian palm civet showed that 27 species of mammals, birds, invertebrates and plants were consumed. Among all recorded species 9 were wild, 3 domestic, and 15 plant species. Consumption of wild prey was
33.68% compared to 10.88% domestic prey species and 53.37% plants. Among wild prey consumption of Indian gerbil was high 9.84% and among domestic prey consumption of poultry was high 6.74%. Among plant species frequency of occurrence of Himalayan wild pear was high (27.46%) (Table 3.11) (Figure 3.6, 3.7,
3.8, 3.11). General linear model fitted explained 59.1% variation in dietary components of Asian palm civet. The model showed that consumption of different dietary components of Asian palm civet differed significantly (F= 4.48, df=28, p=0.000) (Appendix IV).
Seasonal variation in diet composition of Asian palm civet was assessed by performing scats analysis of 30 samples collected during summer, 45 during autumn,
21 during winter and 12 during spring season. Consumption of wild prey was high during winter season (44.9%) and low during autumn season. Consumption of domestic prey was high during autumn season (15.79%) and low during winter season (4.08%). Consumption of plant matter was high during autumn season
(56.58%) and low during winter season (44.9%). According GLM diet of Asian palm civet did not differ statistically between four seasons of the year F=1.969, df=3, p=0.122. Consumption of wild prey species was also not statistically different during different seasons F=1.11, df=3, p=0.369. Similarly, consumption of domestic prey did not differ significantly during different seasons F=2.2, df=3, p=0.16.
Consumption of plant species in diet of Asian palm civet did not differ significantly during different seasons F=0.699, df=3, p=0.557.
3.4.3.5 Diet composition of small Indian civet
Analysis of 44 scats showed that mean percent volume of hairs in scats of small Indian civet was 2.7±2.47%, birds 24.37±4.51%, bones 2.73±1.21%, amphibians 0.47±0.28%, insects 0.005±0.005%, plants 69.54±7.77%, grits
0.05±0.05%, anthropogenic material 0.02±0.02% and sand/clay/unidentified
0.123±-0.07%. During summer season mean percent volume of plant species was high 76.87% followed by birds 19.04% whereas, mean percent volume of anthropogenic items was low 0.09%. During autumn season mean percent volume of plants was high 77.71% followed by birds 21.11% and low was Sand/clay/ unidentified 0.09%. During winter season percent volume of plants was high 46.23% followed by birds 7.85% and lowest was Sand/clay/ unidentified 0.02%. During
Table 3.10 Percent volume of prey items recovered from the scats of Asian Palm
civet
Prey Summer Autumn Winter Spring Total Mean ± SE species/items (n=30) (n=45) (n=21) (n=12) (n=108) recovered Hairs 1.34 0.40 5.15 5.32 12.21 3.05±1.27 Feathers 7.78 7.35 0.23 8.03 23.39 5.84±1.87 Bones 0.26 3 0 0 3.26 0.81±0.73 Snails 0.08 0.46 0.33 0 0.87 0.22±0.1 Insects 0.006 0.015 1.31 0 1.33 0.33±0.32 Plants 83.52 81.1 91.93 86.16 342.71 85.67±2.32 Grits 0.025 0 0.17 0 0.19 0.04±0.04 Anthropogenic 0 0 0.06 0 0.06 0.01±0.01 Sand/clay/ 7 7.67 0.82 0.49 15.96 3.99±1.93 unidentified 100 100 100 100 400
Table 3.11 Percent frequency of occurrence of prey items in diet of Asian palm civet
Prey species/items Summer Autumn Winter Spring Total recovered (n=30) (n=45) (n=21) (n=12) Wild prey House mouse (Mus musculus) 4.17 1.32 4.08 5 3.1 Indian Gerbil (Tetra indica) 10.42 5.26 16.33 10 9.84 Roof or house rat (Rattus rattus) 0 2.63 2.04 5 2.07 Norway rat (Rattus norvegicus) 4.17 2.63 4.08 5 3.63 Rhesus monkey (Macaca mulatta) 4.17 5.26 2.04 5 4.15 Desert hare (Lepus nigricollis dayanus) 2.08 3.95 2.04 0 2.59 Snail (Cornu sp.) 6.25 5.26 0 0 3.63 Insects (Orthoptera) Grasshopper 2.08 1.32 10.2 0 3.63 Insects (Coleoptera) (beetle) 0 0 4.08 0 1.04 Total wild prey 33.33 27.63 44.9 30 33.68 Domestic prey Cow (Bos taurus) 2.08 6.58 2.04 0 3.63 Sheep (Ovis aries) 0 1.32 0 0 0.52
Poultry (Gallus gallus domesticus) 6.25 7.89 2.04 15 6.74 Total domestic prey 8.33 15.79 4.08 15 10.88 Plant species Bara bair (Ziziphus jujube) 4.167 0 2.04 5 2.07 Aakharay (Rubus fruticosus) 10.42 1.32 0 25 5.7 Wild Himalayan pear (Dhandali) (Pyrus pashia) 31.25 31.58 28.57 0 27.46 Choti bairi (Ziziphus oxyphylla) 6.25 6.58 0 10 5.18 Kov (Olea ferruginea) 0 6.58 2.04 0 3.1 Kumlo (Lannea coromandelica) 0 1.32 0 0 0.52 Jaru grass (Themeda anathera) 0 2.63 0 0 1.04 Apple (Pyrus malus) 2.08 0 0 5 1.04 Orange (Citrus reticulate) 0 1.32 0 0 0.52 Khobani (Prunus amrmeniaca) 0 0 0 10 1.04 Maluk (Diospyros lotus) 0 5.26 0 0 2.07 Wild fig (Ficus carica) 0 0 4.08 0 1.04 Water melon (Citrullus lanatus) 0 0 4.08 0 1.04 Dhania (Coriandrum sativum) 2.08 0 2.04 0 1.04 Wheat (Triticum aestivum) 0 0 2.04 0 0.52 Total plant species 56.25 56.58 44.9 55 53.37 Grits 2.08 0 4.08 0 1.55 Anthropogenic 0 0 2.04 0 0.52
30
25
20
15
10 Frequency Frequency prey of species
5
0
Kov
Grits
Cow
Snail
Apple Sheep
Maluk Wheat
Kumlo
Apricot
Poultry Orange
Wild Wild fig
Aakharay
Bara bair Bara
Coriander
Jaru Jaru grass
Orthoptera
Coleoptera
Norway rat Norway
Choti bairi Choti
Desert hare Desert
Water melon Water
Indian Gerbil Indian
House mouse House
Anthropogenic
Rhesus monkey Rhesus Wild Wild Himalayan Roof or rat house Roof or
Figure 3.11 Variation in frequency of prey species in diet of Asian palm civet
spring season percent volume of plants was high 77.33% followed by birds 19.47% and lowest was Sand/clay/ unidentified 0.03% (Table 3.12).
Analysis of 44 scats showed that 17 prey species occurred in diet of small
Indian civet. Among them 8 were wild, 1 domestic, and 8 were plant species.
Frequency of occurrence of wild prey species in diet of small Indian civet was
30.65%, domestic prey 14.52%, and plants 51.61%. Among wild prey frequency of occurrence of Indian gerbil was high (11.29%) followed by house mouse (4.84%).
Only one species of domestic prey was hen and it accounted for (14.52%). Among plant species frequency of occurrence of Himalayan wild pear was high (24.19%)
(Table 3.13) (Figure 3.6, 3.7, 3.8, 3.12). The general linear model explained 46.9% variation in diet of small Indian civet (R squared=0.469). The model showed that consumption of different dietary items varied significantly in diet of small Indian civet (F=2.8, df=18, p=0.002) (Appendix V).
Seasonal variation in diet composition of small Indian civet was assessed by analyzing 44 scat samples collected during four seasons (summer, winter, autumn, spring). Among 44 scats 12 were collected during summer season, 14 in autumn, 10 in winter and 8 scats in spring season. Consumption of wild prey species by small
Indian mongoose was high during winter season (41.67%) and low during autumn season (25%). Consumption of domestic prey species by small Indian was high during winter season (25%) and low during summer and spring season (10% each).
Consumption of plant matter by small Indian mongoose was high during autumn and spring season (60% each) and low during winter season (33.33%). General linear model showed that consumption of different dietary components by small Indian civet did not differ significantly during different seasons F=0.65, df=3, p=0.58.
Table 3.12 Percent volume of prey items recovered from the scats of small Indian
civet
Sr. Prey Summer Autumn Winter Spring Total Mean ± SE No. species/items (n=12) (n=14) (n=10) (n=8) (n=44) 1 Hairs 0.3 0.33 10.1 0.07 10.8 2.7±2.47 2 Feathers 19.04 21.11 37.85 19.47 97.47 24.37±4.51 3 Bones 2 0 5.8 3.1 10.9 2.73±1.21 4 Amphibians 1.13 0.76 0 0 1.89 0.47±0.28 5 Insects 0.02 0 0 0 0.02 0.005±0.005 6 Plants 76.87 77.71 46.23 77.33 278.14 69.54±7.77 7 Grits 0.2 0 0 0 0.2 0.05±0.05 8 Anthropogenic 0.09 0 0 0 0.09 0.02±0.02 9 Sand/clay 0.35 0.09 0.02 0.03 0.49 0.123±-0.07 100 100 100 100 400
Table 3.13 Percent frequency of occurrence of prey items in diet of small Indian civet
Summer Autumn Winter Spring Prey species/items recovered Total (n=12) (n=14) (n=10) (n=8) Wild prey Indian Gerbil (Tatera indica) 10 5 25 10 11.29 House mouse (Mus musculus) 5 10 0 0 4.84 Norway rat (Rattus norvegicus) 0 5 0 10 3.23 Roof or house rat (Rattus rattus) 0 0 8.33 0 1.61 Himalayan bulbul (Pycnonotus 5 0 0 10 3.23 leucogenys) House sparrow (Passer 0 0 8.33 0 1.61 domesticus) Amphibians 5 5 0 0 3.22 Insects (Orthoptera) Grasshopper 5 0 0 0 1.61 Total wild prey 30 25 41.67 30 30.65 Domestic prey Poultry (Gallus gallus 10 15 25 10 14.52 domesticus) Total domestic prey 10 15 25 10 14.52 Plant species Wild Himalayan pear (Dhandali) 25 40 16.67 0 24.19 (Pyrus pashia)
Aakhray (Rubus fruticosus) 5 5 0 30 8.06 Apple (Pyrus malus) 5 0 8.33 10 4.84 Kov (Olea ferruginea) 10 0 0 0 3.23 Choti bairi (Ziziphus oxyphylla) 5 5 0 0 3.23 Water melon (Citrullus lanatus) 0 5 0 0 1.61 Maluk (Diospyros lotus) 0 5 0 0 1.61 Loquat (Eriobotrya japonica) 0 0 8.33 20 4.84 Total plant species 50 60 33.33 60 51.61 Grits 5 0 0 0 1.61 Anthropogenic 5 0 0 0 1.61
30
25
20
15
10 Frequency of prey species prey of Frequency
5
0
Indian Gerbil Choti bairi Wild HimalayanpearWild Poultry Aakhray House mouse Apple Lokat Norway rat Himalayanbulbul Kov Amphibians orRoof rathouse House sparrow Orthoptera melonWater Maluk Grits Anthropogenic
Figure 3.12 Variation in frequency of prey species in diet of small Indian civet
Similarly, consumption of wild prey species did not differ significantly F=0.148, df=3, p=0.93 and consumption of plant species did not differed significantly F=0.56, df=3, p=0.64 (Table 3.13).
3.4.3.6 Diet composition of Indian grey mongoose
We analyzed 57 scats of Indian grey mongoose and percent volume of birds was high 34.25±6.76% in diet of Indian grey mongoose, followed by sand/clay/unidentified 17.86±3.48%, bones 16.81±2.38% and hairs 11.15±1.73%.
Lowest percent volume was contributed by snails 0.4±0.34%. During summer season, highest percent volume was recorded for birds 32%, followed by plants
21.82%, bones 17.43% and lowest was recorded for egg shells 0.3%. During autumn season, highest percent volume was contributed by bones 23.1% followed by plants
21.06% and lowest was contributed by snail 0.3%. During winter season, highest percent volume was contributed by birds 44%, followed by sand/clay/unidentified
26.96%, hairs 13% whereas, lowest was anthropogenic material 0.02%. During spring season, high percent volume was contributed by birds 45%, followed by hairs
15%, bones 14.74% and lowest was anthropogenic material 0.56% (Table 3.14).
Analysis of 57 scat samples showed that 22 prey species occurred in diet of
Indian grey mongoose. Among them 15 were wild, 1 domestic, and 6 plant species.
Diet of Indian grey mongoose consisted of (60%) wild species, (19%) domestic prey species, and (14%) plant species. Among wild prey species consumption of house mouse was high and the only domestic prey species i.e., chicken/poultry accounted for 19%. Among plant species consumption of choti bairi was high (5%). (Table
3.15) (Figure 3.6, 3.7, 3.8, 3.13). General linear model explained 64.7% variation in
consumption of dietary items by Indian grey mongoose (R squared=0.647). The variation in consumption of dietary items significantly differed (F= 5.74, df=23, p=0.000) (Appendix VI).
Seasonal variation in diet composition of Indian grey mongoose was assessed by conducting analysis of 57 samples; 13 in summer, 18 in autumn, 14 in winter and
12 in spring. Consumption of wild prey species was high during summer season
(72%) and low during spring season (52.63%). Consumption of domestic prey species was high during winter season (29.17%) and low during summer season
(8%). Consumption of plant species was high during summer season (16%) and low during winter season (8.33%). GLM showed that diet of IGM did not differ significantly during four seasons F= 1.028, df=3, p=0.38. Consumption of wild prey species did not differ significantly F=0.53, df=3, p=0.66. Similarly, consumption of plant species did not differ significantly F=0.37, df=3, p=0.77 (Table 3.15).
3.4.3.7 Diet composition of small Indian mongoose
Analysis of 69 scats of small Indian mongoose showed that percent volume of birds was high 29.55±7.28% followed by 21.27±3.89, insects 16.08±1.57%, plants
11.91±1.73%, bones 8.09±3.45, reptiles 5.3±2.35%, amphibians 4.09±2.51%,
Sand/clay/ unidentified 2.34±1.08, anthropogenic 1.24±0.37, and grits 0.14±0.09%.
During summer season percent volume of hairs was high 26.59%, followed by birds
15.3% whereas, percent volume of anthropogenic material was low 1.9%. During autumn season percent volume of hairs was high 28.92 % followed by birds 23.83% whereas, percent volume of grits was low 0.35%. During winter season percent volume of birds was high 29.52% followed by hairs 17.1% whereas, percent volume of grits was low 0.22%. During spring season percent volume of birds was high
Table 3.14 Percent volume of prey items recovered from the scats of Indian grey
mongoose
Prey Summer Autumn Winter Spring Total species/items Mean ± SE (n=13) (n=18) (n=14) (n=12) (n=57) recovered Hairs 7.4 9.2 13 15 44.6 11.15±1.73 Feathers 32 16 44 45 137 34.25±6.76 Bones 17.43 23.1 11.95 14.74 67.22 16.81±2.38 Reptiles 4 5 0 3.7 12.7 3.18±1.09 amphibians 0 3.98 0 0 3.98 0.99±0.99 Insects 3.3 1.6 2.8 4.8 12.5 3.13±0.66 Egg shells 0.2 0 0 1.4 1.6 0.4±0.34 snail 0 0.3 0 0 0.3 0.08±0.08 Plants 21.82 21.06 1.2 3.24 47.32 11.83±5.57 Grits 0 0.07 0.07 0 0.14 0.035±0.02 Anthropogenic 0.47 0.15 0.02 0.56 1.2 0.3±0.13 Sand/clay/ 13.38 19.54 26.96 11.56 71.44 17.86±3.48 unidentified 100 100 100 100 400
Table 3.15 Percent frequency of occurrence of prey items in diet of Indian grey
mongoose
Summer Autumn Winter Spring Prey species/items recovered Total (n=13) (n=18) (n=14) (n=12) Wild prey House mouse (Mus musculus) 16 12.5 12.5 5.26 12 Indian Gerbil (Tatera indica) 12 12.5 4.17 10.53 10 Norway rat (Rattus 4 3.13 4.17 5.26 4 norvegicus) Roof or house rat (Rattus 0 3.13 12.5 0 4 rattus) Spotted dove (Streptopelia 4 0 4.17 5.26 3 chinensis) Himalayan bulbul 4 0 0 0 1 (Pycnonotus leucogenys) House sparrow (Passer 4 3.13 0 5.26 3 domesticus) Red-vented bulbul 0 3.13 0 0 1 (Pycnonotus cafer) Reptiles 4 3.13 0 5.26 3
Amphibians 0 3.13 0 0 1 Orthoptera (grasshopper) 4 3.13 12.5 5.26 6 Hymanoptera (ants bees) 12 9.38 4.17 5.26 8 Coleoptera (beetles) 4 0 0 0 1 Snail (Cornu sp.) 0 3.13 0 0 1 Egg shells 4 0 0 5.26 2 Total wild prey 72 59.38 54.17 52.63 60 Domestic prey Poultry (Gallus gallus 8 15.63 29.17 26.32 19 domesticus) Total domestic prey 8 15.63 29.17 26.32 19 Plants Choti bairi (Ziziphus 12 6.25 0 0 5 oxyphylla) Water melon (Citrullus 4 6.25 0 0 3 lanatus) Apple (Pyrus malus) 0 3.13 0 5.26 2 Dhania (Coriandrum sativum) 0 0 4.17 0 1 Jaru grass (Themeda 0 0 4.17 5.26 2 anathera) Khabal grass (Cynodon 0 0 0 5.26 1 dactylon) Total plants 16 15.63 8.33 15.79 14 Grits 0 3.13 4.17 0 2 Anthropogenic 4 6.25 4.17 5.26 5
20
18
16
14
12
10
8
6 Frequency Frequency prey of species 4
2
0
Poultry House mouse Indian Gerbil Hymanoptera Orthoptera Choti bairi Anthropogenic Norway rat orRoof rathouse Spotteddove House sparrow Reptiles melonWater shells Egg Apple grass Jaru Grits Himalayanbulbul Red-vented bulbul Amphibians Coleoptera Snail Coriander Khabal grass
Figure 3.13 Variation in frequency of prey species in diet of Indian grey mongoose
49.55% followed by birds 17.1% and low percent volume was contributed by grits
0.22% (Table 3.16).
Analysis of 69 scat samples of small Indian mongoose showed that 17 prey species occurred in diet of small Indian mongoose. Among them 10 were wild prey species, only 1 domestic prey species and 6 plant species. Frequency of occurrence of wild prey was (59.68%), domestic prey (16.94%) and plant matter (11.29%).
Among wild prey species frequency of occurrence of house rat was high (10.48%) followed by house mouse (9.68%). The only domestic species i.e., hen/poultry accounted for 16.94%. Among plants frequency of occurrence of Jaro grass
(Themeda anathera) was high (4.03%) followed by choti bairi (2.42%) (Table 3.17)
(Figure 3.6, 3.7, 3.8, 3.14). General linear model explained 67.1% variation in diet of small Indian mongoose. There was significant difference in consumption of different dietary items (F= 6.61, df= 18, p=0.000) (Appendix VII).
For assessment of seasonal variation in diet composition of small Indian mongoose we analyzed 69 scats samples. Among these scats 17 were collected during summer season, 21 in autumn, 12 during winter season and 19 during spring season. Consumption of wild prey was high during summer season (66.67%) and low during winter season (54.17%). Consumption of domestic prey was high during spring season (24.24%) and low during summer season (10%). Consumption of plant species was high during summer season (13.33%) and low during autumn season
(8.1%). GLM showed that diet of small Indian mongoose did not differed significantly during four seasons F=0.38, df=3, p=0.76. Consumption of wild prey did not differ significantly F=0.138, df=3, p=0.93. Similarly, consumption of plant species during four seasons was not statistically significant F=0.094, df=3, p=0.96.
Table 3.16 Percent volume of prey items recovered from the scats of small Indian
mongoose
Prey Summer Autumn Winter Spring Total Mean ± species/items (n=17) (n=21) (n=12) (n=19) (n=69) SE recovered Hairs 26.59 28.92 12.48 17.1 85.09 21.27±3.89 Feathers 15.3 23.83 29.52 49.55 118.2 29.55±7.28 Bones 4.63 3.85 18.4 5.47 32.35 8.09±3.45 Reptiles 11.4 5.32 0 4.46 21.18 5.3±2.35 amphibians 6.08 10.27 0 0 16.35 4.09±2.51 Insects 14.22 14.89 20.76 14.43 64.3 16.08±1.57 Plants 14.42 10.85 14.9 7.48 47.65 11.91±1.73 Grits 0 0.35 0 0.22 0.57 0.14±0.09 Anthropogenic 1.9 0.68 1.86 0.5 4.94 1.24±0.37 matter Sand/clay/ 5.46 1.04 2.08 0.79 9.37 2.34±1.08 unidentified 100 100 100 100 400
Table 3.17 Percent frequency of occurrence of prey items in diet of small Indian
mongoose
Prey species/items Summer Autumn Winter Spring Total recovered (n=17) (n=21) (n=12) (n=19) Wild prey Indian Gerbil (Tatera indica) 6.67 5.4 4.17 3.03 4.82 House mouse (Mus 13.33 5.4 12.5 9.09 9.68 musculus) Roof or house rat (Rattus 10 13.51 8.33 9.09 10.48 rattus) House sparrow (Passer 3.33 0 0 3.03 1.61 domesticus) Red-vented bulbul 0 0 4.17 3.03 1.61 (Pycnonotus cafer) Amphibians 3.33 8.1 0 0 3.23 Reptiles 6.67 2.7 0 6.06 4.03 Orthoptera (grasshopper) 16.67 10.81 16.67 6.06 12.1 Hymanoptera (ants, bees) 6.67 8.1 4.17 15.15 8.87 Coleoptera (beetles) 0 5.4 4.17 3.03 3.23 Total wild prey 66.67 59.46 54.17 57.58 59.68 Domestic prey
Poultry (Gallus gallus 10 16.22 16.67 24.24 16.94 domesticus) Total domestic prey 10 16.22 16.67 24.24 16.94 Plants Melon (Cucumis melo) 3.33 0 0 0 0.81 Choti bairi (Ziziphus 6.67 2.7 0 0 2.42 oxyphylla) Water melon (Citrullus 3.33 0 0 0 0.81 lanatus) Khabal (Cynodon dactylon) 0 2.7 0 3.03 1.61 Grass (Themeda anathera) 0 0 8.33 9.09 4.03 Apple (Pyrus malus) 0 2.7 4.17 0 1.61 Total plants 13.33 8.1 12.5 12.12 11.29 Grits 0 5.41 0 3.03 2.42 Anthropogenic 10 10.81 16.67 3.03 9.68
18
16
14
12
10
8
6
Frequency of prey species prey of Frequency 4
2
0
Amphibians Poultry Orthoptera orRoof rathouse House mouse Anthropogenic Hymanoptera Indian Gerbil Reptiles Grass Coleoptera Choti bairi Grits House sparrow Red-vented bulbul Khabal Apple Melon melonWater
Figure 3.14 Variation in frequency of prey species in diet of small Indian mongoose
3.4.4 Diversity Index, Richness and Evenness of Prey Species
Prey species diversity index in diet of common leopard was high during summer season (2.27) and low during winter season (1.85). Prey richness was high during summer (13), and low during winter (7 species). Prey evenness was high during winter season (0.95) and low during spring and summer (0.88) each (Table
3.18) (Figure 3.15, 3.16 & 3.17).
Prey species diversity index for Asiatic jackal was high during summer and winter season (2.22) each, and low during spring (1.51). Prey species richness was high during summer (12) and low during spring (7). Prey species evenness was high during winter (1) and low during spring (0.78) (Table 3.18) (Figure 3.15, 3.16 &
3.17).
Prey species diversity index in diet of Kashmir hill fox was high during autumn season (2.38) and low during spring season (1.83) whereas, prey richness was high during autumn (16) and low during spring (9) species. Prey evenness was high during autumn season (0.86) and low during summer and winter season (0.83)
(Table 3.18) (Figure 3.15, 3.16 & 3.17).
Prey species diversity of Asian palm civet was high during autumn season
(2.48) and low in spring season (2.22). Prey species richness was high during autumn and winter seasons (19) each and low in spring season (11). Prey species evenness was high during spring season (0.93) and low during winter season (0.837) (Table
3.18) (Figure 3.15, 3.16 & 3.17).
Diversity index of prey species in diet of small Indian civet was high during summer season (2.39) and low during winter (1.82). Prey species richness was high during summer (13) species and low during winter and spring (7) species each. Prey
species evenness was high during spring season (0.94) and low during autumn season
(0.84) (Table 3.18) (Figure 3.15, 3.16 & 3.17).
Prey species diversity in diet of Indian grey mongoose was high during autumn season (2.63) and low during winter season (2.2). Prey species richness was high during autumn season (17) and low during winter season (12). Prey species evenness was high during summer and spring season (0.94) each and low during winter season (0.89) (Table 3.18) (Figure 3.15, 3.16 & 3.17).
Prey species diversity index in diet of small Indian mongoose was high during autumn season (2.48) and low during winter (2.23). Prey species richness was high during autumn and spring (14) species and low in winter (11) species. Prey species evenness was high during summer (0.95) and low during spring (0.9) (Table
3.18) (Figure 3.15, 3.16 & 3.17).
3.5 DISCUSSION
In the face of ever shrinking habitat of wildlife due to increasing human population, growing agricultural needs and unsustainable use of wild resources carnivore conservation is challenging. People living in and around protected areas are often dependent on livestock for their livelihood (Mishra et al., 2004).
Throughout the distribution range of leopards, a dietary shift from wild prey species to domestic species has been observed (Judas et al., 2006; Spalton and Al-Hikmani,
2006). Many factors are responsible for such phenomenon. Firstly, lack of wild prey results in leopard’s attack on livestock. Secondly, husbandry practices and penning conditions. Thirdly, guarding strategies play important role in loss of livestock during leopard attacks. Fourthly, seasons/temperature influence the rate of livestock depredation by leopards. Fifth factor which may have impact on rate of livestock
depredation by leopards is time of the day since attacks on livestock can be high during specific time of the day and it is usually when livestock is left unattended.
In Pakistan, common leopard has been reported to predate on snakes, lizards, rodents, Sindh ibex, markhor, urial, rhesus monkeys, porcupines and in the regions where wild prey is limited leopards are known for attacking domestic livestock including; cows, calves, donkeys, ponies, goats and sheep. Although its predation on crop-destroying porcupines, stray village dogs and rhesus monkeys might be considered beneficial to man, however, because of its attacks on domestic livestock and human beings it has been ruthlessly persecuted by local communities whenever encountered and has been always considered symbol of fear and contempt in
Pakistan (Roberts, 1997).
During present study, we recorded both wild as well as domestic prey in diet of common leopard at Pir Lasura National park. However, common leopard consumed more domestic prey as compared to wild prey species. We recorded mammals, birds, and anthropogenic items in diet of common leopard. Among wild prey consumption of rhesus monkey was high. Similar findings have been reported by other studies. Predation of leopard on primates has been reported from Asia and
Africa (Kummer et al., 1981; Cowlishaw, 1994; Isbell, 1994; Nowell and Jackson,
1996; Zuberbühler and Jenny, 2002; Hayward et al., 2006). Leopard predation on rhesus monkey has been reported by (Mukherjee and Mishra, 2001; Lodhi, 2007).
We found that consumption of domestic prey was higher as compared to wild prey and domestic prey species comprised of goat, sheep, cow, buffalo, horse, poultry and dogs. Among domestic prey species consumption of goat was high followed by dog and our findings are in line with other studies conducted in
Table 3.18 Prey species diversity index, prey species richness and evenness in diet of sympatric carnivore species in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan.
