Food Habits of Golden Jackal (Canis aureus) and Striped Hyena

 
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World Journal of Zoology 7 (2): 106-112, 2012
ISSN 1817-3098
© IDOSI Publications, 2012
DOI: 10.5829/idosi.wjz.2012.7.2.63139

                 Food Habits of Golden Jackal (Canis aureus) and Striped Hyena
                    (Hyaena hyaena) in Sariska Tiger Reserve, Western India

                      Pooja Chourasia Krishnendu Mondal, K. Sankar and Qamar Qureshi

                                     Wildlife Institute of India, Dehradun, India
    .
    Abstract: Food habits of golden jackal (Canis aureus) and striped hyena (Hyaena hyaena) were investigated
    using scat analysis between November 2010 and June 2011 in Sariska Tiger Reserve, Western India. Total 104
    scats of golden jackal and 86 scats of striped hyena were collected and analyzed. The frequency of occurrence
    of each prey species was estimated through bootstrapping using program SIMSTAT. Niche breadth of these
    two species was quantified using Levin’s measure. The diet overlap in the two species was assessed using
    Pianka’s index. Twelve food items were identified in golden jackal scats and nine in striped hyena scats.
    Vegetative matter contributed maximum (17.57%) in jackal’s diet followed by rodents (15.77%), chital (10.81%),
    sambar (5.41%) and nilgai (4.05%). Nilgai and domestic cattle contributed maximum (24.76% each) in the diet
    of striped hyena, followed by sambar (17.14%), chital (16.19%) and vegetative matter (10.48%). The estimated
    dietary overlap between striped hyena and golden jackal was 67%. Niche breadth for golden jackal was
    estimated as 0.69 and for striped hyena it was 0.57. The considerable overlap was attributed to mutual
    dependence on ungulates, which indicated high resource competition between the two species.

    Key words: Canis aureus       Dietary Niche Overlap       Food Habits       Hyaena hyaena      Prey Size    Scat

                   INTRODUCTION                                     between predator and prey sizes. A confounding factor in
                                                                    predator-prey studies that include a wide size-range of
     The extent of niche differentiation and resource               mammalian predators is the high variability in diet of
partitioning determines the degree to which different               smaller species, which can switch between insectivore,
species can either coexist or competitively exclude each            omnivore and carnivore. Canid and felid species above
other [1, 2]. An important mode of resource partitioning is         certain threshold weight class predate purely on
the degree of dietary overlap between sympatric species             vertebrates, while those below may be omnivorous or
[3]. This overlap is influenced by each species’ physical           predate on invertebrates and vertebrates. This implies that
ability to obtain food [4, 5] and also by variation in the          large carnivores (>21.5 kg) constitute a distinct functional
spatial and temporal availability of food [6]. For distinct         group from which predator-prey size relationships should
carnivore guilds different mechanisms may be involved,              emerge more clearly than from carnivore assemblages that
such as different body sizes of predators [7] and prey              are distributed across both sides of the body mass
species [8, 9], activity pattern [10] and microhabitat use          threshold [14]. The diet of the golden jackal and the
[11]. However, whereas resources are limited, the strength          striped hyena was studied through scat analysis in India
of competition between sympatric species generally                  [15-18] and other countries [19-22]. Merve et al. [23]
decreases with increased resource partitioning [12].                studied diet overlap between brown hyena (Parahyaena
     Study by Dickman [13] showed that the smaller                  brunnea) and black-backed jackal (Canis mesomelas) in
predators in an insectivorous mammal guild consumed                 Atherstone Nature Reserve, Pilanesberg National Park
smaller prey, but switched over to larger prey when the             and private farm (Mankwe Wildlife Reserve) in South
larger predators were removed. Consequently prey                    Africa. They inferred that, despite the marked body size
selection could depend more on competition among                    differences, brown hyena and black-backed jackal can be
predator guild members than on any inherent relationship            considered as either meso-carnivores or apex predators,

