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Status and conservation of the snow leopard

Panthera uncia in Api Nampa Conservation


Area, Nepal
GOPAL KHANAL, LAXMAN PRASAD POUDYAL, BISHNU PRASAD DEVKOTA
R I S H I R A N A B H A T and P E R W E G G E

Abstract The snow leopard Panthera uncia is globally Introduction


threatened and reliable information on its abundance,
distribution and prey species is a prerequisite for its conser-
vation. In October–November  we assessed the distri- T he snow leopard Panthera uncia, although globally
threatened, is one of the least studied large mammalian
carnivores (Snow Leopard Network, ). Its patchy distri-
bution of the snow leopard in the recently established Api
Nampa Conservation Area in the Nepal Himalayas. bution spans c. . million km of high mountains across 
Within selected blocks we conducted sign surveys and Asian countries (McCarthy et al., ). Based on field stud-
counted the number of bharal Pseudois nayaur, its principal ies, extrapolation of density estimates to non-surveyed areas,
wild prey, along transects totalling  km. We recorded  expert opinion and interviews, the global population is
putative snow leopard signs at an encounter rate of . estimated to be ,–,, and in  the species was re-
signs/km. Generalized linear models of the number of categorized, from Endangered to Vulnerable, on the IUCN
signs detected per transect showed that elevation had a posi- Red List, suggesting that threats to snow leopards have di-
tive influence and human activities a negative influence on minished (McCarthy et al., ). Nevertheless, burgeoning
sign encounter rate; prey abundance had only a weak posi- livestock grazing pressure on its natural prey (Mishra et al.,
tive influence on sign encounter rate. Within the effectively ), poaching (Li & Lu, ; Nowell et al., ), and
surveyed area of c.  km, we counted  bharal at an es- escalating conflicts with herder communities over livestock
timated density of . animals/km. Recruitment of bharal depredation (Bagchi & Mishra, ; Suryawanshi et al.,
was low, estimated at  kids/ adult females, most likely a ) continue to threaten the species. As many as
result of poor or overgrazed rangeland. We estimate – snow leopards are killed annually in retaliation by
the total number of bharal in this conservation area to be herder communities or as a result of becoming accidentally
. ,, a prey base that could sustain – snow leopards. ensnared in traps set for other species (Nowell et al., ).
Based on our field observations, we identified human dis- Habitat loss, fragmentation and degradation from infra-
turbance and habitat degradation associated with extraction structure development (Zaller, ), resource extraction
of non-timber forest products, livestock grazing, and poach- (Farrington, ) and changing climate patterns (Li et al.,
ing as the main threats to the snow leopard. Standardized ) have exacerbated anthropogenic impacts on snow
sign surveys, preferably supplemented by sampling with leopard habitat.
remote cameras or with genetic analysis of scats would In  governments of the snow leopard range countries
provide robust baseline information on the abundance of initiated the Global Snow Leopard Ecosystem Protection
snow leopards in this conservation area. Program to combat threats and strengthen transnational
collaboration for snow leopard conservation (Snow
Keywords Api Nampa Conservation Area, bharal, Nepal, Leopard Working Secretariat, ). Knowledge of the
Panthera uncia, Pseudois nayaur, snow leopard abundance and distribution of the snow leopard across its
range is central to the success of the Programme, but reliable
information on population status and local threats is un-
GOPAL KHANAL* (Corresponding author) Department of Forests, Ministry of available for large parts of the species’ range, both within
Forests and Environment, Government of Nepal, Singhadurbar Kathmandu, and outside protected areas. Less than % of the species’
Nepal. E-mail khanal.joshipur@gmail.com
distribution range has been adequately surveyed (Snow
*Also at: Centre for Ecological Studies, Lalitpur, Nepal
Leopard Network, ). As a result, conservation decisions,
LAXMAN PRASAD POUDYAL and RISHI RANABHAT Department of National Parks and including those of the Program, are based in part on extra-
Wildlife Conservation, Ministry of Forests and Environment, Government of
Nepal, Kathmandu, Nepal polations of density estimates to non-surveyed areas.
BISHNU PRASAD DEVKOTA Institute of Forestry, Pokhara Campus, Pokhara, Nepal
Assessments of snow leopard conservation status in unsur-
veyed regions across the species’ range are thus urgently re-
PER WEGGE Faculty of Environmental Sciences and Natural Resource
Management, Norwegian University of Life Sciences, Ås, Norway quired for informing management decisions.
Received  July . Revision requested  January .
The , km of suitable habitat in the Nepal
Accepted  January . Himalayas has been estimated to support – snow

