The flare-horned Markhor (Capra falconeri

Transcription

The flare-horned Markhor (Capra falconeri
COMPETITION FOR FOOD RESOURCES BETWEEN MARKHOR (Capra
falconeri) AND DOMESTIC UNGULATES IN CHITRAL AREA
NASRA ASHRAF
06-arid-593
Department of Wildlife Management
Faculty of Forestry, Range Management and Wildlife
Pir Mehr Ali Shah
Arid Agriculture University Rawalpindi
Pakistan
2010
ii
COMPETITION FOR FOOD RESOURCES BETWEEN MARKHOR (Capra
falconeri) AND DOMESTIC UNGULATES IN CHITRAL AREA
By
NASRA ASHRAF
(06-arid-593)
A thesis submitted in partial fulfillment
of the requirements for the degree of
Master 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
2010
ii
iii
CERTIFICATION
I hereby undertake that this research is an original one and no part of this thesis
falls under plagiarism. If found otherwise, at any stage, I will be responsible for the
consequences.
Name: Nasra Ashraf
Signature:
Registration No. : 06-arid-593
Date: 06-08-2010
Certified that the contents and form of the thesis entitled “Competition for
Food Resources between Markhor (Capra falconeri) and Domestic Ungulates in
Chitral Area” submitted by Nasra Ashraf have been found satisfactory for the
requirement of the degree.
Supervisor
________________________
(Dr. Maqsood Anwar)
Co-Supervisor
________________________
(Dr. Muhammad Ali Nawaz)
Member
________________________
(Prof. Dr. Iftikhar Hussain)
Member
________________________
(Dr. Irshad A. Khan)
Date of Viva Voce:
Chairman:
Director Advanced Studies:
External Examiner: ____________________
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~Dedicated to
My Caring and
Encouraging
Brothers~
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CONTENTS
Page
i
List of Table
vii
ii
List of Figures
viii
iii
List of Abbreviations
ix
iv
Acknowledgments
x
1
INTRODUCTION
1
2
REVIEW OF LITERATURE
11
3
MATERIALS AND METHODS
18
STUDY AREA
18
3.1.1
Geographical Location
18
3.1.2
Climate
18
3.1.3
Topography
20
3.1.4
Flora
20
3.1.5
Fauna
22
3.1.6
Domestic Livestock
22
3.1.
3.2
3.3
SAMPLING
23
3.2.1
Sample size
23
3.2.2
Sample Collection Procedure
23
3.2.3
Storage and Handling of samples
23
3.2.4
Collection of Reference Materials
24
REFERENCE KEY
24
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3.3.1
3.4
Microphotography
24
Microhistological analysis of faeces (faecal analysis)
27
3.4.1
Slide preparation
27
3.4.2
Slide reading
28
3.5
ANALYTICAL PROCEDURE
28
3.6
DIET COMPOSITION
28
3.7
DIET SELECTION
29
3.8
DIET OVERLAP
29
3.9
STATISTICS ANALYSIS
30
RESULT AND DISCUSSION
31
4.1
DIET COMPOSITION
31
4.2
SEASONAL VARIATION
34
4.3
DIET COMPARISON
38
4.3.1
Spring diet comparison
38
4.3.2
Summer diet comparison
40
4
4.4
4.5
DIET SELECTION
42
4.4.1
Diet selection of markhor and goat during spring season
42
4.4.2
Diet selection of markhor and goat during summer season
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DIET OVERLAP BETWEEN MARKHOR AND DOMESTIC GOAT
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SUMMARY
53
LITERATURE CITED
57
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LIST OF TABLES
Table No.
Page
Table 3.1 h Plant species collected from the study area for reference
26
slides
Table 4.1
Plant species identified in faecal samples of markhor during
32
spring and summer 2009
Table 4.2
Plant species identified in faecal samples of domestic goat
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during spring and summer 2009
Table 4.3
Relative importance values (RIVs) of Plant species in the
36
diet of markhor and domestic goat during spring 2009
Table 4.4
Relative importance values (RIVs) of Plant species in the
37
diet of markhor and domestic goat during summer 2009
Table 4.5
Diet Selection Values (DSV) of markhor and domestic goat
44
during spring 2009
Table 4.6
Diet Selection Values (DSV) of markhor and domestic goat
during summer 2009
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45
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LIST OF FIGURES
Fig. No.
Page
3.1
Map of the Study Area
19
3.2
Temperatures in Chitral during 2009 (-11.8 to 19.85 Celcius)
21
3.3
Rainfall in Chitral during 2009
21
3.4
Feacal sample of markhor
25
3.5
Feacal sample of domestic goat
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4.1
Proportions of plant species in the diets of markhor and
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domestic goat during spring 2009
4.2
Proportions of plant species in the diets of markhor and
41
domestic goat during summer 2009
4.3
Microphotograph of various reference plants
51
4.4
Cells of plant parts found in the faeces of markhor
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4.5
Cells of plant parts found in the faeces of goat
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LIST OF ABBREVIATIONS
°C
Degree Centigrade
CGNP
Chitral Gol National Park
CITES
Convention on International Trade in
Endangered Species of wild fauna and flora
DSV
Diet Selection Value
DPX
A mixture of Distyrene, a Plasticizer, and
Xylene
IUCN
International Union for Conservation of
Nature
Km
Kilometer
m
Meter
mm
Millimeters
NWFP
North West Frontier Province
NCCW
National Council for Conservation of
Wildlife
NaOH
Sodium Hydroxide
PV
Prominence Value
%
Percent
RIV
Relative Importance Value
TGR
Tushi Game Reserve
VCC
Village Conservation Committee
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ACKNOWLEDGMENTS
All praises to Almighty Allah alone, the compassionate and merciful, who
blessed me the courage to get higher education and to complete this manuscript.
Blessing of Allah on Holy Prophet, Muhammad (Peace be upon him) whose
teachings have served as beam of light for humanity in the hours of despair and
darkness.
I would like to take this opportunity to express my deepest sense of gratitude to
the dearest personality, my supervisor Dr. Maqsood Anwar, Associate Professor,
Department of Wildlife Management, PMAS Arid Agriculture University, Rawalpindi,
for his expertise, encouraging behavior and constructive guidance during the
completion of this task.
Very special thanks are due to Dr. Muhammad Ali Nawaz, Country Director,
Snow Leopard Trust Pakistan, for his inspiring guidance, valuable suggestions, and
generous help during the research and procurement of samples. Many thanks to staff of
Snow Leopard Trust Pakistan, for their help in sample collection.
I express countless and earnest thanks to Prof. Dr. Iftikhar Hussain,
Chairman, Department of Wildlife Management, PMAS Arid Agriculture University,
Rawalpindi, for his sincere efforts, and cooperation during this research work.
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I am grateful to all the people in NARC for their support during the work. I also
very thankful to Dr. Rehmatullah Qureshi and Dr. Irshad A. Khan for their
assistance. I would also like to special thanks to my lovely friends, Sara Shabbir and
Misbah Sarwar for their help, co-operation and encouragement through out the study
period.
Finally, I would like to express my greatest gratefulness to my parents and
brothers for their affection, kindness and prayers that encouraged me to achieve
success in every sphere of life.
Nasra Ashraf
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1
Chapter 1
INTRODUCTION
The name “Markhor” apparently was derived from Persian language meaning
snake eater. However, it is mostly considered to be derived from Pashto language word
“Mar Akhkar” in which “Mar” means snake and “Akhkar” means horn. The Markhor
has horns twisting like a snake; therefore it got its name as “Mar Akhkar”. With the
passage of time, it changed to Markhor (Ali, 2008).
Two subspecies of Markhor (Capra falconeri) are recognized: flare-horned
Markhor (Capra falconeri falconeri) which includes the Kashmir (Capra falconeri
cashmiriensis) and Astore (Capra falconeri falconeri) forms, and straight-horned
Markhor (Capra falconeri megaceros) which includes the Kabul (Capra falconeri
megaceros) and Suleiman (Capra falconeri jerdoni) forms. Markhor are typically
associated with steppic mountain conditions and regions of meagre erratic rainfall,
occurring between altitudinal range of 600m to 3600m. They occur in Chitral, north
and east of Quetta, Torghar Range, Suleiman Hills, Dir and Swat Kohistan. A small
population also occurs in Azad Jammu and Kashmir. The Astor Markhor (Capra
falconeri falconeri) is confined to Gilgit region and occurs on both sides of the Astor
River and up the Indus valley as far as the Haramosh range in southwestern Baltistan.
Astor Markhor also occurs on both side of the River Indus in Baghicha area. The
kashmir Markhor (Capra falconeri cashmiriensis), occurs in Chitral town, southwards
into Dir and westwards into Swat kohistan. The kabul Markhor (Capra falconeri
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megaceros) survive precariously in the Khanori hills of the Malakand Agency as well
as the Sakra range just north east of Mardan. The Suleiman Markhor (Capra falconeri
jerdoni) occurs in scattered isolated populations on all the major mountain ranges
immediately to the north and east of Quetta (Roberts, 1997).
Body length of Markhor varies from 132 to 186cm. Weights have been
estimated as varying from 100 to 109kg for an adult male and 32 to 50kg for an adult
female. Horn length is 135 to 143cm in male and 25 to 30cm in female. Gestation
period varies from 135 to 170 days. One or two young per birth is usual, rarely three.
Life span is 12 to 13 years. Males rarely survive beyond 7th or 8th year (Ranjitsinh et
al., 2007).