Corresponding Author: K. Sankar, Wildlife Institute of India. P.O. Box 18,
                      Chandrabani, Dehradun-248 001, (Uttarakhand), India.
                      Mob: +91 9412992784.
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depending on the presence or absence of larger                       carnivores and omnivores such as tiger, leopard, jungle
carnivores [24]. As meso-predators, the potential food               cat (Felis chaus), common mongoose (Herpestes
availability increases as dietary items previously                   edwardsii), small Indian mongoose (H. auropunctatus),
unattainable, are killed by apex predators and                       ruddy mongoose (H. smithi), common palm civet
interspecific competition could decline concomitantly.               (Paradoxurus hermaphroditus), small Indian civet
The diet of black backed jackal comprises high proportion            (Viverricula indica), ratel (Mellivora capensis) and prey
of small and large mammals, corroborating that both                  species like chital (Axis axis), sambar (Rusa unicolor),
scavenge and hunt competently [25]. Brown hyenas,                    nilgai (Boselaphus tragocamelus), common langur
on the other hand, are predominant scavengers [26].                  (Semnopithecus entellus), wild pig (Sus scrofa), porcupine
Thus, jackals may be excluded from carcass sites by                  (Hystrix indica), rufous-tailed Hare (Lepus nigricollis
brown hyenas, eventually forcing them to hunt more                   ruficaudatus) and Indian peafowl (Pavo cristatus)[30].
frequently [27].                                                     There are 32 villages within Sariska TR of which six are
     Both striped hyena (Hyaena hyaena) and golden                   located in the intensive study area of 160 km2. A large
jackal (Canis aureus) are found in semi-arid zone and                number of brahmini cattle (Bos indicus), buffaloes
tropical dry and moist deciduous forests of the country              (Bubalus bubalis), goats (Capra hircus) and sheep (Ovis
and their diet ranges from wild ungulates to domestic                aries) are kept by people living in villages.
livestock and rodents. Although numerous studies have
investigated their diet separately, only a few studies [23]          Scat Analysis: Estimation of food habits of golden jackal
have investigated the overlap and resource partitioning,             and striped hyena was done using scat analysis. Scats
which may vary with the presence or absence of apex                  were collected on transects, trails and roads whenever
predators like tiger (Panthera tigris) and leopard                   encountered within the intensive study area. Each
(Panthera pardus). In the present study, we assessed 1)              collected scat sample was labeled with the name of the
the relative contribution of different prey species to the           species, date and location. The collected scats were sun
diet of golden jackal and striped hyena and 2) the dietary           dried and later broken down and washed under running
overlap between these two sympatric species. In the                  water through a sieve. The scat contents were teased
present study we estimated the food habits of striped                apart and remain of different food items such as hair,
hyena (Hyaena hyaena) and golden jackal (Canis aureus)               feather, scales of reptiles, invertebrate remains and
and compared the dietary overlap between them in Sariska             vegetable matter (grass and fruit seeds) were separated.
Tiger Reserve, Western India from November 2010 to June              The frequency of occurrence of each food item was
2011.                                                                estimated through bootstrapping using program
                                                                     SIMSTAT [31]. This technique measures 95% confidence
            MATERIALS AND METHODS                                    limits of the proportion of each food item in the diet, the
                                                                     limits of which range measure the random sampling errors
Study Area: The study was conducted in Sariska Tiger                 [19]. Niche breadth of the two species was quantified
Reserve (Sariska TR) (27°05'-27°33' N; 76°15'-76°35' E)              using Levin’s measure [26] to measure the uniformity of
between November 2010 and June 2011. Sariska TR is                   resources being utilized by each species. The overlap in
situated in the Aravalli Hill Range in the semi-arid part of         diet of the two species was assessed using Pianka’s [32]
Western India [28]. The total area of the Tiger Reserve is           index.
881 km2, of which 273.8 km2 is notified National Park. The
vegetation of Sariska corresponds to Northern tropical                                       RESULTS
dry deciduous and Northern tropical thorn forests [29].
Open areas are covered with scrub forests dominated by                    Altogether 104 scats of golden jackal and 86 scats of
shrubs such as Zizyphus nummularia, Capparis sepiaria,               striped hyena were collected and analyzed. Hair samples
Capparis decidua, Adathoda vasica, Prosopis juliflora                from scats were identified from their histological structure
and Acacia sp. The valleys are dominated by Zizyphus                 using microscope. In total 12 food items were identified in
mauritiana mixed forest, gentle slopes are dominated by              golden Jackal scats and nine in striped hyena scats. The
Anogeissus pendula forest and the steep slopes are                   frequency of occurrence and percentage frequency of
occupied by Boswellia serrata forest. Other than golden              occurrence of each food item found in golden jackal and
jackal and striped hyena, the Park supports various                  striped hyena scats are given in table 1.

