Assessing changes in distribution of the Endangered snow leopard Panthera uncia and its wild prey over 2 decades in the Indian Himalaya through ...
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Assessing changes in distribution of the Endangered
snow leopard Panthera uncia and its wild prey over 2
decades in the Indian Himalaya through interview-
based occupancy surveys
ABHISHEK GHOSHAL, YASH VEER BHATNAGAR, BIVASH PANDAV
KOUSTUBH SHARMA, CHARUDUTT MISHRA
R . R A G H U N A T H and K U L B H U S H A N S I N G H R . S U R Y A W A N S H I
Abstract Understanding species distributions, patterns of have been used to estimate national and global snow leopard
change and threats can form the basis for assessing the con- ranges.
servation status of elusive species that are difficult to survey.
Keywords Asiatic ibex, blue sheep, carnivore, occupancy,
The snow leopard Panthera uncia is the top predator of the
snow leopard, survey, threat, ungulate
Central and South Asian mountains. Knowledge of the dis-
tribution and status of this elusive felid and its wild prey is Supplementary material for this article can be found at
limited. Using recall-based key-informant interviews we es- https://doi.org/./S
timated site use by snow leopards and their primary wild
prey, blue sheep Pseudois nayaur and Asiatic ibex Capra
sibirica, across two time periods (past: –; recent:
–) in the state of Himachal Pradesh, India. We Introduction
also conducted a threat assessment for the recent period.
Probability of site use was similar across the two time peri-
ods for snow leopards, blue sheep and ibex, whereas for wild
T he distribution and habitats of some species have been
negatively affected by human activities and develop-
mental pressures globally (Sanderson et al., ). Large
prey (blue sheep and ibex combined) overall there was an mammals, especially apex carnivores, are often considered
% contraction. Although our surveys were conducted in to be umbrella or flagship species for ecosystem conserva-
areas within the presumed distribution range of the snow tion (Simberloff, ). Large home ranges and habitat
leopard, we found snow leopards were using only % of specificity tend to make large mammals vulnerable to
the area (, km). Blue sheep and ibex had distinct dis- range contraction and extinction (Ceballos et al., ;
tribution ranges. Snow leopards and their wild prey were not Michalski & Peres, ). Understanding threats to large
restricted to protected areas, which encompassed only % mammals and determining their occurrence and changes
of their distribution within the study area. Migratory live- in distribution are thus important for conservation planning
stock grazing was pervasive across ibex distribution range at large spatial scales (MacKenzie et al., ; Karanth et al.,
and was the most widespread and serious conservation ; Taubmann et al., ).
threat. Depredation by free-ranging dogs, and illegal hunt- Since the s India has experienced a % increase in its
ing and wildlife trade were the other severe threats. Our human population alongside rapid economic growth, which
results underscore the importance of community-based, land- has negatively affected mammal species and their habitats
scape-scale conservation approaches and caution against reli- (Madhusudan & Mishra, ; Das et al., ; Singh &
ance on geophysical and opinion-based distribution maps that Bagchi, ). Studies at national and regional levels have re-
ported declines in distribution ranges of some common large
mammalian species (Karanth et al., ; Pillay et al., ). In
comparison to the large mammals of the Indian peninsula, the
ABHISHEK GHOSHAL* (Corresponding author), YASH VEER BHATNAGAR†, distribution and conservation status of the high-altitude large
KOUSTUBH SHARMA†, CHARUDUTT MISHRA†, R. RAGHUNATH and
KULBHUSHANSINGH R. SURYAWANSHI† Nature Conservation Foundation, 3076/5,
mammalian assemblage have received little scientific attention
IV Cross, Gokulam Park, Mysore, Karnataka 570002, India (Gaston et al., ; Mishra et al., ). Reliable baseline in-
E-mail abhishek@ncf-india.org formation on distribution range is lacking even for charismatic
BIVASH PANDAV Wildlife Institute of India, Dehradun, Uttarakhand, India and threatened species such as the snow leopard Panthera
*Also at: Snow Leopard Trust, Seattle, Washington, USA; Wildlife Institute of uncia and its primary prey, blue sheep Pseudois nayaur and
India, Dehradun, Uttarakhand, India; and Saurashtra University, Rajkot, Asiatic ibex Capra sibirica, in India (Lovari et al., ;
Gujarat, India
†Also at: Snow Leopard Trust, Seattle, Washington, USA Lyngdoh et al., ; Snow Leopard Network, ).
Received February . Revision requested April . The snow leopard, which is categorized as Endangered
Accepted June . First published online October . on the IUCN Red List (Jackson et al., ), is the top
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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. https://doi.org/10.1017/S0030605317001107Snow leopards in the Indian Himalaya 621
predator and flagship species for conservation of the Indian steep cliffs and rocky outcrops. The terrain is rugged, with
Himalaya (Snow Leopard Network, ; Bhatnagar et al., cliffs and sharp ridgelines interspersed with alpine and sub-
). The Indian Greater and Trans-Himalaya form the alpine tracts in the Greater Himalaya. The south-east and
southern limit of the global snow leopard distribution north-west extremes of our study area are characterized by a
range and constitute c. % of the estimated global snow leop- relatively moist climate and tracts of forest. The entire region is
ard range (Snow Leopard Working Secretariat, ). characterized by Palaearctic conditions, with annual tempera-
Amongst the five Indian states where snow leopards occur, ture ranging from c. −°C in winter to c. °C in summer.