Markhor are gregarious in habit forming large groups when there is a
concentration of the population in an area they inhabit (Schaller, 1977). Markhor have
polygynous mating systems; the populations’ overall reproductive rate would be little
affected by the loss of a small number of adult males (Caughley, 1977).
Markhor are diurnal animals with greatest activity in early morning and late
evening but in mid winter they have been observed in Chitral feeding intermittently
during the day (Roberts, 1997). Markhor are found at altitudes where they eat grass,
leaves, and whatever other vegetative matter they can find, often standing on their hind
legs to reach the top leaves of trees (Stahl, 2007). An intermediate forager, it consumes
3
primarily grasses and forbs during spring and summer months, while in the winter it
turns primarily to browse for nourishment (Ranjitsinh et al., 2007).
Chitral Gol National Park (CGNP) in Chitral and Palas valley in Kohistan
support the largest surviving populations of Kashmir Markhor (Capra falconeri
cashmiriensis). Temperate coniferous forests and adjoining alpine meadows provide
habitat for this sub-species (Ali, 2008). Malik (2002) estimated 800-1000 Markhor in
Chitral Division. Survey reports indicated 1400 Markhor in Chitral, Dir Kohistan, and
Swat Divisions (GoNWFP, 2005).
In Chitral area Pir Panjal or Kashmir subspecies occurs from about forty miles
north of Chitral town, southwards into Dir and westwards into Swat Kohistan
(Roberts, 1997). Research studies concluded that Markhor population in Chitral is the
largest single concentration of this species in the world and has been effectively
protected by the NWFP Wildlife Department (IUCN, 2001).
Chitral Gol was declared as a National Park by the NWFP Government in
1984. The park is composed of an alpine river catchment in the Hindu Kush mountains
(Aleem, 1978).
The main trees and shrubs are Quercus ilex, Pinus gerardiana,
Juniperus macropoda, Salix spp., Abies pindrow, Pistacia khinjuk, Rosa spp. and
Viburnum (GoNWFP, 2006). Chitral Gol National Park, encompass several valleys,
and an intact vegetation cover which obstructs observation of the animals, it is likely
4
that the annual counts in Chitral Gol have underestimated the population present in the
park (Aleem, 1977).
Wild goats are distributed in the arid rocky mountainous regions of North
Africa, the Middle East and south-west Asia (Shafique and Barkati, 2002). In Pakistan,
Markhor (Capra falconeri) and Ibex (Capra ibex) are the two main wild relatives of
domestic goats (Capra hircus) (Roberts, 1997).
The basic body pattern of all wild and domestic goats is similar, and moreover,
they can freely interbreed in captivity (Manceau et al., 1999). The close gene pool
relationships among diverse taxa within the genus Capra and the possibility of
(repeated) introgressive hybridization of ancestral or modern populations render the
whole group difficult taxonomically (Hammer et al., 2008). The wild goat (Capra
aegagrus) is a common species of goat, with a distribution ranging from Europe and
Asia Minor to central Asia and the Middle East (Stahl, 2007).
Levels of differentiation between some Capra falconeri lineages and modern
domestic goats were similar to levels between other wild goat species (i.e., Capra
aegagrus, Capra ibex) and domestic goats. Among pure Markhor lineages, paraphyly
was observed for C. f. heptneri, suggesting occurrence of shared ancestral
polymorphism among Markhor subspecies and/or ancient or recent gene exchange
between subspecies (Hammer et al., 2008).
5
Goats are the fastest growing ruminants in Pakistan. Pakistan is rich in goat
genetic resources and is the third largest goat producing country in the world after
China and India. Preference for goat meat is the major reason for its increased demand.
There are 25 goat breeds in the country and two wild relatives such as Markhor and
Ibex (Khan et al., 2008). At present, there are 53.8 million domestic goats in Pakistan
and their population is increasing at the rate of more than 3% per annum (GOP, 2006).
Punjab, Sindh, Balochistan and NWFP have 37, 23, 22 and 18 percent goat population,
respectively. The combined livestock population of Chitral in 1996 was estimated at
173,262 head of cattle, 188,822 sheep and 335,782 goats. As goat population is
dominant over other ungulates in Chitral area so Markhor is likely to have close
competition with domestic goats for food resources in this area (GoNWFP and IUCN,
2004).
Domestic goats are ruminants; that is, they have a four-compartment stomach
designed to digest large quantities of forages. Ruminants eat quickly and swallow their
food at first without much chewing. Later, they regurgitate their food and thoroughly
chew it and swallow. This regurgitated food is called the cud, and healthy ruminants
will spend as much time chewing their cud as they do grazing or eating hay. This is
thought to be a predator avoidance adaptation, as the ruminant can find a sheltered
place to peacefully chew its cud and be less vulnerable to predator attack than while
grazing (Coffey et al., 2004).
Contrary to the popular image of goats thriving on tin cans, goats actually
require a more nutritious diet than do other ruminants. Their shorter digestive system
6
does not retain food for the longer period, and thus does not digest nutrients fully. This
quicker digestion allows them to eat larger quantities of food to make up for their
reduced absorption of nutrients, but it is goats’ unique grazing behavior that really
enables them to thrive on pasture. With their small mouths and flexible lips, grazing
goats are able to select the highly nutritious parts of plants and leave parts that are less
nutritious (Coffey et al., 2004).
The hunting of Markhor remained banned because of it being listed on
Appendix-I of CITES (Convention on International Trade in Endangered Species of
wild fauna and flora). The situation only changed in 1997 when CITES eventually
granted Pakistan with six permits for sport hunting Markhor trophies. Federal
government granted these permits to Balochistan, North West Frontier Province
(NWFP) and Northern Areas. CITES increased Pakistan’s quota from 6 to 12 Markhor
hunting permits in 2003. The trophy fees have increased, between 1988 and 2006,
from USD 15,000 to USD 40,000 for Markhor. The rates are agreed upon by NCCW.
Out of the fee, 20% is paid to the Provincial Government, while 80% is used to fund
the community conservation programmes (Bellon, 2008).
The hunting season for Markhor starts in November and goes on until March.
The animals sought by hunter are exclusively older males with the largest end of
horns. It is the responsibility of the game guards to identify the appropriate animals.
However, the hunter is free to select the animal to be hunted. The sustainability of
Trophy hunting is dependent on the allocations quotas for each subspecies and
7
ensuring rigorous enforcement of these quotas. Hence, the first requirement to initiate
such projects is to survey the animal population and assess the maximum number of
specimens that can be harvested without disrupting the reproduction cycle (Lightner,
2006).
The Government of NWFP strongly discourages poaching and illegal trade of
Markhor and other wildlife species through enforcement of the NWFP Wildlife Act of
1975 by the field staff in major markets and potential areas of wildlife importance.
Legal action has been taken against offenders (Ali, 2008).
Markhors are today struggling for survival in semiarid, cliff side mountain
areas (at an elevation of 600–3,600 m) of Tajikistan, Afghanistan, Pakistan, in the
southern border region of Uzbekistan and Turkmenistan, as well as in northern India.
The majority of their total world population is found in Pakistan and is estimated to
encompass about 3,200–3,700 animals, with numbers generally decreasing (Weinberg
et al., 1997). However, certain conservation measures, such as community-based
conservation or ex-situ breeding, have been implemented in recent years and appear to
have a positive effect on at least some Markhor populations (Virk, 2000).
Most Caprinae species face threats of extinction due to genetic isolation,
specialized habitat requirements, and low reproductive rates, in addition to human
causes. Humans are the primary predators on Markhor. Because Markhor inhabit very
steep and inaccessible mountainous habitat, several strongholds of Markhor
8
populations have been rarely approached by man. Golden eagles (Aquila chrysaetos)
have been reported preying upon young Markhor. Among the wild carnivores,
Himalayan lynx (Felis lynx), leopard cats (Felis bengalensis), snow leopards (Uncia
uncia), wolves (Canis lupus), and black bears (Ursus thibetanus) are the main
predators of Markhor (Ali, 2008).
Habitat loss played a lead role in bringing Markhor to the verge of extinction.
Wild lands are rapidly shrinking due to the ever increasing human population and
subsequent increase in demand for timber and fuel wood (Schackleton 2001).
Conditions outside of Village Conservation Committee (VCC) were exacerbated by an
increased livestock density, overgrazing, lack of alternate livelihood for rural
populations, a decrease in natural dominant plant species, and invasion of alien plant
species. All these factors progressively contribute to depletion of biodiversity and
decrease in productivity of fodder resources (Kleinn, 2005).
Markhors are among the most endangered mammal species and several
conservation measures including ex-situ breeding, are implemented to prevent their
extinction (Hammer et al., 2008). Markhor is classifieded as Endangered globally as
per IUCN Red List of 2005. The Flare-horned Markhor (Capra falconeri falconeri)
which includes Kashmir and Astore form is listed “endangered” in Pakistan (Sheikh
and Molur, 2005).
As a conservation strategy in areas where Markhor are found, Government of
NWFP launched a comprehensive extension program to create awareness among
9
people about wildlife importance in general and conservation of Markhor in particular.
The purpose of the program is dissemination of information about wildlife of the
province, its importance and role in the environment, benefits of sustainable use,
problems in conservation, and the need for protection to ensure the support of the
people and mitigate problems in conservation and management of the resource through
a number of conservation and awareness tools (Ali, 2008).
