Wildlife Biology - 2019 - Akrim - Diet Composition and Niche Overlap of Two Sympatric Carnivores Asiatic Jackal Canis

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Wildlife Biology 2019: wlb.00440
doi: 10.2981/wlb.00440
© 2019 he Authors. his is an Open Access article
Subject Editor: John Ball. Editor-in-Chief: Ilse Storch. Accepted 21 January 2019
Diet composition and niche overlap of two sympatric carnivores:

Asiatic jackal Canis aureus and Kashmir hill fox Vulpes vulpes
griffithii, inhabiting Pir Lasura National Park, northeastern
Himalayan region, Pakistan
Faraz Akrim, Tariq Mahmood, Muhammad Sajid Nadeem, Tashi Dhendup, Hira Fatima and

Shaista Andleeb
F. Akrim, T. Mahmood ([email protected]), H. Fatima and S. Andleeb, Dept of Wildlife Management, Pir Mehr Ali Shah Arid
Agriculture Univ., PK-46300 Rawalpindi, Pakistan. FA also at: Dept of Zoology, Univ. of Kotli, Azad Jammu and Kashmir, Pakistan.
– M. S. Nadeem, Dept of Zoology, Pir Mehr Ali Shah Arid Agriculture Univ., Rawalpindi, Pakistan. – T. Dhendup, Wildlife Biology Program,
Dept of Ecosystem and Conservation Sciences, College of Forestry and Conservation, Univ. of Montana, Missoula, Montana, USA, and: Ugyen
Wangchuvk Inst. for Conservation and Environmental Reserach, Bhumthang, Bhutan.
Studies on dietary habits and niche overlap of sympatric carnivore species can be vital for their conservation. hey can reveal
the potential level of inter-speciic competition and prey species overlap, thus highlighting species speciic conservation
requirements. We investigated diet composition of two such sympatric carnivore species Asiatic jackal Canis aureus and
the Kashmir hill fox Vulpes vulpes griithii in and around Pir Lasura National Park, in northeastern Himalayan region of
Pakistan by using fecal analysis after conirming the carnivore species by genetic analyses. he scats of the two carnivores
were collected during four diferent seasons of the year and analyzed in the laboratory. Results revealed sixteen prey species
in the diet of Asiatic jackal and 21 species in the diet of the Kashmir hill fox. For Asiatic jackal, wild prey contributed
approximately 18%, whereas 60% of the diet comprised of domestic prey. For Kashmir hill fox, the wild prey consumption
was approximately 18%, while consumption of domestic prey was 51% and plants contributed 28%. Niche breadth of
the Asiatic jackal was broader (0.78) compared to Kashmir hill fox (0.31). Niche overlap between these two sympatric
carnivores was found to be a rather high 0.81.
Keywords: diet composition, dietary niche breadth, niche overlap
Knowledge of a predator’s diet is vital to understand its ecol- mucus cells and bacteria (Bang and Dahlström 1975, Bujne
ogy and to predict its efect on the dynamics of prey popula- 2000). However, the quantity and size of carnivore scats
tions (Oli 1993). Scat analysis has been used extensively by can be diferent based upon age of individuals (Akrim et al.
various researchers in various parts of the world for inves- 2018), prey species consumed and absorption capacity (Bang
tigating the diet composition of carnivores, especially the and Dahlström 1975).
elusive species. However, the correct identiication of scats Carnivore species that are widely distributed globally
in the ield, based on morphology of the scats, is vital oth- include golden or Asiatic jackal Canis aureus and the red fox
erwise wrong information may be communicated to readers Vulpes vulpes. he Asiatic jackal has a widespread distribution
(Akrim et al. 2018). herefore, more recently, ield identiied in north and north-east Africa, also occurs in the Arabian
scats of carnivores are being subjected to molecular identii- Peninsula and has expanded its range into Europe, where
cation techniques for conirmation of the carnivore species. the species has patchy distribution. It is regularly found as a
Components of carnivore feces can include feathers, bones, vagrant in Austria, Slovakia, Slovenia and north-eastern Italy
hairs, teeth, claws, scales, arthropod chitin, plant matter, (Krystufek et al. 1997). It is well distributed in central Asia
and the entire Indian subcontinent (Jhala and Moehlman
2008). Red fox Vulpes vulpes is most widespread terrestrial
his work is licensed under the terms of a Creative Commons
Attribution 4.0 International License (CC-BY) < http:// carnivore species (covering nearly 70 million km2) glob-
creativecommons.org/licenses/by/4.0/ >. he license permits ally (Harris and Baker 2001, IUCN 2016). It is distributed
use, distribution and reproduction in any medium, provided the across the whole northern hemisphere from the Arctic circle
original work is properly cited. to southern North America Europe, North Africa, Asiatic
1
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steppes, Pakistan, Indian and Japan (Jenkins and Craig Material and methods
1992, Gloor et al. 2001, Wandeler et al. 2003, Crooks et al.
2010, Soulsbury et al. 2010). Study area
Although, the Asiatic jackal is not found at higher eleva-
tions in the Himalayas (above 3500 m), it has however been his study was conducted in and around Pir Lasura National
