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Acta Theriologica

ISSN 0001-7051

Acta Theriol
DOI 10.1007/s13364-014-0187-8
Population density of striped hyenas
in relation to habitat in a semi-arid
landscape, western India
Randeep Singh, Qamar Qureshi,
Kalyanasundaram Sankar, Paul
R.Krausman, Surendra Prakash Goyal &
Kerry L.Nicholson
1 3
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ORIGINAL PAPER
Population density of striped hyenas in relation to habitat
in a semi-arid landscape, western India
Randeep Singh & Qamar Qureshi &
Kalyanasundaram Sankar & Paul R. Krausman &
Surendra Prakash Goyal & Kerry L. Nicholson
Received: 17 January 2014 / Accepted: 7 April 2014
#Mammal Research Institute, Polish Academy of Sciences, Biaowiea, Poland 2014
Abstract We used camera trapping in conjunction with a
spatial explicit capturerecapture model to estimate striped
hyena (Hyaena hyaena) density and occupancy models to
investigate factors affecting striped hyena detection probabili-
ties in Ranthambhore Tiger Reserve (RTR), Rajasthan, India.
A sampling effort of 4,450 trap days/nights over 75 days yield
68 photo captures of 21 unique striped hyenas (based on
individual markings and visual identification); the estimated
striped hyena density was 5.491.27 individuals/100 km
2
.
Results of our occupancy model suggested that a rugged
terrain is an important factor that influences striped hyena
detection probability. Correlation with striped hyena detection
with human settlement provides evidence of social tolerance of
striped hyena towards humans, and more occurrence of re-
sources allowed coexistence of hyena in a human-dominated
landscape. This elasticity (inhabited areas close to humans)
demonstrated by striped hyenas is an exception among carni-
vore communities living in this semi-arid habitat.
Keywords Abundance
.
Cameratrap
.
Livestock
.
Population
estimation
.
Ranthambhore
.
Scavenging
Introduction
Striped hyenas (Hyaena hyaena) are widely distributed in an
area extending from east and northeast Africa, through the
Middle East, the Caucasus region, and central Asia, to the
Indian subcontinent (Mills and Hofer 1998). In India, the
striped hyena occupies habitats located in arid and semi-arid
regions to the wet zone of the southwestern coast (Rieger
1979), except the moist forests of the northeastern region
(Prater 1971). This nocturnal carnivore is a long-ranging and
solitary forager, living in small groups with unknown compo-
sitions (Mills and Hofer 1998). It feeds on a variety of verte-
brates, invertebrates, and vegetation and plays an important
biological role in consuming carrion (Wagner 2006; Wagner
et al. 2008). Striped hyenas often live close to human habita-
tion and can survive on organic matter and recycling dead
organic components generated by humans (Kruuk 1976;
Mendelssohn and Yom-Tov 1999). Despite the important role
of the striped hyenas in ecosystems, disproportionately little
effort has been expended in studying them. Habitat alteration
across their range is a likely cause of declining populations
(Ripple et al. 2014). The International Union for the
Conservation of Nature (IUCN) has classified the striped
hyenas as near threatened (Arumugam et al. 2008).
Evaluation of the population status and their habitat attributes
at the local and regional level is crucial for planning a conser-
vation strategy to maintain viable populations. Unfortunately,
few studies have looked into the ecology of striped hyenas
(Singh et al. 2010).