Local communities living in and around Markhor habitat have limited
opportunities to earn their livelihoods. They are mostly dependent on natural
resources. The unsustainable use of these resources has resulted in degradation of
wildlife habitat and depletion of Markhor populations in many areas. One of the best
strategies to save Markhor from extinction is the involvement of local communities in
its conservation and protection (Ali, 2008).
Habitat degradation also caused the migration of Markhor to remote and
unsuitable habitats due to the loss of cover, which further accelerated the process of
population decline (Malik, 1993). Markhor occurs mainly in highly fragmented
populations of relatively small size (100 individuals) that are threatened by habitat
loss, uncontrolled poaching, illegal trophy hunting, and forage competition from
domestic livestock (Weinberg et al., 1997).
Central Asia’s wild ungulates are already heavily under pressure from
poaching, habitat competition with livestock, and degradation of vegetation. Under
10
these circumstances the promotion of their utilization may bear the risk of intensifying
the exploitation and thus giving them the final blow for extinction (Michel, 2008).
There are possible future threats to the populations of wild ungulates, including
competition for grazing by uncontrolled numbers of domestic livestock, and the
possibility of disease transmission from livestock, particularly those of the nomads
from Afghanistan, to the wild Caprines (Woodford et al., 2004).
The present study was, therefore, designed to collect data on feeding aspects of
Markhor and domestic goat in Chitral area, in order to examine the extent of resource
competition between two species. The objectives of study were;
¾ To analyze the food composition of Markhor and domestic goat.
¾ To determine the degree of competition between Markhor and domestic
goat in terms of forage consumption.
¾ To provide recommendation for managing Markhor habitat and reducing
competition with domestic livestock.
11
Chapter 2
REVIEW OF LITERATURE
The principal food of Markhor in Chitral during winter is leaves or acorns of
the Holly Oak (Quercus ilex). During spring and summer months they graze
principally on tussocks of grass and in Gilgit the principal species which is grazed
appear to be Pennisetum orientale. When the grasses are dried up they will also
browse on the leaves and twigs of bushes (Roberts, 1997).
Markhors forage up to 12 or 14 hours per day, including a resting period to
chew cud. As is true of other large, mountain-dwelling ungulates, C. falconeri
maintains a strictly herbivorous diet composed of a variety of grasses in the spring and
summer months. During the autumn and winter months, it switches over to leaves,
twigs, and shrubs. Markhor diets include, but are not limited to, Pennisetum orientale,
Enneapogon persicum, Hippophae rhamnoides, and Quercus ilex (Nowak, 1999).
Common plant species found in its habitat of Markhor include Oak (Quercus
ilex), Sagebrush (Artemisia sp.), Indigofera sp., and Ephedra sp. as dominant species
below 2600 m with a few scattered wild Almond (Amygdalus sp.), Pistachio (Pistacio
sp.), Spruce (Picea smithiana), and Fir (Abies pindrow) at higher elevations. The
grasses Cymbopogon, Stipa, and Chrysopogon provide ground cover (Schaller, 1977).
11
12
Flora of Chitral, Kohistan, Swat, and Dir habitat consists of Pinus wallichiana,
Cedrus deodara, Pinus gerardiana, Picea smithiana, Indigofera gerardiana,
Viberburnum cotinifolium, Polygonum alpinum, Chenopodium foliosum, and
Sambucus ebulus (Ali, 2008).
Markhor occasionally climb into Oak trees (Quercus spp.) to consume the
foliage. Food preferences change with the season and availability. The Markhor is a
goat of low elevations as compared to other Caprinae (Schaller, 1977).
Domestic goats may sometimes need supplemental feeding, especially during
the winter. Goats need a proper balance of energy in the form of roughage or grain, as
well as protein, vitamins, minerals, and clean water. Protein and energy requirements
vary, depending on the type of goat and its stage of production (Coffey et al., 2004).
In Indian Trans-Himalaya, domestic goats were recorded feeding on 26 species
in summer and 14 species in winter. Their diet was similar to sheep in that they fed on
both graminoids and herbs in summer (55% graminoids, 42% herbs and 3% shrubs),
with no single species forming the bulk of the diet. They fed mostly on graminoids in
winter (75% graminoids, 20% herbs and 5% shrubs), with E. longe-aristatus forming
the bulk of the diet (Mishra et al., 2004).
Goats prefer browsing (eating woody plants) but will also graze on grasses and
weeds. Goats are known to stand on their hind legs to reach leaves and brush. Since
13
goats, cattle, and sheep prefer different forages, in many pasture situations these
species do not compete for the same food. Therefore, they can be managed quite
successfully in a multispecies grazing system, allowing the land to be used more fully
and generate more income. Land grazed by both goats and cattle returns 25% more
than land grazed only by cattle (Coffey et al., 2004).
Goats are efficient browsers and prefer eating brushy plants along with some
other woody and weedy plants found on the ranges. Goats are able to digest a large
variety of fibre and roughage. The daily feed intake of goats ranges from 3 to 4% of
body weight as expressed in pounds (dry matter/head/day). The daily feed intake is
influenced by body weight, % of dry matter in the feeds eaten (12 to 35% in forages,
86 to 92% in hays and concetrates), palatability, and physiological stage of the goats
(growth, pregnancy, and lactation) (Rashid, 2008).
Each goat is able to consume up to 3 to 5% of its body weight in dry matter
daily (perhaps more if the forage is highly digestible). To consume that amount,
however, goats must be pastured in an area with a large quantity of available vegetative forage. Goats will eat less when they are moved to poor pastures (Coffey et al.,
2004).
A review of information available on the food plants and feeding habits of
Himalayan ungulates revealed that of the 12 alpine ungulate species only four have
been studied in detail. Analyses of the compiled data on food plants show that a total
14
of 140 wild plant species are palatable to different ungulate species. However,
information on palatability of many other plant species is lacking. The information on
the food plants of domestic ungulates is also scarce. Based on whatever information is
available it was found that out of 140 plant species, 13 are common in the diet of wild
and domestic ungulates. Some of the recent studies have given some indications of a
competition existing between wild and domestic Himalayan ungulates for food, hence
it becomes imperative to study their feeding habits and document their food plants
(Awasthi et al., 2003).
Northern Areas of Pakistan comprise large number of livestock populations,
including native cattle, sheep and goats. In the cold winter the animals are kept in the
houses in the valley where they are fed maize stover, wheat straw and hay. In the
summer months the animals are taken up into the mountains to graze on the subalpine
and finally the alpine pastures. Late April or early May animals start trekking through
the mountains. First they move to the edge of the conifer forests and then they move to
the alpine pastures high up in the mountains. Maize stover and some hay and grasses
collected are the main feed of the stall fed large animals (Hashmi and Shafiullah,
2003).
Livestock grazing impacts on native wildlife are an important conservation
concern globally (Fleischner, 1994). There have been few attempts to evaluate
livestock grazing impacts on native wildlife. Consequently, the debate on what kind of
impacts local human resource use has on native wildlife remains ill-informed. The
15
debate on whether local human use of wildlife reserves should be modified, curtailed
or encouraged continues to be fuelled by activism rather than by ecology (Mishra and
Rawat, 1998).
In India (Spiti Valley), the issue of competition between livestock and wild
herbivores has remained contentious. Livestock grazing occurs in pastures during most
of the year except in the extreme winter. The livestock diet is supplemented by stall
feeding in winter. Livestock populations are maintained beyond points of natural
resource limitation through supplemental feeding (Mishra, 2001).
Diet overlap or competition for food between wild herbivores and livestock and
forage availability is known to affect reproductive performance in ungulates (Clutton,
et al., 1982). Forage availability influences the body condition of females and
consequently their fecundity (Leader, 1988). Further, there is evidence for densitydependent mortality in neonates and calves (Saether, 1997).
The role of competition in structuring large-herbivore assemblages remains
especially unclear owing to the logistic difficulty in manipulating populations and
measuring competition at the population level (Forsyth, 2000). Indeed, impacts of
livestock grazing on wildlife have been a subject of much debate in South Asia
(Mishra and Rawat, 1998). Long histories of livestock grazing can further make the
evaluation of grazing impacts on native wildlife and landscape difficult, since the
impacts are often pervasive and therefore can go unnoticed (Fleischner, 1994).
16
Almost a century ago, it has been recorded that the presence or absence of a
species in a given area could be determined by interspecific competition.
Subsequently, other studies demonstrated that the joint placement of pairs of similar
species that compete for the same resource could cause extinctions as a result of
resource competition. Competition has since been a subject of debate regarding its role
in structuring species composition in ecological communities (Mishra et al., 2002).
In a study on patterns in species richness of grazer assemblages in Africa as a
function of resource competition and facilitation, Dayan and Simberloff (1998)
predicted an ‘optimum’ difference in body mass between co-existing members of any
natural grazer assemblage, a pattern expected to be influenced by the long-term effects
of competition and facilitation. Empirical data corroborated this prediction; species in
‘intact’ African grazer assemblages indeed show a constant weight ratio, with each
grazer species on average being a constant proportion larger than the nearest smaller
one. Such morphological relationships (a constant weight ratio in the case of grazers)
between the members of a guild are believed to be brought about by character
displacement (a co-evolutionary response) and species sorting (an ecological
response), both being consequences of competition (Dayan and Simberloff, 1998).
Large herbivores with similar ecological requirements are expected to partition
resources to coexist, especially in areas where the resources are in short supply
(Wiens, 1977). Resource partitioning is accomplished through evolutionary divergence
of resource use by co-occurring species apparently in response to competition (Walter,
17
1991). The issue of forage competition between livestock and wild herbivores,
although long acknowledged as being important for conservation management (Kie et
al., 1991), has remained a contentious one and reviews suggest a worldwide scarcity of
studies aimed at understanding it (Putman, 1996).