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2 G. Khanal et al.

leopards, c. % of the global population (WWF Nepal, and river valleys. Vegetation above , m is largely dry al-
). Studies of the snow leopard in Nepal commenced pine steppe characterized by meadows rich in sedges and
with those of Schaller (), Jackson & Ahlborn (), graminoids such as Stipa, Carex and Kobresia. On drier
and Jackson () in Shey Phoksundo National Park. sites and rugged slopes vegetation is dominated by dry
Recent studies have focused on the western and central grasses and shrubs such as Caragana brevifolia and
Himalayas (Thapa, ; Wegge et al., ; Ale et al., Lonicera spinosa. Below , m mixed broadleaved and
, ; Devkota et al., ), but a large part of potential conifer forest species include blue pine Pinus wallichiana,
habitat remains poorly studied, in particular Api Nampa Himalayan fir Abies spectabilis, birch Betula utilis, burans
Conservation Area in the western Himalayas. Rhododendron arboreum, oak Quercus spp. and juniper
Reliable estimates of species abundance are fundamental Juniperus indica. The fauna includes the Himalayan marmot
for conservation and management. Recently developed Marmota himalayana, woolly hare Lepus oiostolus, musk
non-invasive techniques, including camera trapping and deer Moschus chrysogaster, Himalayan goral Naemorhedus
DNA analysis, are among the best methods to study large goral and bharal (blue sheep) Pseudois nayaur. Bharal is the
elusive carnivores such as the snow leopard (Jackson et al., most common ungulate and a food source for the snow leop-
; Janecka et al., ), but are costly. Camera trapping ard, golden jackal Canis aureus and grey wolf Canis lupus.
can be logistically challenging when information on pres- Within the suitable habitat for the snow leopard
ence/absence of the study species is not available. In these (generally above , m), local communities are mainly
circumstances, information on spatial distribution can be agro-pastoralist. Green peas, potato and barley are the
useful for defining conservation priorities. The presence main agricultural crops. The dominant source of income
and spatial distribution of a species can be assessed based for local communities is animal husbandry and collection
on indirect evidence of species occurrence (e.g. pugmarks, of non-timber forest products, including a valuable
scrapes; Jackson & Hunter, ). medicinal product that is a complex of the parasitic fungus
The goal of this study was to determine snow leopard pres- Cordyceps sinensis and the ghost moth caterpillar Thitarodes
ence and to estimate the abundance of its wild prey in Api spp. (Pant et al., ). Livestock species include sheep,
Nampa Conservation Area, and thus to estimate the number goats, yaks, cattle–yak hybrids (dzo, jhopas), and horses.
of snow leopards that this area could potentially support. We Migratory herders employ a seasonal grazing pattern in
also examined the influence of ecological and anthropogenic which they move large herds of livestock to higher elevations
variables on encounter rate of snow leopard signs. in late April, rotate them among pastures, and then herd
them to lower elevations in mid September.
Study area
Methods
The , km Api Nampa Conservation Area, established
in , lies in the far-western Himalayas of Nepal (Fig. ) Snow leopard sign survey
at altitudes of –, m. Monthly mean maximum tem-
perature is c. °C in summer and in winter the minimum We adopted the Snow Leopard Information Management
temperature can drop to c. −°C. Precipitation above , System (Jackson & Hunter, ) to determine snow leopard
m is mainly in the form of snow, which starts to melt in presence and to estimate the relative encounter rate of snow
March. Topography is characterized by undulating terrain leopard signs. Although this system was not developed spe-
cifically for estimating snow leopard abundance, it is never-
theless a useful method for conducting presence/absence
surveys as it is straightforward for field assistants with little
formal education to use, has minimal impact on the species
being studied, and is inexpensive (Ale et al., ). Based on
anecdotal information of snow leopard occurrence we
selected six study blocks, Tinkar, Nampa, Ghusa,
Khandeshowri, Sitaula and Rapla, with areas of – km.
Within these blocks we located ridgelines, narrow valleys
and human trails (used and unused) where snow leopards
were most likely to leave signs of territory marking and/or
communication (Jackson, ), to serve as transects. No
routes were surveyed more than once. We recorded the
location and sign type (scrape, pugmark, scent marking,
scats) with a GPS, and the elevation at the beginning and
FIG. 1 The location of Api Nampa Conservation Area in Nepal. end of the transect. To quantify potential human