Himachal Pradesh has c. one-fifth of the potential snow leop- The region is drained by three major rivers, the Sutlej in
ard habitat (,–, m altitude; Project Snow Leopard, Kinnaur District, the Spiti in Spiti subdivision and part of nor-
; Snow Leopard Network, ), corresponding to thern Kinnaur, and the Chandra-Bhaga or Chenab in Lahaul
c. , km. Within Himachal Pradesh a large, contiguous and Pangi subdivisions.
patch of potential snow leopard habitat is constituted by the Given the difficult terrain, limited plant productivity and
four regions of Kinnaur (, km at ,–, m altitude), extreme climate, the local agro-pastoralist communities
Lahaul (, km), Spiti (, km) and Pangi (, km). inhabit the landscape at low densities (one individual
These regions (, km) constitute % of the potential per km in Lahaul and Spiti; per km in Kinnaur; per
snow leopard habitat in Himachal Pradesh. Since the s km in Pangi). Kinnaur, Lahaul and Pangi are inhabited pri-
considerable socio-economic changes have occurred in these marily by followers of the Hindu religion, with Buddhists in-
four regions, including improvements in connectivity and the habiting only the uppermost villages, whereas Spiti is
advent and spread of cash crops (e.g. green peas in Spiti and inhabited primarily by Buddhists. Their traditional depend-
Lahaul, and apples in Kinnaur and Spiti) (Mishra, ; Rana ence on natural resources for food (including wildlife), fuel,
et al., ; Basannagari & Kala, ). Increased road connect- livestock grazing (sheep, goats, cattle, yaks, yak–cattle hy-
ivity along international borders has led to a rise in infrastruc- brids), fodder, construction material and medicinal plants
ture projects and unregulated tourism, especially since , is widespread and varies across the landscape (Mishra,
when Kinnaur and Spiti were opened to Indian and inter- ). The regional economy has shifted from a subsistence
national tourists (Snow Leopard Network, ). and barter-based system to being cash-based and market-
We conducted a threat assessment and evaluated changes driven (Mishra, ; Bhatnagar & Singh, ). The advent
in the distribution of the snow leopard, blue sheep and ibex in of the green pea Pisum sativum as a cash crop from on-
these regions over a period of decades. We used multi- wards and the opening up of Kinnaur District and Spiti sub-
season occupancy modelling (MacKenzie et al., ) on division to tourists in have accelerated these changes.
detection/non-detection data collected by interviewing Development and agricultural intensification are providing
local people and corrected for imperfect detection across job opportunities and attracting non-native labourers to the
two time periods (Karanth et al., ; Pillay et al., ; landscape. Additionally, migratory herders from Shimla,
Taubmann et al., ), past (–) and recent (– Kangra and Chamba districts graze livestock in the high-
). The method facilitated analysis of data from past and altitude pastures across most of the study area during sum-
recent time periods based on recall of interviewees. Multiple mer (June–August).
respondents from a sampling unit were treated as replicate Carnivores in the study area include the snow leopard,
surveys for the primary sampling seasons, thereby accounting the wolf Canis lupus and the brown bear Ursus arctos at
for imperfect detection. Thus, sampling difficult-to-detect . , m altitude, and the common leopard Panthera par-
species across a large area and multiple time periods is pos- dus and Himalayan black bear Ursus thibetanus at , , m
sible using this method. We present distribution maps of the altitude (Jayapal & Ramesh, ; Mishra et al., ). Among
snow leopard, blue sheep and ibex for the two time periods, wild herbivores, the Himalayan tahr Hemitragus jemlahicus,
using appropriate covariates. We discuss the implications of Himalayan serow Capricornis thar, musk deer Moschus
our findings in the context of snow leopard conservation pro- leucogaster and Himalayan goral Naemorhedus goral occur
grammes at state, national and global levels. in the Greater Himalaya, and blue sheep and Asiatic ibex
occur in both the Greater and Trans-Himalaya.
Study area
The study area comprised , km at ,–, m Methods
altitude (potential snow leopard habitat, Snow Leopard
Network, ) covering the contiguous potential snow Survey design and data collection
leopard habitat across the Greater and Trans-Himalaya
Mountains in Himachal Pradesh (Fig. ). The vegetation in We used key-informant interviews with a structured ques-
the Trans-Himalaya is largely dry alpine steppe (Champion tionnaire to record occurrences of snow leopards and
& Seth, ), with gentle-rolling uplands interspersed with their prey for two time periods: – and –
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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. https://doi.org/10.1017/S0030605317001107622 A. Ghoshal et al.
FIG. 1 The locations of Kinnaur, Lahaul
and Spiti Districts and Pangi sub-division
of Chamba District, in Himachal Pradesh,
India. The shaded region denotes ,–
, m altitude, the potential snow
leopard Panthera uncia habitat in
Himachal Pradesh.