Species prey species diversity index (H') prey richness (S) prey evenness (E)
Summer Autumn Winter Spring Summer Autumn Winter Spring Summer Autumn Winter Spring
Common leopard 2.27 2.09 1.85 2.03 13 10 7 10 0.88 0.9 0.95 0.88
Asiatic jackal 2.2 2.1 2.2 1.51 12 11 9 7 0.9 0.87 1 0.78
Kashmir hill fox 2.2 2.38 1.87 1.83 14 16 10 9 0.83 0.86 0.81 0.83
Asian palm civet 2.37 2.48 2.46 2.22 16 19 19 11 0.85 0.84 0.837 0.926
Small Indian civet 2.39 1.93 1.82 1.83 13 10 7 7 0.93 0.84 0.935 0.94
Indian grey 2.55 2.63 2.2 2.45 15 17 12 14 0.94 0.93 0.89 0.928 mongoose
Small Indian 2.43 2.48 2.23 2.37 13 14 11 14 0.95 0.94 0.93 0.9 mongoose
Summer Autumn Winter Spring
2.63
2.55
2.48 2.48
2.46
2.45
2.43
2.39
2.38
2.37 2.37
2.27
2.23
2.22
2.2 2.2 2.2 2.2
2.1
2.09
2.03
1.93
1.87
1.85
1.83 1.83
1.82 1.51
C O M M O N A S I A T I C R E D F O X ASIAN PALM S M A L L I N D I A N S M A L L LEOPARD JACKAL CIVET I N D I A N G R E Y I N D I A N CIVET MONGOOSE MONGOOSE
Figure 3.15 Prey species diversity index of dietary habits of sympatric carnivore
species in and around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
Summer Autumn Winter Spring
19 19
17
16 16
15
14 14 14 14
13 13 13
12 12
11 11 11
10 10 10 10
9 9
7 7 7 7
C O M M O N A S I A T I C R E D F O X ASIAN PALM S M A L L I N D I A N S M A L L LEOPARD JACKAL CIVET I N D I A N G R E Y I N D I A N CIVET MONGOOSE MONGOOSE
Figure 3.16 Prey species richness in diets of sympatric carnivore species in and
around Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan
Summer Autumn Winter Spring
1
0.95 0.95
0.94 0.94 0.94
0.935
0.93 0.93 0.93
0.928
0.926
0.9 0.9 0.9
0.89
0.88 0.88
0.87
0.86
0.85
0.84 0.84
0.837
0.83 0.83
0.81 0.78
C O M M O N A S I A T I C R E D F O X ASIAN PALM S M A L L INDIAN GREY S M A L L LEOPARD JACKAL CIVET I N D I A N MONGOOSE I N D I A N CIVET MONGOOSE
Figure 3.17 Prey species evenness index of dietary habits of sympatric carnivore
species in and around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
distribution range of leopard. According to Shehzad et al., (2015) diet of the leopard comprised of mainly domestic prey species and consumption of goat was highest
(64.9%), followed by dog (17.5%) and cow Bos taurus (12.3%). Leopard has been reported to kill massive numbers (n=22) of livestock such as sheep in a single attack
(Sangay and Vernes, 2008). Athreya et al. (2016) reported that diet of leopard consisted of livestock, and 39% domestic dogs. There were few wild species recorded from diet of leopard.
Pir Lasura National park is such a landscape surround by human population and number of livestock in and around the park is over a million. In such a landscape availability of domestic prey is more than that of domestic prey specie therefore it is also logical for such as a species to predate on species which are abundant in the study area and requires less effort and this phenomenon has been reported by other studies. Carnivore density in natural or semi-natural ecosystems is related to biomass of prey species available (Fuller and Sievert, 2001; Karanth et al., 2004; Carbone et al., 2010; Karanth and Nichols, 2010). Recent studies have demonstrated that large carnivores can persist in human-dominated landscapes by predating fully or partially on domestic livestock (Gehrt et al., 2010; Yirga et al., 2012; Athreya et al., 2013).
The potential of human dominated landscape to support large carnivore species must be investigated in terms of availability and abundance of wild as well as domestic prey species (Boitani and Powell, 2012). In human-dominated landscapes biomass of domestic prey species can be higher than wild prey species (Schaller, 1983;
Seidensticker et al., 1990; Mizutani, 1999).
We recorded anthropogenic items in diet of common leopard and our results are in line with other studies. Anthropogenic food items and garbage can also be
consumed by carnivore species (Gehrt et al., 2010). Such food resources can be abundant in such a landscape and may result in higher densities of wild carnivores in human-dominated landscapes when compared against wild areas. In urban areas, higher density of red foxes Vulpes vulpes (Bino et al., 2010) and black bear
(Beckmann and Berger, 2003) was reported in urban as compared to wild habitats due to better feeding opportunities on crops, livestock and garbage. Domestic animals, such as goat and sheep are easier prey for leopard as they lack anti-predatory behavior, unlike their wild counterparts (Diamond, 2002). However, availability does not always indicate easy access to prey since livestock might be guarded by day and penned at night.
Livestock depredation by leopard has resulted in antagonistic interaction between local communities and leopard in Pir Lasura National park. Interaction between felids and human is complex and spectrum of such relation ranges from fascination to fear (Boomgaard, 2001; Loveridge et al., 2010). From conservation of large felids, they are often represented as flagship species (Treves and Karanth,
2003) but antagonistic interaction of large felids occurs in areas where these species live in human-dominated landscapes, where presence of these species can result in livestock depredation and loss of human life (Treves and Karanth, 2003). As a result, large felids are killed in retaliation which is significant cause of their mortality
(Inskip and Zimmermann, 2009). During current study, we found major prey species of leopard was goat followed by dog. This is due to the fact of high availability of goat, poor guarding and penning condition in study area. Dogs are used for guarding livestock and they become easy prey for leopards. Many studies have reported leopard predating on dogs such as (Daniel, 2009). Predation of leopard on goat is
higher and overall consumption of domestic prey is higher than wild prey species which shows that the economic impact of predation is higher. The regression equation of Carbone and Gittleman (2002) estimates that per 90 kg of predator
10,000 kg of prey is required, irrespective of predator size.
Two types of interaction for resources have been recognized which are interference and exploitation (Miller, 1967). Interference competition is defined as any activity which directly or indirectly limits the access of a competitor to a required resource whereas, exploitation competition is defined as reduced availability of a resource because of exploitation of resource by superior competitor (Miller, 1967).
Carnivores compete each other with resource exploitation and also through interspecific killing: a key determinant of distribution and abundance of carnivores.
(Donadio and Buskirk, 2006). As a result, space use, activity pattern also social behavior of the inferior competitor may change (Palomares and Caro, 1999;
Hayward and Slotow, 2009).
Spatial avoidance is one of evolutionary solution for coexistence of carnivores in an area (Hunter and Caro, 2008), however coexisting is also possible by consuming prey species of different weight class or different taxa (Karanth and
Sunquist, 2000; Hayward and Kerley, 2008).
We recorded mammals, birds, insects, and anthropogenic items from diet of
Asiatic jackal at Pir Lasura National Park. Consumption of domestic prey was higher than that of wild prey. Wild prey species included rhesus monkey which was main wild prey species and consumption of four species of rodents. Among domestic prey species, poultry was main prey followed by goat. Whereas consumption of plant matter was low. Studies conducted elsewhere, have reported rodent and plants as
main component of jackal diet. However, during present study we found that this is not the case in our study area. According to results of our study jackal seems to be scavenger and predates on domestic poultry for major part of its diet and then rodents and rhesus monkey. Our results are in line with Roberts wo reported that in Pakistan conception of jackal is as a scavenger and carrion eater (Roberts, 1997). Jackals feed on refuse in premises of villages, however, majority of their diet comprise of rodents, reptiles, fruits, and insects when available (Roberts, 1997). A study conducted in
Bharatpur, Rajastan, India during 1984-1985 showed that diet of jackal comprised of rodents 26.5%, birds 24.1 %, grass 20%, fruits 16.5%, insects 4.1%, snakes 4.1%, chital 2.4%, nilghai 1.2%, and fish 1.2% (Sankar, 1988). Stomach contents of four jackals in Rajasthan Indian revealed that diet of jackals mostly consisted of fruits of
Ziziphus, with small proportion of beetles and scorpions. Another individual consumed desert jirds (Meriones hurrianae) and small mongoose (Prakash, 1959A).
Analysis of 138 scats of Asiatic jackal in sal forests in India showed that 68 percent of diet consisted of rodents followed by plants 27%, 11% reptiles, 8% fish and 9.4% birds (Schaller, 1967). A study conducted in Sariska Tiger Reserve, India showed that diet of Asiatic jackal comprised of plant matter 17.57%, rodents 15.77%, cattle
15.32%, chital 10.81% fruits 9.01%, birds 7.21%, sambar 5.41%, hare 4.05%, nilgai
4.05%, goat 1.80% reptiles 1.8% (Mondal et al., 2012). Diet of jackals usually comprise of small and meso mammals such as rodents, rabits, birds, fishes, insects and vegetation (Aiyadurai and Jhala, 2006). Diet of jackal in Keoladeo national park,
India comprised of mainly plant matter 38.44%, nilgai 8.13%, rodents 10.31%, chital
9.69%, cattle 7.5%, anthropogenic 1.56%(Singh et al., 2016). High proportion of mammals in diet of jackal suggest that they scavenge on deal animals and hunt calves
as hunting large mammals might be difficult (Singh et al., 2016). Rodent are important part of jackal diet (Mukherjee et al., 2004; Jaeger et al., 2007; Majumder et al., 2011; Singh et al., 2016). Diet of Asiatic jackal in Pakistan showed that 46.47
% of its diet is animal matter by volume including rodents, mongooses, wild boar, livestock, birds and domestic poultry and 25.08% plant included wheat tomato, berries, grams, orange, melon and water melon (Mahmood et al., 2013). Most of the species have recorded plants and rodents as major prey species in diet of Asiatic jackal this could be due to the difference in sampling procedures. Jackals living in high mountainous regions can take up mammals only as major part of their diet
(Schaller (1970). In India Schaller (1970) reported that frequency of occurrence of rodents in diet of jackal was 94%, however he also reported snakes and lizards 29% and insects 6.7 %. Previous, studies which have been conducted in Pakistan has also reported large proportions of plant matter in diet of jackal however during present study scats of jackal were confirmed genotypically and it was interesting that scats identified as from jackal have very less or no plant matter and other sympatric carnivores such as fox and civets consumed plant matter in large proportions.
There is paucity of information available on diet composition of fox in
Pakistan and no published literature is available. However, some information is available from Roberts (1997) for comparison. Foxes are adaptable hunters and can hunt hares, rodents, reptiles and small birds. However, when vertebrate prey is not available they can subsist on insects and fruits. According to Roberts foxes feed on fruits of bair tree (Zizyphus mauritiana). In diet of fox mice, rats, desert hare, Indian gerbils, have been reported by Roberts (1997). Studies in Indian Rajasthan showed that foxes feed on wild melon, termites (Prakash, 1959A). Foxes have reported to
feed on scorpions, fruits of Zizyphus nummularia, spiders, cockroaches and some melon seeds in stomach. Roberts (1997) never encountered any sign of foxes feeding on domestic poultry although he has reported such instance of domestic poultry taken up by civets, cats, mongooses and martins.
During present study, we recorded that diet of Kashmir hill fox diet comprised of mammals, birds, invertebrates and plant matter. We also recorded grits and anthropogenic material in diet of fox. Consumption of poultry was higher as compared to any other dietary component followed by goat. We identified 6 species of plants in diet of fox. It has also consumed sheep, cow, buffalo which are large prey species and it seems that it might have fed on carcasses of these animals showing its scavenging behavior. The consumption of different food items showed variation during different seasons based on their availability. Our findings are in line with other studies such as, analysis of 70 stomach contents of red fox in Egypt showed that it feeds on diverse diet including; rodents, carrion, reptiles, birds, fishes, plants, invertebrates and anthropogenic material. Diet showed seasonal variations based on availability of different food items (Basuony et al., 2005; Baker et al., 2006;
Kidawa and Kowalczyk, 2011). Red fox is essentially omnivore having diverse diet which includes invertebrates, small mammals, birds, plant matter and carrion
(Flower, 1932; Englund, 1965; Amores, 1975; Macdonald, 1979; Osborn and
Helmy, 1980; Ciampaloni and Lovari, 1985; Calisti et al., 1990; Basuony, 1998).
Fox has been reported by other studies to feed on plant fruits in southern Europe
(Ciampaloni and Lovari, 1985; Calisti et al., 1990) it is known to feed on plant species or parts of plants having high sugar content (Basuony et al., 2005). Food items rich in proteins such as mammals and birds might be expected in diet of
carnivores during pregnancy period (Ciampaloni and Lovari, 1985). According to study conducted by Bakaloudis et al. (2015) red fox is generalist predator and it consumes, birds, small size carnivores, amphibians and reptiles, invertebrates in a seasonal way.
According to optimal foraging theory which states that selection of prey species by a predator is based on cost and benefits during capturing, handling and consuming (Krebs and Davies, 1993). The diet selection of red fox depends on seasonal variation of foraging costs which is affected by availability of primary prey species (Erlinge, 1981; Raymond et al., 1990). Primary prey species is always taken when available which shows that cost of predation on such species is never high.
During current study consumption of poultry was high than any other prey item which seems to be major prey species for fox in the study area. Similarly, predation on poultry shows fox tendency to choose easy to catch prey species which requires minimum energy expenditure. Furthermore, feeding on carrion requires least energy
(Basuony et al., 2005). Seasonal variation in diet and high variability of diet shows ability of fox to adapt to variable conditions during year (Diaz-Ruiz et al., 2013).
Result of current study shows that Asian palm civet is omnivore. Major component of its diet was plant species followed by wild prey and domestic prey.
Among domestic prey consumption of poultry was high. We also recorded cow and sheep in its diet showing its scavenging behavior. We also recorded invertebrates including snails, beetles, grasshoppers, grits, and anthropogenic matter in diet of
Asian palm civet. Asian palm civet is reported to live in vicinity of villages, and subsists on rats, mice and attracted towards food orchards and plantations (Roberts,
1997). No other published literature is available for Asian palm civet from Pakistan
however, results available from other countries are comparable. Asian palm civets play important role in food web as predator, as prey and as seed dispersers (Aroon et al., 2012). Their diet includes; small mammals, birds, amphibians, reptiles, invertebrates and eggs and fruits. According to results of our study consumption of plant matter was high in diet of Asian palm civet. Our findings are in line with other studies such as Joshi et al. (1995) showed that diet of Asian palm civets mainly comprised of fruits in Nepal (84.5%). Krishnakumar and Balakrishanan (2003) reported that diet of Asian palm civet comprised of 82% fruits. Jothish (2011) reported that 90.1% of diet of Asian palm civet comprised of fruits in India.
Grassman (1998) recorded 18 species of fruits in diet of Asian palm civet in
Thailand. Krishnakumar and Balakrishanan (2003) recorded 10 fruit species in diet of Asian palm civets in India. Similarly, Su and Sale (2007) reported 31 fruit species in diet of Asain palm civet. In Myanmar. Frequency of fruits in diet was high during rainy season than winter season (Rabinowitz, 1991; Kitamura et al., 2002) and it consume more mammalian prey when fruits availability is decreased (Aroon et al.,
2012).
We found scats of Asian palm civit during current study which were completely composed of plant matter and this is also reported from other studies which reported that scats of Asian palm civet are fully composed of plant matter during fruiting seasons (Rabinowitz, 1991; Corlett, 1998). Diet of Asian palm civet comprise of small mammals, arthropods, lizards, snakes, birds, fruits. Small mammals were most abundant 53.7% followed by plants 37.65%. niche breadth was widest during autumn season (B=0.169), followed by summer (B=0.148), and narrowest during winter (B=0.148). Overall dietary niche breadth was 0.412. Asian
palm civet consumed arthropods amphibians and reptiles during rainy seasons corresponding with their availability (Aroon et al., 2012).
Diet of Asian palm civet was variable during different seasons of current study. Diet of Asian palm civet was variable during different seasons in Thailand
(Aroon et al., 2012; Jothish, 2011). Whereas, study conducted in Thailand
(Rabinowitz, 1991) and Nepal (Joshi et al., 1995) there was no seasonal variation.
Niche breadth of civets was narrow during winter when there was low availability of fruits and high during fruiting seasons.
Frequency of occurrence of plants was high in diet of small Indian civet followed by wild prey species, domestic prey, insects, amphibians, grits, and anthropogenic material. Diet of small Indian civet comprised of mainly plants, rodents, birds. Diet of small Indian civet has never been studied and no published literature is available except some observations by Roberts (1997). Small Indian civet is omnivorous, in feeding habits and rely on fruits during their availability and feed on variety of insects and arthropods. In India from stomach of a dead civet remains of beetles, seeds of bairs fruit (Zizyphus jubata) and unidentified finch was recovered (BNHS Survey, 1913). A specimen was caught at Kalabagh feeding on ripe grapes. Small Indian civet hunts on small mammals, birds and reptiles. They have been reported to rob nests of birds for eggs and nestlings. They spent most of the day sleeping in their burrows and hunt at night. They are known to dig for insect larvae and succulent roots (Roberts, 1997). Diet of small Indian civets comprised of rodents (80%) insects (23%) (Wang et al., 1976). According to another study diet of small Indian civet consisted of rodents and shrews 40%, and insects 95% and earthworms 67% (Chuang and Lee, 1997) which is not the case in our study since
we have recorded 47.76% plants in diet of small Indian civet. Diet breadth was found to be 4.46 (Chuang and Lee, 1997). Diet breadth in another study was reported as
2.58 and it consumed 9 prey species. Small Indian civet consumed rodents, birds, invertebrates, and plants matter (Wang and Fuller, 2003). Small Indian civets when live in outskirts of villages and cultivated land they may predate on more diverse diet
(Wang, 1998). Civets are known to consume fruits and (Rabinowitz, 1991) reported
76% fruits in diet of masked palm civet. Joshi et al. (1995) reported that frequency of occurrence of fruits in diet of common palm civet were found to be 85% and during fruiting season whole scats entirely comprised of fruits. Furthermore, during shortage of fruits common palm civets also consumed insects, mollusks, birds, reptiles and small mammals (Joshi et al., 1995).
Diet composition of Indian grey mongoose consisted of mammals, birds, reptiles, amphibians, invertebrates, egg shells, and plant species. We also recorded grits and anthropogenic items. Frequency of occurrence of house mouse was high in diet of Indian grey mongoose. Among birds, frequency of chicken/poultry was high.
Four species/orders of invertebrates were recorded frequency of occurrence of
Hymenoptera (ants,bees) was high and frequency of occurrence of coleoptera
(beetles) and snails was low . Our results are in line with other studies such as Roberts
(1997) reported that Indian mongoose feed on reptiles, birds, and amphibians.
Prakash (1959A) reported in Rajasthan, India that grey mongoose feed on grey partridges, rodents, invertebrates and lizards. Indian grey mongoose is known as an opportunistic hunter. It has been reported to feed on rodents, reptiles, invertebrates and birds, eggs of birds and fruits. We also recorded birds, egg shells invertebrates, plants reptiles and rodents in diet of Indian grey mongoose such results are reported
by other studies. Indian grey mongoose feed on the red jungle fowl its chicks, eggs, peafowl, partridges, small mammals and snakes in Indian and also reported to search for food under stones on the beach side in Hawaii (Santiapillai et al., 2000;
Postanowicz, 2002). It has been reported to feed on grasshoppers, centipedes, fish, frogs, scorpions and crabs (Whitfield, 1978). Hussain et al. (2017) reported that
Indian grey mongoose feeds on rodents, birds, insects, and plants.
During current study, we recorded that the diet of small Indian mongoose consisted of mammals, birds, invertebrates, reptiles, amphibians, and plants. We also recorded anthropogenic matter and grits. Among mammals, frequency of occurrence of house rat was high. This could be due to the reason that small Indian mongoose lives around human habituation where availability of house rat might be high. Small
Indian mongoose feeds on rodents such as Rattus rattus, and Mus musculus when it lives around human settlements (Roberts, 1997). Only one species of domestic prey was recorded i.e., chicken/ poultry. Among 3 species of invertebrates recorded in diet of small Indian mongoose frequency of occurrence of order orthoptera
(grasshoppers) was high followed by Hymenoptera (ant, bees) and low was coleopteran (beetles). Among 6 plant species recorded in diet of small Indian mongoose frequency of occurrence of Themeda anathera grass was high followed by choti bairi and low was melon and water melon. It has been reported to feed on beetles, scorpions, snakes, lizards, spiders and amphibians (Prakash, 1959A). It feeds on birds and their eggs and nestlings (Roberts, 1997). Seaman and Randall (1962) reported that small Indian mongoose consumes small mammals, reptiles, amphibians, birds and plant matter. Some population of mongoose are insectivorous and others may consume fruits during some seasons (Seaman and Randall, 1962).
Prey species of small Indian mongoose are rodents, insects and plant matter (Fruits)
(Mahmood et al., 2011; Siddiqui et al., 2004; Hussain et al., 2017).
3.6 CHAPTER SUMMARY
Knowledge of a predator’s diet is important for understanding its ecology and for predicting its influence on the dynamics of prey populations. We investigated diet composition of seven sympatric carnivore species at Pir Lasura national park,
Azad Jammu and Kashmir, Pakistan. We used molecular identification technique to confirm for the carnivore species. Accuracy for Common leopard scats was found to be 95.8%, Kashmir hill fox 88.9%, Asiatic Jackal 90.3%, Asian palm civet 74.3%, and Small Indian Civet 92.4%. Scats of Indian grey mongoose and small Indian mongoose were not subjected to molecular identification since their scats were collected outside their burrows. Morphological characteristics of scats showed that they overlap greatly among different species based on their diameter and length. Diet of seven carnivore species was studied by analyzing a total of 473 scats samples collected from the field. Among those, were 39 from common leopard, Asiatic jackal
64, Kashmir hill fox 92, Asian palm civet 108, small Indian civet 44, Indian grey mongoose 57, and small Indian mongoose 69 scat samples. Diet of common leopard comprised of 17 prey species (10 wild and 7 domestic). Frequency of occurrence of wild prey was 34.85 % of total leopard diet whereas domestic prey contributed
59.1%. Sixteen species of mammals, birds, insects and plants were identified from diet of Asiatic jackal. Among these species 10 were wild, 5 were domestic and 1 species of plant. Frequency of wild prey in diet of jackal was 18.48% and domestic was 59.78%. We recorded 21 species of mammals, birds, plants and insects in diet of red fox. Among these 21 species 10 were wild, 5 domestic and 6 plant species.
Frequency of occurrence of wild prey was 17.96%, domestic prey 50.9%, plants
27.54%. Analysis of 108 scats samples of Asian palm civet showed that 27 species of mammals, birds, invertebrates and plants were consumed. Among all recorded species 9 were wild, 3 domestic, and 15 plant species. Consumption of wild prey was
33.68% compared to 10.88% domestic prey species and 53.37% plants. Analysis of
44 scats showed that 17 prey species occurred in diet of small Indian civet. Among them 8 were wild, 1 domestic, and 8 were plant species. Frequency of occurrence of wild prey in diet of small Indian civet was 30.65%, domestic prey 14.52%, and plants
51.61%. Analysis of 57 scat samples showed that 22 prey species occurred in diet of small Indian mongoose. Among them 15 were wild, 1 domestic, and 6 plant species.
Diet of Indian grey mongoose consisted of (60%) wild species, (19%) domestic prey species, and (14%) plant species. Analysis of 69 scat samples of small Indian mongoose showed that 17 prey species occurred in diet of small Indian mongoose.
Among them 10 were wild prey species, only 1 domestic prey species and 6 plant species. Frequency of occurrence of wild prey was (59.68%), domestic prey
(16.94%) and plant matter (11.29%).
Chapter 4
ASSESMENT OF CARNIVORE PREY SPECIES ABUNDANCE
AT PIR LASURA NATIONAL PARK
4.1 INTRODUCTION
Carnivores often inflict economic losses to local communities by predation on domestic livestock and as a result carnivores are often persecuted (Treves and
Karanth, 2003; Gusset et al., 2009). Such phenomenon is often observed where humans live near protected areas and livestock losses pose strong impact on small- scale households, thus making biodiversity conservation challenging (Baker et al.,
2008; Dar et al., 2009). Over decades, humans have encroached carnivore habitat because of expanding human population, consequently environment functioning has been impacted negatively at all levels starting from individuals to ecosystems
(Ripple et al., 2014).
Albeit the abundance of domestic livestock exceeds those of wild prey species in many areas, carnivores prefer to kill wild prey to avoid human revenge
(Loveridge et al., 2010). However, when wild prey becomes scarce, carnivores predate on livestock for their survival (Mondal et al., 2011; Zhang et al., 2013).
Carnivores predate on domestic livestock during wet season when wild prey disperses in lush woods to gain more fitness. Thus, wild prey becomes less available to carnivore species. Meanwhile, domestic livestock enters these lush vegetative areas for uncontrolled grazing (Patterson et al., 2004; Kissui, 2008). In many areas livestock depredation is less during winter season when density of prey become high
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(Dar et al., 2009) or depredation is high during dry season when vegetative cover decreases, wild prey migrates and domestic livestock concentrates around limited water resources (Schiess-Meier et al., 2007).
The relation between availability of wild and domestic prey species and their predation by carnivores can vary in different landscapes. Top predators by their predatory behavior control prey populations which in turn impacts whole ecosystem a phenomenon known as tropic cascade. Absence of top predators leads to more medium sized predator. Suryawanshi et al. (2013) reported that livestock depredation by snow leopard may be more intense when wild prey is abundant, as more wild prey will support high density of predator. Therefore, carnivores can kill more livestock.
To evaluate if dichotomy between scarcity of wild prey and increase depredation by carnivores on domestic livestock is true? we designed current study to investigate availability and consumption of wild and domestic prey by sympatric carnivore species in a human-dominated landscape located in north-eastern Himalayan region of Pakistan.
4.2 REVIEW OF LITERATURE
Khorozyan et al. (2015) predicted that large carnivore species kill more livestock when wild prey reaches minimum threshold. According to their results cattle depredation by carnivores is high when prey biomass is < 812.41 Kg/km2.
Similarly, for sheep and goat predation is high when prey biomass is <544.57 kg/km2 regardless of size of study areas, body masses and population density of carnivore species.
Due to species specific ecological characteristic, pattern of livestock
depredation by carnivores may differ. Such as snow leopard is known to kill large number of animals in one attack, similarly lions might kill more livestock as compared to other cat (Jackson et al., 2010). Big cat species are often known to predate on larger prey species such as cattle and buffaloes however, small cat species are known to prefer small prey species such as sheep, goats, and juveniles of cattle and buffaloes (Zarco-González et al., 2013). Jackals and foxes are also reported to scavenge on dead animals.
Predator-prey dynamics is very important as density of predators is positively relative to the density of prey species available. Prey biomass is often used to predict carnivore density and carrying capacity based on current prey (Carbone and
Gittleman, 2002; Carbone et al., 2011). However, this rule is only applicable from bottom to top approach i.e., carnivores controlled by prey. This rule is not applicable when increasing top–down approach such as “carnivores controlled by humans, e.g., via poaching” results in limiting carnivore population whereas, population of prey species remains sufficient (Khorozyan et al., 2015; Bauer et al., 2014).
Study area size is inversely proportional to the density of predators and their prey species and they can mediate the predator-prey relationship and livestock depredation by predators (Carbone and Gittleman, 2002). For example, prey species are often studied in smaller areas having high density such as protected areas which are often areas of low livestock depredation by carnivores (Biswas and Sankar,
2002).
4.3 MATERIALS AND METHODS
Abundance of carnivore prey species in the study area was estimated to
establish their availability to the carnivores. To record the abundance of mammalian prey species, “signs survey method” was applied. Direct and indirect signs like direct sightings, burrows (Begon, 1979) and faecal material (Wood, 1988; White and
Eberhardt, 1980) of prey species were recorded in specified area by area searches at
10 different sampling sites of the Pir Lasura National Park to assess abundance of prey species. Furthermore, trapping was performed for small mammals following
Erlinge (Erlinge et al., 1983; Sutherland, 1998). The numbers of captured animals were divided by the numbers of trapping nights to get index of abundance. We used
10 traps used in 4 days (1 day each season) total trap nights were 40 and area was
100mx100m.
For recording abundance of bird species as prey of carnivores, Line transect
(Burnham et al., 1980) method was used. Twenty transects were established in ten different sites (Two transects in each study site. Each transect was 200 m wide (100 on each side) and 500 m long. Total area of each transect was 0.1 km2. Abundance of amphibians and reptiles was assessed at ten study sites using “Visual Encounter method” along a line transect within each sampling site (Campbell and Chiristman,
1998; Heyer et al., 1988; Fellers and Freel, 1995). Abundance of snails was recorded by area search method and insect abundance was recorded using wet pitfall traps and sweep netting (Luff, 1975; Greenslade, 1964) method in a specified area in sampling sites. For pitfalls we used 5 boxes (each was 22-inch circumference and A= 38.51- inch sq., total area of 5 boxes 192.55, while 1m=1550 sq inch).