recorded at lower elevations in Himalayan valleys such as Park (PLNP; 33°25¢92²N to 33°29.31²N and 74°05¢64²E
Murree hills up to 2150 m (Roberts 1997). he red fox, on to 74°03¢02²E), District Kotli, north-eastern Himalayan
the other hand occurs up to 4500 m asl (IUCN 2016). region, Pakistan. he park is an important protected area
he Asiatic jackal is an opportunistic feeder and it usually in the country regarding some key sympatric carnivores. It
scavenges on garbage, human waste and animal carcasses. encompasses 1580 ha area with elevation ranging between
Jackals can hunt singly and in packs which are more success- 1000 and 2000 m above sea level (a.s.l.). Since the current
ful in hunting. When single, it hunts smaller prey such as study also focused surrounding areas of the Park, the overall
rodents, hares and birds by locating prey by hearing. It can size of the whole study area was 17 183 ha. he valleys of the
dig out Indian gerbils Tatera indica from burrows and can park consist of subtropical pine vegetation, with the tops/
hunt young, old, or sick ungulates 4–5 times larger than its mountains having sub-tropical dry evergreen forest. Average
own size and body weight. Besides vertebrates, the Asiatic annual rainfall in the study area is 1500 mm. he study area
jackal has been reported to feed on invertebrates and plant experiences four diferent seasons during the calendar year
matter. Jackals are omnivorous and the range of their diet including summer (May–July), autumn (August–October),
varies among diferent seasons and habitat (Wyman 1967, winter (November–January) and spring (February–April).
Moehlman 1983 ). hey are generalist feeders and large Major wildlife species in the park include common leopard
quantities of vegetable matter and fruits are included in their Panthera pardus, rhesus monkey Macaca mulatta, Asiatic
diet, however, bulk of their food is comprised of rodents jackal Canis aureus, Kashmir hill fox Vulpes vulpes griithii,
and reptiles. hey also supplement their diet with fruits and small Indian mongoose Herpestes javanicus, Indian grey
insects when available (Roberts 1997). mongoose Herpestes edwardsii, barking deer Muntiacus munt-
he red fox is an omnivore; it has broad diet which jak, Indian pangolin Manis crassicaudata and kaleej pheasant
includes; invertebrates, small mammals, birds, ishes, fruits Lophura leucomelanos (Akrim et al. 2017, 2018).
and carrion (Flower 1932, Macdonald 1979, Osborn and Local people keep a variety of animals including domestic
Helmy 1980). Diet composition of the red fox depends on cows, bufalos, goats, dogs, horses, poultry and rabbits. A
various factors including habitat type, prey availability (Kolb majority of people are associated with professions of doing
and Hewson 1979, Leckie et al. 1998, Sidorovich et al. 2006) agriculture, government jobs, labor and shop-keeping with
and seasonal variation in food availability (Goszczyński average household income per month ranging from US$
1986, Jędrzejewski and Jędrzejewska 1992, Baltrunaite 100 to 200. Farmers, shopkeepers and labor or usually keep
2001, 2002). Data on dietary composition of the red fox livestock for milk and meat production and they depend on
and Asiatic jackal is well documented in other countries, livestock for subsistence.
however very few studies on diet of Asiatic jackal have been
conducted in Pakistan and no research study has focused on Morphological identiication of carnivore scats
the red fox.
he Asiatic jackal and the red fox are both reported he diet composition of Asiatic jackal and Kashmir hill fox
from Pakistan and in many parts of the country are sym- was investigated by analysis of scat samples. We conducted
patric in distribution (Roberts 1997). However, at higher surveys to collect scats of both carnivores during summer,
elevation areas, the sub-species Kashmir hill fox Vulpes autumn, winter and spring seasons during 2014–2017 using
vulpes griithii occurs. Since both species are carnivores area searches on 30 trails in the study area. hree people
and share distribution ranges in the country, their feeding participated in survey but to eliminate one source of varia-
niches may overlap, potentially resulting in interspeciic tion, only a single person (the author) was responsible for
competition for food. Although a few individual studies morphological identiication of the scats in the ield. When
have focused on investigating the diet of the Asiatic jackal any scat was encountered, ield identiication was made
in the country such as Nadeem et al. (2012) and Mah- based on its morphology including diameter, length, shape,
mood et al. (2013), the red fox has been neglected totally color, odor, physical appearance such as characteristic con-
in this regard, with virtually no studies reported from Pak- tents (hairs, bones and plant material) (Seton 1925, Jackson
istan that have investigated its diet and prey species. he and Hunter 1995). Additional criteria included nature of
study of diet composition and niche overlap of the two scat deposit site and presence of tracks or signs of activity
carnivores can reveal the potential level of competition for of the species under study. he diameter at widest point,