(following Pillay et al., and Taubmann et al., ). The study species among images of numerous other carnivore
survey was carried out during the summer season (June– and mountain ungulate species, such as the Tibetan wolf
September) of . The study area was divided into Canis lupus filchneri, brown bear, common leopard,
grid cells (sites) of × km (area km) each, the area Himalayan black bear, Himalayan tahr, Himalayan serow,
being similar to the estimated mean home-range size of the musk deer, Himalayan goral, Kashmir stag or hangul
snow leopard (Johansson et al., ). Cervus elaphus hanglu, markhor Capra falconeri, kiang
The questionnaire (Supplementary Material S) con- Equus kiang, Tibetan gazelle Procapra picticaudata, Tibetan
tained two sections. The first dealt with information related argali Ovis ammon hodgsoni, wild yak Bos mutus and
to the attributes of the respondents that could affect reliable Ladakh urial Ovis vignei vignei. Respondents who failed to
detection of the study species (Table ), and the second con- identify study species from the images were excluded from
tained questions regarding detections of the study species the analyses, as were those who were unable to provide reli-
during the two time periods, including the location, year able information on location of species detection. The iden-
and approximate month of each detection. tity of the respondents has been kept confidential.
Upon entering a village we first created a focus group of We identified conservation threats to the snow leopard
two or three people with thorough knowledge of the site or and its wild prey in Kinnaur, Lahaul, Spiti and Pangi based
grid cell. With their help we prepared resource maps on discussion with the focus groups and personal observa-
(Suryawanshi et al., ), with approximate locations of set- tions. The threats were ranked based on area of spread, inten-
tlements, agricultural areas, pastures, seasonal use of pastures, sity and urgency, following Margoluis & Salafsky (). We
number of livestock and areas used for biomass extraction. identified fourteen threats in four broad categories, namely,
This information was transferred to printed maps of the livestock–wild prey interactions, people–wildlife interactions,
area. During this exercise we did not record species detec- developmental activities and other human disturbances. For a
tion/non-detection, as the reports of one person could be in- particular threat in a region, a score of – was assigned, with
fluenced by another in such a group. These respondents and indicating low and severe. Thus, total ranking of a threat
others were later interviewed individually by AG, specifically (sum of scores for area, intensity and urgency) in a region
to gather data on detection/non-detection of snow leopards could be –. We considered a score of – to indicate
and their prey. The resource mapping facilitated the plotting low threat, – medium threat, and – severe threat.
of detection locations on a map. The respondents included
herders, active and former hunters, cattle herders, medicinal Data analyses
plant collectors, forest department officials, staff and contrac-
tual workers, military personnel, tourist guides, porters and To define the past time period we chose two landmark
photographers. Key informants were shown images of events: the introduction of the green-pea crop in Lahaul &
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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TABLE 1 Site and survey covariates used in occupancy analyses to assess changes in distribution of the snow leopard Panthera uncia and
wild prey (blue sheep Pseudois nayaur and Asiatic ibex Capra sibirica) between two time periods (– and – in the Greater
and Trans-Himalaya of Himachal Pradesh, India (Fig. ), with description, variable type, and expected relationship with site use and de-
tection probability.
Variable Expected relationship with site use and detection
Description type probability
Site covariates
Altitude Mean altitude Continuous Positive correlation with site use by study species
up to c. 5,200 m altitude
Ruggedness Proportion of rugged area, using terrain Continuous Positive correlation with site use by study species
ruggedness index
Normalized difference Proportion of vegetated area Continuous Positive correlation with site use by blue sheep &
vegetation index ibex
Migratory livestock Presence or absence of migratory livestock Binary Negative correlation with site use by blue sheep
grazing grazing and ibex
Livestock population Number of sheep/goats Continuous Negative correlation with site use by blue sheep
and ibex
Survey covariates
Active Whether respondent was active or not during Binary Active respondents are more likely to encounter
the primary survey periods study species
Time spent Number of months in which the respondent Continuous Respondents spending more time in area of
was active in the area of knowledge knowledge are more likely to detect study species
Familiarity Number of years for which the respondent Continuous Respondents with higher familiarity are more
was familiar with the area of knowledge likely to detect study species
Age Age of respondent in years Continuous Mixed effect on encounter of study species
Profession Categorical variable representing sedentary, Categorical Respondents with outdoor work are more likely
semi-outdoor & outdoor work to detect study species
Spiti in , and the opening up of Kinnaur and Spiti to periods. Each interviewee’s report from a particular site
tourists in . We defined the recent time period as the was assigned as a replicate survey within the site. Site
-year period prior to our survey (–). As years covariates that could influence probability of site use by a
is a considerably long period to assume population closure species were modelled using logistic insertions (Table ).