4.4 RESULTS
Abundance of 5 species of rodents were estimated including house mouse,
Table 4.1 Abundance estimation of rodent prey species in the PLNP, AJ&K.
Abundance Total Species Summer Autumn Winter Spring Abundance estimation captured /Km2
House mouse (Mus 2 1 1 3 7 0.175 17.5 musculus)
Indian gerbil (Tatera 3 2 1 2 8 0.2 20 indica)
House rat 2 2 1 1 6 0.15 15 (Rattus rattus)
Norway rat (Rattus 1 1 1 1 4 0.1 10 norvegicus)
Turkistan rat (Rattus 1 0 0 0 1 0.025 2.5 pyctoris)
Table 4.2 Abundance estimation mammalian species in the PLNP, AJ&K.
Abundance Species Summer Autumn Winter Spring estimation /Km2
Barking deer Muntiacus muntjac 10 3 5 7 2.5 (Pellet groups)
Kashmir hill fox Vulpes vulpes 5 3 2 1 1.1 griffithi (Direct)
Rhesus monkey 20 28 44 19 11.1 Macaca mulatta (Direct)
Wild boar Sus scrofa (Direct) 14 7 20 13 5.4
Indian crested Porcupine Hystrix 23 14 23 25 8.5 indica (burrows)
Asian palm civet Paradoxurus 0 4 2 0 0.6 hermaphroditus (Direct)
Desert hare (Lepus nigricollis 3 2 0 2 0.7 dayanus (Direct)
Table 4.3 Abundance estimation of bird species in and around PLNP.
Abundance Species Summer Autumn Winter Spring estimation/Km2
Kalij pheasant 4 3 2 1 5 (Lophura leucomelano)
Indian peafowl 2 2 1 0 2.5 (Pavo cristatus)
Spotted dove 18 13 7 11 24.5 (Streptoptopelia chinensis)
Red-vented bulbul 19 16 9 14 29 (Pycnonatus cafer)
Himalayan bulbul 22 24 10 16 36 (Pycnonatus leucogenys)
House sparrow 79 98 61 78 158 (Passer domesticus)
Indian gerbil, house rat, Norway rat and Turkistan rat. Abundance of Indian gerbil was high (20/km2) followed by house mouse (17.5/km2) and lowest abundance was recorded for Turkistan rat (2.5/ km2) (Table 1). Abundance of Barking deer, red fox, rhesus monkey, wild boar, porcupine, Asian palm civet and desert hare was also recorded. Among these species abundance of rhesus monkey was high (11.1/ km2), followed by porcupine (8.5/ km2), wild boar (5.4/ km2) whereas, Asian palm civet had lowest abundance (0.6/ km2) (Table 2).
Abundance of six species of birds was also recorded including Kalij pheasant,
Indian peafowl, spotted dove, red-vented bulbul, Himalayan bulbul and house sparrow. Among these bird species, abundance of house sparrow was high (158/ km2) followed by Himalayan bulbul (36/ km2), red-vented bulbul (29/ km2) and least was recorded for Indian peafowl (2.5/ km2) (Table 3).
Abundance of all amphibians was (78.1/ km2) and all reptiles was (89.4/ km2)
(Table 4). Abundance of invertebrates showed that members of order hymenopteran were more abundant (2310.3/ m2) followed by orthoptera 42.26/m2 and Coleoptera
(22.14/m2) and abundance of snails was (3.4/km2) (Table 5).
Abundance of domestic prey species were computed for whole district.
Abundance of domestic hen (excluding population in farms) was (548.69/km2) followed by goats (105.17/km2) and cows 43.07/ km2) and least abundant was mule
(0.03/ km2) (Table 6). Abundance of dog was estimated during household survey and it was (22/ km2).
4.5 DISCUSSION
Table 4.4 Abundance of amphibians and reptiles in the PLNP, AJ&K.
Abundance Species Summer Autumn Winter Spring /km2
Amphibians 452 319 0 10 78.1
Reptiles 496 357 7 34 89.4
Table 4.5 Abundance estimation of invertebrates in the PLNP, AJ&K.
Species Summer Autumn Winter Spring Abundance Snails (10 sites) 11 0 23 0 3.4/Km2 Orthoptera 6 4 8 3 42.26/m2 (Grasshoppers) Hymanoptera 340 356 161 291 2310.3/m2 (Ants/beees) Coleoptera (Beetles) 4 5 0 2 22.14/m2
Table 4.6 Abundance of domestic prey species in district Kotli Azad Kashmir. (Data
source report DCR 2017.18-5-2017 data collected from 52 units. Department
of Livestock and dairy development Kotli AJ&K. District area (1862 km sq.)
Species Total population Population Estimation/Km2 Hens (domestic) 807540 548.69 Goats and sheep 195840 105.17 Cows 80210 43.07 Buffalo, Ox 52645 28.27 Horses 289 0.16 Donkeys 12321 6.62 Camel 215 0.12 Mule 50 0.03
The results of our study showed that consumption of wild prey by common leopard was 34.85% while abundance of those wild prey species was estimated to be
57.4/Km2 whereas, consumption of domestic prey species by common leopard was
59.1% and availability of those species was found to be 747.36/Km2. Major wild prey species of common leopard was rhesus monkey and its consumption was
10.61% of its diet interestingly, this species was most abundant wild species in the study area having abundance 11.1/km2. Among domestic prey species consumption of goats and sheep was high and it was more than 30% of leopard diet. Abundance of goats and sheep in the study area was high as compared to other ungulate species and it was 105.17/Km2.
Consumption of wild prey by Asiatic jackal was 18.48% whereas, its availability was 114.2/km2. Among wild prey species consumption and availability of rhesus monkey was high. Among wild prey species consumption and availability of goat and poultry was high.
Consumption of wild prey in diet of Kashmir hill fox was around 18% and availability of same species in habitat was 125.7/ km2. Major prey species in diet were insects 4% and wild boar 3% followed by house mouse 2.4% and their abundance was 2310.3/m2, 5.4/km2 and 17.5/km2 respectively. Among domestic prey species consumption of poultry was high 29.34% and its availability was also high 548.69/km2. Similarly, consumption of goat and sheep was above 19% and their availability was 105.17/km2.
Large portion of plants was recorded in diet of Asian palm civet, however consumption of wild prey was 33.68% and domestic was 10.88%. Despite of high
availability of domestic prey Asian palm civet preferred wild prey especially rodents.
Among domestic species consumption of poultry was high 6.74% and it was quite abundantly available. Among wild prey species consumption of 4 rodent species accounted for 18.64% collectively however, consumption of Indian gerbil was high
9.84% among rodents and its availability was also high 20/km2. Diet of Asian palm civet contained significant percentage of invertebrates 8.3% and availability of invertebrate species was high.
Diet of small Indian civet consisted of plant matter, wild prey and domestic prey species. Consumption of wild prey was 30.65% and wild prey was 14.52%.
Among wild prey species consumption of Indian gerbil was high 11.29% and it was abundant rodent species in the study area having abundance of 20/km2. Other dietary components included birds, amphibians and insects. Only one domestic species domestic chicken was consumed, and it was quite abundant in habitat as well
548.69/km2.
Consumption of wild prey by Indian grey mongoose was 60% among wild prey consumption of house mouse was high 12% followed by Indian gerbil 10% and these species were abundant in the habitat as well having abundance of 17.5/km2 and
20/km2. Consumption of invertebrates was high 16% and their availability was also high. Only one domestic species domestic chicken accounted for 19 % of its diet and its abundance was recorded as 548.69/km2.
Consumption of wild prey by small Indian mongoose was 59.68% and availability of species consumed was also high such as frequency of occurrence of house mouse was high in diet and its availability was also high. Similarly,
consumption of insect was high in diet of small Indian mongoose and invertebrates were abundant in habitat. Consumption of domestic chicken was 16.94% and its availability was high in the study area 548.69/km2.
4.6 CHAPTER SUMMARY
Abundance of carnivore prey species in the study area was estimated to establish their availability to the carnivores using direct and indirect signs. Among rodents, abundance of Indian gerbil was high (20/Km2) followed by house mouse
(17.5/Km2) and lowest abundance was recorded for Turkistan rat (2.5/ Km2).
Abundance of rhesus monkey was (11.1/Km2), followed by porcupine (8.5/Km2), wild boar (5.4/Km2) whereas, Asian palm civet had lowest abundance (0.6/Km2).
Among bird species, abundance of house sparrow was high (158/ Km2) followed by
Himalayan bulbul (36/ Km2), red-vented bulbul (29/ Km2) and least was recorded for Indian peafowl (2.5/ Km2). Abundance of all amphibians was (78.1/ Km2) and all reptiles were (89.4/ Km2). Abundance of invertebrates showed that members of order hymenopteran were more abundant (2310.3/ m2) followed by orthoptera
42.26/m2 and Coleoptera (22.14/m2) and abundance of snails was (3.4/Km2).
Abundance of domestic hen (excluding population in farms) was (548.69/Km2) followed by goats (105.17/Km2) and cows 43.07/ Km2) and least abundant was mule
(0.03/ Km2). Abundance of dog was (22/ Km2).
Chapter 5
NICHE BREADTH AND NICHE OVERLAP AMONG
SYMPATRIC CARNIVORE SPECIES OF PIR LASURA
NATIONAL PARK
5.1 INTRODUCTION
Carnivores belong to order Carnivora which is diverse order of eutherian mammals. Carnivores are primarily meat-eaters, although few species are often omnivorous such as mongooses, foxes and civets (Hinton and Dunn, 1967; Barun et al., 2008; Mahmood et al., 2011). Carnivores range in size from the least weasels to elephant seals. They are known to predate on a variety of prey species. Knowledge of diet composition is prerequisite for conservation of species (Oli, 1993). Diet composition of carnivores is usually investigated using their fecal droppings
(Shehzad et al., 2015). Faecal components of carnivores may include mammalian remains such as hairs and bones, feathers of birds, teeth, claws, invertebrates and plant matter, mucus and bacteria (Bang and Dahlström, 1975; Bujne, 2000).
Whereas, the size and the quantity of feces produced by each individual varies with age, the type of ingested food, and its absorption capacity (Bang and Dahlström,
1975).
Dietary overlap between carnivores can indicate level of interspecific competition. Manipulative studies can help to understand competition (Mac-Nally,
1983; Wiens, 1989), data on niche overlap is essential first step (Carrera et al., 2008;
Glen and Dickman, 2008). High overlap among carnivore species indicate level of intraspecific competition as well as strength of interference competition. Predators
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having similar diet often exhibit aggression, in order to eliminate competitor or predators may often encounter each other while searching for similar prey (Donadio and Buskirk, 2006; Polis et al., 1989). Carnivore living in premises of human occupied landscapes are reported to take up domestic livestock as prey (Sangay and
Vernes, 2008; Shehzad et al., 2015) and such cases usually result in antagonistic human carnivore interaction.
Extent of resource partitioning depends upon difference in body size of the species involved (Schoener, 1974). The divergence in morphological traits is often associated with the divergence in ecological traits, for example, food (Brown and
Wilson, 1956; Sih, 1993; Dayan and Simberloff, 1998). Since larger animals consume larger prey and variety of eaten prey is greater as compared to the smaller animals, larger animals have broader tropical niche breadth as compared to the smaller (Schoener, 1974). In Potohar region of Pakistan, the grey mongoose has a wider food niche breadth as compared to the small Indian mongoose. Food niche breadth was wider during summer season and low during winter season for grey
Indian mongoose and small Indian mongoose. It shows that more numbers of prey species were available during summer season as compared to winter season. High niche overlap between two mongoose species was recorded which was 0.95 (Hussain et al., 2017).
Association between body size and food resource partitioning has been previously reported in fishes (Ross, 1986; Schafer et al., 2002), invertebrates
(Woodward and Hildrew, 2002; Tall et al., 2006), shrews (Brannon, 2000) and carnivores (Martin, 1994; Jones, 1997; Dayan and Simberloff, 1998; Jones and
Barmuta, 2000; Garcia and Virgos, 2007). Sympatric small carnivores consume rodents as their major food source in common and take other secondary food items in size corresponding to their body size.
Knowledge of resource partitioning among sympatric carnivores is very important from conservation point of view. The Pir Lasura National Park in AJ&K is an important area regarding distribution of some important carnivore species including common leopard, Asiatic jackal, red fox and mongooses. Despite of having rich biodiversity, Pir Lasura National Park has not been focused for detailed scientific wildlife studies in the past. There is paucity of information on how resources are shared between different carnivore species found in the Park.
Therefore, the current study was designed to investigate dietary niche breadth and niche overlap among sympatric carnivores’ species at Pir Lasura National Park, Azad
Jammu and Kashmir, Pakistan.
5.2 REVIEW OF LITERATURE
Hussain et al. (2017) studied resource partitioning among two sympatric mongoose species in the Pothwar region, Pakistan. According to results of their study, the grey mongoose had a wider food niche breadth as compared to the small
Indian mongoose. Food niche breadth was wider during summer season and low during winter season for both mongoose species. It showed that more numbers of prey species were available during summer season as compared to winter season.
They found high niche overlap between two mongoose species using Pianka’s Index and niche overlap between the two species was 0.95.
Resource availability affects resource partitioning, generally, the degree of diet overlap among sympatric carnivore will increase with increase in abundance of common food and decrease with lean time (Schoener, 1974; Wiens, 1989; Smith,
1991) as the species will shift their food to different food items during the lean time.
In addition, feeding strategy of different animals may also affect the results of resource partitioning; for example, opportunistic generalists may consume all the remaining food resources. If these available food resources are similar among patches, as is the case usually during lean time, overlap in diet of these species will increase (Serafini and Lovari, 1993; Padial et al., 2002). But in a community, that consists of both generalists and specialist predators, diet overlap may increase or decrease between different species during lean time (Carvalho and Gomes, 2004).
5.3 MATERIALS AND METHODS
Current study was conducted at Pir Lasura national park, Azad Jammu and
Kashmir, Pakistan. We investigated diet composition of seven sympatric carnivore species at Pir Lasura national park, Azad Jammu and Kashmir, Pakistan. We used molecular identification technique to confirm for the carnivore species. Diet of seven carnivore species was studied by analyzing a total of 473 scats samples collected from the field. Among those, were 39 from common leopard, Asiatic jackal 64,
Kashmir hill fox 92, Asian palm civet 108, small Indian civet 44, Indian grey mongoose 57, and small Indian mongoose 69 scat samples. In this chapter, we computed niche breadth and niche overlap among different carnivore species.
5.3.1 Niche Breadth
Niche breadth (L) is a parameter which attempts to measure that in a specific environment, to what extent a species is specialized or unspecialized. Specialist species will have few food sources in their diet and eventually will have small niche breadth. A generalist that feeds on many kinds of food items will have wider niche breadth. We measured dietary niche breadth of seven sympatric carnivore species at
Pir Lasura National park, using niche breadth (L) and standardized Levins index (0-
1) (Lst) (Levins, 1968; Colwell and Futuyma, 1971) as follows:
Where pi is the relative percentage of food item i and n is the number of food items.
Lst is standardized niche breadth and its value ranges from 0 to 1. A higher
Lst indicates broader diet niche of the animal.
5.3.2 Niche Overlap
We used the frequency of occurrence of each prey item to compute dietary overlap between seven sympatric carnivore species (including common leopard,
Asiatic jackal, Kashmir hill fox, Asian palm civet, small Indian civet, Indian grey mongoose and Small Indian mongoose) occurring at Pir Lasura National Park, Azad
Jammu and Kashmir, Pakistan using Pianka’s index. Pianka’s index ranges from zero
(no overlap) to one (complete overlap) (Pianka, 1973). We chosed this index to allow direct comparison of the degree of overlap in similar studies of carnivores conducted
elsewhere in the world (Fedriani et al., 2000; Ray and Sunquist, 2001; Jacomo et al.,
2004). The Pianka’s index was calculated using formula:
Where pij (or pik) is the relative percentage of food item i in diet j (or k).
The Pianka’s index ranges from 0 to 1 and the higher value indicates higher degree of overlap in diet whereas lower values indicate lower degree of overlap in diet of target carnivore species.
5.4 RESULTS
5.4.1 Dietary Niche Breadth
Among seven different carnivores of the study area, niche breadth of Indian grey mongoose was widest 18 (0.72) followed by Asiatic jackal 14.2 (0.78), common leopard 13.88 (0.72), small Indian mongoose 12 (0.64), small Indian civet 10(0.52),
Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31). (Table 5.1; Fig. 5.1)
The dietary niche breadth of common leopard was found broad (Table 5.1;
Fig. 5.1) during spring season 14.76 (0.76) and narrow during winter season
9.33(0.46). Total niche breadth of common leopard was 13.88 (0.72).
Similarly, Niche breadth of Asiatic jackal (Table 5.1; Fig. 5.1) was found broad during summer season 17.25(0.96)) but narrow during spring season 6.67
(0.33). Total niche breadth of the Asiatic jackal was 14.2 (0.78).
Niche breath of Kashmir hill fox (Table 5.1; Fig. 5.1) was wider 9.07 (0.37) during autumn season but narrow during spring season 4.8 (0.17). The overall niche breadth of fox was 7.89(0.31).
Niche breadth of Asian palm civet (Table 5.1; Fig. 5.1) was wide during autumn season 7.82 (0.243) but narrow during winter season 7.53 (0.23). Overall niche breadth of Asian palm civet was 9.69 (0.31).
Niche breadth of small Indian civet (Table 5.1; Fig. 5.1) was wider 9.6 (0.48) during summer season but narrow during autumn season 4.8 (0.21). Total niche breadth of small Indian civet was 10 (0.52).
Niche breadth of Indian grey mongoose (Table 5.1; Fig. 5.1) was broad 20
(0.83) during autumn season but narrow 9.3 (0.36) during winter season. Overall, niche breadth of Indian grey mongoose was 18(0.72).
Table 5.1 Niche breadth of sympatric carnivores occurring in and around Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan
L= Niche breadth; Lst= standardized niche breadth (Value 0-1).
Sr. Species Summer Autumn Winter Spring Total
no. Niche breadth Niche breadth Niche breadth Niche breadth Niche breadth
L (Lst) L (Lst) L (Lst) L (Lst) L (Lst)
1 Panthera pardus 11.68 (0.59) 11.88(0.6) 9.33(0.46) 14.76(0.76) 13.88(0.72)
2 Canis aureus 17.25(0.96) 16.67(0.92) 10.9(0.58) 6.67(0.33) 14.2(0.78)
3 Vulpes Vulpes 6.77(0.26) 9.07(0.37) 5.2(0.19) 4.8(0.17) 7.89(0.31)
4 Paradoxurus hermaphroditus 7.72(0.24) 7.82(0.243) 7.53(0.23) 7.69(0.24) 9.69(0.31)
5 Viverricula indica 9.6(0.48) 4.8(0.21) 8.65(0.43) 6.72(0.32) 10(0.52)
6 Herpestes edwardsii 16(0.64) 20(0.83) 9.3(0. 36) 12(0.47) 18(0.72)
7 Herpestes javanicus 11.64(0.59) 11.02(0.56) 10.41(0.52) 9.35(0.46) 12(0.64)
Summer Autumn Winter Spring Total
0.96
0.92
0.83
0.78
0.76
0.72 0.72
0.64 0.64
0.6
0.59 0.59
0.58
0.56
0.52 0.52
0.48
0.47
0.46 0.46
0.43
0.37
0.36
0.33
0.32
0.31 0.31
0.26
0.243
0.24 0.24
0.23
0.21
0.19 0.17
C O M M O N A S I A T I C K A S H M I R ASIAN PALM S M A L L INDIAN GREY S M A L L L E O P A R D JACKAL H I L L F O X C I V E T INDIAN CIVET MONGOOSE I N D I A N MONGOOSE
Figure 5.1 Niche breadth of sympatric carnivore species occurring at Pir Lasura
National Park, Azad Jammu and Kashmir, Pakistan.
Dietary niche breadth of small Indian mongoose (Table 5.1; Fig. 5.1) was found broad during summer season 11.64(0.59) but narrow during spring season
9.35(0.46). Total niche breadth of small Indian mongoose was 12(0.64) (Table 5.1;
Fig. 5.1).
5.4.2 Niche Overlap
Among the seven different sympatric carnivore species at PLNP, AJ&K, high niche overlap was found between Asian palm civet and small Indian civet (0.9),
Indian grey mongoose and small Indian mongoose (0.89), Asiatic jackal and
Kashmir hill fox (0.81), Kashmir hill fox and small Indian civet (0.66), common leopard and Asiatic jackal (0.62), Kashmir hill fox and Indian grey mongoose (0.62), while lowest niche overlap was recorded common leopard and small Indian civet
(0.04) (Table 5.2; Figure 5.2).
5.5 DISCUSSION
Knowledge of a predator’s diet is important for its conservation and management. We investigated diet composition of seven sympatric carnivore species at Pir Lasura national park, Azad Jammu and Kashmir, Pakistan. Diet of seven carnivore species was studied by analyzing a total of 473 scats samples collected from the field. Among those, were 39 from common leopard, Asiatic jackal 64,
Kashmir hill fox 92, Asian palm civet 108, small Indian civet 44, Indian grey mongoose 57, and small Indian mongoose 69 scat samples. Diet of common leopard comprised of 17 prey species (10 wild and 7 domestic). Frequency of occurrence of wild prey was 34.85 % of total leopard diet whereas domestic prey contributed
59.1%. Sixteen species of mammals, birds, insects and plants were identified from
diet of Asiatic jackal. Among these species 10 were wild, 5 were domestic and
Table 5.2 Niche overlap between sympatric carnivore species occurring in and around Pir Lasura National Park, Azad Jammu and Kashmir,
Pakistan
Canis aureus Vulpes vulpes Paradoxurus Viverricula Herpestes Herpestes
griffithi hermaphroditus indica edwardsii javanicus
Panthera pardus 0.62 0.38 0.09 0.04 0.072 0.086
Canis aureus 0 0.81 0.23 0.35 0.58 0.56
Vulpes vulpes griffithi 0 0 0.55 0.66 0.62 0.55
Paradoxurus hermaphroditus 0 0 0 0.9 0.37 0.29
Viverricula indica 0 0 0 0 0.54 0.43
Herpestes edwardsii 0 0 0 0 0 0.89
Figure 5.2 Niche overlap between sympatric carnivore species at Pir Lasura National
Park, Azad Jammu and Kashmir, Pakistan.
APC = Asian palm civet; SIC = small Indian civet; IGM = Indian grey mongoose;
SIM = small Indian mongoose; Jack = Jackal; Fox = Kashmir hill fox; L = common leopard.
1 species of plant. Frequency of wild prey in diet of the jackal was 18.48% and domestic was 59.78%. We recorded 21 species of mammals, birds, plants and insects in diet of the red fox. Among these 21 species 10 were wild, 5 domestic and 6 plant species. Frequency of occurrence of wild prey was 17.96%, domestic prey 50.9%, plants 27.54%. Analysis of 108 scats samples of Asian palm civet showed that 27 species of mammals, birds, invertebrates and plants were consumed. Among all recorded species 9 were wild, 3 domestic, and 15 plant species. Consumption of wild prey was 33.68% compared to 10.88% domestic prey species and 53.37% plants.
Analysis of 44 scats showed that 17 prey species occurred in diet of small Indian civet. Among them 8 were wild, 1 domestic, and 8 were plant species. Frequency of occurrence of wild prey in diet of small Indian civet was 30.65%, domestic prey
14.52%, and plants 51.61%. Analysis of 57 scat samples showed that 22 prey species occurred in diet of small Indian mongoose. Among them 15 were wild, 1 domestic, and 6 plant species. Diet of Indian grey mongoose consisted of (60%) wild species,
(19%) domestic prey species, and (14%) plant species. Analysis of 69 scat samples of small Indian mongoose showed that 17 prey species occurred in diet of small
Indian mongoose. Among them 10 were wild prey species, only 1 domestic prey species and 6 plant species. Frequency of occurrence of wild prey was (59.68%), domestic prey (16.94%) and plant matter (11.29%).
Knowledge of resource partitioning among sympatric carnivores is very important from conservation point of view. We computed niche breadth and overlap among sympatric carnivore species at Pir Lasura national park. Among seven different carnivores of the study area, niche breadth of Indian grey mongoose was
widest 18 (0.72) followed by Asiatic jackal 14.2 (0.78), common leopard 13.88
(0.72), small Indian mongoose 12 (0.64), small Indian civet 10(0.52), Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31).
Dietary niche breadth of common leopard was broad during spring season which is indicative of the fact that more prey species were available to this top predator in the study area during spring season. But during winter season, the narrow dietary breadth of common leopard indicates availability of less prey species. For
Asiatic Jackal, on contrary, more prey species were available during summer season as indicated by its broad niche breadth and less availability of prey species during spring season, represented by narrow niche breadth.
Niche breath of Kashmir hill fox was wider during autumn season showing availability of more prey species but narrow during spring season, shown by less availability of prey species. A survey of past published literature shows lacking scientific studies regarding quantifying niche breadth of the three above mentioned carnivores (common leopard, Asiatic Jackal, and red fox) in the country. So, now, the results of the current study cannot be compared with other data. The current study, therefore, reports here the niche breadths of the three mentioned carnivores from the country for the first time. Lanszki et al. (2006) reported that trophic niche breadth of golden jackal was (0.06) and red fox was (0.09) in Hungary.
Niche breadth was found broad for the Asian palm civet, during autumn season but narrow during winter season. These findings of the current study are supported by some previous published studies such as Aroon et al. (2012) reported that niche breadth of Asian palm civet was widest during autumn season (B = 0.169),
the current study also recorded high niche breadth of this species during autumn season as more prey species are available during this season. This species is frugivorous in dietary habits and ample quantity of fruits are present during autumn season at Pir Lasura National Park. Aroon et al. (2012) reported narrow niche breadth during winter season which is like the findings of the current study that its niche breadth is narrow during winter season. Overall, they reported dietary niche breadth being 0.412 (Aroon et al., 2012). In the current study, we found overall dietary niche breadth of Asian palm civet being 0.31, which might be because of the variation of habitat and diversity of prey species available in specific study area. Niche breadth of civets was narrow during winter when there was low availability of fruits and high during fruiting seasons.
The overall niche breadth of small Indian civet was 10 (0.52); wider niche breadth of small Indian civet during summer and narrow during autumn season in
Pir Lasura National Park can be compared with some earlier studies; Chuang and
Lee (1997) reported that dietary breadth of small Indian civet was 4.46 in northern
Taiwan. Another study reporting its niche breadth as 2.58 and it consumed 9 prey species; including rodents, birds, invertebrates, and plants matter (Wang and Fuller,
2003). Small Indian civets when living in outskirts of villages and cultivated land, may predate on more diverse diet (Wang, 1998).
Niche breadth of Indian grey mongoose was found broad 18(0.72) during autumn season and narrow 7.8(0.29) during winter season. Hussain et al. (2017) studied niche breadth and niche overlap between two mongoose species (Herpestes javanicus and H. edwardsii) in Potohar region, Pakistan. They reported that there
was seasonal variation in niche breadth of Indian grey mongoose and it was high during summer season and narrow during winter season.
Dietary niche breadth of small Indian mongoose was broad during summer season 11.64 (0.59) and narrow during spring season 9.35 (0.46). Total niche breadth of small Indian mongoose was 12 (0.64). Hussain et al. (2017) reported niche breadth of small Indian mongoose being variable during different seasons and it was highest
(7.2) in summer but lowest (6.1) in winter. The findings of the current study have also shown variation in niche breadth of small Indian mongoose during different seasons being high during summer season. But unlike Hussain et al. (2017), narrow niche breadth was recorded during spring season, which could be attributed to habitat variability and to variation in prey species availability during different seasons.
Regarding comparison of dietary niche overlap among seven different carnivore species sympatric at PLNP, AJ&K, high niche overlap was found between
Asian palm civet and small Indian civet (0.9), indicating that both species consume similar food and they compete strongly for resources at Pir Lasura National Park.
Niche overlap among small Indian mongoose and Indian grey mongoose was 0.89.
There are not many previously published records to compare such findings except
Hussain et al. (2017) who reported that there is high niche overlap among two mongoose species in the Pothwar region, Pakistan and the current study have also recorded similar findings that both mongoose species compete for resources. Niche overlap among Asiatic jackal and Kashmir hill fox was also high (0.81) showing both species competing for same resources in the study area. Lanszki et al. (2006) reported that trophic niche overlap among jackal and fox was 0.73 in Hungary. But
it can be argued that if some food item is shared but the two predators at the moment are not competing for it, however, in future, this resource may become short in environment due to some reason and that time the same two predators will start competing for it, since it is included in the food habits of both the predators.