Communicated by: Matthew W. Hayward
R. Singh (*)
:
Q. Qureshi
:
K. Sankar
:
S. P. Goyal
Wildlife Institute of India, Post Box # 18, Dehradun,
Uttarakhand 248001, India
e-mail: randeep04@rediffmail.com
Q. Qureshi
e-mail: qnq@wii.gov.in
K. Sankar
e-mail: sankar@wii.gov.in
S. P. Goyal
e-mail: goyalsp@wii.gov.in
P. R. Krausman
Boone and Crockett Programin Wildlife Conservation, University of
Montana, Missoula, MT 59812, USA
e-mail: paul.krausman@umontana.edu
K. L. Nicholson
Grims Wildlife Research Station, Department of Ecology, SLU,
SE-730 91 Riddarhyttan, Sweden
e-mail: kernicholson@yahoo.com
Acta Theriol
DOI 10.1007/s13364-014-0187-8
Author's personal copy
In recent years, camera trapping has been widely used to
study small to large vertebrates (Cutler and Swann 1999) and
to gather quantitative information (OConnell et al. 2011), and
it has been an efficient way to understand the biology of
species and habitat relationships (Carter et al. 2012; Schuette
et al. 2013). Striped hyenas, in particular, can be identified
individually from their pelt patterns, and camera traps can be
used to estimate their population density (Gupta et al. 2009;
Harihar et al. 2010).
In the present study, our research objectives were to esti-
mate striped hyena population density using capturerecap-
ture methods and to investigate the influence of environmental
factors and human associated variables (i.e., terrain rugged-
ness, forest cover, and distance from human settlements) on
striped hyena distribution in Ranthambhore Tiger Reserve
(RTR), Rajasthan.
Materials and methods
Study area
The RTR(25 54 to 26 12 N, 76 22 to 76 39 E) is situated
in the semi-arid part of Rajasthan, western India (Fig. 1) and is
a transition zone between the desert and peninsular India
(Rodgers and Panwar 1988), dominated by northern tropical,
dry, deciduous, and thorny forests (Champion and Seth 1968).
The RTR is surrounded by 332 villages within 5 km of the
reserve boundary with more than 150,000 people and live-
stock (Bagchi et al. 2003), and humans are primarily engaged
in cutting grass, lopping trees, and grazing livestock activity.
This area receives 800 mm of annual precipitation (interpola-
tion from Hijmans et al. 2005, http://www.diva-gis.org/
climate), and temperatures range from 2 C in January to a
maximum of 47 C in May (Singh et al. 2014). The RTR is
characterized by a subtropical dry climate with four distinct
seasons: summer (MarchJune), monsoon (JulyAugust),
post - monsoon ( Sept emberOct ober ) , and wi nt er
(NovemberFebruary). The elevation gradient is ranged from
200 to 500 m of RTR, and 76 % of the area is categorized as
undulating (Singh 2011). Besides the hyena, other large pred-
ators in RTR include tigers (Panthera tigris), leopards
(Panthera pardus), and sloth bears (Melursus ursinus). The
RTR also supports five species of wild ungulates: chital (Axis
axis), sambar (Rusa unicolor), nilgai (Boselaphus
tragocamelus), chinkara (Gazella gazelle), and wild pig (Sus
scrofa).
Sampling design
Our research was part of a long-term monitoring program in
RTR (Singh et al. 2013a, b). We designed a camera-trapping
grid system based on distribution maps produced using
indirect sign (i.e., tracks, scat) surveys. Cameras were de-
signed and deployed close to the trails, which were most
frequently used by carnivores to maximize capture and recap-
ture probabilities of the individuals (Wilson and Delahay
2001). The sampling area was divided into 11-km grid,
and cameras were systematically placed across the sampling
grid. We used 60 remotely triggered cameras at one time; 13
were active infrared systems (TrailMaster TM1550;
Goodson and Associates, Inc., Lenexa, KS, USA), and 47
were digital passive infrared systems. The passive infrared
systems were comprised of 17 Wildview systems (Wildview,
Grand Prairie, TX, USA), 9 Stealth Cam units (Stealth Cam,
Grand Prairie, TX, USA), and 21 pair of Moultrie systems
(Moultrie Game Feeders, Alabaster, AL, USA). Each
TrailMaster unit was connected to a fully automatic 35-mm
camera (Canon A1 Mini DX; Canon USA, Inc., Lake Success,
New York) with autoflashes. Due to variable detection zones
of different cameras, we placed the cameras so they would all
get sufficient time to detect an animal and take full-frame
pictures of the subject and three pictures continuously at short
intervals. Both active and passive cameras had white flashes
that could reach up to 3035 m.