Even though conclusive information on habitat degradation, and direct
competition between domestic and wild herbivores from the region has just started
coming, it is evident from some preliminary studies that the present livestock grazing
levels in areas such as eastern Ladakh and Spiti may already be unsustainable. The
potential impacts of excessive grazing by livestock include depletion of the scarce
forage for wildlife, habitat degradation, disease transfer, and reduction in the breeding
performance of both wildlife and domestic stock (Bhatnagar and Mathur, 2001).
In areas, where large and increasing numbers of people put pressure on small
and shrinking ungulate populations and their habitats, these species are at risk being
exterminated within few years (Michel, 2008). Competition for forage between wild
and domestic herbivores and human-wildlife conflicts need to be addressed in a
participatory manner (Bhatnagar and Mathur, 2001).
18
Chapter 3
MATERIALS AND METHODS
3.1 STUDY AREA
3.1.1 Geographical Location
The study was conducted in Chitral Gol National Park (CGNP) and Tushi
Game Reserve (TGR). Chitral Gol National Park is situated in the beautiful valley of
Chitral, northwest in the Khyber-Pakhtunkhwa province of Pakistan at 35o 56'N
latitude to 71'40oE longitude (Fig 3.1). Total area of the Chitral Gol National Park is
7750 hectares. It is accessible through a dirt road from Chitral town; the Park is
bordered by Rombor Valley to the southwest. Tushi Game Reserve has an area of 1545
hectares. The Tushi Game Reserve lies in the foothills of the Hindukush Mountain
Range, at a distance of about a half an hour drive from the Chitral town. The important
buffer zones of the area are Kuju Gol and Gohtan Gol.
3.1.2 Climate
The climate of Chitral Gol National Park and Tushi Game Reserve is dry
temperate, characterized by hot summers in the lower areas and cold summers in the
upper elevations. Precipitation ranges from 200 mm to 800 mm annually and is mostly
received in winter and spring, largely in the form of snow (Rasheed 2007). While
mean annual precipitation is 445 mm, principally in winter and spring (Ali, 2008).
Frost and snow start from September on high peaks and in November in the valley.
Snow stays till March in valleys and June on mountains. Summer and autumn are dry,
receiving barely 10–25 mm of rain per month (GoNWFP and IUCN Pakistan, 2004).
18
19
Fig. 3.1: Map of the Study Area
20
0
The mean annual temperature is 16.8C (Ali, 2008). Figures 3.2 and 3.3 show the
temperature and rainfall, respectively of Chitral district during 2009.
3.1.3 Topography
The Chitral Gol National Park consists of mostly high, rugged, and steep
mountains with slope varying from 450 to 1200. High mountain topography is the
characteristic feature of the study area. Therefore the entry into Chitral is limited to
only few passes i.e., Lowari Pass and Shandoor Pass, which are at an altitude of
3300m and 3800m, respectively. These passes are closed due to heavy snowfall for
almost six months in winter and the area is totally inaccessible (Ali & Qaiser, 2005).
The topography varies consisting of alpine meadows leading to the permanent glaciers.
High mountain and rough topography of the area have given rise to lot of narrow side
valleys. Erosion and glacial drift have contributed to fan-deposits along both the main
and side valleys, which are the main areas of habitation by humans, hence villages and
cultivated fields are found on these alluvial deposits (Ali & Qaiser, 2006).
3.1.4 Flora
The main trees and shrubs growing in Chitral Gol National Park and Tushi
Game Reserve include Quercus ilex, Pinus gerardiana, Juniperus macropoda, Salix
spp., Abies pindrow, Pistacia khinjuk, Viburnum and Rosa spp. (GoNWFP, 2006). At
elevation up to 2,400 metres the vegetation type is dry oak forest (Quercus ilex). At
21
Fig. 3.2: Temperatures in Chitral during 2009 (-11.8 to 19.85
Celcius)
Fig. 3.3: Rainfall in Chitral during 2009
22
higher elevations the type grades into the dry temperate coniferous forest with the
inclusion of Cedrus deodara and Pinus gerardiana (Aleem, N. D).
3.1.5 Fauna
Fauna of Chitral Gol National Park and Tushi Game Reserve has affinities to
Palearctic Faunal Region with only a slight oriental mix from the south. Dry and arid
temperate climate has been compensated for its species richness by great altitudinal
variation. Major wildlife includes; Snow leopard (Uncia uncia), Himalayan lynx (Lynx
lynx), Wolf (Canus lupus), Jackal (Canus aureus), Common red fox (Vulpes vulpes),
Himalayan ibex (Capra ibex sibirica), Hare (Lepus capensis), Long-tailed or Kashmir
marmot (Marmota caudata), snow cock (Tetraogallous himalayensis) and Chukar
partridge (Alectoris chukar) (Din and Nawaz, 2010). Markhor (Capra falconeri) and
black bear (Ursus thibetanus) are also found in the Chitral area (GoNWFP, 2006).
3.1.6 Domestic Livestock
The combined livestock population of Chitral was estimated at 173,262 head of
cattle, 188,822 sheep and 335,782 goats. In addition, an unknown number of animals
belonging to Afghan refugees is also found in the area. Households keep livestock in
small herds of 4 to 10, depending on their capacity to store crop residues, fodder and
hay to feed animals during the winter and spring months. Chitrali cattle are not robust,
weighing only 150 to 200 kg, and generally frail owing to poor nutrition. Cows are
kept for milk, calves for milk let-down and oxen for draught purposes. Goats are kept
23
for meat, milk and hides. Sheep are kept primarily for wool (GoNWFP and IUCN
Pakistan, 2004).
3.2 SAMPLING
3.2.1 Sample size
A total of 120 pellet groups, 30 each of Markhor and domestic goat were
collected from Chitral Gol National Park and Tushi Game Reserve during the spring
and summer seasons.
3.2.2 Sample Collection Procedure
Pellet groups of Markhor and domestic goat were collected between April 2009
and December 2009 which were identified on the basis of pellet dimensions, shape and
structure Fig (3.4, 3.5). All pellet groups were double-bagged, labeled with
information on collector, collecting time, place and conditions. Information of samples
was recorded by using a waterproof pen to minimize the chances of error. The pellet
groups were placed in a paper bag so that it can dry out as quickly as possible. Then it
was taped such that the contents cannot fall out, yet leaving a gap to aid in the air
drying.
3.2.3 Storage and Handling of samples
The pellet groups were placed in a warm, dry place before laboratory analysis
so that they can get dry as soon as possible.
24
3.2.4 Collection of Reference Materials
The collection of potential plant foods of Markhor and goat were made
from the study area i.e Chitral Gol National Park and Tushi Game Reserve. This
collection of reference plants was based on information collected from people
living around Chitral Gol National Park and Tushi Game Reserve. Two specimens
of each plant were collected; one for the reference record and the other for
preparation of reference materials. The plants were dried between the folds of
newspaper. The plants were identified with the help of plant taxonomist. Detail of
specimens collected is given in table 3.1.
3.3 REFERENCE KEY
A herbarium and a photomicrograph key of reference plants were prepared by
collecting plant species from Chitral Gol National Park and Tushi Game Reserve.
Herbarium contains 22 plant species. Diet of Markhor and domestic goat in study area
was identified by developing a reference photographic key of plant species Fig (4.5).
3.3.1 Microphotography:
The diagnostic features of the plant cells such as shape, size, fibres, trichomes,
pores, stomata, from each reference slide were photographed using a camera fitted to
an microscope. Thus, a reference slide library was made of 26 images. This was done
in order to facilitate the screening of photomicrographs for the identification of faecal
fragments during later analysis.
25
Fig. 3.4: Feacal sample of markhor
Fig. 3.5: Feacal sample of domestic goat
26
Table 3.1: Plant species collected from the study area for reference slides
S. No.
Scientific name
Local name
Trees
1
Quercus baloot
Banj
2
Juniperus excelsa
Saroz
3
Pistacia integerrima
Thoak
4
Salix iliensis
Theli
Shrubs
5
Rubus anatolicus
Atchu
6
Origanum vulgare
Ishpain
7
Linum perenne
Shahtili
Grasses and Forbs
8
Astragalus psilocentros
Garmenzu
9
Marsilea minuta
Isqar Basho
10
Capparis spinosa
Kaveer
11
Typha latifolia
Manzhor
12
Prangos pabularia
Moshain
13
Convolvulus arvensis
Polini jusho
14
Rumex lastatus
Shot shakho
15
Bergenia stracheyi
Bisapur
16
Bergenia ciliata
Zakhme Hayat
17
Rumex alpinus
Sirkonzu
18
Hypsicum perforatum
Matali
19
Rheum emodi
Ishpar
20
Berberis calliobotrys
Chowenj
21
Grewia populifolia
Bhovi
22
Artemisia maritime
Dron
27
3.4 MICROHISTOLOGICAL ANALYSIS OF FAECES (FAECAL ANALYSIS)
Examining faecal samples by a microhistological technique (Baumgarther and
Martin, 1939; Sparks and Malechek, 1968) is the most commonly used method for
determining the botanical composition of herbivore diets (Holechek et al., 1982;
Alipayo et al., 1992). Different digestibility among plants, however, may produce
biased estimates (Sanders et al., 1980). The diet of Markhor and domestic goats was
determined through microhistological analysis of their faeces and compared for
resource overlap.
