The status of the Endangered Persian leopard Panthera pardus saxicolor in Bamu National Park, Iran

Abstract

We describe the use of camera-trapping with capture-recapture, occupancy and visitation rate modelling to study the size, demographic structure and distribution of the Persian leopard Panthera pardus saxicolor in Bamu National Park, southern Iran. A total sampling effort of 1,012 trap-nights yielded photo-captures of four adults, two subadult individuals and a cub over 21 sampling occasions. The leopard population size estimated by the M(h) model and jackknife estimator was 6.00 ± SE 0.24 individuals. This gives a density of 1.87 ± SE 0.07 leopards per 100 km2. Detection probability was constant and low and, as a result, estimated occupancy rate was significantly higher than that predicted from photographic capture sites alone. Occupancy was 56% of the protected area and visitation rates were 0.01–0.05 visits per day. The most imminent threats to leopards in Bamu are poaching and habitat fragmentation.

Full text

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The status of the Endangered Persian leopard
Panthera pardus saxicolor in Bamu National Park, Iran
Arash Ghoddousi,Amirhossein Kh. Hamidi,Taher Ghadirian
Delaram Ashayeri and Igor Khorozyan
Abstract We describe the use of camera-trapping with
capture-recapture, occupancy and visitation rate modelling
to study the size, demographic structure and distribution of
the Persian leopard Panthera pardus saxicolor in Bamu
National Park, southern Iran. A total sampling effort of
1,012 trap-nights yielded photo-captures of four adults, two
subadult individuals and a cub over 21 sampling occasions.
The leopard population size estimated by the M(h) model
and jackknife estimator was 6.00 –SE 0.24 individuals. This
gives a density of 1.87 –SE 0.07 leopards per 100 km
2
.
Detection probability was constant and low and, as a result,
estimated occupancy rate was significantly higher than that
predicted from photographic capture sites alone. Occupancy
was 56% of the protected area and visitation rates were 0.01–
0.05 visits per day. The most imminent threats to leopards in
Bamu are poaching and habitat fragmentation.
Keywords Bamu, camera-trapping, density, Iran, leopard,
occupancy, Panthera pardus saxicolor
Introduction
With an area of 1,640,000 km
2
Iran is a vast country
withadiversityoflandscapes,floraandfauna(.8,000
speciesofplantsand.1,674 species of vertebrates; Zehzad
et al., 2002;Firouz,2005;Darvishsefat,2006); c. 7%ofthe
country’s territory is afforded various levels of protection
(Darvishsefat, 2006). Preservation of the biodiversity of Iran
would benefit from the selection and priority conservation of
flagship species, especially carnivores, which can provide
habitat connectivity because of their relatively large home
ranges (Linnell et al., 2000) The leopard Panthera pardus
saxicolor is a flagship species (Breitenmoser et al., 2007)and,
with the extinction of the lion Panthera leo persica and tiger
Panthera tigris virgata, is the only extant large felid in Iran.
Although this subspecies also occurs in neighbouring coun-
tries its stronghold is in Iran; it is categorized as Endangered on
the IUCN Red List (Khorozyan et al., 2005;Khorozyan,2008).
The leopard population in Iran is estimated to be 550–
850 (Kiabi et al., 2002) and its range extends over 850,000
km
2
wherever sufficient prey and protected habitat is
present (Kiabi et al., 2002; Firouz, 2005). It is essential to
count and determine the population structure of this
predator so as to verify its status, monitor population
viability, identify the effects of natural and human factors
on the species and to determine the impact of the decline of
the leopard on the ecosystem.
As leopards are wide-ranging their occupancy, which is
that part of the range (extent of occurrence) actually inha-
bited and used by the species, must be sufficiently large to
fulfil the species’ ecological requirements. To assess the
spatial distribution and viability of the species it is impor-
tant to estimate population occupancy, study the relation-
ship of the species with habitat fragmentation, examine the
effects of study design on occupancy estimation, and to
identify sites visited by leopards (Linkie et al., 2007; Gruber
et al., 2008).
Bamu National Park is one of the most important habitats
for the leopard in Iran. The Park has a long history of
conservation, access for research is relatively easy compared
to other leopard habitat in Iran, and sightings of leopards in
the area are relatively common. However, fragmentation
from human encroachment is ongoing and there is a high
rate of poaching in the area. Here we report the population
size and structure, and occupancy and visitation rates, of the
leopard in Bamu National Park. The study was designed to
provide data for future research on, and conservation of, the
species. This is the first study of a leopard population in Iran
using camera-trapping and modelling, and is one of only
a few carried out on this species worldwide (Henschel & Ray,
2003; Kostyria et al., 2003; Spalton et al., 2006).