Each sampling grid consisted of single cameras positioned
67 mapart froma road to photograph either flanks of passing
hyenas (Karanth and Nichols 1998). The study area was
divided into three consecutive spatially separated blocks and
was systematically sampled in a phased manner under survey
design 4 (Karanth and Nichols 2002; Fig. 1). During the
sampling period, two camera units were stolen in the third
block, so we had 58 sampling stations in that block. The
sampling was conducted from 5 December 2010 to 20
February 2011, with a 25-day sampling period in each block.
After each trapping session (25 days), we took 23-day breaks
to download the photographs from the digital cameras and
placed the cameras in the next block. The camera traps were
functional for 24 h/day/sampling session. Because of the
network of roads, all trapping stations could be checked in
alternate days during sampling periods. The minimum convex
polygon (MCP; Fig. 1) of all the camera-trapping sites cov-
ered an area of 233 km
2
.
Striped hyena density
Individual striped hyenas were identified from photo-
graphs obtained using the camera traps by visually exam-
ining the markings on the pelage of the hind limbs, fore-
limbs, and forequarters (Gupta et al. 2009). Photographs in
which hyenas were individually identified were assigned
with unique identification numbers, and the specific trap
location, sampling period, date, and time of capture were
recorded. We constructed a capture history of striped hy-
enas in spatial explicit capturerecapture (SECR) data for-
mat for analysis that considered a continuous 75-day
Acta Theriol
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sampling occasion (1- to 25-day sessions in block A, 26- to
50-day sessions in block B, and 51- to 75-day sessions in
block C) (OBrien and Kinnaird 2011).
We followed the SECR approach to obtain maximum
likelihood density estimates for striped hyenas using the
camera-trapping data. The likelihood SECR models were
implemented in the R package SECR and DENSITY 5.0
(Efford et al. 2009; Efford 2010; www.otago.ac.nz/density).
The detection probability of each individual was modeled
using the spatial detection function (Efford 2004) and was
explained by two parameters [one-night detection probability
at the center of an individuals home range (g
0
) and a function
of the scale of animal movements (); Efford 2004]. We used
a half-normal detection function because it seemed to be
appropriate for marked recapture data from large carnivores.
We evaluated the log-likelihood function by integrating the
Poisson distribution of the home range centers by adding a
buffer of 10,000 m around the trapping grids (this distance
was chosen to ensure that no individual outside of the buffered
regions had any probability of being photographed by the
camera trap during the survey; Zimmermann et al. 2013).
Environmental factors
We used camera-trapping data within likelihood-based meth-
od (Mackenzie et al. 2002, 2005) to evaluate the factors
influencing the spatial variability of hyena distribution and
their association with habitat attributes. Hyena detection his-
tories (H) were constructed for 178 camera-trapping locations
over the 25 sampling occasions using a standard X matrix
format (Otis et al. 1978). Thus, for each site and occasion,
1 indicated the detection (photograph) of a hyena, while 0
indicated non-detection (Otis et al. 1978). The striped hyenas
detection histories were then used to produce probabilities
(i.e., animal being detected at a specific site in response to
habitat attributes; Carter et al. 2012). We examined the
habitat-attributed terrain features (i.e., terrain ruggedness is
closely associated with availability of den sites), distance to
Fig. 1 Locations of camera traps used to photograph striped hyenas within the core area of Ranthambhore Tiger Reserve, western India, December 2010
to February 2011
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human settlements (i.e., availability of livestock carcasses
near settlements), and forest vegetation (close canopy forest
and scrub vegetation may provide concealed daytime resting
sites for striped hyenas).
The terrain ruggedness was calculated using the extension
Topographical Position Index (TPI) (Jenness 2006) in the
ArcView v.3.2 GIS software package (ESRI, Redlands, CA,
USA) from elevation data at a 30-m resolution derived from
the ASTER Global Digital Elevation Model (http://www.
jspacesystems.or.jp/ersdac/GDEM/E/). The terrain of RTR
was categorized as rugged terrain and flat areas including
valley slopes. Human settlements were extracted from
1:50,000 scale maps produced by the Survey of India and
converted into Euclidian distance maps in ArcView v.3.2.