Niche overlap among Kashmir hill fox and small Indian civet was (0.66) since both carnivore species consume similar prey species like rodents and insects.
Although the difference of body size occurs between both species, however, both species consumes same prey species therefore there is competition among them.
Similarly, because of consuming shared resources niche overlap among common leopard and Asiatic jackal was (0.62). However, this can also be attributed to the scavenging nature of jackal as it feeds on carrion of domestic animals and leopard predate on livestock as indicted by high frequency of domestic prey in leopard diet.
Niche overlap of Kashmir hill fox and Indian grey mongoose was (0.62) which can be result of feeding on rodent species as they were recorded in diets of both carnivores.
Spatial avoidance is one of evolutionary solutions for coexistence of carnivores in an area (Hunter and Caro, 2008), however coexisting is also possible by consuming prey species of different weight class or different taxa (Karanth and
Sunquist, 2000; Hayward and Kerley, 2008). We recorded that though the niche of carnivore species overlaps in the study area, however, their distribution in the study area indicated by relative abundance of their signs was not evenly distributed in all study sites. Signs of different species were found abundant at different sites rather
than at a single site which shows that these carnivore species occupy different areas and they occur different elevations to avoid competition.
5.6 CHAPTER SUMMARY
Studies on dietary niche breadth and niche overlap among sympatric carnivore species are vital from conservation point of view. We measured dietary niche breadth and overlap of seven sympatric carnivore species (including common leopard, Asiatic jackal, Kashmir hill fox, Asian palm civet, small Indian civet, Indian grey mongoose and Small Indian mongoose) occurring at Pir Lasura national park,
Azad Jammu and Kashmir, Pakistan. For measuring dietary niche breadth, we used standardized Levins index and to compute dietary niche overlap we used Pianka’s index. Among seven different carnivores of the study area, niche breadth of Indian grey mongoose was widest 18 (0.72) followed by Asiatic jackal 14.2 (0.78), common leopard 13.88 (0.72), small Indian mongoose 12 (0.64), small Indian civet 10(0.52),
Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31). High niche overlap was found between Asian palm civet and small Indian civet (0.9), Indian grey mongoose and small Indian mongoose (0.89), Asiatic jackal and Kashmir hill fox
(0.81), while lowest niche overlap was recorded common leopard and small Indian civet (0.04). The current study concluded that niche of seven sympatric carnivore species overlaps in Pir Lasura National Park, Azad Jammu and Kashmir, Pakistan.
Chapter 6
ASSESSMENT OF HUMAN-CARNIVORE CONFLICT AT PIR
LASUARA NATIONAL PARK
6.1 INTRODUCTION
In the face of ever shrinking wildlife habitat, increasingly encroached upon by humans and livestock, human-carnivore interactions are often antagonistic. Such interactions have become noteworthy conservation as well as rural livelihood issues since many carnivores have been affected by retaliatory killings in response to livestock depredation (Dar et al., 2009). The main reason for conflict between villagers and conservationists is over livestock depredation (Graham et al., 2005).
The root cause of livestock depredation arises because some carnivore species are specialized to consume ungulates as prey and therefore, they readily kill domestic ungulates (Meriggi and Lovari, 1996; Karanth et al., 1999; Polisar, 2000).
There is conflict between the herders/ farmers and conservationists but not the common leopard; the conflict between the parties arises because common leopard kills livestock (Redpath et al., 2015; Fisher, 2016). Big cats are more susceptible to such issues as they have larger home ranges and broader niche breadth (Linnell et al., 2001; Macdonald and Sillero-Zubiri, 2002 ). Such issues may become worse by factors like increase in human population and loss of carnivores natural habitat
(Athreya and Belsare, 2007; Inskip and Zimmermann, 2009) or reduced wild prey abundance. Moreover, level of livestock depredation by carnivore species is elevated in the regions where humans live in surroundings of protected areas (Linnell et al.,
2001; Mishra, 2001; Conforti and Azevedo, 2003), and such issues often result
157
in mortality of carnivore species in and around protected areas (Gurung et al., 2008).
Predation on livestock during scarcity or absence of natural prey brings carnivores into direct conservation concerns with humans. It happens in areas where habitat and native prey of carnivores have been lost due to anthropogenic activities
(Patterson et al., 2004). Thus livestock depredation is one of the biggest issues due to which populations of big cats are declining (Nowell and Jackson., 1996; Inskip and Zimmermann, 2009). Such livestock depredation by carnivores impacts rural communities financially because people living in close proximity to carnivore habitat are mostly included in the lowest income category. Any loss of their livestock to carnivores places an economic burden for poor villagers. Furthermore, attacks of carnivores even on humans result in severe injuries or deaths (Bibi et al., 2013). As a result, negative perception develops among native people towards carnivores
(Naughton-Treves, 1998; Linkie et al., 2007; Dar et al., 2009; Kabir et al., 2014).
Attempts to mitigate villager-conservationists conflict should be based on explicit understanding of underlying spatio-temporal patterns of livestock depredation (Dar et al., 2009; Kabir et al., 2014). Carnivores were once widely distributed across Azad Jammu and Kashmir (North-East Pakistan) and in other parts of the country but more recently its distribution range has shrunk because of their persecution over livestock depredation behavior. Consequently, they have either declined or vanished over enormous areas of their past known range (Nowell and
Jackson., 1996; Roberts, 1997; Sheikh and Molur, 2004; Dar et al., 2009).
The depletion of big cats makes it critical to seek out mitigation approaches that may increase sustainable co-existence of both local communities and big cats
(Mishra et al., 2003). This situation is important particularly in case of Pir Lasura
National Park (PLNP) in AJ&K with high human density living inside and around the Park. There is paucity of baseline knowledge on different dimensions of human carnivore interactions in this National Park, therefore, the current study aimed at investigating human-carnivore interactions in and around PLNP during 2014-15, with retrieval of previous data over a period of last eight years (2008 to 2015) using questionnaire survey. The particular objectives included investigating the degree of livestock depredation by different carnivores, numbers of carnivores persecuted as a result of livestock depredation, the temporal patterns of livestock depredation by carnivores, economic losses to local community as a result of carnivore attacks, and local perception towards carnivore species in the study area.
We hypothesized that “more livestock may be depredated by common leopard than any other predators in the area owing to its larger body size”. Livestock depredation by carnivores was also likely to be influenced by temporal variation in the study area as well as guarding strategies being practiced, herd size and penning conditions. In addition, livestock depredation by carnivores may have negative impact on local perception resulting in killing of carnivores in retaliation.
6.2 REVIEW OF LITERATURE
Human-carnivore conflict is common in the areas where large carnivores occur in the surrounding areas of human habitation. A few studies have been documented from different parts of the country highlighting the conflict. Dar et al.,
(2009) studied patterns of human-carnivore conflict in the Machiara National park
(MNP), AJ&K. Statistics showed that four different carnivore species were
responsible for livestock losses, among those; common leopard caused greater loss
>90%. Major livestock species depredate by leopard were goats (57.3%) and sheep
(27.8%). Villages deprived of electricity faced more leopard attacks and vice versa.
Leopard attacked larger herds more frequently, and different guarding strategies used did not reduce leopard attacks. Leopard attacks caused less livestock loss as compared to disease. Chattha et al., (2013) reported that in Machiara National Park
AJ&K, leopard mainly attacked goats (50.50%) and sheep (28.53%). Livestock grazing, poor husbandry practices were identified as major cause of human-leopard conflict.
Bibi et al., (2013) studied human-carnivore interaction in Dhirkot Azad
Jammu and Kashmir, Pakistan. Three carnivore species were responsible for livestock killing and human injury. Common Leopard was responsible for majority of livestock losses and including mainly goats (20%) and donkeys (18%). Four persons were injured because of leopard attack which resulted in killing of four leopards by local community.
Kabir et al., (2014) studied human leopard conflict at Machiara Natinal Park
AJ&K. common leopard killed 301 in 2007-2008 and majority of killing was recorded during May. Leopard killed mainly goats and sheeps. More attacks were reported during the night time. Attitudes of the local community towards leopards was negative.
6.3 MATERIALS AND METHODS
6.3.1 Methods
Before conducting the questionnaire survey, written permission was
Figure 6.1 Locations of livestock depredation and retaliatory killing of common
leopard (Panthera pardus) during current study period in and around Pir
Lasura National Park, Azad Jammu and Kashmir, Pakistan.
obtained from the Department of Wildlife Management, PMAS Arid Agriculture
University Rawalpindi, Pakistan, for collecting data from local people regarding livestock depredation by carnivores in the study area. Since the current research involved fieldwork in the area occupied by the people, and it could affect species or ecosystems within which people have de facto or de jure tenure rights or cultural connections, therefore, it was carried out in a way that respected the local beliefs, economic and cultural interests, and rights.
At the start of the current study, before administering the questionnaire surveys, a meeting was held with the Department of Wildlife and Fisheries staff to distinguish the towns that had suffered losses of livestock to carnivores during previous eight years (from 2008 to 2015). To avoid bias during data collection, locals were taken into confidence by clarifying that data will be collected independently and their cooperation will not harm them in future.
6.3.1.1 Questionnaire survey
Information on human-carnivore interactions in the study area was gathered by utilizing both subjective and quantitative techniques (Bibi et al., 2013). The subjective strategies included unstructured meetings, participatory perception and group discussions, while quantitative techniques contained organized close end questionnaire (Appendix-VIII), which was the principle information source. We used 170 questionnaires to collect data during April 2014 to March 2015 and respondents were selected using consecutive sampling method. As there was no previous baseline, we also collected data for past eight years (2008-2015) from respondents’ best recollections. To avoid bias we did not include the data when
people were not able to recall or were not able to provide accurate information required for this study (many respondents were not able to remember correctly the year, month, time and location of attacks; in such cases information was not included in analysis). We spent 25 minutes (on average) to collect information from each respondent. The fundamental goals of the questionnaire were to investigate aspects of human–carnivore interactions, sighting of species and perceptions of local communities towards the primary predator species. To confirm identification of predator, we used pictures of carnivore species which were provided to each respondent to identify the predator species. Information was also gathered on demographic and financial state of respondents, livestock depredation pattern, its resulting economic burden on respondents, incidents of retaliatory killing, local knowledge, and resilience towards carnivore species.
Data on demographic and financial elements, for example, name, sex, age, level of formal training, occupation, sort, number and motivation behind keeping animals were gathered for all respondents. Interviews of respondents were kept confidential. Questions that were asked of respondents included direct sightings of carnivore attacks, and indications of carnivore attack at the site of the livestock mortality (including evidence such as damage to the throat or neck of the killed animal). In addition, respondents were asked for information about the sort and number of animals killed, number of predators that attacked, date, time and period of attack, spot and area of attack either inside village or outside village; in forest or knoll, or grazing near village, action taken against the carnivore species responsible for attack, livestock guarding, and whether fencing exists to prevent predator access.
Additional data included presence of electricity and whether livestock were penned
or not at evening. The areas of leopard depredation on livestock reported during questionnaire survey were visited and their geographic coordinates were recorded using GPS (Garmin etrex 10).
Information was simultaneously collected on indigenous perception concerning: most problematic carnivore species responsible for majority of livestock predation, spatio-temporal attack patterns of major challenging carnivore species, explanation underlying increase or decrease in livestock depredation by carnivores during past few years, relationship between attacks and PLNP, acquaintance of respondents about authority accountable for protecting key predator species in PLNP and also inclination of respondents to conserve these predator species in the Park.
Attitudes towards conservation of the key predator species was assessed using the approach of Treves and Karanth (2003). Following Treves and Karanth
(2003), we gave respondents five conceivable management choices for relieving human-carnivore interaction in PLNP: firstly do nothing; secondly repulse the predator utilizing troublesome stimulants; thirdly habitat improvement for the predator; fourthly translocation of the predator and lastly lethal control.
For every predator type, total numbers and types of livestock killed were recorded using self-designed questionnaires. Human injuries and deaths caused by carnivores and numbers of predators executed were recorded. If carcass of a predator was not found it was considered poaching; but if the carcass of predator was recovered and found confiscated by the authorities, it was considered as retaliatory killing. Data on monthly livestock depredation were then assigned to seasons. The mean monthly temperature and rainfall data of the study area were retrieved from
Pakistan Meteorological department for analysis. The budgetary losses of every respondent because of carnivore attack on livestock were computed considering local market price (in Pakistani Rupee converted into US $; average price of goat = $ 200
(Rs. 21000/-), cow, buffalo and ox = $ 2402.8 each (Rs.252000/-), horse = $ 2884.5
(Rs. 300,000/-), Dog = $ 96.12 (Rs. 10,000/-), rabbit = $4.72 (Rs. 500/-). The livestock prices also considered sex, age and state of livestock killed by predator.
Sites of leopard depredations on livestock and incidences of retaliatory killings were visited in person to confirm information provided by the local people and geographical coordinates of all sites were recorded to construct a GIS-based map using Quantum Geographical system (Q-GIS) software version 1.8.0. Lisboa and
Arc GIS software version 10.1 (Figure 6.1).
6.3.1.2 Statistical analysis
We performed a Spearman’s rank correlation with mean monthly predator attacks against mean monthly temperature of the study area in order to check if there was any seasonality on livestock depredation by carnivores, and whether mean rainfall per month affects livestock depredation by carnivores. Likewise, we tested for a correlation between the numbers of livestock losses to leopard and elevation of study sites with reported livestock depredation. As key prey species such as Rhesus monkey and Barking deer were mostly recorded at higher elevations (1300m –
2000m) we aimed to evaluate if elevation correlated with depredation events by leopard. The positive correlation means that leopard attacks on livestock increased with increasing the particular parameter such as temperature or rainfall while the negative correlation means the vice versa.
We used one-way Analysis of Variance (ANOVA) and post hoc LSD tests to evaluate leopard depredation on livestock during: a) different hours of the day; b) different months; and c) different seasons. For the main livestock predators, one-way
ANOVA and post hoc LSD test were performed to investigate whether total livestock loss outside villages were related to herd size or not.
The impact of penning livestock during the predator attack was tested using
Chi square goodness of fit test. The influence of numbers of livestock losses to main predators, financial loss to main predators on local tolerance towards predator management was tested using Chi square. The effect of different guarding strategies
(none, human only, dog only, both human and dog) used and local perception regarding management of common leopard in their surroundings was tested using
Chi square goodness of fit test. All analysis was conducted in software package SPSS version 23.
We quantified extent of leopard depredation on livestock by means of a kernel density analysis. Kernel density analysis expresses numbers of livestock losses to common leopard per kilometer square of study area. Mapping kernel density allowed us to identify areas suffering from livestock losses because of depredation by leopard and identifying areas where mitigation measures should be set up. To estimate a density value, we used bandwidth of 2000m as a search radius to calculate rate of livestock depredation per square kilometer and cell size was 20m.
This 2000m radius is logical and reasonable to identify priority areas where mitigation measures should be set up (Figure 6.2). Estimated density values were then classified using Jenks methods (Jenks and Caspall, 1971). All analysis was conducted in ARC GIS software version 10.1.
6.4 RESULTS
We interviewed 170 respondents who had lost their livestock, poultry, dogs or pets during last eight years from 2008-2015 in and around PLNP, AJ&K. The majority of respondents (82.94%) belonged to major classes of rural livelihood including; shopkeepers, farmers, government employee and labor. Average number of livestock per respondent was 12.6 livestock heads.
Results showed the respondents had lost a total of 306 animals over 8 years as a result of carnivore attacks. Majority of attacks (70.58%) occurred at six study sites which included; Panag gali (23.52%) followed by Karaila panjal gali (14.11%),
Pir kana (11.17%), Sairi (8.82%), Kallar galla (6.47%) and Chitibakri (6.47%) (Fig.
1). Attacks of leopard were not found correlated with elevation of the study site (r =
0.14). Kernel density analysis showed that Pir Kana, Sairi, Katera khandhar, Pehli gali, and Pana gali were the areas where high livestock depredation/km2 by common leopard was reported (Figure 6.2).
6.4.1 Major Predator
Common leopard (Panthera pardus) emerged as major predator of the livestock and it was responsible for majority of livestock losses (88.56%), followed by Asiatic Jackal (Canis aureus; 5.22%), Small Indian Civet (Viverricula indica;
3.22%) and the Indian grey mongoose (Herpestes edwardsii; 2.94%). Most of the respondents reported (65%) raiding on fruits in their gardens at night by civets. The common leopard preyed upon goats (82.22%), dogs (9.22%), rabbit (4.05%), domestic cow (1.84%), buffalo (1.10%), ox (1.10%) and horse (0.73%; Table 6.1).
Majority of leopard attacks occurred at day time (56.36%) when livestock
Figure 6.2 Kernel density estimation for livestock depredation by common leopard
in and around Pir Lasura National Park. Search radius was 2000 m whereas,
cell size used was 20m. Red areas shows areas facing high livestock
depredation by common leopard where mitigation measures should be set up.
Table 6.1 Livestock, dog, pet and poultry killed by various carnivore species in and around Pir Lasura National Park (PLNP), Azad Jammu
and Kashmir during 2008-2015.
Sr. No. Predator species Goat Cow Buffalo Ox Horse Dog Rabbit Poultry Total % of total
1 Common Leopard 222 5 3 3 2 25 11 - 271 88.56
(Panthera pardus)
2 Asiatic Jackal ------16 16 5.22
(Canis aureus)
3 Small Indian Civet ------10 10 3.26
(Viverricula indica)
4 Grey Indian mongoose ------9 9 2.94
(Herpestes edwardsii)
Total 222 5 3 3 2 25 11 35 306 100
was grazing outside the villages of the study area while 43.63% attacks occurred at night (inside villages).
6.4.2 Daily Attack Pattern
Temporal patterns of common leopard attacks showed that it attacked livestock during 9:00-12:00 h (n=72, 43.63%), 14:00-16:00h (n=36, 21.81%) and
19:00-22:00h (n = 42, 25.45%) (Figure 6.3), showing statistically significant difference (F=1.89, df = 23, p < 0.05). LSD test showed a significant difference (P <
0.05) between daytime predation and predation at other times (evening and night).
6.4.3 Monthly Attack Frequency
Livestock depredation by common leopard were observed to occur throughout the year with highest numbers of livestock being killed during the month of July (n = 36, 21.81%), followed by in May (n = 33, 20%), June (n = 31, 18.78%) and December (n = 18, 10.90%) (Figure 6.4). LSD test showed significant difference between livestock depredation during the month of July (P < 0.05). Monthly data of leopard depredation on livestock was correlated with mean monthly temperature and we found weak correlation (r = 0.45, n=12, P=0.15). However, we found no correlation between mean monthly leopard attacks and monthly rainfall in the study area (r = 0.013, n = 12, P=0.97).
6.4.4 Seasonal And Annual Variation
Majority of leopard attacks (60.60%) on livestock occurred during summer season (May, June, July), followed by winter (November, December, January)
(19.39%), and spring (February, March, April) (15.15%) but least in autumn season
(August, September, October) (4.84%) (Table 6.2).
1 2430 2 23 3 25 22 4 20 21 15 5 20 10 6 5 19 0 7 18 8 17 9 16 10 15 11 14 12 13
Figure 6.3 Daily/hourly trend of common leopard (Panthera pardus) depredation on
livestock in and around PLNP, AJ&K, Pakistan.
70 60 50 40 30 20 10 0
Goat Cow Buffalo Ox
Figure 6.4 Monthly trend of common leopard (Panthera pardus) depredation on
livestock in and around PLNP, AJ&K.
Table 6.2 Monthly variation in attacks of common leopard (Panthera pardus) during 2008-2015 in and around Pir Lasura National
Park, Azad Jammu and Kashmir, Pakistan.
Years January February March April May June July August September October November December Total
2008 - - - - 1 2 - - 1 - - 1 5
2009 - - - - 5 1 2 - - - - - 8
2010 - - - - 2 1 1 1 - - - - 5
2011 - - - 1 1 - - - - - 1 2 5
2012 - - 2 1 - - 1 - - - 1 1 6
2013 - - - - 2 1 1 - - 1 2 5 12
2014 2 - - 1 10 8 8 1 2 - 3 3 38
2015 3 9 5 6 12 18 23 1 - 1 2 6 86
Total 5 9 7 9 33 31 36 3 3 - 9 18 165
One-way ANOVA showed statistically significant difference in leopard depredation during different seasons (F = 41.67, df = 3, p < 0.001). LSD test showed leopard depredation being significantly different (p < 0.001) during summer season as compared to other seasons.
6.4.5 Guarding, Herd Size And Penning
Attacks of common leopard differed with use of different guarding strategies by the locals. The lowest attacks by common leopard occurred when livestock were guarded by both human and dogs (4.84%), followed by when guarded by only dogs
(10.90%), only by human (27.87%) and highest attacks occurred with no guarding of livestock (56.36%). The effect of different guarding strategies to reduce leopard depredation on livestock was found statistically different (X2 = 105.37, df = 3, p <
0.001). Majority of livestock depredation occurred in larger herd size >20 animals
(32.12%), followed by herd size 10-19 animal (13.33%) and herd size 1-9 animals
(9.09%). We found this difference statistically significant (F = 43.54, df = 2, p <
0.001). LSD test showed that leopard depredation significantly different (p < 0.001) in herd size greater than 20 compared to any other herd size. A majority of respondents penned livestock at night (95.15%). However, penning did not reduce mortality due to leopard (X2=134.55, df = 1, p < 0.001).
6.4.6 Local Perception About Conservation Status
A majority of respondents (89.69%) had no idea about the conservation status of carnivores. Local perception was hugely negative and majority of respondents
(86.06%) showed unwillingness to save leopard. A majority of respondents (52.72%) thought that livestock depredation by leopard can be controlled through killing the predator followed by translocation of predator (22.42%), habitat improvement
(15.15%), repel predator (4.84%) and do nothing (4.84%). Local perception for possible management of leopard in their surroundings differed significantly
(X2=118.7, df = 4, p < 0.001). LSD showed that significant number of respondents proposed lethal control (p < 0.001). Majority of respondents (86.66%) had ample knowledge about the authority responsible (wildlife department) for conservation of carnivore species in study area.
6.4.7 Retaliatory Killing
Many incidences of retaliatory killing of common leopard were reported during 2008-2015 period. Fifteen common leopards (n = 15) were killed in response to leopard depredation on livestock. Common leopard also killed one human in
March 2012 and 4 persons were injured as a result of leopard attack. In addition, one
Cub of leopard was poached by locals during 2015.
6.4.8 Financial Losses
Among all key predator species, common leopard was found responsible for majority of the financial losses, approximately estimated at US$ 79,065 for the period of 8 years. Average loss caused by common leopard was US$ ~
60/person/year.
6.5 DISCUSSION
The common leopard (Panthera pardus) is listed as “Vulnerable” with declining population trend (IUCN, 2016). Its broad geographic range, remarkable adaptability, and secretive nature, have contributed to a misconception that the species might not be severely threatened across its range. However, not only its several sub-species and regional populations are “Critically Endangered” but also its overall range loss is greater than the average for terrestrial large carnivores whereby each of its nine sub-species has approximately lost 98% of its historic distribution
range (Jacobson et al., 2016). Negative perception of human- leopard interaction is a major conservation and rural livelihood issue throughout the distribution range of the cat species (Inskip and Zimmermann, 2009). The current study highlights underlying spatio-temporal patterns of human-common leopard interaction, especially in context of livestock depredation and leopard persecution, in the Azad
Jammu and Kashmir part of north-east Pakistan. In the study area, leopard has been found to be heavily persecuted by the local communities in response to livestock depredation. During past 20 years, there has been 17-26% increase in livestock of
Azad Jammu and Kashmir (Termizi and Rafique, 2001) and so livestock depredation by carnivores is also expected to increase.
Since livestock possess reduced anti-predatory skills, they are more vulnerable to predation (Nowell and Jackson., 1996) by wild carnivores. In this context, animals weighting around 25-50 kg are most vulnerable to depredation by leopard as they can be easily killed and dragged to safe place (Dar et al., 2009; Bibi et al., 2013). Khorozyan et al. (2015) reported that big cats kill more livestock when wild prey reaches a minimum threshold’. However, in the current study, since no previous data are available regarding abundance of the prey species of the carnivores in study area, we cannot generalize this fact here in this case. Neither there are any previous studies that could establish the status of the wild prey in the study area.
Therefore, it is hard to conclude that common leopard has depredated more on livestock because of the reason that its wild prey has reached a minimum threshold.
In the current study, common leopard has emerged as the major predator with
88.5% depredation on livestock (total losses 306 livestock during last 8 years), while other carnivores contributing only minor percentages such as Asiatic Jackal (Canis aureus) 5%, small Indian Civet (Viverricula indica) and Indian grey mongoose
(Herpestes edwardsii) 3% each. The most heavily depredated animal has been the domestic goat (82%), followed by domestic dog (9%). Few studies on common leopard have previously reported it as major predator of livestock including goat and sheep in some other areas of the Azad Jammu and Kashmir, Pakistan (Dar et al.,
2009; Qamar et al., 2010; Bibi et al., 2013; Kabir et al., 2014). Sekhar (1998) reported that leopard is responsible for 88% of goats, sheep and calves depredation in Sariska Tiger reserve Rajasthan, India. Similarly, Karanth and Sunquist (1995) reported mean body mass of leopard prey as 23 kg in tropical forests of India. The common leopard preferably preys on species weighing 10- 40 kg (Hayward et al.,
2006). In the current study, after goats, domestic dogs were most depredated species by common leopard. In the study area, domestic dogs are often used by local community for guarding purposes, therefore, these dogs are also more prone to leopard attacks. The findings of the current study are also supported by Dar et al.
(2009) who reported depredation of 34 dogs by common leopard in Machiara
National Park, Azad Jammu and Kashmir, even higher than the n = 24 dogs depredation being reported during the current study.
The current study indicates that there are three peaks of common leopard attacks on livestock in temporal context; majority of attacks occurred at morning time, followed by at night and then at evening time. More leopard depredation on livestock have been reported to occur between 9:00 am to 1200 noon, by the local respondents, and in fact, this is the high time when locals leave their livestock unattended , especially for grazing and thus make their livestock vulnerable for leopard depredation. Woodroffe et al. (2007) had reported that the risk of leopard attack was higher during the daytime for larger herds in Africa. Ahmed et al. (2012) had reported more leopard killing at night in central India. Some earlier studies also
reported high frequency of leopard attacks during night time in other parts of Azad
Jammu and Kashmir (Dar et al., 2009; Kabir et al., 2014).
The common leopard depredated on livestock during whole year with majority of killing occurring in the months of July (22%), May (20%) and June
(18%), respectively. In the study area, these three months are hottest months regarding temperature and are categorized as summer season. Overall, majority
(61%) of livestock depredation by common leopard occurred in summer with only
19% being in winter season. During summer season, locals leave their livestock for grazing in the open fields, in the surrounding of villages in PLNP and thus livestock become an easy prey for common leopard. Another peak of leopard attacks was observed during the month of December, and it seems logical to increase guarding during summer and winter months to reduce negative human-carnivore interaction.
The current stud highlights total financial losses due to livestock depredation by common leopard computed to be approximately US$ 80,000 during last eight years, which itself suggests alarming rate of livestock depredation by leopard.
Results of the current study showed that presence of both, human and dog, were most effective to reduce common leopard attacks on livestock, both inside and outside the village. Either dogs or human alone were not much effective in reducing leopard depredation on livestock and highest attacks were observed when livestock was left unattended. Ogada et al. (2003) had reported that better herding practices can reduce predator attacks on livestock. Earlier on, Kolowski and Holekamp (2006) also showed that dogs were ineffective in deterring leopard attacks in Kenya. Similar findings were reported by Dar et al. (2009) that dogs were ineffective in reducing
leopard attacks on livestock in Machiara National Park, AJ&K. Kabir et al. (2014) reported guarding strategies to be ineffective in Machiara National Park, AJ&K.
Results of the current study showed that penning practices of livestock at night were ineffective in reducing leopard attacks since penning structures were very poor making it easier for common leopard to kill livestock due to its predatory behaviour. Such findings are in line with other studies conducted in Azad Jammu and Kashmir, Pakistan; for example Dar et al. (2009) reported poor penning conditions in Machiara National Park, AJ&K. Improving conditions of pens/corals can help in reducing the livestock losses due to leopard attacks.