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Snow leopards in Nepal 3

disturbance on the snow leopard we recorded presence/ab- models. We assessed collinearity between the predictor vari-
sence of recent human activities/use of the area (e.g. fire- ables, using Pearson’s correlation coefficients. We did not
wood collection, livestock grazing, snares, poaching camps include covariates that were substantially correlated
and fires), as a proxy. A transect was considered to be dis- (Pearson’s |r|. .) in the same model (Dormann et al.,
turbed if direct evidence for any of these activities was ob- ). We ranked models using the Akaike information cri-
served at least at one location along the transect. A distance terion adjusted for small samples (AICc; Burnham &
of at least  m was maintained between consecutive trans- Anderson, ). Models with ΔAICc ,  were considered
ects, to avoid spatial autocorrelation. Transects were at to be strongly supported by the data. We used the R package
,–, m, nearly all above the treeline. Surveys were MuMin to perform model averaging of all candidate models
conducted in , during October–November, when mi- (Barton, ), and computed % confidence intervals of
gratory livestock graze at lower elevations, resident livestock the beta coefficients for each predictor variable. The
are fed in stalls, and human disturbance is minimal. model-averaged beta-coefficients of covariates and their
% confidence intervals were examined to assess the signifi-
cance of their effect on leopard sign encounter rate.
Bharal survey Confidence intervals that included zero indicated no signifi-
cant effect of the covariates. The relative importance of each
While conducting sign surveys along transects we also
covariate was determined by summing the Akaike weights of
counted bharal using binoculars and spotting scopes
the models containing these factors. All analyses were
(Schaller, ; Shrestha & Wegge, ). To determine
carried out in R .. (R Development Core Team, ).
population structure and recruitment we recorded the age
We calculated the maximum number of snow leopards
class and sex of each bharal seen. Following Schaller
that the available bharal population could support following
() and Wegge (), bharal were categorized as adult
the predator–prey ratio suggested by Oli (). We esti-
males (+ years old; horns curved and .  cm in length),
mated the number of bharal that the suitable snow leopard
subadult males (– years; horns curved backwards, with
habitat (c.  km; G. Khanal, unpubl. data) could support
a length of – cm), young males (– years; horn length
based on bharal counts in six blocks (a total of c.  km),
– cm), adult females (.  years), yearlings (both sexes,
and then converted the estimated number into biomass.
– years) and young (,  year). Additional information on
Because there is little reliable empirical information on
group size, particular characteristics and composition of the
body weights for each category of bharal, we considered
group (e.g. male groups), location, time and behaviour were
adult male, subadult male, young male, adult female, year-
recorded for cross checking and to avoid double counting.
ling and young age classes to have a mean weight of , ,
For the same reason we also surveyed adjacent transects
, ,  and  kg, respectively (Schaller, ). We then
that were within the potential daily movement distance of
used the predator–prey weight ratio ( : –; Oli, )
bharal groups (– km). Surveys were conducted in ,
to estimate the number of snow leopards that Api Nampa
during October–November, when bharal herds congregate
Conservation Area could potentially support. This ratio
for courtship and mating and are therefore easiest to count.
was based on findings (Jackson & Ahlborn, ) that an
adult snow leopard requires – kg of meat annually
Data analysis (– bharal) after accounting for energy expenditure in a
wild snow leopard and the inedible parts of its prey (Jackson
Density estimates for bharal within each block were calcu- & Ahlborn, ).
lated by dividing the number of animals recorded by the ef-
fectively sampled area (the area covered by scanning from
the survey transects). We expected that mean elevation Results
(mean of elevation at beginning and end of transect), prey
availability (number of bharal encountered per transect) We surveyed  transects, covering a total distance of  km,
and human disturbance (presence/absence of recent with lengths of –, m and a mean length of ,
human activities/use) would influence the sign encounter m. We recorded  snow leopard signs on  transects
rate (Wolf & Ale, ; Sharma et al., ; Alexander (Table ), with a mean encounter rate of . signs/km.
et al., a; Suryawanshi et al., ). To test the influence We excluded scats from the analysis because we were uncer-
of these potential predictor variables on the number of signs tain of identification to species. Scrapes were the most
encountered along each transect, we used generalized linear frequently detected sign (n = , ./km), followed by pug-
models with a Poisson error distribution and log link func- marks (n = , ./km). The highest encounter rate of all
tion. The predictor variables were standardized to z-scores types of sign was in the Tinkar block (./km; Table ).
prior to analysis, which allowed us to interpret the model The predictor variables elevation, human disturbance
coefficients and to compare effect sizes between alternative and prey availability were not highly intercorrelated

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4 G. Khanal et al.