3.4.1 Slide preparation:
In the laboratory the faecal samples were ground with mortar and pestle.
The ground material was sieved through cotton cloth to remove large
unidentifiable particles and dust. Faecal Samples were washed in flowing water
and soaked in a soaking solution (1 part distilled water, 1 part Ethyl alcohol, 1 part
glycerine) for over night. Then the samples were again ground in Virtis
Homogenizer at National Agricultural Research Centre, Islamabad. Fifty percent of
the samples was transferred to a labeled test tube and five percent warm sodium
hydroxide solution was added to it. The test tube was heated in a boiling water bath
for 4 to 6 minutes. The particles were allowed to settle before removing the
supernatant dark fluid and this treatment was repeated 3 to 7 times until a relatively
clear solution was produced. Then the material was washed four times with warm
distilled water. It was dehydrated through a series of 25%. 50%, 75% and 100%
alcohol treatments, each for 10 minutes. Then alcohol was removed through a
28
series of xylene and alcohol mixtures (25%. 50%, 75% and 100% xylene) each for
10 minutes, except 100% for overnight. The next day the material was transferred
to a clean glass slide and was evenly spread and mounted in DPX mounting
medium under a cover slip. The same procedure was followed for preparation of
slides of reference plant collection, except for using 10 % NaOH solution.
3.4.2 Slide reading
The slides were studied in detail by recording the specific cell
characteristics of plants. These characteristics were used as key features for
comparison and identification of the materials found in faeces. The images of
faecal fragments were compared with the plant reference photomicrographs at the
similar level of magnification, exposure, brightness, and colour conditions.
3.5 ANALYTICAL PROCEDURE
Each faecal sample was analysed to identify plant species composition by the
Microhistological analysis. The faecal samples were examined by microhistological
technique. The fragments were identified on the basis of shape and relationships of the
short cells, shape of the wall of the long cells, stomatal cells, cell wall shape, pattern
and distribution of epidermal cells adjacent to guard cells, nature and arrangement of
epidermal hairs.
3.6 DIET COMPOSITION
Plant species found in faecal sample were ascertained after a detailed analysis
of all cell characteristics and compared with reference photographic key. The relative
29
frequency of a plant species in the faeces was calculated and expressed as the Relative
Importance Value (RIV), which is the total number of fragments identified for a given
food species divided by the total number of all counts made in the sample, multiplied
by 100 (Jnawali, 1995).
3.7 DIET SELECTION
Diet Selection Value (DSV) was calculated using the following equation,
reflecting the consumption (Relative Importance Value) in relation to the availability
(Prominence Value) of food plants (Jnawali, 1995):
Where
is the Prominence Value (PV) for species x. Prominence value reflects the
relative availability of plant species in Markhor habitat.
is the Relative
Importance Value (RIV) for species x. Relative importance value reflects the relative
frequency of a plant species in the faeces.
3.8 DIET OVERLAP
The modified Morisita index of overlap (Horn, 1966) was calculated to
estimate the diet overlap between the ungulates and thus to measure their overall diet
similarity. The index
varies from 0.0 for completely distinct pairs (no food species
in common) to 1.0 for complete overlap:
30
Where x and y are the proportion of plant group in the total diet of Markhor and
domestic goat respectively.
3.9 STATISTICAL ANALYSIS
Paired chi-square test was applied to determine the significance of any
differences in consumption found within each plant.
31
Chapter 4
RESULTS AND DISCUSSION
4.1 DIET COMPOSITION
The collection of faecal samples of Markhor and domestic goat were carried
out from the study area during two seasons of the year i.e. spring and summer.
Therefore, the analyses were made separately for each collection with respect to the
season and animal species. A total of 14 plant species were recorded in the diet of
Markhor out of which leaves of 13 species and stem parts of one species were
consumed (Table 4.1). Similarly domestic goat consumed 13 plants species. Of these,
the leaves of 11 species and stem parts of two species were consumed (Table 4.2).
Thus the leaves were the most common food item found in faeces regardless of the
season. The results of present study support the earlier investigations by Coffey et al.
(2004) who reported that both wild and domestic goat prefer browsing. Goats are
known to stand on their hind legs to reach leaves.
Plant species such as Prangos pabularia and Linum perenne were found in the
diet of Markhor and domestic goat in both the spring and summer seasons. In the
spring seasons Rumex lastatus was dominant while in the summer seasons Hypsicum
perforatum was dominant in Markhor food. In the spring seasons Salix iliensis was
dominant while in the summer seasons Linum perenne was dominant in domestic goat
food.
31
32
Table 4.1: Plant species identified in faecal samples of markhor during spring
and summer 2009
S. No. Plant Species
Part Eaten
No. (%) of faeces containing plant parts
Spring-2009
Summer-2009
1
Salix iliensis
leaves
8 (26.6)
-
2
Origanum vulgare
leaves
8 (26.6)
6 (20.0)
3
Rumex lastatus
leaves
9 (30.0)
-
4
Astragalus psilocentros
leaves
8 (26.6)
-
5
Juniperus excelsa
leaves
7 (23.3)
-
6
Prangos pabularia
leaves
8 (26.6)
6 (20.0)
7
Linum perenne
leaves
8 (26.6)
7 (23.3)
8
Rubus anatolicus
leaves
4 (13.3)
-
9
Artemisia maritime
leaves
-
6 (20.0)
10
Pistacia integerrima
leaves
-
6 (20.0)
11
Rheum emodi
leaves
-
8 (26.6)
12
Hypsicum perforatum
stem
-
8 (26.6)
13
Quercus baloot
leaves
-
7 (23.3)
14
Bergenia ciliate
leaves
-
5 (16.6)
33
Table 4.2: Plant species identified in faecal samples of domestic goat during
spring and summer 2009
S. No.
Plant Species
Part Eaten
No. (%) of faeces containing plant parts
Spring-2009
Summer-2009
1
Salix iliensis
leaves
8 (26.6)
-
2
Juniperus excelsa
leaves
6 (20.0)
-
3
Prangos pabularia
leaves
7 (23.3)
7 (23.3)
4
Linum perenne
leaves
8 (26.6)
7 (23.3)
5
Capparis spinosa
stem
3 (10.0)
-
6
Artemisia maritime
leaves
-
6 (20.0)
7
Pistacia integerrima
leaves
-
6 (20.0)
8
Origanum vulgare
leaves
-
7 (23.3)
9
Rheum emodi
leaves
-
8 (26.6)
10
Hypsicum perforatum
stem
-
7 (23.3)
11
Quercus baloot
leaves
-
6 (20.0)
12
Prangos pabularia
leaves
7 (23.3)
7 (23.3)
13
Bergenia ciliate
leaves
-
6 (20.0)
34
4.2 SEASONAL VARIATION
Eight plant species were recorded in the faeces of Markhor and five in the
faeces of domestic goat from spring season samples (Table 4.3). Among those, four
plant species were found common in the diet of both Markhor and domestic goat,
including Salix iliensis, Juniperus excelsa, Prangos pabularia, Linum perenne. Hence,
there was a considerable diet overlap among the both animal species. Due to more or
less complete digestion of some plant species unidentified material consisted of 6.4%
and 17.1% in the diet of Markhor and domestic goat respectively (Table 4.3).
In summer samples, nine plant species were identified in the faeces of both the
Markhor and domestic goat (Table 4.4). In summer, the overlap of diet of Markhor
with domestic goat was higher than spring season. As in summer, all nine plant species
were found common in the diet of both Markhor and domestic goat. Unidentified
material consisted of 6.29 % and 6.31% in the diet of Markhor and domestic goat,
respectively. Current study indicated that the extent of overlap between Markhor and
domestic goat for summer forage was high than in spring, thus competition was likely
to occur during this season. The results of earlier study conducted in the Indian TransHimalayas by Mishra et al. (2004) are in line with the results of present study, who
reported that there was considerable diet overlap among the herbivore species in
summer and livestock grazing caused a significant reduction in the standing crop of
forage. Similarly, Bagchi et al. (2003) found that wild and domestic goats utilize the
same habitat types and do not differentiate in their diets. Wild and domestic goats are
intermediate feeders with associated opportunistic feeding behaviour, diet overlap
35
seems to be inevitable. Hence, they are expected to compete for the same forage when
feeding on the same range (Mishra et al., 2004).
Paired chi-square tests showed that during spring season, consumption of all
plants (Salix iliensis, Origanum vulgare, Rumex lastatus, Astragalus psilocentros,
Juniperus excelsa, Linum perenne, Capparis spinosa, Rubus anatolicus) except
Prangos pabularia was significantly different (P <0.05, χ² =4.09-15.89, df =1) among
the two species. While consumption of Prangos pabularia was equal both in Markhor
and domestic goat (P > 0.05, χ² =3.35, df =1).
Paired chi-square tests showed that during summer season, consumption of all
plants (Artemisia maritime, Pistacia integerrima, Origanum vulgare, Rheum emodi,
Hypsicum perforatum, Quercus baloot, Prangos pabularia, Linum perenne, Bergenia
ciliate) were equal both in Markhor and domestic goat (P > 0.05, χ² = 0.02-0.47, df
=1), having non-significant difference in their diets.