within sampled units, we assumed that the occupancy esti- Topographic covariates such as mean altitude above mean
mates would reflect the probability of the sites being used by sea level, and proportions of rugged area in a site (terrain
the study species in the respective time periods. This inter- ruggedness index) and vegetated area in a site (normalized
pretation of the conventional occupancy estimate as the difference vegetation index) were generated using Shuttle
probability of site use allows for changes within the sam- Radar Topographic Mission (SRTM) and Landsat data, re-
pling units and relaxation of the closure assumption spectively. As snow leopards are known to occur between
(MacKenzie et al., ; Taubmann et al., ). This design , m (above the tree line) and , m (approximate
assumes that the changes in probability of site use within lower limit of permafrost and limit of vegetation growth)
each sampling unit during each sampling period were ran- in the Western Himalaya (Project Snow Leopard, ;
dom and described by the specific covariates applied. Snow Leopard Network, ), we used mean altitude and
Probability of colonization (γ) was estimated as the prob- its square to model probability of site use as a linear and
ability of a site that was not used by a species during the quadratic function of altitude. We expected probability of
past time period being used during the recent time period, site use to be higher in sites with a greater proportion of ter-
whereas probability of extinction (ε) was estimated as the rain ruggedness (Snow Leopard Network, ; McCarthy &
probability of a site that was used by a species during the Mallon, ). A higher proportion of vegetated area was ex-
past time period not being used during the recent time per- pected to favour the distribution of wild prey. Presence of
iod (MacKenzie et al., ). The probabilities of local col- migratory livestock grazing reflected areas that were used
onization and local extinction can be interpreted as for grazing only during the peak summer (mid June–mid
probability of distribution expansion and contraction, re- August) and were far from villages, and hence not grazed
spectively, between the two periods. by local livestock. Presence of migratory livestock grazing
Data on sightings of snow leopards, blue sheep and ibex was expected to lower the probability of site use by wild
gathered during interviews were arranged in a detection/ prey. The livestock population (i.e. the number of sheep/
non-detection (/) framework for past and recent time goats in a site) was expected to negatively influence
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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. https://doi.org/10.1017/S0030605317001107624 A. Ghoshal et al.
probability of site use by wild prey. If the confidence interval TABLE 2 Number of detections, and number of respondents report-
of the coefficient of a variable included zero we interpreted it ing detections, of snow leopards, wild prey (cumulative detections
as a statistically non-significant association. of blue sheep and ibex), blue sheep and ibex in the Greater and
Trans-Himalaya of Himachal Pradesh, India (Fig. ). Multiple re-
Covariates that might have influenced the probability of
spondents reported both blue sheep and ibex from more than one
detection of a species and its reporting to the researcher grid, and therefore the detections of wild prey sum to more than
were used as survey covariates to model detection probabil- the detections of individual species.
ity (Table ). These included respondent age, profession,
duration (in years) of familiarity with the area for which No. of
they were providing information, time spent in that area an- No. of detections respondents
nually, and a binary variable indicating whether or not a re- Past Recent Past Recent
spondent visited snow leopard and prey habitat during each Snow leopard 54 110 24 30
time period. Wild prey 330 351 27 33
We modelled probability of site use and detection Blue sheep 51 144 27 33
Ibex 55 214 27 33
probability through single-species multi-season models for
snow leopards, blue sheep and ibex, using Presence .
(MacKenzie et al., ; Hines, ). We parameterized
our models to estimate empirically historical and recent ., respectively. Naïve estimates of the proportion of sites
probabilities of site use and probability of distribution ex- used by wild prey in the past and recent time periods were
pansion. Estimates of probabilities of distribution contrac- . and ., respectively. When analysed individually,
tion were derived. Models were ranked in ascending order naïve estimates of the proportion of sites used by blue
based on the Akaike information criterion (AIC, Akaike, sheep in the past and recent time periods were . and
; Burnham & Anderson, ). ., and by ibex . and ., respectively.
Our analyses yielded multiple models with similar AIC va-
lues, thus indicating that no one model, by itself, could ex-
plain the observed variation in the data adequately. Thus, Detection probability
following a multi-model inference approach (Burnham & According to null models the detection probabilities of snow
Anderson, ), we considered a set of models with a cumu- leopards, blue sheep, ibex and wild prey were . ± SE .,
lative Akaike weight of $. to yield the best approximating . ± SE ., . ± SE . and . ± SE ., respective-
model by using model averaging (Burnham & Anderson, ly. When modelled with survey covariates, detection prob-
; Symonds & Moussalli, ). Models that did not con- abilities of snow leopards, wild prey, blue sheep and ibex
verge were dropped from the model set before model aver- varied between the past (–) and recent (–)
aging. We plotted the model-averaged estimates of past and time periods (Table ). Age of respondent showed a weak
recent occupancy to develop probabilistic maps of past and negative effect on detections of snow leopards, blue sheep
recent distributions of the snow leopard and its wild prey. and wild prey. Familiarity with the area of knowledge was
an important factor for ibex detection. Time spent each
year by a respondent in the area of survey had little effect
Results on detection probability. The profession of the respondent
had a positive effect on detection probability for all species.
We carried out systematic key-informant interviews
across Kinnaur, Lahaul, Spiti and Pangi. For analyses we
used of these interviews, having excluded two respon- Site use, and expansion or contraction of distribution:
dents who provided unreliable information. Respondents snow leopard
reported detection/non-detection of snow leopards and
their wild prey from a mean of grids per respondent The model-averaged estimate of probability of site use by
(range –). The mean number of respondents who reported snow leopards was . ± SD . (range . ± SE . to
sighting snow leopards per grid was . ± SD . (range . ± SE .) for the past time period and . ± SD .