TABLE 1 Results of transect surveys for snow leopard Panthera uncia signs conducted at six sites within Api Nampa Conservation Area
(Fig. ) during October–November, .

Total transect No. of No. of signs No. of scrapes No. of pugmarks No. of scats
Site length (km) transects (per km) (per km) (per km) (per km)
Tinkar 39.14 21 92 (2.35) 40 (1.02) 31 (0.79) 16 (0.41)
Nampa 23.62 11 43 (1.82) 19 (0.8) 14 (0.59) 8 (0.34)
Ghusa 20.62 8 36 (1.75) 15 (0.72) 12 (0.58) 8 (0.39)
Sitaula 9.57 5 16 (1.67) 8 (0.83) 4 (0.41) 3 (0.31)
Khandeswori 5.64 3 8 (1.42) 4 (0.71) 3 (0.53) 1 (0.17)
Rapla 7.62 3 8 (1.05) 3 (0.39) 3 (0.39) 2 (0.26)
Total 106.21 51 203(1.91) 89 (0.84) 67 (0.63) 38 (0.35)

(Pearson correlation coefficients were all , .) and hence available for bharal and snow leopards in Api Nampa.
all were used in the regression analyses. Elevation was the With an estimated density of . bharal/km in the sur-
most important variable in determining the encounter veyed blocks, we estimate that the suitable habitat (% or
rate of snow leopard signs along transects, followed by a  km of the whole , km protected area) could sup-
negative effect of human use (Tables  & ). Snow leopard port a minimum of , bharal. Considering  kg as the
signs were also positively associated with number of bharal mean weight of an individual bharal, this total population is
detected from the transect, but the relationship was not sta- c. , kg of biomass, which, based on the predator–prey
tistically significant (Table ). ratio (Oli, ), could sustain – snow leopards.
We counted a total of  individual bharal in  groups,
comprising % males, % females, % yearlings and %
young (Table ), with a mean group size of . (range –). Discussion
The highest density of bharal was in the Tinkar block and
the lowest in Sitaula, and the greatest numbers of bharal Our study provides the first evidence of snow leopard pres-
were sighted at ,–, m (Fig. ). The overall density ence in Api Nampa Conservation Area, Nepal. The infor-
of bharal within the effectively surveyed area of c.  km mation on the estimated number of snow leopards and
was estimated to be . ± SE ./km. The adult sex ratio bharal biomass and density reported here will help in setting
was female biased, with  males/ adult females. Overall up priorities for conservation, research and management.
recruitment rate was  young/ females. The ratio of
yearlings to young was . : , indicating a % mortality Snow leopard distribution
rate of kids, assuming a stationary age structure and similar
recruitment rate to that of the previous  years. We recorded snow leopard presence in five of the six blocks
Approximately , kg of bharal biomass was estimated surveyed. Although the rate at which we encountered snow
to be available within the effectively sampled area. leopard signs (./km for all blocks combined) was lower
This biomass could support – snow leopards. Based on than reported in other protected areas of Nepal
our occupancy surveys, c.  km of suitable habitat is (Kanchenjunga Conservation Area: . signs/km, Thapa,

TABLE 2 Poisson regression models describing the occurrence of the snow leopard in Api Nampa Conservation Area (Fig. ) during
October–November , ranked according to the Akaike information criterion adjusted for small sample size (AICc).

Model1 K2 AICc ΔAICc3 loglink Wi4


Elevation + HumanDist 3 241.47 0.00 −117.48 0.62
Elevation + HumanDist + Prey 4 242.46 0.99 −146.79 0.38
Elevation 2 252.95 11.48 −124.35 0.00
Prey + Elevation 3 254.17 12.70 −123.83 0.00
Prey + HumanDist 3 258.27 16.80 −125.88 0.00
HumanDist 2 267.10 25.63 −131.43 0.00
Prey 2 290.46 44.99 −143.11 0.00
Invariant/null 1 305.60 64.13 −151.76 0.00

Elevation (m); HumanDist, presence/absence of recent human use of the area; Prey, number of bharal encountered.

Number of parameters.

Difference between the AICc value of the best-supported model and successive models.

Akaike model weight.

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Snow leopards in Nepal 5

TABLE 3 Model-averaged parameter estimates and % confidence interval describing snow leopard occurrence in Api Nampa
Conservation Area (Fig. ) during October–November .