their prey and the species-speciic habitat requirements length, disjoint segments and weight of each scats sample
of the two carnivores wherever the two are sympatric in was measured and samples were preserved in 95% ethanol
their distribution. he Pir Lasura National Park (PLNP), for molecular identiication and further analysis.
is an important protected area and has both the Asiatic
jackal and the Kashmir hill fox. Here, we investigate the Molecular identiication of carnivore scats
diet composition and the niche overlap between these
two sympatric meso-carnivores in the human-dominated he morphologically-identiied scats of the two carnivore
landscape in and around Pir Lasura National Park, north species were subjected to molecular identiication for further
Himalayan region, Pakistan. conirmation. For this purpose, we extracted fecal DNA in
2
the Non-invasive and Environmental DNA Lab (NIEL), Scale replication
Conservation Genomics group (CGG) at the University of
Montana, Missoula, USA which is dedicated to DNA extrac- Cuticular scale patterns of mammalian hair were identiied
tions. We used QIAamp DNA Stool Mini Kits (Qiagen, by using a slightly modiied procedure after Lavoie (1971).
Inc., Valencia, CA) for extraction of DNA from scats. We Two to three drops of transparent nail polish were spread
used negative controls to keep track of cross contamina- evenly on a glass slide. A small hair was placed in vertical
tion during extraction (Beja-Pereira et al. 2009). he total position along axis of slide so as one end of hair projected
volume of DNA extracts from each scat sample was 100 µl. out of slide. After the nail polish was dry the end of hair pro-
he PCR for all scats samples were carried out in a total jecting out was plucked with a single motion using forceps to
volume of 50 µl. he recipe of our master mix (MM) per get a cast of the hair’s surface in the nail polish. he hair cast
sample was 20.375 µl H2O, 5 µl bufer (7 µl MgCl2, 0.375 µl was studied under a microscope against a reference collection
BSA, 2 µl dNTP, 2.5 µl 12S/V5 primer F, 2.5 µl 12S/V5 for identiication.
primer R, 0.25 µl Taq polymerase and 10 µl DNA as extract
template for each scat sample. he PCR condition was dena- Identiication of plant matter
turation 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 Plant matter recovered form scats of two carnivore species was
elongation at 72 °C for 1:30 min. A inal elongation at 72°C mainly comprised of seeds and fruit parts. Recovered seeds
for 5 min at the end and 4°C until the product was removed and fruit remains were compared with reference material
from PCR. All PCRs were conducted on Eppendorf vapo. collected from the ield to identify them. Seeds were also
protect Master cycler pro and all reactions included a nega- sown and germinated in pots for plant species identiication.
tive and positive control. All sequences were then run on a
3130 genetic analyzer and sequences were read using Finch Dietary niche breadth
TV software. he sequences were then subjected to NCBI
We measured dietary niche breadth of two sympatric
Blast for species identiication. All failed samples were dis-
carnivore species using niche breadth (L) and standardized
carded and not included in the inal analysis.
Levins index (0–1) (Lst) (Levins 1968, Colwell and Futuyma
1971) as follows:
Scat analysis
-1
æ n ö L -1
he collected scat samples were assigned to each of the carni- L = ç å pi2 ÷ and Lst =
vore species based on molecular identiication for investigat- è i =1 ø n -1
ing diet composition of the two carnivores. During analysis, where pi is the relative percentage of food item i and n is the
scat samples were sun dried irst and then morphological number of food items.
characteristics of scats such as length, breadth and weight Lst is the standardized niche breadth and its value ranges
were recorded. For disintegration, scat samples were soaked from 0 to 1. A higher Lst indicates a broader diet niche.
in warm water and then washed under tap water in a sieve to
remove dust and mucus and segregated diferent prey items Niche overlap
such as hairs, bones, insects, bird feathers and plant parts
(Mahmood et al. 2013). hese prey parts were dried and We used the frequency of occurrence of each prey item to
divided into diferent groups such as plant-based diet and compute the dietary overlap between jackal and fox occur-
animal-based diet. ring in the study area using Pianka’s index, the value of
which ranges from zero (no overlap) to one (complete over-
Whole mount preparation lap) (Pianka 1973). We chose this index to allow comparison
of the degree of overlap to similar studies of carnivores con-
We used hairs recovered from the scats for identiication ducted elsewhere in the world (Fedriani et al. 2000, Ray and
of mammalian prey species of the two carnivores. For this Sunquist 2001, Jacomo et al. 2004). he Pianka’s index was
purpose, light microscopic slides of the hairs of prey species calculated using formula:
were prepared. Hairs were washed in carbon tetrachloride
15–20 min), long hairs were cut into small pieces and jum- n
bled up hairs were separated. For whole mount preparation, åp ij pik