36
Table 4.3: Relative importance values (RIVs) of plant species in the diet of
markhor and domestic goat during spring 2009
S. No.
Markhor
Plants
RIV
Domestic goat
SD
RIV
SD
1
Salix iliensis
13
11.46
22.69
9.92
2
Origanum vulgare
13.32
10.45
0.0
-
3
Rumex lastatus
15.89
11.29
0.0
-
4
Astragalus psilocentros
12.08
12.29
0.0
-
5
Juniperus excelsa
10.94
16.84
17.63
12.76
6
Prangos pabularia
12.89
7.78
18.4
7.90
7
Linum perenne
12.32
10.09
19.64
6.48
8
Rubus anatolicus
3.1
8.95
0.0
-
9
Capparis spinosa
0.0
-
4.49
1.0
10
Unidentified plants
6.4
2.84
17.11
4.40
37
Table 4.4: Relative importance values (RIVs) of plant species in the diet of
markhor and domestic goat during summer 2009
S. No.
Markhor
Plants
RIV
Domestic goat
SD
RIV
SD
1
Artemisia maritime
9.29
7.62
9.82
7.63
2
Pistacia integerrima
8.94
6.86
9.41
6.67
3
Origanum vulgare
10.68
3.88
8.7
7.11
4
Rheum emodi
12.65
7.11
13.22
6.08
5
Hypsicum perforatum
13.1
7.80
11.9
7.91
6
Quercus baloot
10.28
9.44
9.51
6.71
7
Prangos pabularia
9.16
6.73
11.24
9.53
8
Linum perenne
12.52
4.33
13.37
4.46
9
Bergenia ciliate
7.03
9.62
6.47
12.34
10
Unidentified plants
6.29
2.44
6.31
2.61
38
4.3 DIET COMPARISON
4.3.1 Spring diet comparison
Rumex lastatus was the dominant plant species in the spring diet of Markhor
with 15.89% composition, followed by Origanum vulgare (13.32%), Salix iliensis
(13%), Prangos pabularia (12.89%), Linum perenne (12.32%), Astragalus
psilocentros (12.08%), Juniperus excels (10.94%,) and Rubus anatolicus (3.1%), and
unidentified plants 6.4% (Fig 4.1). The dominant plant in the diet of domestic goat was
Salix iliensis (22.69%), followed by Linum perenne (19.64%), Prangos pabularia
(18.4%), Juniperus excelsa (17.63%), and Capparis spinosa (4.49%). All these plant
species were common with Markhor diet except Capparis spinosa (4.49%). Capparis
spinosa was present only in the diet of domestic goat. Unidentified plant material was
17.11% in goat’s diet. Four plant species i.e. Origanum vulgare, Rumex lastatus,
Astragalus psilocentros and Rubus anatolicus were present only in the diet of
Markhor.
The dominant plant in the diet of Markhor was Rumex lastatus during spring.
This finding is supported by earlier investigations by Aleem (1976), who reported that
during spring season, Markhor preferentially utilize Rumex lastatus. During the
present investigation it was noted that Capparis spinosa was present only in the diet of
goat. Field observations of PAMP (2008) have supported that the frequency of
occurrence of Capparis spinosa is very low in Chitral Gol National Park because local
population collect this plant for fuel wood and herding livestock.
domestic goat during spring 2009
Fig. 4.1: Proportions of plant species in the diets of markhor and
39
40
4.3.2 Summer diet comparison
Hypsicum perforatum was the dominant plant species in the summer diet of
Markhor with 13.1% composition, followed by Rheum emodi (12.65%), Linum
perenne (12.52%), Origanum vulgare (10.68%), Quercus baloot (10.28%), Artemisia
maritime (9.29%), Prangos pabularia (9.16%), Pistacia integerrima (8.94%),
Bergenia ciliate (7.03%) and unidentified plants 6.29% (Fig 4.2). The dominant plant
in the diet of domestic goat was Linum perenne (13.37%), followed by Rheum emodi
(13.22%), Prangos pabularia (11.24%), Artemisia maritime (9.82%), Quercus baloot
(9.51%), Pistacia integerrima (9.41%), Origanum vulgare (8.7%) and Bergenia ciliate
(6.47%). All these plant species were common with Markhor diet. Unidentified plant
matters were 6.31% in domestic goat’s diet.
The present study revealed that during summer, Markhor and domestic goat
share their diets. This information is supported by field observations of PAMP (2008),
who reported that the households of Chitral send their livestock to different zones of
Chitral Gol National Park that are also utilized by Markhor during summer. The
herders of Thingshen, Jang Bazaar, Mughlandeh utilize Utras and Dooni areas for
livestock grazing and which share it with Markhor during summer. Therefore, the
competition for forage between Markhor and domestic goat is high in Chitral Gol
National Park because both species mainly occupy the steeper slopes with more
browsing potentials.
goat during summer 2009
Fig. 4.2: Proportions of plant species in the diets of markhor and domestic
41
42
4.4
DIET SELECTION
4.4.1 Diet selection of Markhor and domestic goat during spring season
During the spring season, Markhor strongly preferred Linum perenne (DSV=
2.64) and had a neutral preference for Salix iliensis (DSV= 1.39), Rumex lastatus
(DSV=1.25), Origanum vulgare, (DSV= 1.18) and Juniperus excelsa (DSV=1.03).
While Astragalus psilocentros (DSV=0.62), Rubus anatolicus (DSV=0.55) and
Prangos pabularia (DSV=0.30) was least preferred forage of Markhor. Similarly,
domestic goat strongly preferred Linum perenne (DSV= 4.22) and Salix iliensis
(DSV= 2.42) and had a neutral preference for Juniperus excelsa (DSV= 1.66) and
Capparis spinosa (DSV= 1.56). While Prangos pabularia (DSV= 0.43) was the least
preferred forage of domestic goat. During spring season, both Markhor and domestic
goat showed strong preference for Linum perenne (Table 4.5).
4.4.2 Diet selection of Markhor and domestic goat during summer season
During summer season, Markhor strongly preferred Linum perenne (DSV=
2.69), and had a neutral preference for Pistacia integerrima (DSV=1.19). While
Origanum vulgare (DSV=0.95), Rheum emodi (DSV= 0.81), Bergenia ciliate
(DSV=0.64 ), Hypsicum
perforatum (DSV=0.60), Quercus baloot
(DSV=0.23),
Artemisia maritime (DSV=0.22) and Prangos pabularia (DSV=0.21) were the least
preferred forage of Markhor. Similarly, domestic goat strongly preferred Linum
perenne (DSV =2.87) and had a neutral preference for Pistacia integerrima (DSV
=1.25). While Rheum emodi (DSV=0.85), Origanum vulgare (DSV= 0.77), Bergenia
ciliate (DSV= 0.59), Hypsicum perforatum (DSV= 0.55), Prangos pabularia (DSV=
43
0.26), Artemisia maritime
(DSV=0.24) and Quercus baloot (DSV=0.21) were the
non preferred plants by domestic goat (Table 4.6).
The present study revealed that Linum perenne was the most important
forage species and preferred food both for Markhor and domestic goat in Chitral area
during both the spring and summer seasons. The results of this study are in agreement
with a study conducted under PAMP (2008), which stated that both Markhor and
domestic goat preferentially utilize Linum perenne.
4.5 DIET OVERLAP BETWEEN MARKHOR AND DOMESTIC GOAT
During summer season, diet overlap factor was 0.99 (
=0.99), almost equal
to 1.0, which indicates very high diet overlap between Markhor and domestic goat.
During the spring season, the value of diet overlap factor was 0.69 (
=0.69),
indicating that the diets of Markhor and domestic goat were not completely overlap.
During summer season, a very high diet overlap between Markhor and
domestic goat in the study area revealed by the present investigation indicates a
substantial degree of niche overlap between Markhor and domestic goat. Because in
early summer, goat population move to high elevation for grazing and browsing.
Where they remains from August till the beginning of snowfall in early December.
This high elevation area is also utilized by Markhor during summer.
44
Table 4.5: Diet selection values (DSV) of markhor and domestic goat during
spring 2009
Plant species
Availability of plants (%)
Markhor
Domestic goat
DSV
DSV
Salix iliensis
9.35
1.39
2.42
Origanum vulgare
11.21
1.18
-
Rumex lastatus
12.68
1.25
-
Astragalus psilocentros
19.41
0.62
-
Juniperus excelsa
10.57
1.03
1.66
Prangos pabularia
42.58
0.30
0.43
Linum perenne
4.65
2.64
4.22
Rubus anatolicus
5.56
0.55
-
Capparis spinosa
2.86
-
1.56
45
Table 4.6: Diet selection values (DSV) of markhor and domestic goat during
summer 2009
Markhor
Domestic goat
Availability of plants (%)
DSV
DSV
Artemisia maritime
40.59
0.22
0.24
Pistacia integerrima
7.5
1.19
1.25
Origanum vulgare
11.21
0.95
0.77
Rheum emodi
15.52
0.81
0.85
Hypsicum perforatum
21.55
0.60
0.55
Quercus baloot
43.77
0.23
0.21
Prangos pabularia
42.58
0.21
0.26
Linum perenne
4.65
2.69
2.87
Bergenia ciliate
10.89
0.64
0.59
Plant species
46
The results of present study are similar as of earlier investigations by Aleem
(N.D) reporting that wild animal come in direct competition with livestock. The food
taken by domestic goats is almost the same which is relished by Markhor in Chitral
area. In a study conducted in eastern Ladakh by Bhatnagar and Mathur (2001), it was
indicated that the potential impact of excessive grazing by livestock include depletion
of the scarce forage for wild animals.