Parameter1 Estimate2 Adj. SE Confidence interval z P Relative variable importance


Elevation 0.12 0.02 0.08–0.15 4.69 0.00 1.00
HumanDist −0.09 0.02 −0.05–−0.13 3.49 0.04 0.98
Prey 0.02 0.02 −0.02–0.06 1.15 0.24 0.38

Elevation (m); HumanDist, presence/absence of recent human use of the area; Prey, number of bharal encountered.

Averaged from all models.

; Sargarmatha National Park: . signs/km, Ale et al., species. In the Qilian Shan region of China there was a
; Annapurna Conservation Area: . signs/km, Ale weak association between prey and snow leopard site use
et al., ; Manaslu Conservation Area: . signs/km, at a fine scale ( km; Alexander et al., b), but a strong
Devkota et al., ), the prey biomass was sufficiently positive relationship at a landscape scale (Alexander et al.,
large to support a breeding population of snow leopards. a). It is also possible that other factors, such as terrain
Survey times may influence sign encounter rate. There ruggedness, which we could not measure (our sampling
may be a higher detection rate in winter and post-winter be- units were linear transects and it was not possible to
cause of accumulation of snow and minimal human pres- compute a terrain ruggedness index adequately for such
ence (Jackson & Hunter, ). Devkota et al. (), for features), could have better explained the variability in
example, surveyed both pre-winter (November– sign encounter rate, as in Sagarmatha National Park,
December) and post-winter (May–April). Our surveys, Nepal (Wolf & Ale, ).
however, were conducted just after the migratory livestock In Api Nampa Conservation Area snow leopards were
grazing and fungal caterpillar collection season (October– less likely to leave signs in areas of active human presence.
November). In addition, we did not consider putative Generally, large carnivores avoid using such areas, especially
snow leopard scats in our analysis, which could have low- where hunting or harassment is common (Kerley et al.,
ered the overall sign encounter rate. ). In our study area, although illegal resource extraction
and hunting are prohibited, local herders disclosed that il-
legal activities, such as use of snares to hunt bharal and
Factors influencing sign encounter rate pheasants Lophophorus impejanus, and collection of timber
and non-timber forest products, occur frequently. This
Contrary to our initial expectation, there was only a weak could be causing significant disturbance to snow leopards.
positive association between the availability of bharal and In Sagarmatha National Park snow leopards were less likely
the number of snow leopard signs. Although abundance to use trails used by tourists, guides or porters and their yak
of wild prey is a key determinant of habitat use and abun- caravans (Wolf & Ale, ).
dance of snow leopards (Sharma et al., ; Suryawanshi We were unable to distinguish whether the greater sign
et al., ), it was not the main predictor variable in our encounter rate that we found at higher elevations reflects
study. This could be because the length of the individual greater site use or increased sign detection probability as a
transects (– km) was insufficient to encompass the vari- result of more snow accumulation and greater retention
ability required for the model to detect the relationship, time of signs. Higher elevations are often more remote
given the wide-ranging nature of snow leopards and less visited by people. A study of the fine scale occur-
(Johansson et al., ). A temporal mismatch between rence of snow leopards in Qilian Shan, China, found that
bharal presence and the time when leopards left signs may the probability of site use by snow leopards increased with
also have obscured a direct relationship between the two altitude (Alexander et al., a).

TABLE 4 Bharal population structure in Api Nampa Conservation Area (Fig. ) in October–November .

Study block Adult male Subadult male Young male Female Yearling Young Total Area (km2) Density (per km2)
Tinkar 14 20 21 73 27 35 190 50 3.8
Nampa 12 12 8 51 16 25 124 40 3.1
Ghusa 15 12 9 39 18 18 111 37 3
Sitaula 1 1 0 3 2 2 9 15 0.6
Khandewori 7 5 6 21 7 5 51 31 1.6
Rapla 6 4 3 15 7 7 42 27 1.5
Total 55 54 47 202 77 92 527 200 2.28

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6 G. Khanal et al.

local community leaders and herders, and data available at


the Api Nampa Conservation Area office, suggest that a
minimum of , livestock graze on the rangelands in
summer, c. × higher than our estimate of , bharal.