–), whereas the mean number who reported sighting (range . ± SE . to . ± SE .) for the recent time
wild prey was . ± SD . (range –). Detection data period (Fig. a,b). Probability of site use by snow leopards
for each species are summarized in Table . We developed showed weak positive correlation with the terrain rugged-
competing single-species multi-season occupancy models ness index (βtri = . ± SE .; % CI −.–.) and
for the snow leopard, for all wild prey combined, for altitude (βalt.mean = . ± SE .; % CI −.–.)
blue sheep and for ibex (Supplementary Table S). (Table ); however, the coefficients were not statistically sig-
Naïve estimates of the proportion of sites used by snow nificant. In the % of the area used by snow leopards, the
leopards in the past and recent time periods were . and probability of site use by snow leopards was relatively high
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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TABLE 3 Untransformed estimates of coefficients (β) from the top models of snow leopard, wild prey, blue sheep and ibex model sets.
Snow leopard Blue sheep Ibex Wild prey
Model* β ± SE β ± SE β ± SE β ± SE
Intercept (ψ)_psi1 1.27 ± 0.42 0.97 ± 0.42 −0.77 ± 0.32 3.44 ± 0.66
Intercept (ψ)_psi2 1.27 ± 0.42 0.97 ± 0.42 −0.77 ± 0.32 2.26 ± 0.46
Altitude 0.73 ± 0.73 −0.36 ± 0.34 −1.13 ± 0.52
Altitude_squared −0.49 ± 0.19
Migratory livestock grazing −2.67 ± 0.61 4.26 ± 1.01
Ruggedness 0.89 ± 0.52
Intercept (γ) −0.75 ± 0.96 −3.54 ± 0.98 −1.53 ± 0.52 −1.39 ± 1.29
Intercept (p1) −2.59 ± 0.26 −1.89 ± 0.23 −1.30 ± 0.16 0.54 ± 0.13
Intercept (p2) 0.58 ± 0.21 1.21 ± 0.21 1.21 ± 0.19 −0.52 ± 0.13
Age −0.06 ± 0.02 −0.06 ± 0.01 −0.03 ± 0.01
Familiarity 0.04 ± 0.02 −0.05 ± 0.01 0.01 ± 0.01
Time spent 0.02 ± 0.01
Prof1 1.30 ± 0.27 0.32 ± 0.29 0.28 ± 0.17
Prof2 1.36 ± 0.25 1.12 ± 0.23 0.39 ± 0.14
*Ψ, Probability of site use; γ, probability of distribution expansion; p, probability of detection for the past time period; p, probability of detection for the
recent time period; psi, past probability of site use; psi, recent probability of site use. Site covariates: altitude, mean altitude of a site; altitude_squared,
square of mean altitude; Ruggedness, proportion of rugged area in a site, derived from terrain ruggedness index; Migratory livestock grazing, presence ()/
absence () of migratory livestock grazing in a site; Age, age of respondent in years; Familiarity, no. of years or months for which a respondent is familiar with
the area of knowledge; Time spent, time spent in months or days by a respondent in the area of knowledge per year; Prof, sedentary profession (e.g. office
worker); Prof, sedentary and outdoor profession (e.g. forest guard).
(. .). A relatively higher probability of site use (. .) contraction of blue sheep distribution was . ± SD .
was recorded in Spiti and in the majority of Kinnaur. (range . ± SE . to . ± SE .), indicating no clear
Central and western Lahaul and Pangi had relatively low evidence for overall contraction (Supplementary Fig. S).
probabilities of site use by snow leopards (Fig. a,b).
There was no clear evidence of expansion of the snow leo-
pard’s distribution (γ = . ± SE .; % CI −.–.). Site use, and expansion or contraction of distribution:
The estimated probability of contraction of the snow leo- Asiatic ibex
pard’s distribution was . ± SD ., indicating no clear The estimated probability of site use by ibex was the same for
evidence for overall contraction, and ranged between . ± past and recent time periods (ψ = . ± SD ± .; range: .
SE . and . ± SE . (Supplementary Fig. S). ± SE . to . ± SE .; Fig. c,d). Probability of site use
by ibex showed a positive correlation with migratory livestock
grazing (βmigrh = . ± SE .; % CI .–.; Table ).
Site use, and expansion or contraction of distribution: The relationship between ibex site use and mean altitude
blue sheep was unclear (βalt.mean = −. ± SE .; % CI −.–.)
The model-averaged estimate of probability of site use by blue (Table ). High probability of site use (. .) by ibex was
sheep was . ± SD . (range . ± SE . to . ± SE found in Lahaul and Pangi, and in south-western parts of
.) for the past time period and . ± SD . (range . Spiti and Kinnaur. Central, south and eastern Spiti, and nor-
± SE . to . ± SE .) for the recent time period thern, eastern and south-eastern Kinnaur showed low
(Fig. a,b). Probability of site use by blue sheep showed a probabilities of site use (, .) by ibex (Fig. c,d). There
negative correlation with migratory livestock grazing was no clear evidence for expansion of ibex distribution
(βmigrh = −. ± SE .; % CI −. to −.; Table ). across the study area (γ = . ± SE .; % CI −.–
There were no clear relationships between probability of .). The estimated probability of contraction of ibex distri-
site use by blue sheep and altitude (βalt.mean = −. ± bution was . ± SD . (range . ± SE . to . ±
SE .; % CI −.–.) or ruggedness (βtri = . ± SE SE .), indicating no clear evidence for overall contraction
.; % CI −.–.; Table ). A high probability of site (Supplementary Fig. S).
use (. .) by blue sheep was estimated in the majority of
Spiti and Kinnaur. South-western Spiti and all of Lahaul and Site use, and expansion or contraction of distribution:
Pangi showed low probabilities of site use (, .) by wild prey
blue sheep (Fig. a,b). We found no clear evidence of prob-
ability of expansion of blue sheep distribution (γ = . ± SE The model-averaged estimate of probability of site use
.; % CI −.–.). The estimated probability of by wild prey (blue sheep and ibex combined) was
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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. https://doi.org/10.1017/S0030605317001107626 A. Ghoshal et al.