Study area
The 486 km
2
Bamu (also transliterated as Bamoo or
Bamou) National Park is in Fars Province, north-east of
Shiraz (Fig. 1; Darvishsefat, 2006). Established in 1967 and
upgraded to National Park in 1970, it encompasses three
parallel mountain ridges extending in an east-west di-
rection and the hilly plains between (Plate 1). Topograph-
ically Bamu is confined to the northern macro-slope of the
Zagros Mountains. Elevations are 1,600–2,700 m. Climate is
semi-arid temperate and continental (Darvishsefat, 2006).
A
RASH
G
HODDOUSI
* (Corresponding author), A
MIRHOSSEIN
K
H
.H
AMIDI
,
T
AHER
G
HADIRIAN
and D
ELARAM
A
SHAYERI
Plan for the Land Society,
Tehran, Iran. E-mail ghoddousi@plan4land.org
I
GOR
K
HOROZYAN
Zoological Institute, St Petersburg, Russia, and WWF
Armenia, Yerevan, Armenia
*Current address: 106 John Smith Hall, Silwood Park, Buckhurst Road, Ascot,
Berkshire, SL5 7PY, UK
Received 6January 2009. Revision requested 4February 2009.
Accepted 12 March 2009.
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Mean annual precipitation and temperature are 400 mm
and 16
o
C, respectively. The general vegetation type is arid
scrubland dominated by almonds Amygdalus spp. and
thorns Crataegus spp.. The flora comprises 350 vascular
plant species, including 51 endemics, and the fauna includes
143 species of vertebrates (Darvishsefat, 2006). The western
part of Bamu is separated by the Isfahan–Shiraz highway
and its large mammalian fauna has been depleted by
poaching (Area 6in Fig. 1). Only the eastern part (356
km
2
) is effectively protected (Nowzari et al., 2007). The
leopard prey species in eastern Bamu are wild sheep Ovis
spp., wild or bezoar goat Capra aegagrus, wild boar Sus
scrofa, Indian porcupine Hystrix indica and Cape hare
Lepus capensis; all are relatively common. The goitered
gazelle Gazella subgutturosa is confined to the 60-km
2
Chahmahaky Plain (Nowzari et al., 2007).
Methods
Camera-trapping was carried out in eastern Bamu during
28 September–20 October 2007,2–23 November 2007,19
December 2007–11 January 2008,4–24 February 2008 and
25 February–17 March 2008 for a total of 106 days, using
passive camera-traps (Stealth Cam MC2-GV; Stealth Cam
LLC, Grand Prairie, USA) with 35 mm film. In total we used
30 camera-traps but two failed and eight were stolen. For
convenience the area was divided into five topographically
distinct areas and these were camera-trapped sequentially
(Areas 1–5in Fig. 1), as in other camera-trapping studies
(Henschel & Ray, 2003; Karanth et al., 2004; Soisalo &
Cavalcanti, 2006). To maximize capture probabilities over
the largest possible area, camera-traps were set up along
established leopard trails on ridge tops and in valleys as
evenly and closely as possible so as to capture all leopards
(Fig. 1). The spacing between camera-traps was 2–2.5km,
which corresponds to the diameter of the smallest leopard
home range (8km
2
; Marker & Dickman, 2005). Cameras
were mounted at c. 40 cm above the ground on posts made
of flat stones and sometimes on trees. Each camera-trap
station consisted of 2camera-traps placed on the opposite
sides of a trail so as to photograph both flanks of leopards
(Henschel & Ray, 2003). The camera-traps were set for
24-hour operation, two photographs per sensing, and with
a1-minute delay between subsequent photographs. Sites of
FIG. 1 The location of the camera-trap stations in Areas 1–5. White circles are the stations with captures of leopards Panthera pardus
saxicolor, with individual IDs, and black circles are the stations without captures. Leopard IDs: M1, adult male; M2, subadult male; F1,
female with cub; F2 and F3, adult females; F4, subadult female. The circle on the inset indicates the location of Bamu National Park in
southern Iran.
A. Ghoddousi et al.552
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all camera-traps were recorded with a global positioning
system, and a map of locations drawn using the geogra-
phical information system ArcGIS v.
9.0
(ESRI, Redlands,
USA).
The design of our study was identical to that described in
Karanth et al. (2004). As we had 20 camera-traps and had to
cover five areas with similar sampling effort, we set up the
camera-traps in 20 sites (10 camera-trap stations, with 2
cameras per station) within each area, for 21 successive days,
which corresponded to battery life. Thus there were 21
sampling occasions each of which combined captures from
5days of camera-trapping (1day from each area).