we used transparent nail polish. O jk = i
n n
Prey species of carnivores were identiied using medullary
pattern and cuticular cast pattern of the hairs recovered from åp åp
i
2
ij
i
2
ik
scat samples as described by Moore et al. (1997). Prepared

light microscopic slides were then compared with reference where pij (or pik) is the relative percentage of food item i in
hair slides for identiication. diet j (or k).
Similarly, other parts recovered from scat samples such he prey species diversity index (H′), prey richness (S)
as bones, feathers of birds, invertebrates such as insects and and prey evenness (E) indices were calculated for each of the
plant matter including seeds were also identiied. he hairs two-carnivore species during diferent seasons. Prey species
of prey species were identiied using a light microscope with richness (S) was the total number of prey species consumed
objectives of 10×, 40× and 100× magniication. by each predator during each season.
3
Diversity index (H′) was calculated by using the follow- mammals, birds, insects and plants (Table 1). Among
ing formula: these species, 10 were wild, 5 domestic and 1 plant species.
Frequency of occurrence of wild prey in the diet of jackal
H¢ = -å éë pi ´ ln pi ùû was 18% but that of domestic prey was approximately
60%. Among wild prey, rhesus monkey was most consumed
where pi is prey index. (5.43%), followed by Norway rat (3.26%). Among domestic
he evenness index (E) was calculated by using the formula: prey, consumption of poultry was high (27.17%), followed
H¢ by domestic goat (20.65%). he plant matter represented
E= 3.26% of the jackal’s diet (Table 1). Bones and hairs of the
ln of S
prey remains dominated the scats of Asiatic jackal by percent
where, S represents the prey species richness and H′ volume occurrence followed by bird feathers. A general linear
represents the diversity index. model itted explained 82.6% of the variation in prey item
All data were analyzed using SPSS (ver. 23) software and consumption of Asiatic jackal. he consumption of dietary
Excel statistics. items varied signiicantly F = 15.02, df = 17, p < 0.0001.
We compared total frequency of dietary items con- he analysis of seasonal variation in diet composition
sumed by each carnivore species for statistical diferences. revealed a higher consumption of wild prey during the
To compare seasonal variation in diet composition of each autumn season (21.88%) but low during spring (12.5%).
carnivore species we used a generalized linear model (GLM). Similarly, consumption of domestic prey was high during
Similarly, we compared seasonal variation in consumption the winter season (66.67%) and low during autumn and
of wild prey species, domestic prey species and plant mat- spring (56.25% each) (Table 1). GLM showed no statisti-
ter. We repeated GLM for variation in consumption of each cally signiicant diference in seasonal diet consumption
dietary item consumed by each carnivore species. Frequen- of Asiatic jackal during the study period F = 0.946, df = 3,
cies of dietary components of four seasons were our response p = 0.423. Similarly, consumption of wild prey species did
(dependent) variables while carnivore species were explana- not difer signiicantly during four seasons F = 1.082, df = 3,
tory (independent) variables. he distribution of our data p = 0.371. Consumption of domestic prey species did not
was normal p > 0.05. Our scale response was linear which difer during diferent seasons F = 0.346, df = 3, p = 0.792.
speciies normal as the distribution and identity as the link
function. All analysis was conducted in SPSS ver. 23.
Kashmir hill fox
Analysis of n = 92 scat samples of Kashmir hill fox revealed
Results 21 prey species including mammals, birds, plants and
insects in the diet. Among prey species, 10 were wild, 5
Diet composition were domestic and there were 6 plant species. Frequency
of wild prey was approximately 18%, domestic prey 51%
Asiatic jackal and plants 28% (Table 2). Among wild prey species, wild
Analysis of n = 64 scat samples of the Asiatic jackal collected boar was most consumed (2.99%) followed by house mouse
during four diferent seasons (summer, autumn, winter and (2.4%). Among domestic prey, the consumption of poul-
spring) of the year revealed sixteen prey species including try was high (29.34%), followed by domestic goat (14%)
Table 1. Percent frequency (%F) of occurrence of prey items in diet of Asiatic jackal in the study area.
Prey species Summer (n = 17) Autumn (n = 23) Winter (n = 14) Spring (n = 10) % Total freq.
Wild prey
Rhesus monkey Macaca mulatta 4.35 9.38 0 6.25 5.43
Wild boar Sus scrofa 4.35 3.13 0 0 2.17
Desert hare Lepus nigricollis dayanus 0 3.13 0 0 1.09
Indian gerbil Tetra indica 4.35 0 0 0 1.09
Roof or house rat Rattus rattus 0 3.13 0 0 1.09
House mouse Mus musculus 0 0 4.76 0 1.09
Norway rat Rattus norvegicus 0 0 9.52 6.25 3.26
Kalij pheasant Lophura leucomelanos 0 3.13 0 0 1.09
Spotted dove Streptopelia chinensis 4.35 0 0 0 1.09
Insects (Orthoptera) grasshopper 0 0 4.76 0 1.09
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 domesticus 21.74 25 28.57 37.5 27.17
Total domestic prey 60.87 56.25 66.67 56.25 59.78
Jaro grass Thameda anathera 4.35 6.25 0 0 3.26
Grits 4.35 3.13 4.76 6.25 4.35
Anthropogenic 13.04 12.5 9.52 25 14.13
4
(Table 2). A general linear model explained 73.9% varia- autumn (0.86) and low during summer and winter seasons
tion in consumption of diferent dietary items by Kashmir (0.83) (Fig. 1).
hill fox (R2 = 0.739). he consumption of diferent dietary
items also varied signiicantly in the diet of Kashmir hill fox Niche breadth and niche overlap
(F = 8.86, df = 22, p < 0.0001).
Consumption of wild prey species was found to be high When we compared the two sympatric carnivore species in
during spring season (25%) but low during summer season habiting the PLNP study area, the Asiatic jackal was found
(9.43%), whereas consumption of domestic prey was high to have a much broader niche breath compared to the
during winter (66.67%) but low during autumn (40.35%). Kashmir hill fox (Fig. 2). Individually, the niche breadth of
Consumption of plant species was high during summer and Asiatic jackal was found broad during summer (17.25; 0.96)
autumn (35.85% and 35.09%, respectively) but lowest dur- but narrow during spring season (6.67; 0.33). Total niche
ing winter (9.09%). Analysis of the seasonal variation in breadth of the Asiatic jackal was found to be 14.2 (0.78).
the diet composition of Kashmir hill fox by GLM showed On the contrary, the niche breath of Kashmir hill fox was
no signiicant diference across diferent seasons (F = 1.14, wider 9.07 (0.37) during the autumn but narrow during
df = 3, p = 0.337). Consumption of wild prey species also did spring season 4.8 (0.17). he overall niche breadth of fox
not difer across all four seasons (F = 1.025, df = 3, p = 0.395), was 7.89 (0.31) (Fig. 2). Niche overlap between these two
similarly consumption of domestic prey did not difer among sympatric carnivores was found to be 0.81.
all four seasons (F = 0.201, df = 3, p = 0.894).
Prey species diversity, richness and evenness Discussion