Food habits studies of animals by microscopic analysis of faecal material rank
the forage species according to their importance in the diet. The quantification of diets
from faecal analysis rests on the assumption that species composition of epidermal
fragments identified in the faeces corresponds closely to the proportion of each species
ingested. Although faecal analysis has been reported to under-estimate the proportion
of some species, especially herbs, in the diet, Alipayo et al. (1992) suggested that this
is in line with the poor methodology, and good correspondence can be obtained if
appropriate attention is given to the preparation and identification of epidermal
material.
The microphotographs of reference plants are given in Fig. 4.3 and plants
found in the faeces of Markhor and domestic goat are given in Fig. 4.4 and 4.5,
respectively.
47
1
2
3
5
4
6
Continued
48
Fig. ‘page 2’
11
7
8
9
10
12
Continued
49
Fig. ‘page 3’
13
15
17
14
16
18
18
Continued
50
Fig. ‘page 4’
19
21
20
22
22
23
24
Continued
51
Fig. ‘page 5’
25
26
Fig. 4.3: Microphotograph of reference plants
1. Leaves cells of Rumex alpines
2. Stem cells of Artemisia maritime
3. Leaves cells of Artemisia maritime
4. Stem cells of Convolvulus arvensis
5. Leaves cells of Convolvulus arvensis
6. Leaves cells of Origanum vulgare
7. Leaves cells of Capparis spinosa
8. stem cells of Capparis spinosa
9. Leaves cells of Typha latifolia
10. Leaves cells of Rumex lastatus
11. Leaves cells of Rubus anatolicus
12. Leaves cells of Quercus baloot
13. Leaves cells of Prangos pabularia
14. Leaves cells of Pistacia integerrima
15. Leaves cells of Rheum emodi
16. Leaves cells of Astragalus psilocentros
17. Leaves cells of Linum perenne
18. Leaves cells of Hypsicum perforatum
19. Stem cells of Hypsicum perforatum
20. Leaves cells of Berberis calliobotrys
21. Leaves cells of Bergenia stracheyi
22. Leaves cells of Salix iliensis
23. Leaves cells of Juniperus excelsa
24. Leaves cells of Grewia populifolia
25. Leaves cells of Marsilea minuta
26. Leaves cells Bergenia ciliata
52
Fig. 4.4: Cells of Origanum vulgare and Astragalus psilocentros found
in the faeces of markhor
Fig. 4.5: Cells of Origanum vulgare found in the faeces of domestic
goat
53
SUMMARY
Markhor is typically associated with steppic mountain conditions and regions
of meagre rainfall, occurring between altitudinal range of 600m to 3600m. Chitral Gol
National Park (CGNP) supports the largest surviving populations of Kashmir Markhor
(Capra falconeri cashmiriensis) in Pakistan. Markhor is classifieded as Endangered
globally as per IUCN Red List of 2005. The present study was designed to determine
the extent of food overlap between Markhor and domestic goat in Chitral area. For this
purpose, microscopic analysis of faecal material of both the species was done. A total
of 120 pellet groups, 30 each of Markhor and domestic goat were collected from
Chitral Gol National Park and Tushi Game Reserve during the spring and summer
seasons. Different characteristics of plant cells were used to determine the plant
species consumed by Markhor and domestic goat. An herbarium containing 22 plant
species and a photomicrograph key of reference plants were prepared by collecting
plant species from the study area. Diets of Markhor and domestic goat were identified
by matching the slides of faecal sample with the reference photographic key of plant
species. During spring season eight plant species were recorded in the faeces of
Markhor and five in the faeces of domestic goat. Among those, four plant species were
found common in the diet of both Markhor and domestic goat, including Salix iliensis,
Juniperus excelsa, Prangos pabularia and Linum perenne. Rumex lastatus was the
dominant plant species in the spring diet of Markhor. While the dominant plant in the
diet of domestic goat was Salix iliensis. In summer samples, nine plant species were
identified in the faeces of both the Markhor and domestic goat. Among those, all nine
53
54
plant species (Artemisia maritime, Pistacia integerrima, Origanum vulgare, Rheum
emodi, Hypsicum perforatum, Quercus baloot, Prangos pabularia, Linum perenne,
Bergenia ciliate) were found common in the diet of both Markhor and domestic goat.
Hypsicum perforatum was the dominant plant species in the summer diet of Markhor
while the dominant plant in the summer diet of domestic goat was Linum perenne.
This study revealed that the competition between Markhor and domestic goat for food
resources was very high in the study area during the summer season. As the diets of
Markhor and domestic goat were almost same during summer.
Conclusions
The microhistological technique based on the frequency of occurrence of plant
fragments on a microscope field (presence or absence) has become one of the most
popular methods of determining food habits, although it faces several limitations as
reported by the researchers evaluating its accuracy (Sparks and Malechek, 1968). One
of the limitations refers to the fact that the relation between identifiable epidermal
tissue and unidentifiable tissue is not similar for all species, which could result in overestimation or under-estimation some items/species (Santosh et al., 2005).
Markhor consumed eight plant species during the spring season including;
Salix iliensis, Origanum vulgare, Rumex lastatus, Astragalus psilocentros, Juniperus
excelsa, Prangos pabularia, Linum perenne and Rubus anatolicus while five plant
species were recorded in the faeces of domestic goat (Salix iliensis, Juniperus excelsa,
Prangos pabularia, Linum perenne. Capparis spinosa). In summer samples, nine plant
55
species were identified in the faeces of both Markhor and domestic goat which were
common in their diet. These included; Artemisia maritime, Pistacia integerrima,
Origanum vulgare, Rheum emodi, Hypsicum perforatum, Quercus baloot, Prangos
pabularia, Linum perenne and Bergenia ciliate.
During the spring season, Markhor strongly preferred Linum perenne while
domestic goat strongly preferred Linum perenne and Salix iliensis. During summer
season, both Markhor and domestic goat strongly preferred Linum perenne.
Diet overlap factor was 0.99 (
=0.99) during summer season which indicates
very high diet overlap between Markhor and domestic goat. During the spring season,
the value of diet overlap factor was 0.69 (
=0.69), indicating that the diets of
Markhor and domestic goat were not completely overlapped.
Tree species utilized both by Markhor and domestic goat included; Theli (Salix
iliensis), Saroz (Juniperus excelsa), Thoak (Pistacia integerrima) and Banj (Quercus
baloot ). Among shrubs, Ishpain (Origanum vulgare) and Shahtili (Linum perenne)
and among grasses and forbs, Moshain (Prangos pabularia), Dron (Artemisia
maritime), Ishpar (Rheum emodi), Matali (Hypsicum perforatum) and Zakhme Hayat
(Bergenia ciliate) were the species utilized both by Markhor and domestic goats.
Shahtili (Linum perenne) was the key preferred species by Markhor and domestic goat
in both summer and spring seasons.
56
During the summer season, Markhor and domestic goat showed high
overlapping in habitat use as indicated by their diet composition in this season. The
diet preferences of Markhor and domestic goat were found to be quite similar during
summer which lead to competition for food between them.
Recommendations
¾ Livestock grazing in the core habitat of Markhor in Chitral Gol National Park
should be completely banned.
¾ Livestock grazing should be regulated in the way that Markhor habitats
outside of the national park are least affected or not depleted.
¾ Habitat areas of Markhor, especially the preferred plant species for forage in
these areas should be protected. Efforts should also be made for the plantation
of these plants species in the degraded habitat of Markhor.
¾ Awareness of public about importance of biodiversity in general and threatened
species in particular should be raised to win their support and cooperation in
conservation efforts.
¾ Local communities should be involved in conservation and protection of
Markhor habitat. Economic incentive including alternate income sources
should be provided to the people living around Chitral Gol National Park and
Tushi Game Reserve.
57
LITERATURE CITED
Aleem, A. 1977. Population dynamics of markhor in Chitral Gol. Pak. J. Forestry, 27:
86- 92.
Aleem. A. 1976. Markhor in Chitral Gol. Pakistan Journal of Forestry, 26(2): 117128.
Aleem, A.1978. Markhor population dynamics and food availability in Chitral Gol
Game Sanctuary. Pak. J. Forestry, 28: 159-165.
Aleem, A. (N.D). The ecological impact of domestic stock on Chitral Gol Game
Sanctuary, Pakistan. p, 26-29.
Ali, H. and M. Qaiser. 2005. Sustainable use of plant wealth of Chitral and preparation
of Red Data List (Unpublished report submitted to HEC-Pakistan), 70 pp.
Ali, H. and M. Qaiser. 2006. Sustainable use of plant wealth of Chitral and preparation
of Red Data List (Unpublished report submitted to HEC-Pakistan), 85 pp.
Alipayo, D., R. Valdez, J. L. Holechek and M. Cardenas. 1992. Evaluation of
microhistological
analysis
for
determining
composition. J. Range. Manage., 45: 148-152.
57
ruminant
diet
botanical
58
Ali, S. 2008. Conservation and Status of Markhor (Capra falconeri) in the Northen
Parts of North West Frontier Provience, Pakistan. The University of Montana
Missoula, MT. 127 pp.
Awasthi, A., S. K. Uniyal, G.S. Rawat and S. Sathyakumar. 2003. Food plants and
feeding habits of Himalayan ungulates. Current Science, 85(6): 719-723.
Baumgartner, L. L. and A. C. Martin. 1939. Plant histology as an aid in squirrel food
habit studies. J. Wildl. Manage., 3: 266-268.
Bagchi, S., C. Mishra, Y. V. Bhatnagar. 2003. Conflicts between traditional
pastoralism and conservation of Himalayan ibex (Capra sibirica) in the TransHimalayan mountains. Animal Conservation, 7: 121–128.