The local community at PLNP heavily depends upon livestock for subsistence and any livestock depredation by leopard may result in its retaliatory killing. We recorded higher numbers of leopards persecuted (n = 15) in retribution to livestock losses than other studies conducted earlier in the region (Dar et al., 2009;
Bibi et al., 2013; Kabir et al., 2014) with 4 humans being injured and 1 human killed from 2008-2015 due to common leopard attack. The rate of retaliatory killing suggests that negative human-leopard interaction might have increased (86%) in
PLNP compared to some other protected areas of the region. Dar et al. (2009) had reported that people think leopard is not dangerous which is contrary to current study findings. We found local perception hugely negative and majority of respondents were not willing to conserve leopard in their surroundings. Baral and Henien (2007) had earlier on reported that leopard depredation on livestock resulted in negative attitudes of local communities towards leopard. Previous studies had also reported negative perception of local communities towards leopard in other protected areas of
Azad Kashmir (Bibi et al., 2013; Kabir et al., 2014). At current, there is no
compensation for local communities in the PLNP which might be resulting in negative attitudes towards leopard and other carnivores. Local compensation schemes for livestock can improve local tolerance towards main challenging carnivore species (Bagchi and Mishra, 2006; Dar et al., 2009). However, the compensation should be according to the local rates and be delivered expeditiously in order to be effective (Madhusan, 2003; Naughton-Treves et al., 2003).
6.6 CHAPTER SUMMARY
Antagonistic human–carnivore interactions have been a major conservation issue since many carnivore species depredate on livestock resulting in provoked retaliatory killing of carnivores by local communities. Mitigation of such threats requires explicit understanding of underlying spatio-temporal patterns. We collected data on livestock depredation by carnivores and retaliatory killing of leopards at Pir
Lasura National Park (PLNP) AJ&K during 2014-15, also retrieving data about previous years from 2008, using a questionnaire survey. Results showed that 170 people lost 306 livestock, poultry, dogs, and pets to four different carnivore species; common leopard being the major predator killing 88.5% livestock, and minor contributions from Asiatic Jackal (5.2%), small Indian Civet (3.2%) and Indian grey mongoose (2.9%). The depredated livestock by leopard included mainly goats
(82.2%) and dogs (9.2%), and the leopard attacks were found significantly higher during the daytime, Majority of leopard attacks occurred during summer season. A positive correlation was found between monthly leopard depredation on livestock and mean monthly temperature. A significant effect of different strategies used for guarding livestock was found with least attacks occurring when livestock were guarded by both human and dogs. Common leopard was responsible for majority of the financial loss estimated approximately at US$ 80,000 for the current study period
and it negatively affected local tolerance. As a result, local perception was hugely negative (86%) and included killing of fifteen leopards. We suggest further detailed studies, compensation programs, and awareness education campaigns for conservation of common leopard in the PLNP, AJ&K.
Chapter 7
GENERAL DISCUSSION
Mammalian scats have been commonly used in biological studies to estimate population size (Kohn et al., 1999; Webbon et al., 2004), distribution patterns or species richness (Dalén et al., 2004), as they are abundant and easily found (Sanz et al., 2007). In many cases, it is assumed that scats are correctly identified, but it is difficult using scat morphology alone (Davison et al., 2002; Prugh and Ritland,
2005). It becomes more difficult when sympatric species have similar body features, behavior and feeding habits, and so the visual identification of scats becomes error prone (Ruiz-González et al., 2008). Knowledge of a predator’s diet is vital to understand its ecology and to predicting its effect on the dynamics of prey populations (Oli, 1993). The diets of red foxes and dogs have been studied in
Australia (Triggs et al., 1984; Brown and Triggs, 1990; Lunney et al., 1990; Glen,
2005) and both are opportunistic predators, and diet comprised of mammals.
Although, the diets of both species can vary in different habitats and seasons
(Mitchell and Banks, 2005). Knowledge of distribution is prerequisite, to understand how the individuals of a species interact with each other and with their habitat.
Distribution of a species can be affected by many biotic and abiotic factors which include; food availability, vegetation characteristics, competition (interspecific and intraspecific), power of animal dispersions, geographic barriers, climate and weather, and habitat quality. These factors and many others can determine distribution of a species and can limit distribution in a country or in a region (Sclater and Sclater, 1899; Roberts, 1997).
181
During current study, we recorded common leopard in all (100%) surveyed sites and direct and indirect signs of common leopard were recorded between 757 m-
1891 m. The leopard was found evenly distributed in the study area. No previous published studies are available that report on the distribution of the leopard in the study area. Roberts (1997), however, described that four sub-species of Panthera pardus are known to occur in Pakistan including; firstly, Panthera pardus saxicolor
(Pocock, 1927) found in Balochistan (also found in Persia), secondly, Panthera pardus sindica (Pocock, 1930) occurring in Kirthar, Sindh, thirdly, Panthera pardus fusca found in whole of India and fourth sub species Panthera pardus millardi
(Pocock, 1930) is known to occur in state of Kashmir. Siddiqui (1961) stated that all above four sub-species occur in Pakistan although at present it might be hard to distinguish individual specimens into sub-species while population from Sindh and
Balochistan are so small as compared to northern Himalayan population. Throughout its range, there is considerable variation in the pattern and density of rosettes or spots on the body of the leopards since the leopards in Swat and Hazara districts have longer and more luxuriant pelage during winter season. Roberts (1997) described the distribution of the leopards in Pakistan as it is confined to the forest of Himalayan region up to tree line or at lower altitudes in valleys which are more arid hilly regions in the north. It also occurs in hilly areas associated with Acacia modesta and Acacia senegal scrub forests of Waziristan, Balochistan and Sindh Kohistan. It was once inhabitant of the” Salt Range” and still survives in Kala Chitta Hills but is not found in human settlement areas, cultivated lands, riverain tracts for many decades. But during the current study, signs of the leopard have been recorded near human habitations. This might be because of the reason that scarcity of wild prey in forests is now resulting in the leopards’ interaction with human populations and it is known
to predate on their livestock. It is also distributed in Kirthar Hills, Kalat and Mekran,
Ziarat, Murree Hills, Margallah Hills, Chitral, and the Chilas district of Gilgit-
Baltistan.
The second sympatric carnivore, the Asiatic jackal was found distributed in all (100%) surveyed sites during the current study period. No previously published on distribution of the jackal is available for comparison from the study area, however, there are few reports from the other parts of the country. Roberts (1997) described
Asiatic jackal as very adaptable animal and readily found in mountainous regions, forest plantations and riverine tracks. It is well adapted to dry, open habitat and avoids dense forests. During the current study, direct and indirect signs of the jackal were recorded near human settlements, cultivated lands, and in open and close forest areas. Roberts (1997) had reported Asiatic jackal occurring at 2150m elevation in
Hazara district, Shogran, and Murree hills and reported that it did not penetrate high mountain region. During the current study, we have recorded distribution of the jackal at elevation ranging from 721 m to 1690 m asl. However, signs of jackal were not recorded at top of the mountains at elevation around 2000 m.
The fox is among the least studied carnivores in Pakistan. During current study, we recorded direct and indirect signs of Kashmir hill fox (a sub-species of red fox that occurs in AJ&K) from all (100%) surveyed sites in and around Pir Lasura
National Park. Altitudinal range of its distribution was 815m-1646 m a. s. l. in the study area. No previous published studies are available for comparison form study area however, few studies do report on distribution of species in Pakistan. According to Roberts (1997) the species avoids dense forests and it can be found in open areas in the country. In Indus plain, it is reported to prefer extensive uncultivated tracts with sand dunes. During the present study, signs of the fox species were recorded in
all habitat types including; near human dwellings, cultivated lands, open forests as well as close forests. Roberts (1997) also had reported that it occurs throughout the mountainous regions of Balochistan, Khyber Pakhtunkhwa and Himalayas, both in valleys and hilly mountainous regions. We also recorded signs of fox in valleys as well as in the hilly mountains regions.
We recorded direct as well as indirect signs of Asian palm civet in all (100%) study sites. Initially, during surveys we found no information regarding distribution of this species in the study area. Initially, people from local community helped us identify scats of yellow throated martin in the field. However, after molecular identification of those scats, they were confirmed to originating from Asian palm civet and later on we also found road killed specimens of Asian palm civet in the study area. No yellow throated martin was found in the study area based on the direct signs as well as indirect signs viz., molecular identification of scats. Distribution of
Asian palm civet was recorded between 896 m – 1922m a.s.l. It is a wide spread species and its scats were encountered easily near human habitations on boundary walls of settlements, stones and hills. Scats of this species were usually encountered at rides of the mountains. The species was recorded at all the sampling sites surveyed and often encountered by local people especially at night raiding on fruits.
Distribution of Asian palm civet/ common palm civet in Pakistan was previously described by Roberts (1997). According to Roberts, it occurs in extreme north- western boundary in Pakistan and is believed to have very local and restricted distribution in Pakistan. It has been recorded in Rawalpindi, Lahore, Sargodha,
Multan, Sahiwal, Bahawalpur, and Dera Ghazi Khan districts (Ahmad and Ghalib,
1979). It occurred in sub-Himalayan tracts, Mansehra in district Hazara at an
elevation of 2670m and Rawalpindi District. It was also trapped from Margalla hills at Chak Jabri site.
Distribution of small Indian civet was recorded at seven sampling sites (23%) in the study area during surveys, at an altitudinal range between 1171m-1857m asl.
Since no previous studies have focused this species in Pir Lasura national park so far, therefore, current study results cannot be compared. However, according to
Roberts (1997) small Indian civet is best adapted species among all civets in sub- continent to terrestrial hunting and semidesert landscapes. Published literature reports that the small Indian civet occurs in variety of habitats in Indus basin such as, riverine jungles and sand-dune deserts. It is also found in irrigated forest plantations and avoids high human settlements and cultivated areas as well as mountainous regions. In Pakistan, it is decidedly uncommon and erratic in its distribution. In the current study, we recorded signs of small Indian civet from few sampling sites with single no direct sighting but with recovery of road killed specimens. Signs of this species were also less common in the study area while its scats were found in dense or open areas having high understory vegetative cover, also in areas having sparse human settlements surrounded by dense forests.
Distribution of Indian grey mongoose was recorded at fifteen different sampling sites (50%) in the study area, with an elevational range from 699m – 1559 m a.s.l. According to Roberts (1997), the grey mongoose is better adapted to arid conditions, it occurs in the Salt Range and is plentiful around Rawalpindi district.
The grey mongoose is reported to prefer more arid tracts and so the evidences of its high occurrence in natural vegetated area and in medium human activity areas, cultivated lands and avoidance to live near human dwellings is consistent with
previous published literature such as by Roberts (1997), Santiapillai et al. (2000),
Francis (2008), and Hussain et al. (2017). Roberts (1997) had reported that it does not penetrate into the Murree foothills, however, in the current study, we recorded its distribution at 1559m elevation in Azad Kashmir which is almost the same elevation as that of Murree hills. Hussain et al. (2017) reported that in Potohar region mongooses are distributed at elevational range of 203m to 874m elevation. Both
Roberts (1997) and Hussain et al. (2017) reported that Indian grey mongoose avoids human settlement areas with high disturbance. However, in the current study, we recorded distribution of Indian grey mongoose in areas having human settlements surrounded by wild areas or cultivated land. Indian grey mongooses were found distributed in nullas having undisturbed semi-wild habitat near dense human habituations. They were often sighted in or near human settlements predating on domestic chickens. During present study, local community reported, and the author/researcher also sighted that Indian grey mongooses enter in their homes through sewerage pipelines. However, in highly populated areas having no undisturbed habitats around, we did not record direct or indirect signs of the Indian grey mongoose.
Distribution of small Indian mongoose was recorded at all 30 (100%) sampling sites surveyed in the current study. It was found distributed at elevational range between 691 m- 1624 m asl. Earlier on, Roberts (1997) had reported small
Indian mongoose as one of the commonest small carnivore in Southern Sind and
North-Eastern Punjab, having its distribution around Jhelum, Gujranwala and the
Salt Range. The species is well adapted to live in rocky areas with stunted thorn scrub typical of the Salt range. During the present study, this species was recorded in all study sites even in dense human populations. Distribution of the small Indian
mongoose within or near the high human habitation and medium human activity areas or in cultivated lands had also been reported earlier (Roberts, 1997; Mahmood et al., 2011; Hussain et al., 2017), where they recorded the occurrence of the small
Indian mongoose near human habitations, near poultry farms and in cultivated lands.
Roberts (1997) had also reported that the small Indian mongoose is well adapted to live in the outskirts of villages and towns.
Spatial avoidance is one of evolutionary solution for coexistence of carnivores in an area (Hunter and Caro, 2008), however coexisting is also possible by consuming prey species of different weight class or different taxa (Karanth and
Sunquist, 2000; Hayward and Kerley, 2008). We recorded that though the distribution of all carnivore species overlaps in the study area however, relative abundance of signs was not even in all study sites. Signs of different species were found abundant at different study sites rather than at a single site which shows that these carnivore species occupy different areas to avoid competition.
Baseline data on diet composition of carnivore species is prerequisite fro their conservation and management. In the face of ever shrinking habitat of wildlife due to increasing human population, growing agricultural needs and unsustainable use of wild resources carnivore conservation is challenging. People living in and around protected areas are often dependent on livestock for their livelihood (Mishra et al.,
2004). Throughout the distribution range of leopards, a dietary shift from wild prey species to domestic species has been observed (Judas et al., 2006; Spalton and Al-
Hikmani, 2006). Many factors are responsible for such phenomenon. Firstly, lack of wild prey results in leopard’s attack on livestock. Secondly, poor husbandry practices and penning conditions. Thirdly, guarding strategies play important role in loss of
livestock during leopard attacks. Fourthly, seasons/temperature influence the rate of livestock depredation by leopards. Fifth factor which may have impact on rate of livestock depredation by leopards is time of the day since attacks on livestock can be high during specific time of the day and it is usually when livestock is left unattended.
In Pakistan, the leopard has been reported to predate on snakes, lizards, rodents, Sindh ibex, markhor, urial, rhesus monkeys, porcupines and in the regions where wild prey is limited the leopards are known for attacking domestic livestock including; cows, calves, donkeys, ponies, goats and sheep. Although its predation on crop-destroying porcupines, stray village dogs and rhesus monkeys might be considered beneficial to man, however, because of its attacks on domestic livestock and human beings it has been ruthlessly persecuted by local communities whenever encountered and has been always considered symbol of fear and contempt in
Pakistan (Roberts, 1997).
Diet of the common leopard comprised of mainly domestic prey as compared to wild prey species, including mammals, birds, and anthropogenic items. We found that consumption of domestic prey was high as compared to wild prey and domestic prey species included; goat, sheep, cow, buffalo, horse, poultry and dogs. The consumption of goat was high followed by dog and our findings are in line with other studies conducted in distribution range of leopard. According to Shehzad et al.,
(2015) diet of the leopard comprised of mainly domestic prey species and consumption of goat was highest (64.9%), followed by dog (17.5%) and cow Bos taurus (12.3%). Leopard has been reported to kill massive numbers (n=22) of livestock such as sheep in a single attack (Sangay and Vernes, 2008). Athreya et al.
(2016) reported that diet of leopard consisted of livestock, and 39% domestic dogs.
There were few wild species recorded from diet of the leopard.
Among wild prey species, rhesus monkey was more frequently and heavily consumed. Similar findings have been reported by some other studies such as predation of common leopard on primates has been reported from Asia and Africa
(Kummer et al., 1981; Cowlishaw, 1994; Isbell, 1994; Nowell and Jackson, 1996;
Zuberbühler and Jenny, 2002; Hayward et al., 2006). Leopard predation on rhesus monkey has also been reported by (Mukherjee and Mishra, 2001; Lodhi, 2007).
Pir Lasura National park is such a landscape surround by human population and the number of livestock in and around the park is over a million. In such a landscape availability of domestic prey is more than that of domestic prey species, therefore, it is quite logical for such as a species to predate on prey which is abundant in the area and requires less effort. Carnivore density in natural or semi-natural ecosystems is related to biomass of prey species available (Karanth and Nichols,
2010). Recent studies have demonstrated that large carnivores can persist in human- dominated landscapes by predating fully or partially on domestic livestock (Yirga et al., 2012; Athreya et al., 2013). The potential of human dominated landscape to support large carnivore species must be investigated in terms of availability and abundance of wild as well as domestic prey species (Boitani and Powell, 2012). In human-dominated landscapes biomass of domestic prey species can be higher than wild prey species (Mizutani, 1999) .
Livestock depredation by the leopard has resulted in antagonistic interaction between local communities and the leopard in Pir Lasura National Park. Interaction between felids and human is complex and spectrum of such relation ranges from fascination to fear (Boomgaard, 2001; Loveridge et al., 2010). From conservation of
large felids, they are often represented as flagship species (Treves and Karanth,
2003) but antagonistic interaction of large felids occurs in areas where these species live in human-dominated landscapes, where presence of these species can result in livestock depredation and loss of human life (Treves and Karanth, 2003). As a result, large felids are killed in retaliation which is significant cause of their mortality
(Inskip and Zimmermann, 2009). During current study, we found major prey species of the leopard was goat followed by dog. This is due to the fact of high availability of goat, poor guarding and penning condition in study area. Dogs are used for guarding livestock and they become easy prey for the leopards. Studies have reported the leopard predating on dogs such as (Daniel, 2009). Predation of the leopard on goat is higher and overall consumption of domestic prey is higher than wild prey species which shows that the economic impact of predation is higher.
During this study we recorded mammals, birds, insects, and anthropogenic items from the diet of Asiatic jackal. Consumption of domestic prey was higher than that of wild prey. Wild prey species included rhesus monkey (which was the main wild prey species) and four species of rodents. Among domestic prey species, poultry was the main prey followed by goat, whereas consumption of plant matter was low.
Studies conducted elsewhere, have reported rodents and plants as main components of the jackal diet. However, we found that this was not the case in our study area, where the jackal seems to be more of a scavenger and predates on domestic poultry for a major part of its diet, followed by rodents and rhesus monkeys. Our results are in line with Roberts who reported that in Pakistan the jackal is perceived as a scavenger and carrion eater (Roberts, 1997). The jackal feed on refuse in villages, however the majority of its diet is comprised of rodents, reptiles, fruits, and insects when available (Roberts, 1997). A study conducted in Bharatpur, Rajastan, India
during 1984-1985 showed that diet of the jackal comprised of rodents 26.5%, birds
24.1 %, grass 20%, fruits 16.5%, insects 4.1%, snakes 4.1%, chital 2.4%, nilghai
1.2%, and fish 1.2% (Sankar, 1988). Stomach contents of four jackals in Rajasthan
Indian revealed that diet of the jackals mostly consisted of fruits of Ziziphus, with small proportion of beetles and scorpions. Another individual consumed desert jirds
(Meriones hurrianae) and small mongoose (Prakash, 1959). Analysis of 138 scats of
Asiatic jackal in Sal forests in India showed that 68 percent of their diet consisted of rodents followed by plants 27%, 11% reptiles, 8% fish and 9.4% birds (Schaller,
1967). A study conducted in Sariska Tiger Reserve, India showed that diet of the
Asiatic jackal was comprised of plant matter 17.57%, rodents 15.77%, cattle 15.32%, chital 10.81% fruits 9.01%, birds 7.21%, sambar 5.41%, hare 4.05%, nilgai 4.05%, goat 1.80% reptiles 1.8% (Mondal et al., 2012). Diet of the jackal is comprised of small and medium-sized mammals such as rodents, rabbits, birds, fishes, insects and vegetation (Aiyadurai and Jhala, 2006). The diet of jackal in Keoladeo national park,
India was comprised of mainly plant matter 38.44%, nilgai 8.13%, rodents 10.31%, chital 9.69%, cattle 7.5%, anthropogenic 1.56% (Singh et al., 2016). This high proportion of mammals in the diet of the jackal suggest that they scavenge on dead animals and hunt calves as hunting larger mammals might be difficult (Singh et al.,
2016). Rodent are an important part of the jackal diet (Mukherjee et al., 2004; Jaeger et al., 2007; Majumder et al., 2011; Singh et al., 2016). Diet of Asiatic jackal in
Pakistan showed that 46.47 % of its diet is animal matter by volume including rodents, mongooses, wild boar, livestock, birds and domestic poultry and 25.08% plant included wheat, tomato, berries, grains, orange, melon and water melon
(Mahmood et al., 2013). Most of the species have recorded plants and rodents as major prey species in diet of Asiatic jackal this could be due to the difference in
sampling procedures. The jackals living in high mountainous regions can take up mammals only as major part of their diet (Schaller, 1970). In India Schaller (1970) reported that frequency of occurrence of rodents in the diet of jackal was 94%, however he also reported snakes and lizards 29% and insects 6.7 %. Previous, studies which have been conducted in Pakistan has also reported large proportions of plant matter in the diet of jackal however during present study scats of the jackal were confirmed genotypically and it was interesting that scats identified as originating from the jackal have very little or no plant matter and other sympatric carnivores such as fox and civets consumed plant matter in large proportions.
There is paucity of information available on diet composition of the fox in
Pakistan and no published scientific literature is available. However, some information is reported by Roberts (1997) who states that the foxes are adaptable hunters and can hunt hares, rodents, reptiles and small birds but when vertebrate prey is not available they can subsist on insects and fruits. According to Roberts, foxes feed on fruits of ber tree (Zizyphus mauritiana). In the diet of the fox, mice, rats, desert hare, Indian gerbils, have been reported by Roberts (1997). Studies in Indian
Rajasthan showed that the foxes feed on wild melon, termites (Prakash, 1959). Thef oxes have reported to feed on scorpions, fruits of Zizyphus nummularia, spiders, cockroaches and also had some melon seeds in stomach. Roberts (1997) never encountered any sign of the foxes feeding on domestic poultry although he has reported instances of domestic poultry being eaten by civets, cats, mongooses and martins.
During the present study, we recorded that diet of Kashmir hill fox diet was comprised of mammals, birds, invertebrates and plant matter. We also recorded anthropogenic material in the diet of the fox. Consumption of poultry was higher
than any other dietary component, followed by goat. We identified 6 species of plants in the diet of the fox. The Kashmir hill fox also consumed sheep, cow, buffalo which likely indicate the fox’s scavenging behavior. The consumption of different food items varied among during different seasons, in agreement with other studies
(Basuony et al., 2005; Baker et al., 2006; Kidawa and Kowalczyk, 2011). The red fox is essentially an omnivore having a diverse diet which includes invertebrates, small mammals, birds, plant matter and carrion (Flower, 1932; Englund, 1965;
Amores, 1975; Macdonald, 1979; Osborn and Helmy, 1980; Ciampaloni and Lovari,
1985; Calisti et al., 1990; Basuony, 1998). The fox has been reported by other studies to feed on plant fruits in southern Europe (Ciampaloni and Lovari, 1985;
Calisti et al., 1990) it is known to feed on plant species or parts of plants having high sugar content (Basuony et al., 2005).
Asian palm civet and the small Indian civet are omnivorous and thus play vital role to limit the prey populations as well as in the seed dispersal of some important wild plants. Both the civet species are protected under provincial, federal and AJ&K acts and rules in Pakistan, yet they are persecuted in response to the damage they cause to the orchard fruit trees in AJ&K. The present study revealed that both species mainly depend on wild food and they caused relatively little damage to orchard fruits, cultivated crops and domestic animals. In the diet of Asian palm civet only 1.6% orchard fruits, 2.6% crops and 6.74% poultry included. Small Indian civet relatively consumed more orchard fruits (10%), crops (1.61%) and poultry
(14.52%). Though the pest role of both the civets appeared to be limited, however, the general perception and attitude of farmers for the civets were found always be negative. Therefore, farmers and orchard owners were killing these civets, whenever
encountered. Some management strategies are required in the study area to reduce the conflict and for the conservation of civets.
Results of the current study show that Asian palm civet mainly consumed plants, wild and domestic prey species. The Asian palm civet is reported to live in the vicinity of villages, and subsists on rats, mice and is also attracted towards food orchards and plantations (Roberts, 1997). No previous published literature is available for Asian palm civet diet from Pakistan. However, results available from few other countries are comparable. Asian palm civet plays an important role in the food web as a predator, as prey and as seed dispersers (Aroon et al., 2012). Their diet includes; small mammals, birds, amphibians, reptiles, invertebrates, eggs and fruits. In the current study also, consumption of plant matter was high in the diet of
Asian palm civet. Our findings are in line with other studies but we found relatively lower consumption of orchard fruits as compare to other regions. Joshi et al. (1995) reported 84.5% fruits in the diet in Nepal. Similarly Krishnakumar and
Balakrishanan (2003) recorded 82% fruits and Jothish (2011) reported 90.1% of the diet of Asian palm civet comprised of fruits in various localities of India. The findings of the current study hereby report about 53% plant-based diet of the Asian palm civet in the PLNP, AJ&K. Grassman (1998) recorded 18 species of fruits in the diet of Asian palm civet in Thailand. Krishnakumar and Balakrishanan (2003) recorded ten fruit species in India; Su and Sale (2007) reported 31 fruit species in its diet. In Myanmar, the frequency of fruits in the diet was high during the rainy season than winter season (Rabinowitz, 1991; Kitamura et al., 2002) and it consumes more mammalian prey when fruits availability is decreased (Aroon et al., 2012). In the current study, Asian palm civet consumed 15 plant species overall, including nine wild species, three important orchard fruit trees, and cultivated Coriander
(Coriandrum sativum), water melon (Citrullus lanatus) and wheat (Triticum aestivum). As far as consumption of wild plants is concerned, the civet species is playing a a vital role in their seed dispersal in the study area. However, feeding on orchard fruit trees results in the persecution of the civet species by local people, which raises a conservation concern. The natives do confess that they kill Asian palm civet when it visits their orchard for feeding on fruit trees.
We found some scats of Asian palm civet that were completely composed of plant matter, and this is also verified from other studies which reported that scats of
Asian palm civet are fully composed of plant matter during fruiting seasons
(Rabinowitz, 1991; Corlett, 1998). The findings of the current study showed that diet of Asian palm civet comprised of small mammals, arthropods, lizards, snakes, birds, and fruits. Small mammals were most abundant (53.7%), followed by plants
(37.65%).
The frequency of occurrence of plants was high in the diet of the other civet species, the small Indian civet. This civet species also consumed wild prey species, domestic prey, insects, amphibians, as well as grits, and anthropogenic material. Diet of small Indian civet in the current study comprised of mainly plants, rodents, and birds. Previously no published literature is available about the dietary habits of the small Indian civet for comparison from Pakistan except some field observations by
Roberts (1997). Small Indian civet is omnivorous, in feeding habits and relies on fruits during their availability but also feeds on a variety of insects and arthropods.
In India, from the stomach of a dead civet were recovered remains of beetles, seeds of a bairs fruit (Zizyphus jubata) and unidentified finch (Bnhs, 1913). A specimen was caught at Kalabagh feeding on ripe grapes. Small Indian civet hunts on small mammals, birds, and reptiles. They have been reported to rob nests of birds for eggs
and nestlings. They spent most of the day sleeping in their burrows and hunt at night.
They are known to dig for insect larvae and succulent roots (Roberts, 1997).
Similarly, another study highlighted that the diet of small Indian civets comprised of rodents (80%) insects (23%) (Wang et al., 1976). According to another study, its consists of rodents and shrews 40%, and insects 95% and earthworms 67% (Chuang and Lee, 1997), which is not the case in the current study since we have recorded approximately 55% plants in the diet of small Indian civet. Diet breadth was found to be 4.46 (Chuang and Lee, 1997). Diet breadth in another study was reported as
2.58, and it consumed nine prey species. Small Indian civet consumed rodents, birds, invertebrates, and plants matter (Wang and Fuller, 2003). Small Indian civets when living in outskirts of villages and the cultivated land they may predate on a more diverse diet (Wang, 1998). Civets are known to consume fruits and (Rabinowitz,
1991) reported 76% fruits in the diet of masked palm civet. Joshi et al. (1995) reported that frequency of occurrence of fruits in the diet of common palm civet was found to be 85% and during fruiting season whole scats entirely comprised of fruits.
Furthermore, during a shortage of fruits, common palm civets also consumed insects, mollusks, birds, reptiles and small mammals (Joshi et al., 1995). Thus, our findings highlight that small Indian civet is omnivorous in its dietary habits by consuming
52% plant matter and 45% animal-based diet. Also, our study also highlights that this civet species play important role in seed dispersal of some wild plant species but on the other hand, damages fruit orchards for which is being persecuted in the study area even being given protection through wildlife acts and rules.