FIG. 2 (a) Past and (b) recent probability
of site use (occupancy) by snow leopards
across the Greater and Trans-Himalaya
mountains of Kinnaur, Lahaul, Spiti and
Pangi, Himachal Pradesh, India.
. ± SD . (range . ± SE . to . ± SE .) vegetated area (normalized difference vegetation index)
for the past time period and . ± SD . (range . ± SE (βndvi = −. ± SE .; % CI −.–.) or presence/
. to . ± SE .) for the recent time period. absence of migratory livestock grazing (βmigrh = −. ± SE
Probability of site use by wild prey varied as a quadratic .; % CI −.–.; Table ). The probability of site use
function of altitude (βalt.mean = −. ± SE ., % CI by wild prey was high (. .) in most of the study area.
−. to −.; βalt.sq = −. ± SE .; % CI −. to Central Lahaul, parts of northern Spiti and parts of
−.), and increased with ruggedness (βtri = . ± SE .; Kinnaur (especially along the Sutlej River valley) showed
% CI .–.). There was no clear relationship between relatively lower probabilities of site use by wild prey
probability of site use by wild prey and proportion of (, .). There was no clear evidence for expansion of the
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FIG. 3 Past and recent probability of site use (occupancy) by blue sheep (a & b) and Asiatic ibex (c & d) across the Greater and
Trans-Himalaya mountains of Kinnaur, Lahaul, Spiti and Pangi, Himachal Pradesh, India.
distribution of wild prey (γ = . ± SE .; % CI −.– and depredation of wildlife and livestock by free-ranging
.). The estimated probability of contraction of the distri- dogs.
bution of wild prey was . ± SD . (Supplementary In Kinnaur the severe threats identified were reduced
Fig. S), indicating evidence for a marginal overall contrac- prey as a result of competition with local livestock and sub-
tion of distribution (range . ± SE . to . ± SE .). sistence hunting by the local community, illegal hunting and
The probability of contraction of the distribution of wild wildlife trade by immigrant labourer and migratory com-
prey was relatively high (ε . .) in % of the sites. munities, and depredation by dogs. Local livestock grazing
These areas were in western Lahaul, northern and eastern was practised in and around villages all across Kinnaur,
Spiti, south-western Spiti and Kinnaur (Supplementary whereas migratory livestock grazing was prevalent in
Fig. S). south-eastern and south-western Kinnaur (Supplementary
Fig. S). Southern and south-western Kinnaur also experi-
enced hunting and wildlife trade by local and migratory
Conservation threats herder communities and immigrant labourers. The depre-
dation of wildlife and livestock by free-ranging dogs was
Threat assessment (Table ; Supplementary Fig. S) indi- found to be widespread.
cated the major threats across the study area were reduced In Spiti reduced prey as a result of competition with local
prey as a result of competition with both local and migratory and migratory livestock, and depredation by dogs, were se-
livestock, reduced prey as a result of subsistence hunting by vere threats. Local livestock grazing was widespread except
local communities, illegal hunting and wildlife trade by both in a few villages in eastern Spiti on the left bank of the Spiti
local and immigrant labourer and migratory communities, River. Migratory livestock grazing was prevalent in north-
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628
A. Ghoshal et al.
TABLE 4 Threat ranking ( = low threat; = severe threat) based on area (how widespread is the threat; = least widespread; = most widespread), intensity (how severe is the threat; = low
severity; = high severity) and urgency (how immediate/urgent is the threat; = least urgent; = most urgent), following Margoluis & Salafsky (), for Kinnaur, Lahaul, Spiti and Pangi,
Himachal Pradesh, India (Fig. ). Light grey shading indicates a low level of threat, medium grey indicates a medium level of threat, and dark grey indicates a severe level of threat.
Kinnaur Spiti Lahaul Pangi (Chamba)
Total Total Total Total
Threat Area Intensity Urgency ranking Area Intensity Urgency ranking Area Intensity Urgency ranking Area Intensity Urgency ranking
Livestock–wild prey interactions
Prey reduction as a result 14 13 5 32 13 13 4 30 7 3 4 14 3 4 5 12
of competition with local
livestock
Prey reduction as a result 7 14 6 27 12 14 9 35 14 14 10 38 14 14 10 38
of competition with mi-
gratory livestock
Prey reduction as a result 1 2 3 6 5 6 3 14 2 2 2 6 5 3 4 9
of disease transmitted by
livestock
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
People–wildlife interactions
Retaliatory killing of 2 8 7 17 6 5 12 23 3 8 3 14 4 8 9 21
snow leopards & wild
prey by local community
Prey reduction as a result 13 9 8 30 3 2 10 15 8 9 9 26 9 11 12 32
of subsistence hunting
by local community
Illegal hunting & wildlife 6 10 13 29 4 4 11 19 10 10 13 33 10 12 13 35
trade by local
community
Illegal hunting & wildlife 12 11 14 37 7 8 13 28 12 13 14 39 12 13 14 39
trade by immigrant
labourer & migratory
communities
Developmental activities
Impacts of roads on 4 7 10 21 8 9 8 25 4 7 6 17 7 6 7 20
snow leopard habitat
Impacts of hydro-electric 5 6 11 22 2 1 6 9 9 5 11 25 6 5 6 17
projects on snow leopard
habitatSnow leopards in the Indian Himalaya 629
western, western and south-western Spiti (Supplementary
ranking
Total
Fig. S). Depredation of wildlife and livestock by free-
16
28
34
6
5
ranging dogs is an emerging conservation threat in Spiti.