he prey species diversity index of the diet of the Asiatic Knowledge of a predator’s diet is vital to understand its
jackal was found to be high during summer and winter sea- ecology and to predict its efect on the dynamics of prey
sons (2.22 each), but low during spring (1.51). Prey species populations (Oli 1993). No previous studies exist on diet
richness was high during summer (12) but low during spring composition of Asiatic jackal and Kashmir hill fox from
(7). Prey species evenness was high during winter (1) but Azad Jammu and Kashmir particularly and diet composition
low during spring 0.78 (Fig. 1). Whereas for the Kashmir of the Kashmir hill fox have not been studied in Pakistan so
hill fox, prey species diversity index in diet was high during far. During this study we recorded mammals, birds, insects
autumn season (2.38) but low during spring (1.83) whereas, and anthropogenic items from the diet of Asiatic jackal at
prey richness was high during autumn (16 species) and low Pir Lasura National Park. Consumption of domestic prey
during spring (9 species). Prey evenness was high during was higher than that of wild prey. Wild prey species included
Table 2. Percent frequency (%F) of occurrence of prey items in diet of Kashmir hill fox in the study area.
Prey species/items recovered Summer (n = 27) Autumn (n = 31) Winter (n = 19) Spring (n = 15) % Total freq.
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
5
18
Asiatic Jackal Red Fox 16
16
14
14
12
12 11
Value of index
10
10 9 9
8 7
4 2.2 2.1 2.2