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Table 1:     Frequency of occurrence and percentage frequency occurrence of each food item found in golden jackal and striped hyena scats in Sariska Tiger
             Reserve, Western India between November 2010 and June 2011
                     Golden jackal                                                                            Striped hyena
                     --------------------------------------------------------------------------------------   ------------------------------------------------------------------------------------
Food item            Frequency of occurrence (±SD) Percentage frequency of occurrence                         Frequency of occurrence (±SD) Percentage frequency of occurrence
Scales of reptiles               4±1.9                                            1.80                                     -                                           -
Vegetation                       37.5±4.7                                         17.57                                    12.8±3.4                                    10.48
Fruit                             19.1±3.8                                        9.01                                     -                                           -
Bird                              15.5±3.6                                        7.21                                     2.3±1.5                                     1.90
Insect                           15.3±3.8                                         7.21                                     -                                           -
Rodent                           34±4.5                                           15.77                                    1.2±1.1                                     0.95
Hare                              8.9±2.9                                         4.05                                     1.2±1.1                                     0.95
Sambar                           11.3±3.1                                         5.41                                     20.9±4.4                                    17.14
Chital                            23.1±3.9                                        10.81                                    19.8±4.0                                    16.19
Nilgai                           8.7±2.8                                          4.05                                     30.2±4.8                                    24.76
Cattle                            32.9±4.5                                        15.32                                    30.2±4.8                                    24.76
Goat                             3.7±1.9                                          1.80                                     3.5±2.1                                     2.86

     Vegetative matter contributed maximum (17.57%) in                                            jackal predating on goats in the study area but cattle
golden jackal’s diet followed by rodents (15.77%), cattle                                         predation by them was not recorded. The result of scat
Bos indicus 15.32%), chital (10.81%), fruits (9.01%), birds                                       analysis supports the typical diet generalist of golden
(7.21%), sambar (5.41%), hare (4.05%), nilgai (4.05%), goat                                       jackal, feeding on a variety of food source available to
(1.80%) and reptile scales (1.80%). No attempt was made                                           them in the study area.
to identify the rodent and reptile species in golden                                                   Out of 86 striped hyena scats, three scats had three
jackal’s scat. Out of 104 golden jackal scats, one had six                                        food items; 40 scats had two food items and 43 scats had
food items, one had five food items, ten had four food                                            single food items. Nilgai and cattle (Bos indicus)
items, 23 had three food items, 34 had two food items and                                         contributed maximum (24.76%) in the diet of striped hyena
35 had single food item. Zizyphus mauritiana seeds were                                           followed by sambar (17.14%), chital (16.19%), vegetation
the only fruit species found in golden jackal’s diet.                                             (10.48%), goat (2.86%), unidentified bird (1.90%), hare
Beetles (Class: Insecta), centipedes (Class: Chilopoda),                                          (0.95%) and unidentified rodent (0.95%). The remains of
millipedes (Class: Diplopoda) in golden jackal’s diet were                                        larger ungulates such as sambar, nilgai and chital were
identified on the basis of locomotory organs, i.e. shiny                                          observed more in scat samples as compared to the
elytra of beetles and round or oval shields of centipedes/                                        remains of goat, bird, rodent and hare. The above
millipedes. Feathers of grey partridge (Francolinus                                               mentioned large ungulates are also the major food items
pondicerianus) and peafowl (Pavo cristatus) were                                                  of tiger and leopard in the study area [33, 34]. This may
identified in golden jackal’s diet. Narrow ventral scales                                         lead to the inference that the occurrence of larger
and translucent round scales were also identified to be of                                        ungulates in hyena diet is perhaps the result of
lizards (Calotes sp.) and skinks (family: Scincidae) but not                                      scavenging.
at species level.                                                                                      The estimated dietary overlap between striped hyena
     In some incidences (n=9) golden jackals were                                                 and golden jackal was 67% (Table 2). Niche breadth for
found to prey upon chital fawns in the study area and this                                        golden jackal was estimated as 0.69 and for striped hyena
may be attributed to higher occurrence of chital in their                                         it was 0.57 (Table 3). MacArthur and Levin’s [35] niche
scats. Golden Jackal was never found to prey on sambar                                            overlap was calculated as an overlap measure of species
or nilgai (adult or fawn) in the study period, but they                                           A on species B and vice-versa. The niche overlap of
scavenge on sambar and nilgai kills made by tiger or                                              golden jackal on striped hyena was 66% (0.66) i.e. out of
leopard in the study area, which is reflected in lower                                            total diet range of striped hyena, 66% of food items are
occurrence in their scats. Golden Jackal were also found                                          common with golden jackal and the niche overlap of
scavenging on cattle carcasses at the dump sites outside                                          striped hyena on golden jackal was 47% (0.47) i.e. out of
villages in the study area, which is attributed to higher                                         total diet range of golden Jackal, 47% of food items are
occurrence of cattle in their diet. There were records of                                         common with striped hyena.