In Lahaul reduced prey as a result of competition with
Area Intensity Urgency
migratory livestock, illegal hunting and wildlife trade by
1
8
11
3
2
the local community, and immigrant labourer and migra-
tory communities, and depredation by dogs, were the severe
Pangi (Chamba)
threats. Migratory livestock grazing was pervasive at rela-
tively high densities (Supplementary Fig. S). Areas grazed
7
9
1
2
10
by migratory livestock were also prone to illegal hunting
by migratory herders and depredation of wildlife by dogs
8
11
1
2
13
(migratory herders are accompanied by guard dogs).
ranking
Hunting was prevalent among the local community of west-
Total
ern Lahaul. Immigrant labour camps were abundant along
12
28
10
17
36
major roadways in Lahaul, and our respondents consistently
Area Intensity Urgency
reported labourers to be involved in illegal hunting and
wildlife trade.
1
5
8
7
12
In Pangi the severe threats were reduced prey as a result
of competition with migratory livestock and subsistence
hunting by the local community, illegal hunting and wildlife
6
12
1
4
11
trade by the local community and immigrant labourer and
Lahaul
migratory communities, and depredation by dogs. Similar
5
11
1
6
13
to Lahaul, migratory livestock grazing was widespread in
Pangi and these areas experienced illegal hunting and dep-
ranking
redation of wildlife by free-ranging dogs. Hunting for sub-
Urgency Total
19
22
27
40
9
sistence and for wildlife trade, by both local people and
immigrant labourers, was prevalent.
1
2
5
7
14
Discussion
Intensity
We set out to examine changes in distribution of the snow
7
10
3
11
12
leopard and its wild prey, and their current conservation sta-
Area
Spiti
tus across the Greater and Trans-Himalaya Mountains of
11
10
1
9
14
Himachal Pradesh between two time periods, past
(–) and recent (–). We used a recall-based
ranking
Total
key-informant survey to collect information on detection/
14
15
17
13
35
non-detection of snow leopards and their wild prey over
Area Intensity Urgency
the two periods. These data were analysed using a single-
species, multi-season site-occupancy framework accounting
1
2
9
4
12
for imperfect detection. We recommend caution in inter-
preting the effects of variables on site use as causation. For
example, normalized difference vegetation index data were
3
4
5
1
12
used to represent the proportion of vegetated area in a site;
Kinnaur
however, these data need not represent palatable forage for
wild prey, and may explain the absence of a significant posi-
10
9
3
8
11
Other human disturbances
tive relationship between proportion of vegetated area and
tion by local & migratory
Depredation of wildlife
Medicinal plant collec-
site use by wild prey.
communities & immi-
(off-roading/camping/
Unregulated tourism
hiking in wilderness,
Our results suggest that snow leopard distribution re-
military personnel
collection by local
Fodder/fuel wood
Illegal hunting by
free-ranging dogs
mismanagement)
mained largely unchanged between the two time periods,
grant labourers
Table 4 (Cont.)
& livestock by
as we did not find any evidence of overall expansion or con-
community
traction of snow leopard distribution in the study area,
garbage
Threat
whereas there was a marginal contraction in the distribution
of wild prey (blue sheep and ibex combined). Snow leopard
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. https://doi.org/10.1017/S0030605317001107630 A. Ghoshal et al.
distribution essentially overlapped with wild prey distribu- Lahaul, which experienced intense migratory livestock
tion, with blue sheep and ibex showing mutually exclusive grazing, widespread presence of free-ranging dogs, and
distributions with marginal overlap. illegal hunting and wildlife trade by local and migratory
Within our study area we recorded high probabilities of herder communities and immigrant labourers (Table ;
snow leopard site use across large patches in Spiti and Supplementary Fig. S).
Kinnaur (Fig. ). These were areas where migratory livestock Across Pangi, Lahaul and northern and south-western
grazing either did not occur or occurred at low intensity, and Spiti, the ibex is the only large wild prey of the snow leopard.