1.51
2.2 2.38 0.9 0.78
1.87 1.83 0.87
2 1
0.83 0.86 0.81 0.83
0
Summer Autumn Winter Spring Summer Autumn Winter Spring Summer Autumn Winter Spring
prey species diversity index (H') prey richness (S) prey evenness (E)
Figure 1. 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.
rhesus monkey (which was the main wild prey species) and grass 20%, fruits 16.5%, insects 4.1%, snakes 4.1%, chital
four species of rodents. Among domestic prey species, poul- 2.4%, nilghai 1.2% and ish 1.2% (Sankar 1988). Stomach
try was the main prey followed by goat, whereas consump- contents of four jackals in Rajasthan Indian revealed that diet
tion of plant matter was low. Studies conducted elsewhere of jackals mostly consisted of fruits of Ziziphus, with small
have reported rodents and plants as main components of proportion of beetles and scorpions. Another individual con-
jackal diets. However, we found that this was not the case in sumed desert jirds Meriones hurrianae and small mongoose
our study area, where the jackal seems to be more of a scav- (Prakash 1959). Analysis of 138 scats of Asiatic jackal in sal
enger and predates on domestic poultry for a major part of forests in India showed that 68 percent of their diet con-
its diet, followed by rodents and rhesus monkeys. Our results sisted of rodents followed by plants 27%, 11% reptiles, 8%
are in line with Roberts who reported that in Pakistan the ish and 9.4% birds (Schaller 1967). A study conducted in
jackal is perceived as a scavenger and carrion eater (Roberts Sariska Tiger Reserve, India showed that the diet of the Asi-
1997). Jackals feed on refuse in villages, however the major- atic jackal was comprised of plant matter 17.57%, rodents
ity of their diet is comprised of rodents, reptiles, fruits and 15.77%, cattle 15.32%, chital 10.81%, fruits 9.01%, birds
insects when available (Roberts 1997). A study conducted in 7.21%, sambar 5.41%, hare 4.05%, nilgai 4.05%, goat
Bharatpur, Rajastan, India during 1984–1985 showed that 1.80% and reptiles 1.8% (Mondal 2012). Diet of jackals
diet of jackal comprised of rodents 26.5%, birds 24.1%, usually are comprised of small and medium-sized mammals
17.25
16.67
Asiatic Jackal Kashmir Hill fox