Diet composition of Indian grey mongoose consisted of mammals, birds, reptiles, amphibians, invertebrates, egg shells, and plant species. We also recorded grits and anthropogenic items. Frequency of occurrence of house mouse was high in
diet of Indian grey mongoose. Among birds, frequency of chicken/poultry was high.
Four species/orders of invertebrates were recorded frequency of occurrence of hymanoptera (ants, bees) was high and frequency of occurrence of coleoptera
(beetles) and snails was low. Our results are in line with other studies such as,
Roberts (1997) reported that Indian mongoose feed on reptiles, birds, and amphibians. Prakash (1959) reported in Rajasthan, India that the grey mongoose feed on grey partridges, rodents, invertebrates and lizards. The Indian grey mongoose is known as an opportunistic hunter. It has been reported to feed on rodents, reptiles, invertebrates and birds, eggs of birds and fruits. We also recorded birds, egg shells invertebrates, plants reptiles and rodents in diet of Indian grey mongoose such results are reported by other studies. Indian grey mongoose feed on the red jungle fowl its chicks, eggs, peafowl, partridges, small mammals and snakes in Indian and also reported to search for food under stones on the beach side in Hawaii (Santiapillai et al., 2000; Postanowicz, 2002). It has been reported to feed on grasshoppers, centipedes, fish, frogs, scorpions and crabs (Whitfield, 1978). Hussain et al.
(2017) reported that Indian grey mongoose feeds on rodents, birds, insects, and plants.
We recorded that the diet of the small Indian mongoose consisted of mammals, birds, invertebrates, reptiles, amphibians, and plants. We also recorded anthropogenic matter and grits. Among mammals frequency of occurrence of house rat was high. This could be due to the reason that small Indian mongoose lives around human habituation where availability of house rat might be high. The small Indian mongoose feeds on rodents such as Rattus rattus, and Mus musculus when it lives around human settlements (Roberts, 1997). Only one species of domestic prey was recorded i.e., chicken/ poultry. Among 3 species of invertebrates recorded in diet of
small Indian mongoose frequency of occurrence of order orthoptera (grasshoppers) was high followed by hymanoptera (ant, bees) and low was coleopteran (beetles).
Among 6 plant species recorded in diet of small Indian mongoose frequency of occurrence of Themeda anathera grass was high followed by choti bairi and low was melon and water melon. It has been reported to feed on beetles, scorpions snakes lizards, spiders and amphibians (Prakash, 1959). It feeds on birds and their eggs and nestlings (Roberts, 1997). Seaman and Randall (1962) reported that small Indian mongoose consumes small mammals, reptiles, amphibians, birds and plant matter.
Some population of mongoose are insectivorous and others may consume fruits during some seasons (Seaman and Randall, 1962). Small Indian mongoose feeds on rodents and insects and plant matter (Mahmood et al., 2011; Siddiqui et al., 2004;
Hussain et al., 2017).
The results of our study showed that consumption of wild prey by the common leopard was 34.85% while abundance of those wild prey species was estimated to be 57.4/Km2 whereas, consumption of domestic prey species by common leopard was 59.1% and availability of those species was found to be
747.36/Km2. Major wild prey species of common leopard was rhesus monkey and its consumption was 10.61% of its diet. Interestingly, this species was most abundant wild species in the study area having abundance 11.1/Km2. Among domestic prey species consumption of goats and sheep was high and it was more than 30% of leopard diet. Abundance of goats and sheep in the study area was high as compared to other ungulate species and it was 105.17/Km2.
Consumption of wild prey by Asiatic jackal was 18.48% whereas, its availability was 114.2/Km2. Among wild prey species consumption and availability
of rhesus monkey was high. Among wild prey species consumption and availability of goat and poultry was high.
Consumption of wild prey in diet of Kashmir hill fox was around 18% and availability of same species in habitat was 125.7/ Km2. Major prey species in diet were insects 4% and wild boar 3% followed by house mouse 2.4% and their abundance was 2310.3/m2, 5.4/Km2 and 17.5/km2 respectively. Among domestic prey species consumption of poultry was high 29.34% and its availability was also high 548.69/km2. Similarly, consumption of goat and sheep was above 19% and their availability was 105.17/km2.
Large portion of plants was recorded in diet of Asian palm civet, however consumption of wild prey was 33.68% and domestic was 10.88%. Despite of high availability of domestic prey Asian palm civet preferred wild prey especially rodents.
Among domestic species consumption of poultry was high 6.74% and it was quite abundantly available. Among wild prey species consumption of 4 rodent species accounted for 18.64% collectively however, consumption of Indian gerbil was high
9.84% among rodents and its availability was also high 20/km2. Diet of Asian palm civet contained significant percentage of invertebrates 8.3% and availability of invertebrate species was high.
Diet of small Indian civet consisted of plant matter, wild prey and domestic prey species. consumption of wild prey was 30.65% and wild prey was 14.52%.
Among wild prey species consumption of Indian gerbil was high 11.29% and it was abundant rodent species in the study area having abundance of 20/km2. Other dietary components included birds, amphibians and insects. Only one domestic species
domestic chicken was consumed, and it was quite abundant in habitat as well
548.69/km2.
Consumption of wild prey by Indian grey mongoose was 60% among wild prey consumption of house mouse was high 12% followed by Indian gerbil 10% and these species were abundant in the habitat as well having abundance of 17.5/km2 and
20/km2. Consumption of invertebrates was high 16% and their availability was also high. Only one domestic species domestic chicken accounted for 19 % of its diet and its abundance was recorded as 548.69/km2.
Consumption of wild prey by small Indian mongoose was 59.68% and availability of species consumed was also high such as frequency of occurrence of house mouse was high in diet and its availability was also high. Similarly, consumption of insect was high in diet of small Indian mongoose and invertebrates were abundant in habitat. Consumption of domestic chicken was 16.94% and its availability was high in the study area 548.69/km2.
We computed niche breadth and overlap among sympatric carnivore species at Pir Lasura national park. Among seven different carnivores of the study area, niche breadth of Indian grey mongoose was widest 18 (0.72) followed by Asiatic jackal
14.2 (0.78), common leopard 13.88 (0.72), small Indian mongoose 12 (0.64), small
Indian civet 10(0.52), Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31).
Dietary niche breadth of common leopard was broad during spring season which is indicative of the fact that more prey species were available to this top predator in the study area during spring season. But during winter season, the narrow dietary breadth of common leopard indicates availability of less prey species. For
Asiatic Jackal, on contrary, more prey species were available during summer season
as indicated by its broad niche breadth and less availability of prey species during spring season, represented by narrow niche breadth.
Niche breath of Kashmir hill fox was wider during autumn season showing availability of more prey species but narrow during spring season, shown by less availability of prey species. A survey of past published literature shows lacking scientific studies regarding quantifying niche breadth of the three above mentioned carnivores (common leopard, Asiatic Jackal, and red fox) in the country. So, now, the results of the current study cannot be compared with other data. The current study, therefore, reports here the niche breadths of the three mentioned carnivores from the country for the first time. Lanszki et al. (2006) reported that trophic niche breadth of golden jackal was (0.06) and red fox was (0.09) in Hungary.
Niche breadth was found broad for the Asian palm civet, during autumn season but narrow during winter season. These findings of the current study are supported by some previous published studies such as Aroon et al. (2012) reported that niche breadth of Asian palm civet was widest during autumn season (B = 0.169), the current study also recorded high niche breadth of this species during autumn season as more prey species are available during this season. This species is frugivorous in dietary habits and ample quantity of fruits are present during autumn season at Pir Lasura National Park. Aroon et al. (2012) reported narrow niche breadth during winter season which is like the findings of the current study that its niche breadth is narrow during winter season. Overall, they reported dietary niche breadth being 0.412 (Aroon et al., 2012). In the current study, we found overall dietary niche breadth of Asian palm civet being 0.31, which might be because of the variation of habitat and diversity of prey species available in specific study area. Niche breadth
of civets was narrow during winter when there was low availability of fruits and high during fruiting seasons.
The overall niche breadth of small Indian civet was 10 (0.52); wider niche breadth of small Indian civet during summer and narrow during autumn season in
Pir Lasura National Park can be compared with some earlier studies; Chuang and
Lee (1997) reported that dietary breadth of small Indian civet was 4.46 in northern
Taiwan. Another study reporting its niche breadth as 2.58 and it consumed 9 prey species; including rodents, birds, invertebrates, and plants matter (Wang and Fuller,
2003). Small Indian civets when living in outskirts of villages and cultivated land, may predate on more diverse diet (Wang, 1998).
Niche breadth of Indian grey mongoose was found broad 18(0.72) during autumn season and narrow 7.8(0.29) during winter season. Hussain et al. (2017) studied niche breadth and niche overlap between two mongoose species (Herpestes javanicus and H. edwardsii) in Potohar region, Pakistan. They reported that there was seasonal variation in niche breadth of Indian grey mongoose and it was high during summer season and narrow during winter season.
Dietary niche breadth of small Indian mongoose was broad during summer season 11.64(0.59) and narrow during spring season 9.35(0.46). Total niche breadth of small Indian mongoose was 12(0.64). Hussain et al. (2017) reported niche breadth of small Indian mongoose being variable during different seasons and it was highest
(7.2) in summer but lowest (6.1) in winter. The findings of the current study have also shown variation in niche breadth of small Indian mongoose during different seasons being high during summer season. But unlike Hussain et al. (2017), narrow
niche breadth was recorded during spring season, which could be attributed to habitat variability and to variation in prey species availability during different seasons.
Regarding comparison of dietary niche overlap among seven different carnivore species sympatric at PLNP, AJ&K, high niche overlap was found between
Asian palm civet and small Indian civet (0.9), indicating that both species consume similar food and they compete strongly for resources at Pir Lasura National Park.
Niche overlap among small Indian mongoose and Indian grey mongoose was 0.89.
There are not many previously published records to compare such findings except
Hussain et al. (2017) who reported that there is high niche overlap among two mongoose species in the Pothwar region, Pakistan and the current study have also recorded similar findings that both mongoose species compete for resources. Niche overlap among Asiatic jackal and Kashmir hill fox was also high (0.81) showing both species competing for same resources in the study area. Lanszki et al. (2006) reported that trophic niche overlap among jackal and fox was 0.73 in Hungary.
Niche overlap among Kashmir hill fox and small Indian civet was (0.66) since both carnivore species consume similar prey species like rodents and insects.
Although the difference of body size occurs between both species, however, both species consumes same prey species therefore there is competition among them.
Similarly, because of consuming shared resources niche overlap among common leopard and Asiatic jackal was (0.62). However, this can also be attributed to the scavenging nature of jackal as it feeds on carrion of domestic animals and leopard predate on livestock as indicted by high frequency of domestic prey in leopard diet.
Niche overlap of Kashmir hill fox and Indian grey mongoose was (0.62) which can
be result of feeding on rodent species as they were recorded in diets of both carnivores.
The common leopard (Panthera pardus) is listed as “Vulnerable” with declining population trend (IUCN, 2016). Its broad geographic range, remarkable adaptability, and secretive nature, have contributed to a misconception that the species might not be severely threatened across its range. However, not only its several sub-species and regional populations are “Critically Endangered” but also its overall range loss is greater than the average for terrestrial large carnivores whereby each of its nine sub-species has approximately lost 98% of its historic distribution range (Jacobson et al., 2016). Negative perception of human- leopard interaction is a major conservation and rural livelihood issue throughout the distribution range of the cat species (Inskip and Zimmermann, 2009). The current study highlights underlying spatio-temporal patterns of human-common leopard interaction, especially in context of livestock depredation and leopard persecution, in the Azad
Jammu and Kashmir part of north-east Pakistan. In the study area, leopard has been found to be heavily persecuted by the local communities in response to livestock depredation. During past 20 years, there has been 17-26% increase in livestock of
Azad Jammu and Kashmir (Termizi and Rafique, 2001) and so livestock depredation by carnivores is also expected to increase.
Since livestock possess reduced anti-predatory skills, they are more vulnerable to predation (Nowell and Jackson., 1996) by wild carnivores. In this context, animals weighting around 25-50 kg are most vulnerable to depredation by leopard as they can be easily killed and dragged to safe place (Bibi et al., 2013; Dar et al., 2009). Khorozyan et al. (2015) reported that big cats kill more livestock when
wild prey reaches a minimum threshold’. However, in the current study, since no previous data are available regarding abundance of the prey species of the carnivores in study area, we cannot generalize this fact here in this case. Neither there are any previous studies that could establish the status of the wild prey in the study area.
Therefore, it is hard to conclude that common leopard has depredated more on livestock because of the reason that its wild prey has reached a minimum threshold.
In the current study, common leopard has emerged as the major predator with
88.5% depredation on livestock (total losses 306 livestock during last 8 years), while other carnivores contributing only minor percentages such as Asiatic Jackal (Canis aureus) 5%, small Indian Civet (Viverricula indica) and Indian grey mongoose
(Herpestes edwardsii) 3% each. The most heavily depredated animal has been the domestic goat (82%), followed by domestic dog (9%). Few studies on common leopard have previously reported it as major predator of livestock including goat and sheep in some other areas of the Azad Jammu and Kashmir, Pakistan (Dar et al.,
2009; Qamar et al., 2010; Bibi et al., 2013; Kabir et al., 2014). Sekhar (1998) reported that leopard is responsible for 88% of goats, sheep and calves depredation in Sariska Tiger reserve Rajasthan, India. Similarly, Karanth and Sunquist (1995) reported mean body mass of leopard prey as 23 kg in tropical forests of India. The common leopard preferably preys on species weighing 10- 40 kg (Hayward et al.,
2006). In the current study, after goats, domestic dogs were most depredated species by common leopard. In the study area, domestic dogs are often used by local community for guarding purposes, therefore, these dogs are also more prone to leopard attacks. The findings of the current study are also supported by Dar et al.
(2009) who reported depredation of 34 dogs by common leopard in Machiara
National Park, Azad Jammu and Kashmir, even higher than the n = 24 dogs depredation being reported during the current study.
The current study indicates that there are three peaks of common leopard attacks on livestock in temporal context; majority of attacks occurred at morning time, followed by at night and then at evening time. More leopard depredation on livestock have been reported to occur between 9:00 am to 1200 noon, by the local respondents, and in fact, this is the high time when locals leave their livestock unattended , especially for grazing and thus make their livestock vulnerable for leopard depredation. Woodroffe et al. (2007) had reported that the risk of leopard attack was higher during the daytime for larger herds in Africa. Ahmed et al. (2012) had reported more leopard killing at night in central India. Some earlier studies also reported high frequency of leopard attacks during night time in other parts of Azad
Jammu and Kashmir (Dar et al., 2009; Kabir et al., 2014).
The common leopard depredated on livestock during whole year with majority of killing occurring in the months of July (22%), May (20%) and June
(18%), respectively. In the study area, these three months are hottest months regarding temperature and are categorized as summer season. Overall, majority
(61%) of livestock depredation by common leopard occurred in summer with only
19% being in winter season. During summer season, locals leave their livestock for grazing in the open fields, in the surrounding of villages in PLNP and thus livestock become an easy prey for common leopard. Another peak of leopard attacks was observed during the month of December, and it seems logical to increase guarding during summer and winter months to reduce negative human-carnivore interaction.
The current stud highlights total financial losses due to livestock depredation by
common leopard computed to be approximately US$ 80,000 during last eight years, which itself suggests alarming rate of livestock depredation by leopard.
Results of the current study showed that presence of both, human and dog, were most effective to reduce common leopard attacks on livestock, both inside and outside the village. Either dogs or human alone were not much effective in reducing leopard depredation on livestock and highest attacks were observed when livestock was left unattended. Ogada et al. (2003) had reported that better herding practices can reduce predator attacks on livestock. Earlier on, Kolowski and Holekamp (2006) also showed that dogs were ineffective in deterring leopard attacks in Kenya. Similar findings were reported by Dar et al. (2009) that dogs were ineffective in reducing leopard attacks on livestock in Machiara National Park, AJ&K. Kabir et al. (2014) reported guarding strategies to be ineffective in Machiara National Park, AJ&K.
Results of the current study showed that penning practices of livestock at night were ineffective in reducing leopard attacks since penning structures were very poor making it easier for common leopard to kill livestock due to its predatory behaviour. Such findings are in line with other studies conducted in Azad Jammu and Kashmir, Pakistan; for example Dar et al. (2009) reported poor penning conditions in Machiara National Park, AJ&K. Improving conditions of pens/corals can help in reducing the livestock losses due to leopard attacks.
The local community at PLNP heavily depends upon livestock forsubsistence and any livestock depredation by leopard may result in its retaliatory killing. We recorded higher numbers of leopards persecuted (n = 15) in retribution to livestock losses than other studies conducted earlier in the region (Dar et al., 2009; Bibi et al.,
2013; Kabir et al., 2014) with 4 humans being injured and 1 human killed from 2008-
2015 due to common leopard attack. The rate of retaliatory killing suggests that negative human-leopard interaction might have increased (86%) in PLNP compared to some other protected areas of the region. Dar et al. (2009) had reported that people think leopard is not dangerous which is contrary to current study findings. We found local perception hugely negative and majority of respondents were not willing to conserve leopard in their surroundings. Baral and Henien (2007) had earlier on reported that leopard depredation on livestock resulted in negative attitudes of local communities towards leopard. Previous studies had also reported negative perception of local communities towards leopard in other protected areas of Azad Kashmir (Bibi et al., 2013; Kabir et al., 2014). At current, there is no compensation for local communities in the PLNP which might be resulting in negative attitudes towards leopard and other carnivores. Local compensation schemes for livestock can improve local tolerance towards main challenging carnivore species (Bagchi and Mishra,
2006; Dar et al., 2009). However, the compensation should be according to the local rates and be delivered expeditiously in order to be effective (Madhusan, 2003;
Naughton-Treves et al., 2003).
7.1 RECOMMENDATIONS
In diet composition studies of carnivores, identification of scats should be based on their genotype to reduce errors resulting from morphological identification of carnivore scats. Wild prey of carnivores should be conserved in the study area.
Habitat of the study area should be conserved since many carnivore species are omnivorous/ frugivorous. Size of Pir Lasura national park should be increased and areas such as Supply, and Panagali should be included in National Park area since many carnivore species were abundant in these areas. There is great need of
compensating local communities for their losses to change their negative perception from killing of common leopard and other carnivores to their conservation. In addition, awareness campaign may also be launched in the study area, to educate people about ecological importance of the leopard and other carnivores so that they could show willing to conserve the carnivore predator. Moreover, they also need to be made aware about modern structures of pens/corals to reduce livestock depredation by common leopard. Also, when livestock is grazing near or outside of villages, guarding by both humans and dogs can help reduce the chances of leopard attack. Increase guarding practices during the months of May, June, July and
December can also help in reducing the livestock losses. Mitigation measures need to be set up, especially for sampling sites such as Pir Kana, Sairi, Katera khandhar,
Pehligali, and Panaggali which were identified as hotspot areas facing livestock depredation by common leopard.
7.2 FUTURE PROSPECTIVE
Modern techniques like next generation sequencing can help to identify predator as well as prey species from carnivore scats, future studies should use such advance methods for assessing dietary habits of carnivore. More detailed studies should be conducted in other parts of the country to provide baseline data prerequisite for conservation and management of wild carnivores.
SUMMARY
Knowledge of a predator’s diet is important for understanding its ecology and for predicting its influence on the dynamics of prey populations. Diet composition of carnivores is usually investigated using their fecal droppings. Species having similar ecological niches often shift their uses of resources in habitats where they are sympatric and such a phenomenon is known as “resource partitioning”. Interspecific competition is important force in structuring resource composition of carnivore community. The current study aimed at investigating resource partitioning among sympatric carnivore species inhabiting Pir Lasura National Park, AJ&K. The main objectives included documenting the diversity and distribution of carnivores at
PLNP, to investigate the diet composition of carnivores in the Park, niche breadth and niche overlap among sympatric carnivores and the human- carnivore conflict at
PLNP.
The study recorded seven different species of carnivores in the Park including; common leopard (Panthera pardus), Asiatic jackal (Canis aureus),
Kashmir hill fox (Vulpes vulpes griffithi), Asian palm civet (Paradoxurus hermaphroditus), small Indian civet (Viverricula indica), Indian grey mongoose
(Herpestes edwardsii), and the small Indian mongoose (Herpestes javanicus). Direct and indirect signs of common leopard, Asiatic jackal, Kashmir hill fox, Asian palm civet and small Indian mongoose were recorded in all 30 sampling sites having different elevational range. Indian grey mongoose was found at 15 different sampling sites while signs of small Indian civet were recorded at 7 sites. The scats were subjected to molecular identification for species identification. Accuracy for common leopard scats was found to be 95.8%, Kashmir hill fox 88.9%, Asiatic jackal
211
90.3%, Asian palm civet 74.3%, and small Indian civet 92.4%. Morphological characteristics of scats showed that they overlap greatly among different species based on their diameter and length. Scat analysis showed seventeen prey species in the diet of common leopard (10 wild and 7 domestic), total frequency of wild prey being 24 % while domestic prey contributed for 41 %. The diet of Asiatic Jackal comprised of 15 prey species (10 wild and 5 domestic), consumption of wild prey was 12.58% and domestic prey contributed 40.74%. A total of 21 prey species were recorded in diet of Kashmir hill fox (10 wild and 5 domestic) in addition to 6 plant species. Total consumption of wild prey was 17%, domestic 48.5% and plants 26 %.
The diet of Asian palm civet comprised of 27 prey items, consumption of wild prey was 33%, domestic prey 11 %, and plants 52%. In the diet of small Indian civet, 17 prey species were recorded, consumption of wild prey being 28.37%, domestic
13.43% and plants 47.76%. The diet of Indian grey mongoose consisted of 15 prey species (14 wild and only 1 domestic) in addition to 6 plant species. The consumption of wild prey was 47.56%, while domestic prey contributed 15.57%, and plants
11.48%. A total of 18 dietary items were recorded in diet of small Indian mongoose, including 11 wild, one domestic and 6 plant species. The abundance of different vertebrate and invertebrate prey species was also estimated in the study area to confirm the availability of wild prey.
Niche breadth of Indian grey mongoose was widest 18 (0.72) followed by
Asiatic jackal 14.2 (0.78), common leopard 13.88 (0.72), small Indian mongoose 12
(0.64), small Indian civet 10(0.52), Asian palm civet 9.69 (0.31), and Kashmir hill fox 7.89(0.31). High niche overlap was found between Asian palm civet and small
Indian civet (0.9), Indian grey mongoose and small Indian mongoose (0.89), Asiatic jackal and Kashmir hill fox (0.81), while lowest niche overlap was recorded common
leopard and small Indian civet (0.04). Data on livestock depredation by carnivores showed that 170 people lost 306 livestock, poultry, dogs, and pets to four different carnivore species; common leopard being the major predator killing (88.5%) livestock, with minor contributions from Asiatic Jackal (5.2%), small Indian Civet
(3.2%) and Indian grey mongoose (2.9%). The depredated livestock by leopard mainly included goats (82.2%) and dogs (9.2%). Common leopard was responsible for majority of the financial loss estimated approximately at US$ 80,000 and it negatively affected local people tolerance towards this top predator. The study concludes, niche of seven sympatric carnivore species overlaps in the study area.
LITERATURE CITED
Ahmad, M. F. and S. A. Ghalib. 1979. A checklist of mammals of Pakistan. Records
Zool. Surv. Pakistan, 7: 1-34.
Ahmed, K. and J. A. Khan. 2008. Food habits of leopard in tropical moist deciduous
forest of Dudhwa National Park, Uttar Pradesh, India. International J. Ecol.
Environ. Sci., 34(2): 141-147.
Ahmed, R. A., K. Prusty, J. Jena, C. Dave, K. R. D. Sunit, K. S. Hemanta and S. D.
Rout. 2012. Prevailing human carnivore conflict in Kanha–Achanakmar
Corridor, Central India. World J. Zool., 7(2): 158-164.
Aiyadurai, A. and Y. V. Jhala. 2006. Foraging and habitat use by golden jackal
(Canis aureus) in the Bhal region, Gujarat, India. J. Bombay Nat. Hist. Soc.,
103(1): 5-12.
Amores, F. 1975. Diet of the red fox (Vulpes vulpes) in the Western Sierra Morena
(South Spain). Donana Acta Vert., 2(2): 221-239.
Andheria, A. P., K. U. Karanth and N. S. Kumar. 2007. Diet and prey profiles of
three sympatric large carnivores in Bandipur Tiger Reserve, India. J. Zool.,
273(2): 1-7.
Aragona, M. and E. Z. Setz. 2001. Diet of the smaned wolf, Chrysocyon brachyurus
(mammalia: Canidae), during wet and dry seasons at Ibitipoca State Park,
Brazil. J. Zool., 254(1): 131-136.
Arivazhagan, C., R. Arumugam and K. Thiyagesan. 2007. Food habits of leopard
(Panthera pardus fusca), dhole (Cuon alpinus) and striped hyaena (Hyaena
hyaena) in a dry thorn forest of Southern India. J. Bombay Nat. Hist. Soc.,
104(1): 178-87.
214
Aroon, S., T. Artchawakom, J. G. Hill and N. Thanee. 2012. Seasonal variation in
the diet of common palm civet (Paradoxurus hermaphroditus) at Sakaerat
Bioshpere Reserve, Thailand. International Conference Proceedings, p. 191-
199.
Athreya, V., M. Odden, J. D. C. Linnell, J. Krishnaswamy and K. U. Karanth. 2013.
Big cats in our backyards: Persistence of large carnivores in a human
dominated landscape in India. PLOS ONE, 8(3): 57872.
Athreya, V., M. Odden, J. D. C. Linnell, J. Krishnaswamy and K. Karanth. 2016. A
cat among the dogs: Leopard Panthera pardus diet in a human-dominated
landscape in western Maharashtra, India. Oryx, 50(1): 156-162.
Athreya, V. R. and A. V. Belsare. 2007. Human–leopard conflict management
guidelines. Kaati Trust, Pune, 63 pp.
Bagchi, S. and C. Mishra. 2006. Living with large carnivores: Predation on livestock
by the snow leopard (Uncia uncia). J. Zool., 268(3): 217-224.
Bailey, T. N. 1993. The African leopard. Ecology and behaviour of a solitary felid.
Columbia University Press. 429 pp.
Bakaloudis, D. E., A. V. A. Bontzorlos, C. G. Vlachos, M. A. Papakosta, E.
N. Chatzinikos, S. G. Braziotis and V. J. Kontsiotis. 2015. Factors affecting
the diet of the red fox (Vulpes vulpes) in a heterogeneous mediterranean
landscape. Turk. J. Zool., 39(6): 1151-1159.
Baker, P., M. Furlong, S. Southern and S. Harris. 2006. The potential impact of red
fox Vulpes vulpes predation in agricultural landscapes in lowland Britain.
Wildlife Biol., 12(1): 39-50.
Baker, P. J., L. Boitani, S. Harris, G. Saunders and P. C. L. White. 2008. Terrestrial
carnivores and human food production: Impact and management. Mammal
Rev., 38(2): 123-166.
Balakrishnan, M. and M. B. Sreedevi. 2007. Captive breeding of the small Indian
civet viverricula indica (é. Geoffroy saint-hilaire, 1803). Small Carnivore
Conserv., 36: 5-8.
Baltrunaite, L. 2002. Diet composition of the red fox (Vulpes vulpes), pine marten
(Martes martes) and raccoon dog (Nyctereutes procyonoides) in clay plain
landscape, Lithuania. Acta Zool Litu., 12(4): 362-368.
Baltrunaite, L. 2001. Feeding habits, food niche overlap of red fox (Vulpes vulpes)
and pine marten (Martes martes) in hilly Moraine highland, Lithuania.
Ecologija, 2(1): 27-31.
Bang, P. and P. Dahlström. 1975. Huellas y señales de los animales de Europa.
Omega, Barcelona. 239 pp.
Baral, N. and J. T. Henien. 2007. Resources use, conservation attitudes, management
intervention and park-people relations in the western Terai landscape of
Nepal. Environ. Conserv., 34(1): 64-72.
Barun, A., I. Budinski and D. Simberloff. 2008. A ticking time-bomb? The small
Indian mongoose in Europe. Aliens, 26(1): 14-16.