Photo-captured animals were sexed from external gen-
italia (males), presence of cubs (females) and general
appearance (much larger body size, plump muzzle, wider
chest and front limbs in males). Individuals were recog-
nized from unique spot and rosette patterns on flanks and
limbs (Henschel & Ray, 2003).
Analysis
We constructed an X-matrix of capture histories for
individual leopards, excluding the dependent cub (05no
captures, 15captures) and used the software CAPTURE v.
2.0
(Colorado State University, Fort Collins, USA) to
estimate leopard abundance and check the hypothesis of
population closure (Karanth, 1995). Population density was
estimated by dividing the estimator of population size by
the effective sampled area that included the area confined
within the outer camera-trap stations and the boundary
strip. The boundary strip was calculated as half of the mean
maximum distance moved (MMDM), i.e. the arithmetic
mean of the maximum distances moved (MDM) by
individuals between recaptures (Henschel & Ray, 2003;
Karanth et al., 2004; Jackson et al., 2006; Soisalo &
Cavalcanti, 2006).
Independent captures were defined as (1) consecutive
photographs of different individual leopards; (2) consecu-
tive photographs of individual leopards taken .0.5hours
apart; and (3) non-consecutive photographs of individual
leopards. A relative abundance index was calculated as the
ratio of independent captures to 100 trap-nights of sam-
pling effort. Sampling effort was calculated as the sum of
days that all camera-trap stations operated (O’Brien et al.,
2003).
To estimate the minimum values of sampling effort
(trap-nights), sampling efficiency (number of independent
pictures) and study area required to obtain an accurate
estimate of leopard density, we plotted these variables
against density across the progressive sum of the land mass
of the sampling areas (Yasuda, 2004; Maffei & Noss, 2008).
The sequence of increasing areas was: Area 1(78.8km
2
),
Areas 1–2(157.3km
2
), Areas 1–3(202.1km
2
), Areas 1–4
(279.8km
2
), and Areas 1–5(356.1km
2
). Correlations
between sampling effort, sampling efficiency and study
area were examined over the individual areas to check for
any collinearity.
We determined the naı
¨ve and actual estimates of leopard
occupancy (w) as described by Linkie et al. (2007). For this,
we used the single-season subprogramme of the software
PRESENCE v.
2.0
(Proteus, Dunedin, New Zealand). In the
naı
¨ve estimate non-detections mean true absence whereas
in the actual estimate non-detections mean either true
absence or non-detection at presence (false absence). In the
data input matrix we inserted 1s (leopard captures 5
detections) and 0s (no captures 5non-detections) across
the 21 sampling occasions (see above) and the 50 camera-
trap stations (10 stations per area 35areas, see above). We
used six pre-defined models that consider detection prob-
ability (p) either constant or survey-specific and the
sampled population as consisting of 1–3arbitrary groups
(MacKenzie et al., 2006).
PRESENCE was run with 15,000 bootstraps, with at least
10,000 required for the best performance (D. MacKenzie,
pers. comm.). The best output models were those that had
the lowest value of Akaike’s information criterion (AIC)
and the highest AIC weight (sum of AIC weights of all
PLATE 1 A typical landscape in Bamu National Park. Photo:
Mani Kazerouni.
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models 51; Table 1). Weighted means of pand wwere
calculated as:
p5X
N
n51
AIC weightn3pnð1Þ
w5X
N
n51
AIC weightn3wnð2Þ
where n51,2,...Nindicates the number of the best output
models (MacKenzie et al., 2006; Linkie et al., 2007). In this
study N54(Table 1).
To calculate the number of camera-trap stations (s) that
need to be set up to reach the best precision of w(SE 5
0.05), we used the equation:
s5w
SE2ð1wÞþ ð1pÞ
pKpð1pÞK1
"#
ð3Þ
where wand pare the weighted mean wand weighted
mean p, respectively, SE is the desired standard error of w,
Kis the optimum number of days that a given camera-trap
station operates and p*51–(1–p)
K
(MacKenzie & Royle,
2005; MacKenzie et al., 2006; Linkie et al., 2007). We
compared the number of days a camera-trap station
operated in this study (21 days, see above) and Kfrom
the reference table in MacKenzie & Royle (2005) and
MacKenzie et al. (2006) to check the closeness of these
parameters to each other.