14.2
Niche breadth
10.9
9.07
7.89
6.77
6.67
5.2
4.8
0.96
0.92
0.78
0.58
0.37
0.33
0.31
0.26
0.19
0.17
L LST L LST L LST L LST L LST

Summer niche Autumn niche Winter niche Spring niche Total niche
Breadth Breadth Breadth Breadth Breadth
Figure 2. 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).
6
such as rodents, rabbits, birds, ishes, insects and vegetation he Kashmir hill fox also consumed sheep, cow and bufalo
(Aiyadurai and Jhala 2006). he diet of jackal in Keoladeo which likely indicate the fox’s scavenging behavior. he con-
National Park, India was comprised of mainly plant mat- sumption of diferent food items varied among during difer-
ter 38.44%, nilgai 8.13%, rodents 10.31%, chital 9.69%, ent seasons, in agreement with other studies (Basuony et al.
cattle 7.5%, anthropogenic 1.56% (Singh 2016). his high 2005, Baker et al. 2006, Kidawa and Kowalczyk 2011). he
proportion of mammals in the diet of jackal suggest that red fox is essentially an omnivore having a diverse diet which
they scavenge on dead animals and hunt calves as hunting includes invertebrates, small mammals, birds, plant matter
larger mammals might be diicult (Singh 2016). Rodents and carrion (Flower 1932, Englund 1965, Amores 1975,
are an important part of jackal diets (Mukherjee et al. 2004, Macdonald 1979, Osborn and Helmy 1980, Ciampaloni
Jaeger et al. 2007, Majumder et al. 2011, Singh 2016). Diet and Lovari 1985, Calisti et al. 1990, Basuony 1998). Fox has
of Asiatic jackal in Pakistan showed that 46.47% of its diet been reported by other studies to feed on plant material in
is animal matter by volume including rodents, mongooses, southern Europe (Ciampaloni and Lovari 1985, Calisti et al.
wild boar, livestock, birds and domestic poultry and 25.08% 1990) and it is known to feed on plant species or parts of
plant included wheat, tomato, berries, grains, orange, melon plants having high sugar content (Basuony et al. 2005).
and water melon (Mahmood et al. 2013). Most of these Niche overlap among Asiatic jackal and Kashmir hill fox
previous studies have recorded plants and rodents as major was high (0.81) indicating a potential for competition in
prey species in diet of Asiatic jackal, but this could be due the study area. Lanszki et al. (2006) reported that trophic
to diference in methodology; ours is the irst study in the niche overlap among jackal and fox was 0.73 in Hungary. Of
region to use genetic methods to conirm the identity of the course, shared food resources are only indicative of competi-
species which produced a given scat. Jackals living in high tion as food could be super abundant. However, it can be
mountainous regions can take up mammals only as major argued that if some food items are shared but the two preda-
part of their diet (Schaller 1970). In India Schaller (1970) tors are not competing for it now, they might in future if this
reported that frequency of occurrence of rodents in the diet resource declines, or one (or both) of the predators increase.
of jackal was 94%, however he also reported snakes and liz- We used scat analysis for documenting diet composition
ards 29% and insects 6.7%. Previous, studies which have and niche overlap of both carnivore species which is most
been conducted in Pakistan has also reported large propor- widely-used method for such purposes. Results of scat analysis
tions of plant matter in the diet of jackal. However unlike only reveal undigested parts recovered from the scats hence we
previous studies, we conirmed scats by genetic analysis; it cannot obtain direct information about digested material using
was interesting that scats identiied as being from jackal had scat analysis. We did not try to calculate biomass consumed,
very little or no plant matter, and that other sympatric carni- because of substantial uncertainties which could afect these
vores such as fox and civets consumed plant matter in large calculations (Chakrabarti et al. 2016, Lumetsberger et al.