Limitations and further studies

The information on snow leopard distribution and factors


influencing the sign encounter rate reported here are
based on limited survey data, and sign encounter rate is a
less accurate metric for assessing snow leopard habitat use
than movement data from radio telemetry. Our analyses
of encounter rates may reflect where we were more likely
to detect signs, rather than habitat use per se. Detection of
snow leopard signs may be influenced by survey season,
search effort, observer experience and fatigue, weather
(snow fall), substrate (e.g. snow cover), sign type and lon-
FIG. 2 Total number of bharal Pseudois nayaur observed across
gevity, terrain type, site selection (e.g. ridgeline vs river
four bands of altitude in Api Nampa Conservation Area (Fig. ) bank trails) and human and livestock use of the area
in autumn. (McCarthy et al., ).
Our estimate that – snow leopards could be supported
in Api Nampa Conservation Area is based on the predator–
Wild prey bharal density estimates prey ratio (Oli, ), which does not take into account the
contribution of livestock and alternative prey such as the
Our density estimates and recruitment rate of bharal were marmot (Mallon et al., ). Livestock can contribute up
lower than in other protected areas in Nepal (Table ). to % of annual snow leopard diet (Wegge et al., ),
Schaller () attributed the low recruitment rate in Shey and smaller mammals such as the marmot can contribute
Phoksundo National Park, Nepal, to excessive livestock as much as % of the summer food of snow leopards
grazing, but high predation on young animals could also (% of annual requirements; Schaller et al., ).
have been a contributing factor. In Spiti Valley, Himachal Although the total prey biomass in the Conservation Area
state, India, high grazing pressure by livestock led to a low could therefore potentially support more than – snow
bharal reproductive rate (Mishra et al., ). However, be- leopards, we observed a relatively low abundance of mar-
cause of fine-scale habitat partitioning, forage competition mots, and local residents indicated there is little livestock
between livestock and bharal may not be as direct as com- depredation by snow leopards. Use of the predator–prey
monly assumed (Shrestha & Wegge, ). We did not col- ratio to estimate snow leopard numbers also assumes
lect information on habitat relationships or on forage there is no competition from other predators for the same
availability but based on our general observations and infor- prey biomass. Because we rarely encountered wolf signs
mation on livestock numbers, we suggest that high grazing and local herders reported that wolves were seldom ob-
pressure by seasonally grazing livestock may be a contribut- served, it is unlikely that other predators consume a sub-
ing factor to the low recruitment of bharal. Accounts from stantial proportion of the bharal biomass.

TABLE 5 Bharal density and population recruitment estimates for Nepal.

Recruitment/young
Location Density (per km2) per 100 females Year Source
Manang District, Annapurna Conservation Area 6.6–10.2 1990 Oli (1994)
Kanchenjunga Conservation Area 4.2 40 2006 Thapa (2006)
Shey Phoksundo National Park 2.1 50 2006 Thapa (2006)
Phu Valley, Manang 9.4 56 2008 Shrestha & Wegge (2008)
Yak Kharka, Manang 2.09 2009 Aryal et al. (2014)
Manaslu Conservation Area 3.8 2011 Devkota et al. (2017)
Annapurna & Manaslu Conservation Areas 5.97 ± SE 0.10 2012 Chetri et al. (2017)
Annapurna Conservation Area 2.3 60 2013 WWF Nepal (2013)

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Snow leopards in Nepal 7

Our method for surveying the bharal population does Inc., USA, for providing the research equipment; Man Bahadur
not account for imperfect detection and therefore probably Khadka, Maheshwar Dhakal, Sher Singh Thagunna, Koustubh Sharma,
Hem Raj Acharya, Ashok Kumar Ram, Saroj Panthi, Gokarna Jung
underestimated the density of this key prey species, thus also
Thapa, Pemba Sherpa, Dipendra Nath and Indra Nagari for their sup-
affecting the inference from the predator–prey ratio. The port; Rachel Moseley for help in improving the text; and the editor and
double observer method, which allows the detection prob- two anonymous reviewers for their constructive comments.
ability to be estimated, for correcting the counts, and also
allows estimation of the precision of the estimates, could
Author contributions Study design: GK, PW; data collection and
have produced more accurate estimates of the bharal popu-
analysis: GK; writing: GK, PW, LPP, BPD, RR.
lation (Suryawanshi et al., ). We counted bharal from
transects that often ran along ridgelines. Bharal are wary
of disturbance and retreat when they detect humans on Conflicts of interest None.
ridgelines, even far away. We estimated bharal density by
dividing the number of animals seen by the area effectively
Ethical standards This research abided by the Oryx Code of
sampled, which was based on the sum of areas that we con- Conduct.
sidered to have been scanned. This method is subjective and
may have overestimated the total sampling area, leading to
an underestimation of density. Overall, the potential for References
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