Vegetation characteristics (close canopy, scrubland, and
grassland) were extracted from a forest cover map available
from RTR (Singh 2011). We carried out the estimates using
the PRESENCE 3.0 software package (Proteus Wildlife
Research Consultants, New Zealand; http://www.proteus.co.
nz/) with the single-species, single-season option (Mackenzie
et al. 2006) with site-specific covariates (Donovan and Hines
2007). To identify the important habitat attributes, we used
detection probability as a constant (.) and then allowed the
habitat attributes () to vary with covariates. We used the delta
Akaike information criterion (AIC
c
) value and Akaike
weights (w
i
) to rank candidate models in order to identify
the most parsimonious model (Burnham and Anderson
2002). We discussed the influence of covariates from models
within two units of delta AIC
c
, which have equal support.
Results
Striped hyenas density
The sampling effort consisted of 4,450 active trap days be-
tween December 2010 and February 2011 based on 178 active
trapping stations. Striped hyenas were photographed at 32.6 %
of the camera-trapping sites. We obtained 157 photo captures
of striped hyenas, of which 118 were suitable for individual
identification (68 left flanks and 50 right flanks). The left flank
was used for further analysis as they had a larger number of
recognizable individuals and equal detectability. The 21
unique adult striped hyena individuals were stabilized at the
16th day of sampling with 68 photo captures (Fig. 2). The g
0
at the center of an individuals home range was estimated at
0.003 (SE=0.006), and the scale of animal movements from
the center of the home range () was 2.1 (SE=0.2)km. The
density of hyenas was estimated to be 5.49 (SE=1.27)
individuals/100 km
2
.
Habitat factors influencing detection probability
The distance to human settlements and terrain ruggedness influ-
enced hyena detection [(distance to human settlements+terrain
0
5
10
15
20
25
0
10
20
30
40
50
60
70
80
1 4 7 10 13 16 19 22 25
U
n
i
q
u
e

i
n
d
i
v
i
d
u
a
l
s
P
h
o
t
o
-
c
a
p
t
u
r
e
s
Sampling occassions
Photo-captures Unique striped hyenas
Fig. 2 Cumulative number of unique striped hyena and camera-trapped photo captures in Ranthambhore Tiger Reserve, western India, December 2010
to February 2011
Table 1 Estimated habitat parameters influencing striped hyena in
Ranthambhore Tiger Reserve, western India, December 2010 to February
2011
Model AIC
c

i
Model
likelihood
nPars
(distance to human settlement+
slope/elevation) (.)
0.00 0.31 1 2
(slope/elevation+close
canopy forest) (.)
1.12 0.19 0.7 2
(.), (.) 2.48 0.09 0.6 2
(.), (slope/elevation+close
canopy forest+scrubland)
6.59 0.007 0.04 3
is the probability of a site used by striped hyena, and is the probability
of detecting striped hyenas in the ith survey; change in Akaike informa-
tion criterion is the difference in AIC
c
values between each model with
the lowest AIC
c
model
AIC
c
change in Akaike information criterion,
i
AIC
c
model weight,
nPars number of parameters in the model
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rugged) and (terrain rugged+close canopy forest (.))]
(Table 1). The photo capture rate of striped hyenas was higher
(71.2 %)<3 km from human settlements (Fig. 3). Striped hyena
photo captures were recorded at 61.0 % of the time in rugged
terrain, and they were captured at 39.0 %of the time in flat areas
including valleys.