Basuony, M., M. Saleh, A. Riad and W. Fathy. 2005. Food composition and feeding
ecology of the red fox Vulpes vulpes (linnaeus, 1758) in Egypt. Egyptian J.
Biol., 7(1): 796-102.
Bauer, D., M. Schiess-Meier, D. R. Mills and M. Gusset. 2014. Using spoor and prey
counts to determine temporal and spatial variation in lion (Panthera leo)
density. Can. J. Zool., 92(2): 97-104.
Beckmann, J. P. and J. Berger. 2003. Rapid ecological and behavioural changes in
carnivores: The responses of black bears (Ursus americanus) to altered food.
J. Zool., 261(2): 207-212.
Begon, M. 1979. Investigating animal abundance: Capture-recapture for biologists.
University Park Press, Baltimore, Maryland. 97 pp.
Beja-Pereira, A., R. Oliveira, P. C. Alves, M. K. Schwartz and G. Luikart. 2009.
Advancing ecological understandings through technological transformations
in noninvasive genetics. Mol. Ecol. Resour., 9(5): 1279-1301.
Bertram, B. C. B. 1999. Leopard. In: D. W. Macdonald, (eds.), The encyclopedia of
mammals. Andromeda Oxford Limited, Oxford, UK. 44-48 pp.
Bibi, S. S., R. A. Minhas, M. S. Awan, U. Ali and N. I. Dar. 2013. Study
of ethno-carnivore relationship in dhirkot, Azad Jammu and Kashmir. The
JAPS., 23(3): 854-859.
Bino, G., A. Dolev, D. Yosha, A. Guter, R. King, D. Saltz and S. Kark. 2010. Abrupt
spatial and numerical responses of overabundant foxes to a reduction in
anthropogenic resources. J. Appl. Ecol., 47(6): 1262-1271.
Biswas, S. and K. Sankar. 2002. Prey abundance and food habit of tigers (Panthera
tigris tigris) in Pench National Park, Madhya Pradesh. India. J. Zool.,
(Lond.). 256(3): 411-420.
BNHS, 1913. Survey of Kathiawar sept. 1912- Feb 1913. JBNHS 23(3): 464-486.
Boitani, L. and R. A. Powell. 2012. Carnivore ecology and conservation: A
handbook of techniques. Oxford University Press. 491 pp.
Boomgaard, P. 2001. Frontiers of fear: Tigers and people in the Malay world, 1600-
1950. Yale University Press, New Haven, USA. 352 pp.
Brannon, M. P. 2000. Niche relationships of two syntopic species of shrews, Sorex
fumeus and S. cinereus, in the Southern Appalachian mountains. J. Mammal.,
81(4): 1053-1061.
Brown, G. W. and B. E. Triggs. 1990. Diets of wild canids and foxes in east
Gippsland 1983-1987, using predator scat analysis. Aust. Mammal., 13(1):
209-213.
Brown, W. L. and E. O. Wilson. 1956. Character displacement. Syst. Zool., 5(2): 49-
64.
Bujne, A. E. 2000. Pollen analysis of faeces as a method of demonstrating seasonal
variations in the diet of Svalbard Reindeer (Rangifer tarandus
platyrhynchus). Polar. Res., 19(2): 183-192.
Burnham, K. P., D. R. Anderson and J. L. Laake. 1980. Estimation of density from
line transect sampling of biological populations. Wildlife Monographs,
72(1): 1-200.
Busby, G. B. J., D. Gottelli, T. Wacher, L. M. F. Belbachir, K. D. Smet, A. B. A.
Fellous, M. Belghoul and S. M. Durant. 2009. Genetic analysis of scat reveals
leopard panthera pardus and cheetah Acinonyx jubatus in southern Algeria.
Oryx, 43(3): 412-415.
Calisti, M., B. Ciampalini, S. Lovari and M. Lucherini. 1990. Food habits and trophic
niche variation of the red fox Vulpes vulpes in a mediterranean coastal area.
Rev. Ecol., 45(1): 309-320.
Camardella, A. R., M. F. Abreu and E. Wang. 2000. Marsupials found in felids scats
is southern Brazil, and a range extension of Monodelphis theresa. Mammalia,
64(3): 379-382.
Campbell, H. W. and S. P. Chiristman. 1998. Herpetological communities, wildlife
research report 13. Washington, D. C.: U.S. Fish and Wildlife Service. 193-
200.
Carbone, C. and J. L. Gittleman. 2002. A common rule for the scaling of carnivore
density. Sci. Justice., 295(5563): 2273-2276.
Carbone, C., N. Pettorelli and P. A. Stephens. 2010. The bigger they come, the harder
they fall: Body size and prey abundance influence predator–prey ratios. Biol.
Lett., 7(11): 312-315.
Carrera, R., W. Ballard and P. Gipson. 2008. Comparison of Mexican wolf and
coyote diets in Arizona and New Mexico. J. Wild. Manage., 72(2): 376-381.
Carvalho, J. C. and P. Gomes. 2004. Feeding resource partitioning among four
sympatric carnivores in the Peneda-Gerês National Park (Portugal). J. Zool.,
263(3): 275-283.
Charoo, S. A., L. K. Sharma, S. Sathyakumar and R. Y. Naqash. 2010. First record
of small Indian civet Viverricula indica in the Kashmir Himalaya, India.
Small Carnivore Conserv., 43: 42-43.
Chattha, S. A., S. M. Hussain, A. Javid, M. N. Abbas, S. Mahmood, M. G. Barq and
M. Hussain. 2015. Seasonal diet composition of leopard (Panthera pardus)
in Machiara National Park, Azad Jammu and Kashmir, Pakistan. Pakistan J.
Zool., 47(1): 201-207.
Chaves, P. B., V. G. Graeff, M. L. B. Lion, L. R. Oliveira and E. Eizirik. 2012. DNA
barcoding meets molecular scatology: Short mtDNA sequences for
standardized species assignment of carnivore noninvasive samples. Mol.
Ecol. Resour., 12(1): 18-35.
Chen, M., M. E. Tewes, K. Pei and L. I. Grassman. 2009. Activity patterns and
habitat use of sympatric small carnivores in southern Taiwan. Mammalia,
73(1): 20-26.
Chinchilla, F. A. 1997. La dieta del jaguar (Panthera onca), El puma (Felis concolor)
Yel manigordo (Felis pardalis) (carnivora: Felidae) En El Parque Nacional
Corcovado, Costa Rica. Rev. Biol. Trop., 45(3): 1223-1229.
Choudhury, A. 2013. The mammals of north east India. Gibbon Books and the Rhino
Foundation for Nature in N. E. India, Guwahati. 431 pp.
Chuang, S. and L. Lee. 1997. Food habits of three carnivores species (Viverricula
indica, Herpestes urva, and Melogale moschata) in Fushan forest, northern
Taiwan. J. Zoo., 243(1): 71-79.
Chutipong, W., N. Tantipisanuh, D. Ngoprasert, A. J. Lynam, R. Steinmetz, K. E.
Jenks, L. I. J. Grassman, M. Tewes, S. Kitamura, M. C. Baker, W. Mcshea,
N. Bhumpakphan, R. Sukmasuang, G. A. Gale, F. K. Harich, A. C. Treydte,
P. Cutter, P. B. Cutter, S. Suwanrat, K. Siripattaranukul, H. B. W. R. Station,
W. R. Division and J. W. Duckworth. 2014. Current distribution and
conservation status of small carnivores in Thailand: A baseline review. Small
Carnivore Conserv., 51: 96-136.
Ciampaloni, B. and S. Lovari. 1985. Food habits and trophic niche overlap of the
badger (Meles meles) and the red fox (Vulpes vulpes) in a Mediterranean
coastal area. Z. Saugetierk, 50(1): 226-234.
Clark, E. L. and J. Munkhbat. 2006. Mongolian red list of mammals. Zoological
Society of London, London, UK. 1206 pp.
Colwell, R. K. and D. J. Futuyma. 1971. On the measurement of niche breadth and
overlap. Ecol., 52(4): 567-576.
Conforti, V. A. and C. C. D. Azevedo. 2003. Local perceptions of jaguars (Panthera
onca) and pumas (Puma concolor) in the Iguacu National Park area, south
Brazil. Biol. Conserv., 111(2): 215-221.
Corbet, G. B. and J. E. Hill. 1992. The mammals of the indo-malayan region. Natural
History Museum publications, Oxford University Press, Oxford, U. K. 496
pp.
Corlett, R. T. 1998. Frugivory and seed dispersal by the vertebrates in the Oriental
(Indomalayan) region. Biol. Rev., 73: 414-448.
Cowlishaw, G. 1994. Vulnerability to predation in baboon populations. Behav.,
131(3): 293-304.
Crooks, K. R., S. P. D. Riley, S. D. Gehrt, T. E. Gosselink and T. R. V. Deelen. 2010.
Community ecology of urban carnivores. In: S. D. Gehrt, S. P. D. Riley, and
B. L. Cypher, (edit..), p. 185-196.
Dalén, L., A. Gotherstrom and A. Angerbjorn. 2004. Identifying species from pieces
of faeces. Conserv. Genet., 5(1): 109-111.
Daniel, J. C. 2009. The leopard in India. A Natural History. Natraj Publishers. 271
pp.
Dar, N. I., R. A. Minhas, Q. Zaman and M. Linkie. 2009. Predicting the patterns,
perceptions and causes of human–carnivore conflict in and around Machiara
National Park, Pakistan. Biol. Conserv., 142(10): 2076-2082.
Davison, A., J. D. S. Birks, R. C. Brookes, T. C. Braithwaite and J. E. Messenger.
2002. On the origin of faeces: Morphological versus molecular methods for
surveying rare carnivores from their scats. J. Zool., 257(2): 141-143.
Dayan, T. and D. Simberloff. 2005. Ecological and community-wide character
displacement: The next generation. Ecol. Lett., 8(8): 875-894.
Dayan, T. and D. Simberloff. 1998. Size patterns among competitors: Ecological
character displacement and character release in mammals, with special
reference to island populations. Mamm. Rev., 28(3): 99-124.
Delibes, M. 1983. Interspecific competition and the habitat of the stone marten
Martes foina. Acta. Zool. Fenn., 174(1): 229-231.
Diamond, J. 2002. Evolution, consequences and future of plant and animal
domestication. Nature, 418(6898): 700-707.
Diaz-Ruiz, F., M. Delibes-Mateos, J. L. Garcia-Moreno, J. M. Lopez-Martin, C.
Ferreira and P. Ferreras. 2013. Biogeographical patterns in the diet of an
opportunistic predator: The red fox Vulpes vulpes in the Iberian Peninsula.
Mammal Rev., 43(1): 59-70.
Donadio, E. and S. W. Buskirk. 2006. Diet, morphology, and interspecific killing in
carnivora. Am. Nat., 167(4): 524-536.
Englund, J. 1965. Studies on food ecology of red fox (Vulpes vulpes) in Sweden.
Viltrevy, 3(1): 377-385.
Erlinge, S. 1981. Food preferences, optimal diet and reproductive output in stoats
Mustela erminea in Sweden. Oikos, 36(3): 303-315.
Erlinge, S., G. Goransson, G. Hansson, L. Hogstedt, G. H. Liberg, O. Nilsson, I.
N. Nilsson, T. Von-Schantz and M. Sylven. 1983. Predation as a regulating
factor on small rodent populations in southern Sweden. Oikos, 40(1): 35-52.
Ewer, R. F. 1973. The carnivores. Ithaca, Cornell University Press, NY. 500 pp.
Farrell, L. E., J. Roman and M. E. Sunquist. 2000. Dietary separation of sympatric
carnivores identified by molecular analysis of scats. Mol. Ecol., 9(10): 1583-
1590.
Fedriani, J. M., T. K. Fuller, R. M. Sauvajot and E. C. York. 2000. Competition and
intraguild predation among three sympatric carnivores. Oecologia, 125(2):
258-270.
Fisher, M. 2016. Whose conflict is it anyway? Mobilizing research to save lives.
Oryx, 50(3): 377-378.
Flower, M. S. S. 1932. Notes on the recent mammals of Egypt, with a list of the
species recorded from that Kingdom. J. Zool., 102(2): 369- 450.
Fragoso, J. M. V. and J. M. Huffman. 2000. Seed-dispersal and seedling recruitment
patterns by last neotropical megafaunal element in Amazon, the Tapir. J.
Trop. Ecol., 16(3): 369-385.
Francis, C. M. 2008. A field guide to the mammals of south-east Asia. Princeton
University Press. Princeton, New Jersey and Oxford, United Kingdom. 392
pp.
Fuller, T. K. and P. R. Sievert. 2001. Carnivore demography and the consequences
of changes in prey availability. In: J. L. Gittleman, S. M. Funk, D. W.
Macdonald and R. K. Wayne, (eds.), Carnivore conservation. Cambridge
University Press. p. 163-178.
Garcia, N. and E. Virgos. 2007a. Evolution of community composition in several
carnivore palaeoguilds from the European pleistocene: The role of
interspecific competition. Lethaia, 40(1): 33-44.
Garcia, N. and E. Virgos. 2007b. Evolution of community composition in several
carnivore palaeoguilds from the European pleistocene: The role of
interspecific competition. Lethaia, 40(1): 33-44.
Gehrt, S. D., S. P. D. Riley and B. L. Cypher. 2010. Urban carnivores: Ecology,
conflict and conservation. The Johns Hopkins University Press. 261 pp.
Gibbs, J. P. 2000. Monitoring populations. In: L. Boitani & T. K. Fuller, (eds.),
Research Techniques in Animal Ecology, Controversies and Consequences.
Columbia University Press, New York, 422 pp.
Gilchrist, J. S., A. P. Jennings, G. Veron and P. Cavallini. 2009. Family herpestidae.
In: D. E. Wilson & R. A. Mittermeier, (eds.), Handbook of the mammals of
the World. Vol. 1: Carnivores. Lynx Editions, Barcelona, Spain, 656 pp.
Gittleman, J. L., S. M. Funk, D. W. Macdonald and R. K. Wayne. 2001. Why
“carnivore conservation”? In: J. L. Gittleman, S. M. Funk, D. W. Macdonald,
& R. K. Wayne, (eds.), Carnivore conservation. Cambridge University Press,
United Kingdom. p. 1-8.
Gittleman, J. L. and M. E. Gompper. 2005. Plight of predators. The importance
of carnivores for understanding patterns of biodiversity and extinction
risk. In: P. Brbosa & I. Castellanous, (eds.), Oxford University Press, United
Kingdom. 827 pp.
Glen, A. S. and C. R. Dickman. 2008. Niche overlap between marsupial and
eutherian carnivores: Does competition threaten the endangered spotted-
tailed quoll? J. Appl. Ecol., 45(2): 700-707.
Gloor, S., F. Bontadina, D. Hegglin, P. Deplazes and U. Breitenmoser. 2001. The
rise of urban fox populations in Switzerland. Mamm Biol., 66: 155-164.
Goszczyński, J. 1986. Diet of Foxes and Martens in Central Poland. Acta Theriol.,
31(36): 491-506.
Graham, K., A. Beckerman, S. Thirgood. 2005. Human–predator–prey conflicts:
Ecological correlates, prey loss and patterns of management. Biol. Conserv.,
122(2): 159-171.
Grassman, L. I. 1998. Movements and fruit selection of two paradoxurinae species
in a dry evergreen forest in southern Thailand. Small Carnivore Conserv,. 19:
25-29.
Greenslade, P. J. M. 1964. Pitfall trapping as a method for studying populations of
Carabidae. J. Anim. Ecol., 33(2): 301-310.
Gurung, B., J. L. D. Smith, C. Mcdougal, J. B. Karki and A. Barlow. 2008.
Factors associated with human-killing tigers in Chitwan National Park,
Nepal. Biol.Conserv., 141(12): 3069-3078.
Gusset, M., M. J. Swarner, L. Mponwane, K. Keletile and J. W. Mcnutt. 2009.
Human–wildlife conflict in northern Botswana: Livestock predation by
endangered African wild dog Lycaon pictus and other carnivores. Oryx,
43(1): 67-72.
Hanfee, F. and A. Hhmed. 2000. Some observations on India’s illegal trade in
mustelids, viverrids, and herpestids. In: S. A. Hussain (edit.), Wildlife and
protected areas, mustelids, viverrids, and herpestids of India. Envis Bulletin
p. 113-115.
Harris, S. and P. Baker. 2001. Urban foxes.British Natural History Series. 152 pp.
Hassinger, J. 1968. Introduction to the mammal survey of the 1965 street expedition
to Afghanistan. Fieldiana Zool., 55(1): 1-18.
Hayward, M. W., P. Henschel, J. O. Brien, M. Hofmeyr, G. Balme and G. H. I.
Kerley. 2006. Prey preferences of leopard (Panthera pardus). J. Zool.,
270(2): 298-313.
Hayward, M. W. and G. I. H. Kerley. 2008. Prey preferences and dietary overlap
amongst Africa’s large predators. S. Afr. J. Wild. Res., 38(2): 93-108.
Hayward, M. W. and R. Slotow. 2009. Temporal partitioning of activity in large
African carnivores: Tests of multiple hypotheses. S. Afri. J. Wild. Res.,
39(2): 109-125.
Heaney, L. R., D. S. Balete, M. L. Dolar, A. C. Alcala, A. T. L. Dans, P. C. Gonzales,
N. R. Ingle, M. V. Lepiten, W. L. R. Oliver, P. S. Ong, E. A. Rickart, B. R.
Tabaranza and R. C. B. Utzurrum. 1998. A synopsis of the mammalian fauna
of the Philippine Islands. Fieldiana. Zoology, new series, 88: 1-61.
Hersteinsson, P. and D. W. Macdonald. 1992. Interspecific competition and the
geographical distribution of red and artic foxes (Vulpes vulpes and Alopex
lagopus). Oikos, 64(3): 505-515.
Heyer, W. R., C. A. Cruz and O. L. Peixoto. 1988. Decimations, extinctions and
colonization of frog populations in southeast Brazil and their evolutionary
implications. Biotropica, 20(3): 230-235.
Hinton, H. E. and A. M. S. Dunn. 1967. Mongooses oliver and llayd, London. 144
pp.
Hunter, J. S. and T. M. Caro. 2008. Interspecific competition and predation in
American carnivore families. Ethol. Ecol. Evol., 20(4): 295-324.
Hunter, L. T. B. 2013. Panthera pardus. In: J. Kingdon & M. Hoffmann, (eds.),
Mammals of Africa. Volume V: Carnivora, pangolins, equids and
rhinoceroses. Bloomsbury Publishing. p. 159-168.
Hussain, R., T. Mahmood, F. Akrim, H. Fatima and M. S. Nadeem. 2017. Human
activity mediates reciprocal distribution and niche separation of two
sympatric mongoose species in Pothwar plateau, Pakistan. Turk J Zool.,
Accepted paper.
Inskip, C. and A. Zimmermann. 2009. Human–felid conflict: A review of patterns
and priorities worldwide. Oryx, 43(1): 18-34.
Isbell, L. A. 1994. Predation on primates: Ecological patterns and evolutionary
consequences. Evol. Anthropol., 3(2): 61-71.
IUCN. 2006. Http://www.Iucnredlist.Org/details/15954/0 accessed on 14 august,
2017.
Jackson, R. and D. O. Hunter. 1995. Snow leopard survey and conservation. 3rd ed.,
International snow leopard trust and U. S. National Biological Service,
Seattle. 120 pp.
Jackson, R. M., C. Mishra, T. M. Mccarthy and S. B. Ale. 2010. Snow leopards:
Conflict and conservation. In: D. W. Macdonald & A. J. Loveridge, (eds.),
Biology and conservation of wild felids. Oxford Univ. Press, Oxford, p. 417-
430.
Jacobson, A. P., P. Gerngross, J. R. Lemeris-Jr., R. F. Schoonover, C. Anco, C. B.
Wu¨Rsten, S. M. Durant, M. S. Farhadinia, P. Henschel, J. F. Kamler, A.
Laguardia, S. Rostro-Garcı´A, A. B. Stein and L. Dollar. 2016. Leopard
(Panthera pardus) status, distribution, and the research efforts across its
range. PeerJ, 1-28.
Jacomo, A. T. A., L. Silveira and J. A. Diniz-Filho. 2004. Niche separation between
the maned wolf (Chrysocyon brachyurus), the crab-eating fox (Dusicyon
thous) and the hoary fox (Dusicyon vetulus) in Central Brazil. J. Zool.,
(London). 262(1): 99-106.
Jaeger, M. M., E. Haque, P. Sultana and R. L. Bruggers. 2007. Daytime cover, diet
and space use of golden jackals (Canis aureus) in agro- ecosystems of
Bangladesh. Mammalia, 71(1): 1-10.
Jędrzejewski, W. and B. Jędrzejewska. 1992. Foraging and diet of the red fox Vulpes
vulpes in relation to variable resources in Białowieża National Park, Poland.
Ecography, 15(2): 212-220.
Jenkins, D. J. and N. A. Craig. 1992. The role of foxes Vulpes vulpes in the
epidemiology of Echinococcus granulosus in urban environments. Med. J.
Aust., 157(11): 754-756.
Jenks, G. F. and F. C. Caspall. 1971. Error on choroplethic maps: Definition,
measurement, reduction. Annals of the Association of American
Geographers, 61(12): 217-244.
Jennings, A. P. and G. Veron. 2011. Predicted distributions and ecological niches of
8 civet and mongoose species in Southeast Asia. J. Mammal., 92(2): 316-
327.
Jones, M. E. 1997. Character displacement in Australian dasyurid carnivores: Size
relationships and prey size. Ecology, 78(8): 2569-2587.
Jones, M. E. and L. A. Barmuta. 2000. Niche differentiation among sympatric
Australian dasyurid carnivores. J. Mammal., 81(2): 434-447.
Joshi, A. R., J. D. Smith and F. J. Cuthbert. 1995. Influene of food distribution and
predation pressure in spacing behavior in palm civets. J. Mammal., 76(4):
1205-1212.
Jothish, P. S. 2011. Diet of common palm civet Paradoxurus hermaphroditus in a
rural habitat in Kerala, India and its possible role in seed dispersal. Small
Carnivore Conserv., 45: 14-17.
Judas, J., P. Paillat, A. Khoja and A. Boug. 2006. Status of the Arabian leopard in
Saudi Arabia. Cat News, 1: 11-19.
Kabir, M. 2011. Food composition of common leopard (Panthera pardus) in
Machiara National park, Azad Jammu and Kashmir. (Unpublished) M. Phil.
thesis Department of Wildlife Management, Pir Mehr Ali Shah, Arid
Agriculture University Rawalpindi. 59 pp.
Kabir, M., A. Ghoddousi, M. S. Awan and M. N. Awan. 2014. Assessment of
human–leopard conflict in Machiara National Park, Azad Jammu and
Kashmir, Pakistan. Eur. J. Wildl. Res., 60(2): 291-296.
Karanth, K. U. and J. D. Nichols. 2010. Non-invasive survey methods for assessing
tiger populations. In: R. L. Tilson & P. J. Nyhus, (eds.), Tigers of the world:
The science, politics and conservation of Panthera tigris. Elsevier, p. 241-
262.
Karanth, K. U. and M. E. Sunquist. 2000. Behavioural correlates of predation by
tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus)
in Nagarahole, India. J. Zool., 250(2): 255-265.
Karanth, K. U. and M. E. Sunquist. 1995. Prey selection by tiger, leopard, and dhole
in tropical forests. J. Anim. Ecol., 64(4): 439-450.
Kauhala, K. and M. Auniola. 2001. Diet of raccoon dogs in summer in the Finnish
Archipelago. Ecography, 24(2): 151-156.
Khorozyan, I., A. Ghoddousi, M. Soofi and M. Waltert. 2015. Big cats kill more
livestock when wild prey reaches a minimum threshold. Biol. Conserv., 192:
268-275.
Kidawa, D. and R. Kowalczyk. 2011. The effect of sex, age, season and habitat on
diet of the red fox V. vulpes in Northeastern Poland. Acta Theriol., 56(3):
209-218.
Kissui, B. M. 2008. Livestock predation by lions, leopards, spotted hyenas, and their
vulnerability to retaliatory killing in the Maasai Steppe, Tanzania. Anim.
Conserv., 11(5): 422-432.
Kitamura, S., T. Yumoto, P. Poonswad, P. Chuailua, P. Plongmai, K. T. Maruhashi
and N. Noma. 2002. Interactions between fleshy fruits and frugivores in a
tropical seasonal forest in Thailand. Oecologia, 133(4): 559-572.
Kohn, M. H., E. C. York, D. A. Kamradt, G. Haugt, R. M. Sauvajot and R. K. Wayne.
1999. Estimating population size by genotyping faeces. Proc. R. Soc. B.,
266(1420): 657-663.
Kolb, H. H. and R. Hewson. 1979. Variation in the diet of foxes in Scotland. Acta
Theriol., 24(6): 69-83.
Kolowski, J. M. and K. E. Holekamp. 2006. Spatial, temporal and physical
characteristics of livestock depredation by large carnivores along a Kenyan
reserve border. Biol. Conserv., 128(4): 529-541.
Krebs, J. R. and N. B. Davies. 1993. An introduction to behavioural ecology.
Blackwell Scientific Publications. 520 pp.
Krishnakumar, H. and M. Balakrishanan. 2003. Feeding ecology of the common
palm civet Paradoxurus hermaphroditus (pallas) in semi-urban habitats of
Trivandrum, India. Small Carnivore Conserv., 28: 10-11.
Krystufek, B., D. Murariu and C. Kurtonur. 1997. Present distribution of the
golden jackal Canis aureus in the Balkans and adjacent regions. Mammal
Rev., 27(2): 109-114.
Lanszki, J., M. Heltai and L. Szabo. 2006. Feeding habits and trophic niche
overlap between sympatric golden jackal (Canis aureus) and red fox (Vulpes
vulpes) in the Pannonian ecoregion (Hungary). Can. J. Zool., 84(11): 1647-
1656.
Lau, M. W. N., J. R. Fellowes and B. P. L. Chan. 2010. Carnivores (mammalia:
Carnivora) in South China: A status review with notes on the commercial
trade. Mammal Rev., 40(4): 247-292.
Lavoie, G. K. 1971. Food habits: A technique for slide preparation. Range science
department, US International Biological P. Technical Report, No. 69. p.1-5.
Leckie, F., S. J. Thirgood, R. May and S. M. Redpath. 1998. Variation in the diet of
the red foxes on Scottish Moorland in relation to prey abundance. Ecography,
21(6): 599-604.
Levins, R. 1968. Evolution in changing environments. Princeton University press,
Princeton, N. J. 120 pp.
Linkie, M., Y. Dinata, A. Nofrianto and N. Leader-Williams. 2007. Patterns and
perceptions of wildlife crop raiding in and around Kerinci Seblat National
Park, Sumatra. Anim. Conserv., 10(1): 127-135.
Linnell, J. D. C., J. E. Swenson and R. Andersen. 2001. Predators and people:
Conservation of large carnivores is possible at high human densities if
management policy is favorable. Anim Conserv., 4(4): 345-349.
Lotka, A. J. 1932. The growth of mixed populations: Two species competing for a
common food supply. J. Wash. Acad. Sci., 22(16): 461-469.
Loveridge, A. J., S. W. Wang, L. G. Frank and J. Seidensticker. 2010. People and
wild felids: Conservation of cats and management of conflicts. In: D. W.
Macdonald & A. J. Loveridge, (eds.), Biology and conservation of wild
felids. Oxford Univ. Press, Oxford. p. 161-195.
Luff, M. L. 1975. Some factors influencing the efficiency of pitfall traps. Oeologia,
19(4): 345-357.
Lunney, D., B. Triggs, P. Eby and E. Ashby. 1990. Analysis of scats of dogs Canis
familiaris and foxes Vulpes vulpes (canidae: Carnivora) in coastal forests
near Bega, New South Wales. Aust. Wildlife Res., 17(1): 61-68.
Mac-Nally, R. 1983. On assessing the significance of interspecific competition to
guild structure. Ecology, 64(6): 1646-1652.
Macdonald, D. 1984. The encyclopedia of mammals. Facts on File. Inc., New York.
895 pp.
Macdonald, D. W. 2006. The encyclopedia of mammals. Oxford University Press,
Oxford. 936 pp.,
Macdonald, D. W. 1979. Helpers' in fox society. Nature, 282(5734): 69-71.
Macdonald, D. W. and C. Sillero-Zubiri. 2002. Large carnivores and conflict: Lion
conservation in context. Wildlife conservation research unit. Oxford
University. p. 1-8.