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Table 2: Dietary overlap between golden jackal and striped hyena as shown by scat analysis in Sariska Tiger Reserve, Western India between November 2010
             and June 2011
                                 Percentage frequency of              Percentage frequency of
Food Item                        occurrence (striped hyena) (PiA)     occurrence (golden jackal) (PiB)   PiA*PiB (C)          PiA2 (D)              PiB2 (E)
Vegetation                               10.48                                 17.57                      184.04              109.75                308.62
Bird                                     1.90                                  7.21                       13.73               3.63                  51.94
Rodent                                   0.95                                  15.77                      15.02               0.91                  248.56
Hare                                     0.95                                  4.05                       3.86                0.91                  16.44
Sambar                                   17.14                                 5.41                       92.66               293.88                29.22
Chital                                   16.19                                 10.81                      175.03              262.13                116.87
Nilgai                                   24.76                                 4.05                       100.39              613.15                16.44
Cattle                                   24.76                                 15.32                      379.24              613.15                234.56
Goat                                     2.86                                  1.80                       5.15                8.16                  3.25
Scales of reptiles                       0.00                                  1.80                       0.00                0.00                  3.25
Fruit                                    0.00                                  9.01                       0.00                0.00                  81.16
                                                                                                          969.11(F)           1905.67 (G)           1110.30 (H)
                                                                                                                              43.65 (I)             33.32 (J)
                                                                                                                                                    0.67 (O)
PiA -% of food item i in the diet of striped hyena; PiB-% of food item i in the diet of golden jackal; C = PiA * PiB; D = PiA ; E = PiB ; F=
                                                                                                                              2           2
                                                                                                                                               C; G =   D and
H = E; I = G0.5; J = H0.5, O = F/ (I*J).

Table 3: Dietary niche breadth between Golden Jackal and Striped Hyena as                    broader niches would be more closely linked to
             shown by scat analysis in Sariska Tiger Reserve November 2010                   characteristics of favorable elements (cover or food) and
             to June 2011                                                                    the landscapes surrounding an element, because the
Variable                               Golden jackal1               Striped hyena2           combined effects of large size and a broad niche on the
n                                         12                           10                    species would impose fewer constraints on its element
B                                         8.604                        6.157                 and landscape use compared to smaller, more specialized
BA                                        0.691                        0.573                 species [36]. In Sariska TR, golden jackals are found to be
M MacArthur and Levins measure            0.663                        0.475                 confined in the valley areas where a range of smaller prey
                                                                                             species as well as fruiting trees (Zizyphus sp.) are
                                       DISCUSSION                                            available, while distribution of striped hyena was wider
                                                                                             including valley habitats and hilly areas [37]. In the
      In Sariska TR, the striped hyena occupies all the                                      present study, broader dietary niche was observed for
major habitat types, compared to golden jackal that is                                       golden jackal as compared to striped hyena, because
mainly distributed in the open scrub land and valley                                         golden jackal assumed to compensate their limited
habitat. There are predictions that body size and                                            distribution by utilizing broader spectrum of food.
behavioral flexibility would influence the response of                                            Many studies have explained dietary differences
predators and this response would be evidenced in                                            within the context of alternative feeding strategies (i.e.
individual use of habitat elements and in the spatial                                        generalist vs. specialist foraging) or intrinsic niche
distribution of predators [36]. Specifically, larger species                                 differentiation (i.e. disparity in habitat requirements or
(i.e. striped hyena) would be more equitably distributed                                     body size) [38-41]. However, not all species fall into such
among all spatial habitat elements compared to smaller                                       discrete categories [42]. Indeed, within a given
species (i.e. golden jackal), because large size would                                       community, the relative breadth of a carnivore’s diet (and
confer an increased ability to traverse the habitat matrix                                   by extension levels of interspecific dietary overlap) is best
with minimal risk of predation. Similarly, species with a                                    defined in relation to the feeding habits manifested by its
broader niche would be more equitably distributed among                                      tropical counterparts [43]. However, when prey
spatial habitat elements because broader diet breadth                                        abundance fluctuates dramatically over a period of time,
would allow a species to use the majority of spatial                                         even alleged specialist species may exhibit more generalist
elements and exploit a wider range of resources.                                             feeding patterns [44]. Moreover, a review of dietary
Additionally, presence of larger species and species with                                    breadth among herbivorous insects revealed that while