the intensity of other threats, such as hunting and natural re- These are also the areas where migratory livestock grazing,
source extraction, was also low (Table ; Supplementary depredation by free-ranging dogs, and illegal hunting and
Fig. S). Among areas that showed a relatively high probability wildlife trade by local and migratory herder communities
of contraction of snow leopard distribution (Supplementary and immigrant labourers are pervasive and intense
Fig. S), % were in south-western parts of Kinnaur and (Table ; Supplementary Fig. S). Migratory livestock graz-
Spiti that experienced migratory livestock grazing and hunting ing is a long-standing, widespread and continually changing
by local and migratory people as well as by immigrant practice (Saberwal, ; Axelby, ) in parts of Kinnaur
labourers. Migratory grazing was prevalent in c. % of and Spiti and all of Lahaul and Pangi, covering almost the
the snow leopard’s distribution within our study area entire estimated ibex distribution. The use by migratory her-
(Supplementary Fig. S), and was relatively intense (c. live- ders of these relatively steep areas, which are also the
stock per km) in c. % of the area, especially in Lahaul and habitats of the ibex, resulted in a positive correlation be-
Pangi (Supplementary Fig. S). Intense migratory livestock tween probability of site use by ibex and the presence of
grazing, along with hunting by migratory herders and immi- migratory livestock grazing. There is evidence of interfer-
grant labourers in Lahaul continue to be serious threats to the ence competition between migratory livestock and ibex
otherwise contiguous snow leopard habitat, from Pangi to the (Bagchi et al., ). Migratory livestock displace ibex
west, Spiti to the east and Kinnaur to the south-east. Our study from pastures through forage removal and direct disturb-
area is also contiguous with the snow leopard habitats of ance. Understanding impacts of migratory livestock grazing
Hemis in Ladakh (Jammu and Kashmir) to the north. on forage availability and ibex densities (exploitative com-
We found that within our study area % of the area trad- petition) at a finer spatial scale would be useful.
itionally considered to be potential snow leopard habitat Blue sheep occur extensively and at relatively high dens-
(Project Snow Leopard, ; Snow Leopard Network, ities in Spiti, especially along the eastern bank of the Spiti
) had a negligible probability of use by snow leopards. River and parts of the western bank (Suryawanshi et al.,
This is despite the fact that areas above , m altitude ), where the terrain is relatively gentle and ibex are
(mainly rocky areas with snow and ice) and below largely absent. These are also areas used only for local live-
, m (forested zone) were excluded from our survey. stock grazing rather than migratory livestock grazing (USL,
Large-scale projections of snow leopard distributions and ), hence the negative correlation between site use by blue
populations have relied on extrapolation of information sheep and migratory livestock grazing.
based on opinions and geophysical modelling (e.g. Snow Snow leopards and wild prey occurred in large parts of
Leopard Working Secretariat, ; Snow Leopard Network, our study area and were not restricted to the seven protected
; McCarthy & Mallon, ). Our results caution against areas that cover % (, km) of the area. Himachal
such extrapolations, as they are likely to substantially overesti- Pradesh thus mirrors a global pattern (Johansson et al.,
mate snow leopard occurrence. ) in which large parts (c. , km) of snow leopard
Blue sheep and ibex distributions in our study area were and wild prey distribution lie outside protected areas. This
largely mutually exclusive, overlapping marginally along the makes landscape-scale participatory conservation ap-
Sutlej and Spiti Rivers (Fig. ). Blue sheep occurred in the proaches, such as India’s Project Snow Leopard () im-
eastern and south-eastern parts of the study area, which plemented in the Upper Spiti Landscape of Himachal
are less rugged, with rolling mountains (Fig. a,b). Ibex oc- Pradesh (USL, ), and the Global Snow Leopard and
curred in the western and south-western parts of the study Ecosystem Protection Program (Snow Leopard Working
area, which are more rugged, with steeper mountains Secretariat, ) currently being implemented, more ap-
(Fig. c,d). Thus, in Himachal Pradesh, blue sheep and propriate in the context of snow leopard and wild ungulate
ibex provide an important prey base for snow leopards, in conservation in the Indian Himalaya.
mostly mutually exclusive areas (Suryawanshi et al., ).
We found that the overall distribution of wild prey appeared
to have declined by % between the two study periods. Areas Acknowledgements
where contraction occurred (Supplementary Fig. S), simi-
lar to areas with contraction of snow leopard distribution, We thank the Himachal Pradesh Forest Department
were in south-western Kinnaur, and central and western (Wildlife Wing) for financial and logistical support; James
Oryx, 2019, 53(4), 620–632 © 2017 Fauna & Flora International doi:10.1017/S0030605317001107
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. https://doi.org/10.1017/S0030605317001107Snow leopards in the Indian Himalaya 631
E. Hines for valuable inputs at various stages of the work Http://www.mbr-pwrc.usgs.gov/software/presence.html [accessed
and during preparation of the manuscript; our survey re- August ].
J AC K S O N , R., M A L LO N , D., M C C A R T H Y , T., C H U N D AWAY , R.A. &
spondents for their kind cooperation; and Lobzang
H A B I B , B. () Panthera uncia. In The IUCN Red List of
Gialson for invaluable assistance during field work. The Threatened Species : e.TA. Http://dx.doi.org/.
Whitley Fund for Nature has been a long-term supporter /IUCN.UK..RLTS.TA.en [accessed July
of our research and conservation work, and we are thankful ].
for support from the Whitley–Fondation Segré Partnership J AY A P A L , R. & R A M E S H , K. () Management Plan for Rupi-Bhaba
Wildlife Sanctuary, Himachal Pradesh [April –March ].
Fund.
Wildlife Institute of India, Dehradun, India.
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and Ecosystem Protection Program. Bishkek, Kyrgyz Republic. interested in theoretical and applied research and community-based
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, –. community-based conservation.
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