proportions. he diference between the results of our analy- 2017). For example, hairs can identify the prey species, but it is
sis and other studies in the region raises a question as to how impossible to know a) whether a young/subadult/male/female
much of this diference is might due to diferences in diets has been preyed upon and these diferent classes vary greatly in
in diferent places or times, or to what extent previous stud- body mass) or b) whether a predator scavenged from a carcass
ies may have misidentiied some unknown fraction of scats. which was killed by another predator.
We suggest that future studies of these predators conirming
them by genetic analysis now that costs make this practical. Conclusion/recommendations
here is a paucity of information available on diet com-
position of fox in Pakistan and no published scientiic lit- his study has provided baseline data on dietary habits and
erature is available. However, some information is available niche overlap of two sympatric carnivore species, the Asiatic
from a book by Roberts (1997) for comparison who states jackal and Kashmir hill fox inhabiting Pir Lasura National
that foxes are adaptable hunters and can hunt hares, rodents, Park, north-eastern Himalayan region of Pakistan. he
reptiles and small birds but when vertebrate prey is not avail- dietary niche breadth of Asiatic jackal was wider than that
able they can subsist on insects and fruits. According to Rob- of Kashmir hill fox, however, the dietary niche of the two
erts, foxes feed on fruits of ber tree Zizyphus mauritiana. In carnivores overlapped 81% in the study area, suggesting a
the diet of fox, mice, rats, desert hare, Indian gerbils, have high level of potential competition between these two car-
been reported by Roberts (1997). Studies in Indian Rajast- nivore species. he diet of both carnivores showed that they
han showed that foxes feed on wild melon, termites (Prakash predated on domestic poultry which leads to conlict, and
1959). Foxes have reported to feed on scorpions, fruits of thus raises concerns for conservation. his human–carnivore
Zizyphus nummularia, spiders, cockroaches and also had interaction, as well as a feeding study on captive animals
some melon seeds in stomach. Roberts (1997) never encoun- should be performed in the future. Our results are important
tered any sign of foxes feeding on domestic poultry although in the context of determining species-speciic conservation
he has reported instances of domestic poultry being eaten by requirements, not only in the study area, but also in other
civets, cats, mongooses and martins. parts of the world where these two carnivores overlap.
During the present study, we recorded that diet of
Kashmir hill fox diet was comprised of mammals, birds, Acknowledgements – he authors are highly thankful to ‘Higher
invertebrates and plant matter. We also recorded anthropo- Education Commission Pakistan’ for providing funding (no. 1-8/
genic material in the diet of fox. Consumption of poultry HEC/HRD/2016/5884). We are also thankful to Prof. Dr. L. Scott
was higher than any other dietary component, followed by Mills and Tamara Max (Univ. of Montana, USA), for their support
goat. We identiied six species of plants in the diet of fox. and guidance.
7
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1903220x, 2019, 1, Downloaded from https://onlinelibrary.wiley.com/doi/10.2981/wlb.00440 by INASP/HINARI - PAKISTAN, Wiley Online Library on [12/01/2023].

Diet composition and niche overlap of two sympatric carnivores:

Faraz Akrim, Tariq Mahmood, Muhammad Sajid Nadeem, Tashi Dhendup, Hira Fatima and

Keywords: diet composition, dietary niche breadth, niche overlap

bled up hairs were separated. For whole mount preparation, åp ij pik

scat samples as described by Moore et al. (1997). Prepared

Prey species diversity, richness and evenness Discussion

4 2.2 2.1 2.2

Asiatic Jackal Kashmir Hill fox

L LST L LST L LST L LST L LST

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