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generalization characterized the diet of many insect                 presence of apex predators like tiger and leopard. This
species, the dietary choices of individuals within                   suggests that niche partitioning in protected areas in the
particular populations were often specialized (according             presence of an apex predator results in a form of
to the traditional framework described above), suggesting            competitive exclusion of jackal at carcasses by hyena,
that in many cases dietary breadth may be a local (i.e.              resulting in a greater level of hunting by the jackals,
system-specific) phenomenon rather than an immutable                 connatural to the study in Africa [23].
species characteristic [42]. The considerable overlap                     Review of the studies showed marked interspecific
(67%) characterizing the diets of striped hyena and golden           difference in plant-matter consumption by golden jackal.
jackal, may be attributed to mutual dependence on                    Whereas, plants did not play an important role in the
ungulates, which intern indicates that resource                      feeding of golden jackals in Hungary [20], compared with
competition between these two species may be high.                   the feeding of jackals in southern areas of Asia [48]. In the
However, striped hyena relied more on large ungulates,               present study vegetative matter i.e. grass and Zizyphus
whereas golden jackal preyed largely on small ungulates              fruits comprised a higher proportion 26.58% in golden
and rodents; these dietary specializations may have                  jackal’s diet.
alleviated overall levels of competition between two                      Seasonal dominance of small mammals was
species. This pattern of exclusive use of a particular prey          reported in golden jackal’s diet [49-51]. In the present
items, despite the overall dietary similarity, characterizes         study no attempt was made to quantify the seasonal diet
the relationships between sympatric coyotes Canis                    of golden jackal. Study by Lanszki et al. [20] on feeding
latrans and gray foxes Vulpes velox [45] and coyotes and             habits and tropical niche overlap between golden jackal
red foxes Vulpes vulpes [46].                                        (Canis aureus) and red fox (Vulpes vulpes) in the
     Depending on food availability, golden jackals may              Pannonia eco-region, Hungary revealed that functional
be solitary hunters, co-operating in pairs or hunting as             response of the golden jackal to a limited but favored
groups (mainly when the parents teach the offspring to               food item was more rapid than that of the red fox.
hunt). The differences in large mammal and small mammal              The golden jackal shifted from small prey to other food
proportions between the diets resulted in a significant              items earlier when the availability of small mammals
difference between golden jackal and striped hyena diets.            declined and also returned to rodent hunting earlier than
However, at 67%, the niche overlap between the species               did the red fox. Long term ecological studies including
was still high. Result of the present study is comparable            seasonal food habits on golden jackal and striped hyena
with the studies by Nowell and Jackson [46] and Walton               are much desired across different landscapes in the
and Joly [25] on black-backed jackal (Canis mesomelas) in            country.
Africa. Studies by Owens and Owens [26, 27] on brown
hyenas showed that they are predominantly scavengers                                ACKNOWLEDGEMENTS
and black-backed jackals are generally excluded from
carcass sites by the much larger brown hyenas, forcing                    We thank Rajasthan Forest Department for
them to hunt more frequently. The similar case may likely            facilitation of this work in Sariska, as a part of ‘Ecology of
happen in Sariska TR where striped hyenas occur in high              Leopard’ research project conducted by the Wildlife
density i.e. 24.5 individual/ 100 sq. km [47]. Study by              Institute of India. We thank Director and Dean, WII, for
Merwe et al. [23] on assessment of diet overlap between              their encouragement and support provided for the study.
brown hyena and black-backed jackal suggested that in                We thank our field assistants Jairam, Omprakash and
black backed jackal’s diet, more large mammals were                  Ramesh for their assistance in field.
consumed outside protected areas whereas more small
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