Madhusan, M. D. 2003. Living amidst large wildlife: Livestock and crop depredation
by large mammals in the interior villages of Bhadra Tiger Reserve, South
India. Environ. Manage., 31(4): 466-475.
Mahmood, T., I. Hussain and M. S. Nadeem. 2011. Population estimates, habitat
preference and the diet of small Indian mongoose (Herpestes javanicus) in
Potohar plateau. Pak. J. Zool., 43(1): 103-111.
Mahmood, T., F. Niazi and M. S. Nadeem. 2013. Diet composition of Asiatic jackal
(Canis aureus) in Margallah Hills National Park, Islamabad, Pakistan. The
JAPS., 23(2): 444-456.
Majumder, A., K. Sankar, Q. Qureshi and S. Basu. 2011. Food habits and temporal
activity patterns of the golden jackal Canis aureus and the jungle cat Felis
chaus in Pench tiger reserve, Madhya Pradesh, India. J. Threat. Taxa., 3(11):
2221-2225.
Martin, S. K. 1994. Feeding ecology of American martens and fisheries. Martens,
sables and fishers: Biology and conservation. Cornell University Press,
Ithaca, New York. p. 297-315.
Meiri, S. 2005. Small carnivores on small islands: New data based on old skulls.
Small Carnivore Conserv., 33: 21-23.
Meriggi, A. and S. Lovari. 1996. A review of wolf predation in Southern Europe:
Does the wolf prefer to livestock? J. Appl. Ecol., 33(6): 1561-1571.
Miller, R. S. 1967. Pattern and process in competition. Adv. Ecol. Res., 4: 1-74.
Mishra, C., P. Allen, T. Mccarthy, M. D. Madhusudhan, A. Bayarjagal and H. H. T.
Prins. 2003. The role of incentive programs in conserving the snow leopard.
Conser. Biol., 17(6): 1512-1520.
Mishra, C., S. E. V. Wieren, P. Ketner, I. M. A. Heitkonig and H. H. T. Prins. 2004.
Competition between domestic livestock and wild bharal Pseudois nayaur in
the Indian Trans-Himalaya. J. Appl. Ecol., 41(2): 344-354.
Mitchell, B. D. and P. B. Banks. 2005. Do wild dogs exclude foxes? Evidence for
competition from dietary and spatial overlaps. Aust. Ecol., 30(5): 581-591.
Mizutani, F. 1999. Biomass density of wild and domestic herbivores and carrying
capacity on a working ranch in Laikipia District, Kenya. African J. Ecol.,
37(2): 226-240.
Moehlman, P. D. 1983. Socioecology of silverbacked and golden jackals (Canis
mesomelas and Canis aureus). In: J. F. Eisenberg & D. G. Kleiman, (eds.),
Recent advances in the study of mammalian behaviour. American Society of
Mammologists Spec. Publ. No. 7, p. 423-453.
Mondal, K., S. Gupta, Q. Qureshi and K. Sankar. 2011. Prey selection and food
habits of leopard (Panthera pardus fusca) in Sariska Tiger Reserve,
Rajasthan, India. Mammalia, 75(2): 201-205.
Mondal, P. C. K., K. Sankar and Q. Qureshi. 2012. Food habits of golden jackal
(canis aureus) and striped hyena (hyaena hyaena) in Sariska Tiger Reserve,
western India. W. J. Zool., 7(2): 106-112.
Mudappa, D. 2013. Herpestids, viverrids and mustelids. In: A. J. T. Johnsingh & N.
Manjrekar, (eds.), Mammals of South Asia: Ecology, behaviour and
conservation, Universities Press. p. 471-498.
Mukherjee, S., S. P. Goyal, A. J. T. Johnsingh and M. L. Pitman. 2004. The
importance of rodents in the diet of jungle cat (Felis chaus), caracal (Caracal
caracal) and golden jackal (Canis aureus) in Sariska Tiger Reserve,
Rajasthan, India. J. Zool., 262(4): 405-411.
Mukherjee, S. and C. Mishra. 2001. Predation by leopard Panthera pardus in
Majhatal Harsang wildlife sanctuary, western Himalaya. JBNHS., 98(2):
267-268.
Nadeem, M. S., R. Naz, S. I. Shah, M. A. Beg, A. R. Kayani, M. Mushtaq and T.
Mahmood. 2012. Season and locality-related changes in the diet of Asiatic
jackal (Canis aureus) in Potohar, Pakistan. Turk. J. Zool., 36(6): 798-805.
Naughton-Treves, L. 1998. Predicting patterns of crop damage by wildlife around
Kibale National park, Uganda. Conserv. Biol., 12(1): 156-168.
Naughton-Treves, L., R. Grossberg and A. Treves. 2003. Paying for tolerance? The
impact of livestock depredation and compensation payments on rural
citizens’ attitudes toward wolves. Conserv. Biol., 17(6): 1500-1511.
Nijman, V., D. Spaan, E. J. Rode-Margono, P. D. Roberts, Wirdateti and K. A. I.
Nekaris. 2014. Trade in common palm civet Paradoxurus hermaphroditus in
Javan and Balinese markets, Indonesia. Small Carnivore Conserv., 51: 11-
17.
Nowak, R. M. 2005. Walker's carnivores of the World. Johns Hopkins Univ. Press,
Baltimore, Maryland. 313 pp.
Nowell, K. and P. Jackson. 1996. Wild cats: Status survey and conservation action
plan. IUCN, Gland, Switzerland. 406 pp.
Ogada, M. O., R. Woodroffe, N. O. Oguge and L. G. Frank. 2003. Limiting predation
by African carnivores: The role of livestock husbandry. Conserv. Biol.,
17(6): 1521-1530.
Oli, M. K. 1993. A key for the identification of the hair of mammals of a snow
leopard (Panthera uncia) habitat in Nepal. J. Zool. Lond., 231(1): 71-93.
Osborn, J. and I. Helmy. 1980. The contemporary land mammals of Egypt (including
Sinai). Field. Zool., Series 5. 618 pp.
Padial, J. M., E. Avila and J. M. Gil-Sanche. 2002. Feeding habits and overlap among
red fox (Vulpes vulpes) and stone marten (Martes foina) in two mediterranean
mountain habitats. Mammal. Biol., 67(3): 137-146.
Page, L. K., R. K. Swihart and K. R. Kazacos. 2001. Seed preferences and foraging
by granivores at raccoon latrines in the transmission dynamics of raccoon
roundworm (Baylisascaris procyonis). Can. J. Zool., 79(4): 616-622.
Palomares, F. and M. Caro. 1999. Interspecific killing among mammalian
carnivores. Amer. Nat., 153(5): 492-508.
Patou, M. L., A. Wilting, P. Gaubert, J. A. Esselstyn, C. Cruaud, A. P. Jennings, J.
Fickel and G. Veron. 2010. Evolutionary history of the Paradoxurus palm
civets – a new model for Asian biogeography. J. Biogeogr., 37(11): 2077-
2097.
Patterson, B. D., S. M. Kasiki, E. Selempo and R. W. Kays. 2004. Livestock
predation by lions (Panthera leo) and other carnivores on ranches
neighboring Tsavo National parks, Kenya. Biol. Conserv., 119(4): 507-516.
Pianka, E. R. 1988. Evolutionary ecology. New York, Harper and Row Press. 356
pp.
Pianka, E. R. 1973. The structure of lizard communities. Annu. Rev. Ecol. Syst.,
4(1): 53-74.
Pocock, R. I. 1930. Mammalia: Faunna of British India, Families Felidae and
Viverridae. Carnivora, Sub-orders Aeluroidae and Arctoidea, vol. II. Taylor
and Francis, London. 279 pp.
Polis, G., C. Myers and R. Holt. 1989. The ecology and evolution of intraguild
predation: Potential competitors that eat each other. Annu. Rev. Ecol. Syst.,
20(1): 297-330.
Prakash, I. 1959. Food of some Indian Desert Mammals. J. Biol. Sci., 2(2): 100-
109.
Prugh, L. R. and C. E. Ritland. 2005. Molecular testing of observer identification of
carnivore feces in the field. Wildl. Soc. Bull., 33(1): 189-194.
Qamar, Q. Z., N. I. Dar, U. Ali, R. A. Minhas, J. Ayub and M. Anwar. 2010. Human–
leopard conflict: An emerging issue of common leopard conservation in
Machiara National Park, Azad Jammu and Kashmir, Pakistan. Pak. J. Wild.,
1(2): 50-56.
Rabinowitz, A. R. 1991. The behavior and movement of sympatric civet species in
Huai Kha Khaeng wildlife sanctuary, Thailand. J. Zool., 223(2): 281-298.
Ray, J. and M. Sunquist. 2001. Trophic relations in a community of African
rainforest carnivores. Oecologia, 127(3): 395-408.
Ray, J. C., L. Hunter and J. Zigouris. 2005. Setting conservation and research
priorities for larger African carnivores. Wildlife Conservation Society, New
York, USA. 203 pp.
Raymond, M., J. F. Robitaille, P. Lauzon and R. Vaudry. 1990. Prey-dependent
profitability of foraging behaviour of male and female ermine, Mustela
erminea. Oikos, 58(3): 323-328.
Redpath, S. M., M. Bhatia and J. Young. 2015. Tilting at wildlife: Reconsidering
human–wildlife conflict. Oryx, 49(2): 222-225.
Riaz, T., W. Shehzad, A. Viari, F. Pompanon, P. Taberlet and E. Coissac. 2011.
Ecoprimers: Inference of new DNA barcode markers from whole genome
sequence analysis. Nucleic Acids Res., 39(21): 145.
Ripple, W. J., J. A. Estes, R. L. Beschta, C. C. Wilmers, E. G. Ritchie and M.
Hebblewhite. 2014. Status and ecological effects of the world's largest
carnivores. Sci. Justice., 343(6167): 1241484.
Roberts, T. J. 1997. The mammals of pakistan. Oxford University Press. 525 pp.
Ross, S. T. 1986. Resource partitioning in fish assemblages: A review of field
studies. Copeia, (1986)2: 352-388.
Ruiz-González, A., J. O. Rubines-Berdión and B. J. Gómez-Moliner. 2008. A non-
invasive genetic method to identify the sympatric mustelids pine marten
(Martes martes) and stone marten (Martes foina): Preliminary distribution
survey on the northern Iberian Peninsula. Eur. J. Wildl. Soc., 54(2): 253-261.
Sahajpal, V., S. P. Goyal, R. Raza and R. Jayapal. 2009. Identification of mongoose
(genus: Herpestes) species from hair through band pattern studies using
Discriminate Functional Analysis (DFA) and microscopic examination. Sci.
Justice., 49(3): 205-209.
Sangay, T. and K. Vernes. 2008. Human–wildlife conflict in the kingdom of Bhutan:
Patterns of livestock predation by large mammalian carnivores. Biol.
Conserv., 141(5): 1272-1282.
Sankar, K. 1988. Some observations in Food Habits of Jackal (Canis aureus) in
Keoladeo National Park, Bharatpur, as shown by Scat Analysis. JBNHS.,
85(1): 185-186.
Sankar, K. and A. J. T. Johnsingh. 2002. Food habits of tiger (Panthera tigris) and
leopard (Panthera pardus) in Sariska Tiger Reserve, Rajasthan, India, as
shown by scat analysis. Mammalia, 66(2): 285-289.
Santiapillai, C., M. Desilva and S. Dissanayake. 2000. The status of mongooses
(family: Herpestidae) in Ruhuna National park, Sri Lanka. J. Bomb. Nat.
Hist. Soc., 97(2): 208-214.
Santos, M. F. M. and S. M. Hartz. 1999. The food habits of procyon cancrivorus
(carnivora, procyonidae) in the Lami Biological Reserve, Porto Alegre,
Southern Brazil. Mammalia, 63(4): 525-530.
Sanz, B., J. V. Turón and A. Balmorí. 2007. Huellas Y Rastros De Los Mamíferos
Ibéricos, second ed. Ediciones Muskari, Zaragoza. 304 pp.
Schafer, L. N., M. E. Platell, F. J. Valesini and I. C. Potter. 2002. Comparisons
between the influence of habitat type, season and body size on the dietary
compositions of fish species in near Shore Marine waters. J. Exp. Mar. Biol.
Ecol., 278(1): 67-92.
Schaller, G. B. 1967. The Deer and the Tiger. A study of Wildlife in India. University
of Chicago Press, Chicago and London. 370 pp.
Schaller, G. B. 1970. Observations on the Nilgiri Tahr Hemitragus hylocrius.
JBNHS., 67(3): 365-89.
Schaller, G. B. 1983. Mammals and their biomass on a Brazilian ranch. Arquivos de
Zoologia, 31(1): 1-36.
Schiess-Meier, M., S. Ramsauer, T. Gabanapelo and B. König. 2007.
Livestock predation, insights from problem animal control registers in
Botswana. J. Wildl. Manag., 71(4): 1267-1274.
Schoener, T. W. 1982. The controversy over interspecific competition. Amer. Sci.,
70(6): 586-595.
Schoener, T. W. 1974. Resource partitioning in ecological communities. Sci. Justice,
185(4145): 27-39.
Schoener, T. W. 1986. Resource partitioning. In: D. J. Anderson & J. Kikkawa,
(eds.), Community ecology; pattern and process. Blackwell Scientific
Publications; Melbourne, Australia, p. 91-126.
Sclater, W. L. and P. L. Sclater. 1899. The geography of mammals. London: K. Paul,
Trench, Trübner & co. 377 pp.
Seaman, G. and J. Randall. 1962. The mongoose as a predator in the Virgin Islands.
J. Mammal, 43(4): 544-546.
Seidensticker, J., M. Sunquist and C. Mcdougal. 1990. Leopards living at the edge
of the Royal Chitwan National Park, Nepal. In: J. C. Daniel & J. S. Serrao,
(eds.), Conservation in developing countries: Problems and prospects.
Oxford University Press, Bombay, India. p. 415-423.
Sekhar, N. U. 1998. Crop and livestock depredation caused by wild animals in
protected areas: The case of Sariska Tiger Reserve Rajasthan, India. Environ.
Conserv., 25(2): 160-171.
Serafini, P. and S. Lovari. 1993. Food habits and trophic niche overlap of the red fox
and the stone marten in a Mediterranean rural area. Acta Theriol., 38(3): 233-
244.
Seton, E. T. 1925. On the study of scatology. J. Mammal., 6(1): 47-49.
Shabbir, S., M. Anwar, I. Hussain and M. A. Nawaz. 2013. Food habits and diet
overlap of two sympatric carnivore species in Chitral, Pakistan. The JAPS.,
23(1): 100-107.
Shehzad, W., M. A. Nawaz, F. Pompanon, E. Coissac, T. Riaz, S. Shah and P.
Taberlet. 2015. Forest without prey: Livestock sustain a leopard Panthera
pardus population in Pakistan. Oryx, 49(2): 248-253.
Shekar, K. S. 2003. The status of mongooses in central India. Small Carnivore
Conserv., 29: 22-23.
Shepherd, C. R. 2012. Observations of small carnivores in Jakarta wildlife markets
, Indonesia, with notes on trade in Javan ferret badger Melogale orientalis
and on the increasing demand for common palm civet Paradoxurus
hermaphroditus for civet coffee production. Small Carnivore Conserv., 47:
38-41.
Siddiqui, M. S. 1961. Checklist of Mammals of Pakistan with particular reference
to the Mammalian Collection in the British Museum (Natural History)
London, Biologia, 7(2): 93-225.
Siddiqui, M. J. I., N. Rana and S. A. Rana. 2004. Analysis of the scats of small Indian
mongoose (Herpestes auropunctatus) with special reference to the insect
fauna in croplands of Faisalabad, Pakistan. Entomol., 26(1): 95-99.
Sidorovich, V. E., A. A. Sidorovich and I. V. Izotova. 2006. Variations in the diet
and population density of the Red fox Vulpes vulpes in the mixed woodlands
of Northern Belarus. Mammal. Biol., 71(2): 74-89.
Sih, A. 1993. Effects of ecological interactions on forager diets: Competition,
predation risk, parasitism and prey behaviour. In: R. N. Hughes, (eds.), Diet
selection. Blackwell Scientific Publications, Oxford, p. 182-211.
Singh, A., A. Mukherjee, S. Dookia and H. N. Kumara. 2016. High resource
availability and lack of competition have increased population of a meso-
carnivore—a case study of golden jackal in Keoladeo National Park, India.
Mammal Res., 61(3): 209-219.
Smith, T. B. 1991. Inter and intra-specific diet overlap during lean times between
Quetea erythrops and bill morghs of Pyrenestes ostrinus. Oikos, 60(1): 76-
82.
Soulsbury, C. D., G. Iossa, P. J. Baker and S. Harris. 2010. Red fox. In: S. D. Gehrt,
S. P. D. Riley, & B. L. Cypher, (eds.), Urban carnivores: Ecology, conflict,
and conservation. Johns Hopkins University Press, p. 63-75.
Spalton, J. A. and H. M. Al-Hikmani. 2006. The leopard in the Arabian Peninsula—
distribution and subspecies status. Cat News, 1: 4-8.
Stein, A. B. and V. Hayssen. 2013. Panthera pardus (Carnivora: Felidae).
Mammal. Spp., 47(1): 30-48.
Stevens, K., A. Dehgan, M. Karlstetter, F. R. F, I. T. Muhammad, S. Ostrowski, M.
A. Jan and A. Rita. 2011. Large mammals surviving conflict in the eastern
forests of Afghanistan. Oryx, 45(2): 265-271.
Su, S. and J. Sale. 2007. Niche differentiation between common palm civet
Paradoxurus hermaphroditus and small indian civet Viverricula indica in
regenerating degraded forest, Myanmar. Small Carnivore Conserv., 36: 30-
34.
Suryawanshi, K. R., Y. V. Bhatnagar, S. Redpath and C. Mishra. 2013. People,
predators and perceptions: Patterns of livestock depredation by snow
leopards and wolves. J. Appl. Ecol., 50(3): 550-560.
Sutherland, W. J. 1998. Ecological census techniques. Cambridge University Press.
336 pp.
Tall, L., L. Cloutier and A. Cattaneo. 2006. Grazer-diatoms size relationships in an
epiphytic community. Limnol. Oceanogr., 51(2): 1211-1216.
Terborgh, J., J. A. Estes, P. Paquet, K. Ralls, D. Boydheger, B. J. Miller and R. F.
Noss. 1999. The role of top carnivore in regulating terrestrial ecosystems. In;
M. Soule & J. Terborgh, (eds.), Continental conservation. The Island Press,
E.U.A., p. 39-64.
Than, Z., H. Saw, S. Htoo, T. Po, M. Maung, A. J. Lynam, T. L. Kyaw and J. W.
Duckworth. 2008. Status and distribution of small carnivores in Myanmar.
Small Carnivore Conserv., 38: 2-28.
Treves, A. and K. U. Karanth. 2003. Human–carnivore conflict and perspectives
on carnivore management worldwide. Conserv. Biol., 17(6): 1491–1499.
Triggs, B., H. Brunner and J. M. Cullen. 1984. The food of fox, dog and cat in
Croajingalong National Park, South Eastern Victoria. Aus. Wild. Res., 11(3):
491-499.
Van, R. H. and M. N. Jayakumar. 2003. The stripe-necked mongoose, Herpestes
vitticollis. Small Carnivore Conserv., 28: 14-17.
Veron, G., M. L. Patou, M. Toth, M. Goonatilake and A. P. Jennings. 2015. How
many species of paradoxurus civets are there? New insights from India and
Sri Lanka. J. Zoolog. Syst. and Evol. Res., 53(2): 161-174..
Walker, C. 1996. Signs of the wild. Struik Publish, Cape Town. 215 pp.
Walter, G. H. 1991. What is resource partitioning? . J. Theor. Biol. 150 (2): 137-143.
Wandeler, P., S. M. Funk, C. R. Largiader, S. Gloor and U. Breitenmoser. 2003. The
city-fox phenomenon: Genetic consequences of a recent colonization of
urban habitat. Mol Ecol. Res., 12(3): 647-656.
Wang, H. and T. K. Fuller. 2003. Food habits of four sympatric carnviores in
Southeastern China. Mammalia, 67(4): 513-519.
Wang, H. and T. K. Fuller. 2001. Notes on the ecology of sympatric small carnivores
in Southeastern China. Mamm. Biol., 66(4): 251-255.
Wang, P., H. Sheng and H. Lu. 1976. The analysis on the food habits of small Indian
civet and its use in captivity breeding. Chinese J. Zool., 20(2): 39-40.
Wang, S. and Y. Xie. 2004. China species Red list. Vol. 1 red list. Higher Education
Press. 224 pp.
Webbon, C. C., P. J. Baker and S. Harris. 2004. Faecal density counts for monitoring
changes in red fox numbers in rural Britain. J. Appl. Ecol., 41(4): 768-779.
Wemmer, C., T. H. Kunz, G. Lundie-Jenkins and W. Mcshea. 1996. Mammalian
sign. In: D. E. Wilson, F. R. Cole, J. D. Nichols, R. Rudran & M. S. Foster,
(eds.), Measuring and monitoring biological diversity - standard methods for
mammals. Smithsonian Institution Press, Washington, p.157-176.
White, G. C. and L. E. Eberhardt. 1980. Statistical analysis of deer and elk pellet
group data. J. Wildlife Manag., 44(1): 121-131.
White, P. J., K. Ralls and R. A. Garrott. 1994. Coyote-kit fox interactions as revealed
by telemetry. Can. J. Zool., 72(10): 1831-1836.
Whitfield, P. 1978. The hunters. New York: Simon and Schuster. United States
Department of Agriculture Circular. p. 118: 1-4.
Wiens, J. A. 1989. The ecology of bird communities. Cambridge University Press,
Cambridge. 543 pp.
Williams, P. A., B. J. Karl, P. Bannister and W. G. Lee. 2000. Small mammals as
potencial seed disperses in New Zealand. Austral Ecol., 25(5): 523-532.
Wood, D. H. 1988. Estimating rabbit density by counting dung pellets. Aust. Wildlife
Res., 15(6): 665-671.
Woodroffe, R., L. G. Frank, P. A. Lindsey, M. K. Symon, O. Ranah and S.
Romanach. 2007. Livestock husbandry as a tool for carnivore conservation
in Africa’s community rangelands: A case-control study. Biodivers. and
Conserv., 16(4): 1245-1260.
Woodward, G. and A. G. Hildrew. 2002. Body-size determinants of niche overlap
and intraguild predation within a complex food web. J. Anim. Zool., 71(6):
1063-1074.
Wozencraft, W. C. 2005. Order carnivora. In: D. E. Wilson & D. M. Reeder mammal
species of the world. Smithsonian Institution Press, p. 532-628.
Wyman, J. 1967. The jackals of the Serengeti. Animals, 10: 79-83.
Yirga, G., H. H. D. E. Iongh, H. Leirs, K. Gebrihiwot, J. Deckers and H. Bauer. 2012.
Adaptability of large carnivores to changing anthropogenic food sources:
Diet change of spotted hyena (Crocuta crocuta) during christian fasting
period in northern Ethiopia. J. Anim. Ecol., 81(5): 1052-1055.
Yu, J. N., S. H. Han, B. H. Kim, A. P. Kryukov, S. Kim, B. Y. Lee and M. Kwak.
2012. Insights into Korean red fox (Vulpes vulpes) based on mitochondrial
cytochrome b sequence variation in East Asia. Zool. Sci., 29(11): 753-760.
Zarco-González, M. M., O. Monroy-Vilchis and J. Alaniz. 2013. Spatial model of
livestock predation by jaguar and puma in Mexico: Conservation planning.
Biol. Conserv., 159: 80-87.
Zhang, C., M. Zhang and P. Stott. 2013. Does prey density limit Amur tiger Panthera
tigris Altaica recovery in northeastern China? Wildl. Biol., 19(4): 452-461.
Zuberbühler, K. and D. Jenny. 2002. Leopard predation and primate evolution. J.
Hum. Evol., 43(6): 873-886.
APPENDICES
Appendix I Dietary items of common leopard General Linear Model results Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Source of Squares df Mean Square F Sig. Corrected 86.000a 17 5.059 9.260 .000 Model Intercept 60.500 1 60.500 110.746 .000 species 86.000 17 5.059 9.260 .000 Error 29.500 54 .546 Total 176.000 72 Corrected Total 115.500 71 a. R Squared = .745 (Adjusted R Squared = .664)
Appendix II Dietary items of Asiatic jackal General Linear Model result output
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 203.444a 17 11.967 15.029 .000 Model Intercept 117.556 1 117.556 147.628 .000 speceis 203.444 17 11.967 15.029 .000 Error 43.000 54 .796 Total 364.000 72 Corrected 246.444 71 Total a. R Squared = .826 (Adjusted R Squared = .771)
246
Appendix III Dietary items of Kashmir hill fox General Linear Model results
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 667.609a 22 30.346 8.863 .000 Model Intercept 303.141 1 303.141 88.537 .000 species 667.609 22 30.346 8.863 .000 Error 236.250 69 3.424 Total 1207.000 92 Corrected 903.859 91 Total a. R Squared = .739 (Adjusted R Squared = .655)
Appendix IV Asian palm civet dietary items General Linear Model results
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 680.638a 28 24.308 4.488 .000 Model Intercept 321.112 1 321.112 59.282 .000 species 680.638 28 24.308 4.488 .000 Error 471.250 87 5.417 Total 1473.000 116 Corrected 1151.888 115 Total a. R Squared = .591 (Adjusted R Squared = .459)
Appendix V Small Indian civet prey species comparison GLM General Linear Model results
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 57.921a 18 3.218 2.800 .002 Model Intercept 50.579 1 50.579 44.015 .000 species 57.921 18 3.218 2.800 .002 Error 65.500 57 1.149 Total 174.000 76 Corrected 123.421 75 Total a. R Squared = .469 (Adjusted R Squared = .302)
Appendix VI Indian grey mongoose dietary items General Linear Model results
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 107.333a 23 4.667 5.744 .000 Model Intercept 104.167 1 104.167 128.205 .000 Species 107.333 23 4.667 5.744 .000 Error 58.500 72 .813 Total 270.000 96 Corrected 165.833 95 Total a. R Squared = .647 (Adjusted R Squared = .535)
Appendix VII Small Indian mongoose dietary items General Linear Model results
Tests of Between-Subjects Effects Dependent Variable: freq Type III Sum Mean Source of Squares df Square F Sig. Corrected 147.184a 18 8.177 6.611 .000 Model Intercept 202.316 1 202.316 163.574 .000 Species 147.184 18 8.177 6.611 .000 Error 70.500 57 1.237 Total 420.000 76 Corrected 217.684 75 Total a. R Squared = .676 (Adjusted R Squared = .574)
Appendix VIII Questionnaire for study of human carnivore interaction Questionnaire no.______
Date: ______Site______GPS Location______
Demographic and socio-economic characteristics of respondents
Name______
Sex______
Age______
Level of formal education______
Occupation______
Livestock holding (type and number of livestock kept) ______
Purpose of keeping livestock______
A- Human carnivore conflict experience 1- Direct sightings a) at day b) at night (alerted by dogs) 2- Indirect signs left near the kill______
3- If still remaining, marks left on the carcass, e.g., nape or throat bite indicating a leopard______B- Financial loss of livestock 1- Type and number of livestock killed______2- Number of predator individuals and livestock herd size at time of attack______3- Place ______4- Location (inside village or outside village; in forest, meadow, or grazing near village) ______5- Date of attack______6- Time of attack______7- Season of attack______C- Action taken against the predators 1- Livestock guarding a) none b) humans only c) dogs only d) human and dogs 2- Available habitat cover for leopard a) open b) close 3- Carnivore attack place of occurance a) inside village b) outside of village D- Information on local conditions 1- Presence of electricity a) yes b) no 2- Livestock penned at night a) yes b) no 3- Geographic position of carnivore attack______4- Elevation of carnivore attack______E- Perceptions towards carnivores 1- The temporal attack trends of the main conflict species______2- Reasons for any perceived increase in attacks______3- The relationship between attacks and PLNP______4- Knowledge about the authority responsible for conserving the main conflict species______5- Conservation status of predators______
F- Tolerance towards carnivores 1- Respondents willingness to conserve the main conflict species______2- Possible management options for mitigating predator attacks on livestock (1) do nothing
(2) repel the predator using disruptive stimulants
(3) habitat improvement for the predator
(4) translocation
(5) lethal control
Notes______