Discussion
Estimates of striped hyena densities in human-dominated
landscape range from 3.67 to 5.03 individuals/100 km
2
(Singh et al. 2010; Athreya et al. 2013), whereas estimates
for protected areas [i.e., Rajaji National Park (Rajaji), RTR,
Gir National Park (Gir), and Sariska Tiger Reserve (Sariska)]
were 3.91, 5.49, 6.50, and 15.1 individuals/100 km
2
, respec-
tively, in India (Alam et al. 2009; Gupta et al. 2009; Harihar
et al. 2010). All of these protected areas are dominated by
humans and have established tiger populations except Gir,
where lions (Panthera leo) exist. Estimated tiger densities in
Rajaji and RTR were 2.25 and 8.09 individuals/100 km
2
,
respectively (Jhala et al. 2011). In Sariska, three tigers were
reintroduced during the time of sampling (Sankar et al. 2010).
The Sariska had the highest wild (105 animals/km
2
) and
domestic (222 animals/km
2
) prey densities (Avinandan et al.
2008; Sankar et al. 2009). It may be the underlying reasons for
the high density of hyenas in Sariska. Being a specialized
scavenger, it was implicit that the striped hyenas distribution
and density may be related to livestock abundance (i.e., car-
casses) (Singh et al. 2010).
Ranthambhore is surrounded by 332 villages within 5 km
of the reserve boundary with 143,468 head of livestock (http://
projecttiger.nic.in/ranthambhore.htm). Every month, an
average of 2030 domestic livestock die due to various
causes (i.e., starvation, inadequate veterinary care,
depredation by tiger or leopard, disease; personal
observation) were not buried or consumed by local people
due to religious views (Chhangani 2009), which provides food
resources (i.e., carcasses) for striped hyenas. Availability of
livestock carcasses may be a reason for higher hyena occur-
rences in the vicinity of human settlements. As it is obvious,
an abundance of large predators may be related to availability
of food resources (Karanth and Sunquist 2000; Karanth et al.
2004). Our results are, in general, agreement with this pattern;
with a decreasing distance of camera traps from villages, the
rate of capture of hyenas increases, because some striped
hyenas lead to optimally choosing disturbed/fringe habitat
for easy access to food resources. So, the striped hyena density
may be expected to be higher in those areas having a large
number of livestock.
The terrain ruggedness provides optimal refuges and den-
ning sites (Kruuk 1976; Rieger 1979; Singh et al. 2010) that
are relatively free of anthropogenic activity (i.e., not used by
livestock, humans, or guard dogs). The extend of our study
area comes under 76 % of rugged terrain categories, and
striped hyenas were photo captured during the study period
61.0 %of the time in rugged terrain, which may provide better
forested habitat and also suggests that disturbance-free habi-
tats may provide optimal conditions for breeding and raising
pups. Additionally, close canopy forest and scrub vege-
tation are important factors, which are associated with
detection of striped hyenas. The presence of the hyena
may have been associated with such habitats because
they provide concealed daytime resting sites. During
field surveys, we flushed hyenas during the daytime in
dense shrub. These observations support our findings
regarding the positive effect of dense vegetation as
hiding or resting places during the daytime for hyenas.
The study shows that the ecology of the striped hyena
is closely linked with human occupation in such a case.
Conservation and management of such a species that
exists beyond protected areas in the long run will de-
pend on species associations with human-dominated
landscapes.
Acknowledgments We are thankful to the Director and Dean of the
Wildlife Institute of India for their continuous support. We are also
thankful to the Wildlife Institute of India, Dehradun (Ministry of Envi-
ronment and Forests, Government of India) for providing financial sup-
port for this work. We offer our humble thanks to the Rajasthan Forest
Department and the park officials and field staff at RTR for permissions
and for facilitating this work. We thank, in particular, our field assistants,
M. S. Gurjar and S. Sharma, for providing valuable and continuous
support during the field work.
0
5
10
15
20
25
30
35
<1000 1000-2000 2000-3000 3000-4000 4000-5000 >5000
P
e
r
c
e
n
t
a
g
e
Distance from human settlements in meters
Percent camera trap Percent photo-capture of striped hyena
Fig. 3 Percentage of photo
captures of striped hyenas and
camera traps and distance from
human settlements (m) in
Ranthambhore Tiger Reserve,
western India, December 2010 to
February 2011
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