Bhatnagar, Y.V. and V.B. Mathur. 2001. A Regional Perspective for Snow Leopad
Conservation in The Indian Trans-Himalaya. International Snow Leopard
Trust, Seattle, p. 26-38.
Bellon, L. 2008. Sustainable Conservation and Grassroots Realities Lessons From The
Conservation Programme in Torghar, Balochistan, Pakistan. Best Practices in
Sustainable Hunting (Case Study), p. 27–31.
Caughley, G. 1977. Analysis of vertebrate populations. John Wiley and Sons, New
York, 234 pp.
59
Clutton, B. T. H., F. E. Guinness, and S. D. Albon. 1982. Red Deer: Behaviour and
Ecology of Two Sexes. Wildlife Behaviour and Ecology Series. The University
of Chicago Press, Chicago, IL. 200 pp.
Coffey, L., M. Hale and A. Wells. 2004. Goats: Sustainable Production Overview.
National Sustainable Agriculture Information Service, p. 2-12.
Dayan, T. and D. Simberloff. 1998. Size patterns among competitors: ecological
character displacement and character release in mammals, with special
reference to island populations. Mam. Rev., 28(3): 99–124.
Din, J. U. and M. A. Nawaz. 2010. Assessment of the status of Himalayan Lynx (Lynx
Lynx
isabellina) in District Chitral, NWFP, Pakistan. J. Animal and Plant
Sci., 20(1): 40-43.
Fleischner, T. L. 1994. Ecological costs of livestock grazing in western North
America. Conserv. Biol., 8(3): 629–644.
Forsyth, D. M. 2000. Habitat selection and coexistence of the Alpine chamois
(Rupicapra rupicapra) and Himalayan tahr (Hemitragus jemlahicus) in the
eastern Southern Alps, New Zealand. J. Zool., Lond. 252: 215–225.
GOP. 2006. Pakistan Livestock Census. Agricultural Census Organization, Statistics
Division, Government of Pakistan, Lahore.
60
GoNWFP and IUCN Pakistan. 2004. Chitral – An Integrated Development Vision
(Chitral Conservation Strategy). IUCN-Pakistan and Government of NWFP,
Karachi, Pakistan, p. 14-103.
GoNWFP. 2006. Chitral Gol National Park Management Plan (Unpublished). Wildlife
Department Government of NWFP. 85pp.
GoNWFP. 2005. Survey Reports of wild mammals and birds in Chitral, Dir Kohistan,
and Swat. Wildlife Department, Government of NWFP.
Hashmi, A. A. and Shafiullah. 2003. Northern Areas Strategy for Sustainable
Development. Agriculture and Food Security. IUCN Pakistan, Northern Areas
Progamme, Gilgit, p. 59-136.
Hammer, S. E., H. M. Schwammer and F. Suchentrunk. 2008. Evidence for
introgressive hybridization of captive markhor (Capra falconeri) with domestic
goat. Biochemical Genetics, 46 (3-4): 216-226.
Horn, H. S. 1966. Measurement of overlap in comparative ecological studies. Am.
Nat., 100: 429-424.
Holechek, J. L., M. Vavra and R. D. Pieper. 1982. Botanical composition
determination of range herbivore diets. J. Range Manage., 35: 309 -315.
61
IUCN. 2001. Status of Flare-horned Markhor in Chitral Gol National Park and Tushi
Game Reserve, Pakistan. Caprinae, p. 1-9.
Jnawali, S. R. 1995. Population ecology of greater one-horned rhinoceros (Rhinoceros
unicornis) with particular emphasis on habitat preference, food ecology and
ranging behavior of a reintroduction population in Royal Bardia National Park
in lowland Nepal. Ph.D thesis, Agricultural University of Norway.
Khan, M. S., M. A. Khan and S. Mahmood. 2008. Genetic resources and diversity in
Pakistani goats. Int. J. Agri. Biol., 10 (2): 227-231.
Kleinn, E. 2005. Development initiatives versus environmental protection? In.
trategies for Development and food Security in Mountain Areas of Central
Asia. International workshop Dushanbe, Tajikistan. 20pp.
Kie, J. G., C. J. Evans, E. R. Loft and J. W. Menke. 1991. Foraging behaviour by mule
deer: the influence of cattle grazing. J. Wildl. Mgmt., 55: 665–674.
Leader, W. N. 1988. Reindeer in South Georgia: The Ecology of an Introduced
Population. Cambridge University Press, Cambridge, UK.
Lightner, J. K. 2006. Identification of species within the sheep-goat kind (Tsoan
monobaramin). J. Creation., 20(3): 61-65.
62
Malik, M. M. 1993. Wildlife Conservation in NWFP. Problems and Prospects.
Unpublished report. 19 pp.
Malik, M. M. 2002. Markhor trophy hunting in NWFP, Pakistan. 3 pp.
Manceau, V., L. Despres , J. Bouvet and P. Taberlet. 1999. Systematics of the genus
Capra inferred from mitochondrial DNA sequence data. Mol Phylogenet Evol.,
13: 504–510.
Mishra, C. and G. S. Rawat. 1998. Livestock grazing and biodiversity
conservation.Conserv. Biol., 12: 712–714.
Mishra, C. 2001. High altitude survival: conflicts between pastoralism and wildlife in
the Trans-Himalaya. Ph. D. Thesis, Wageningen Univ. Wageningen, the
Netherlands.
Mishra, C., S. E. V. Wieren, I. M. A. Heitkönigl and H. H. T. Prins. 2002. A
theoretical analysis of competitive exclusion in a Trans-Himalayan largeherbivore assemblage. Animal Conservation, 5: 251–258.
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. Applied Ecology, 41: 344–354.
63
Michel. S. 2008. Conservation and use of Wild Ungulates in Central Asia- Potentials
and Challenges. Nature Protection Team, Tajikistan, p. 32–40.
Nowak, R. 1999. Walker's Mammals of the World, Sixth Edition. Baltimore and
London: The Johns Hopkins University Press.
PAMP. 2008. Range management study in CGNP. upublished report. 30-37.
Putman, R. J. 1996. Competition and resource partitioning in temperate ungulate
assemblies. London, UK: Chapman and Hall.
Ranjitsinh, M. K., C. M. Seth, R. Ahmad, Y. V. Bhatnagar and S. S. Kyarong. 2007. A
Rapid Assessment of the Pir Panjal Markhor in Jammu and Kashmir:
Distribution, Status and Threats. A Schaller Conservation Survey. Wildlife
Trust of India, p. 42-50.
Rasheed, T. 2007. Values of wildlife with special reference to Northern Mountains.
Unpublished. 12 pp.
Rashid. M. 2008. Goats and their Nutrition. Manitoba Goat Association, p. 3-20.
Roberts, T. J. 1997. Mammals of Pakistan. Revised edition., Oxford University Press,
Karachi, Pakistan, 525 pp.
Saether, B. E. 1997. Environmental stochasticity and population dynamics of large
herbivores. Trends in Ecology and Evolution, 12: 143–149.
64
Sanders, K. D., B. E. Dahl and G. Scott. 1980. Bite count vs. faecal analysis for range
animal diets. J. Range. Manage., 33: 146-149.
Shackleton, D. M. 2001. A review of the community-based trophy hunting programs
in Pakistan. Prepared for the Mountain Area Conservancy Project with the
collaboration of IUCN-Pakistan, NCCW, MoELGRD., 59 pp.
Schaller, G. B. 1977. Mountain monarchs. University of Chicago Press, Chicago,
USA., 198 pp.
Shafique, C. M. and S. Barkati. 2002. Status and ecology of chiltan wild goat, Capra
Aegagrus Chialtanensis (Caprinae). Rec.Zool. Surv. Pakistan, 14: 81-93.
Santosh, S. A., C. Costa and A. Port. 2005. Test of a conversion equation to increase
the accuracy of micro histological analysis of herbivore diets in the Pantanal.
In: Alvarez, J. M. (ed.) Proceedings of 19th International Grasslands Congress;
Dublin, Ireland, pp. 507-514.
Sheikh, K. M. and S. Molur. 2005. Status and red list of Pakistan’s mammals, based on
conservation assessment and management plan for mammals. IUCN, Pakistan.,
344 pp.
Sparks, D. R. and J. C. Malechek. 1968. Estimating percentage dry weight in diets
using a microscopic technique. J. Range Manage., 21: 264-265.
65
Stahl, E. 2007. Summer of the Goats. The Farthest North Goat Association Newsletter,
30 pp.
Virk, A. T. 2000. Integrating wildlife conservation with community-based
development in northern areas, Pakistan. Caprinae newsletter of the IUCN/SSC
Caprinae Specialist Group, 70 pp.
Walter, G. H. 1991. What is resource partitioning? J. Theor. Biol., 150: 137-143.
Wiens, J. A. 1977. On competition and variable environments. Am. Sci., 65: 590-597.
Woodford. M. H., M. R. Frisina and G. A. Awan. 2004. The Torghar Conservation
Project: Management of the livestock. Suleiman Markhor (Capra falconeri)
and Afghan urial (Ovis orientalis) in the Torghar hills, Pakistan. Game Wildl.
Sci., 21 (3): 177-187.
Weinberg, P. I., R. Valdez and A. K. Fedosenko. 1997. Status of the Heptner´s
Markhor (Capra falconeri heptneri) in Turkmenistan. J. Mammal., 78: 826–
829.