Visitation rates were estimated by modelling in Double-
Track Excel workbook (Gruber et al., 2008). This simulates
visitation rates to particular sites based on occurrence
of fresh and/or aged signs (faeces, tracks); this can be
extended to photo-captures. To estimate the area-specific
visitation rates we inserted 1s for captures and 0s for no
captures across the 10 observations (camera-trap stations)
and the time interval of 21 days for each of the five study
areas. Statistical analysis was carried out with Excel 2003
(Microsoft Corp., Santa Rosa, USA) and SPSS v.
13.0
(SPSS
Inc., Chicago, USA).
Results
The total sampling effort of 1,012 trap-nights yielded 31
independent leopard pictures (22% of all wildlife photo-
graphs), resulting in a relative abundance index of 3.06
captures per 100 trap-nights. The total number of leopard
photographs was 72 but only 27 independent captures were
used in the X-matrix because of recaptures within an
occasion. We identified seven individual leopards across
the 21 sampling occasions: one adult male, one subadult
male, one adult female with cub, two adult females and one
subadult female (Plate 2).
Sampling efforts in each of the five areas differed
significantly (v
2
514.51,df54,P50.006) but this varia-
tion did not affect the numbers of individuals captured
(r
2
50.39,F
1,3
51.95,P50.257) or the numbers of in-
dependent leopard photographs obtained in each area
(r
2
50.25,F
1,3
51.02,P50.387). These differences in sam-
pling effort were caused by difficult access to some parts of
the study area, trails closed in winter, theft and malfunc-
tioning of some camera-traps.
The model M(o), implying constant capture probabili-
ties for individual leopards, had the best fit (model selection
criterion 51.0) and the model M(h) of heterogeneity in
capture probabilities was ranked second (0.97). We chose
M(h) because its population estimator is robust and most
relevant to solitary felids in comparison with M(o) (Karanth
et al., 2004;Maffeietal.,2004). The wide-ranging adult male
had a much higher chance of being photographed (12 out of
21 sampling occasions, 57.1%) in comparison with his
conspecifics (females on 2–4occasions, 9.5–19.0%; subadult
male on three occasions, 14.3%). The goodness-of-fit of M(h)
was statistically significant (v
2
527.13,df520,P50.13).
The jackknife was the best estimator of population abun-
dance. The assumption of population closure was not
violated (z5-0.22,P50.41).
The number of leopards in Bamu estimated by the M(h)
model and jackknife estimator was 6.00 –SE 0.24 individ-
uals (95% confidence interval 6–6). The narrow confidence
interval is probably an artefact of the small sample size
(Karanth, 1995; Haines et al., 2006). Average capture
TABLE 1Results of occupancy modelling (see text for details) of the leopard Panthera pardus saxicolor population in Bamu National
Park.
Model AIC
1
AIC
1
weight Model likelihood p
2
–SE w
3
–SE
One group, constant p278.03 0.80 1.00 0.05 –0.01 0.56 –0.13
Two arbitrary groups, constant p282.03 0.11 0.14 0.05 –0.01 0.56 –0.13
One group, survey-specific p
4
282.66 0.08 0.10 0.05 –0.04 0.54 –0.12
Three arbitrary groups, constant p286.03 0.01 0.02 0.05 –1.86 0.56 –2.44
Weighted mean value 0.05 0.56
1
Akaike’s information criterion
2
Detection probability
3
Occupancy
4
Calculated as the arithmetic mean of the survey-specific pvalues
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probability for individual leopards in a sampling occasion
(p
ˆ)was0.21. The MDMs were 0.62–12.38 km and the
MMDM was 5.01 –SE 1.72 km. The boundary strip was
2.50 –SE 0.86 km. The effective sampled area was 321.12 km
2
and thus the leopard density was 1.87 –SE 0.07 individuals per
100 km
2
. This density was attained at a minimum sampling
effort of 400 trap-nights, minimum sampling efficiency of
seven independent pictures and a minimum study area of
150 km
2
(Fig. 2). Sampling effort, sampling efficiency and
study area were uncorrelated (P was 0.25 to 0.93).
The best-fit occupancy models show that detection
probability for leopards at camera-trap stations was con-
stant; the population was represented by a single group and
leopard occupancy was similar across the models (Table 1).
Weighted mean occupancy was 0.56 and therefore leopards
occupied c. 56% of the study area in Bamu. Because of low
detection probability, estimated occupancy was, at 47%,
higher than the naı
¨ve estimate of occupancy (19 out of 50
camera-trap stations, i.e. 38%).
The 21-day duration of camera-trapping at each camera-
trap station was almost the same as the Kthat equals 20 daily
surveys per site with p50.1and w50.6, the tabulated ad hoc
values of pand wclosest to the empirical ones estimated in
this study (MacKenzie & Royle, 2005; MacKenzie et al., 2006).
Therefore in equation (3)weusedK521 days. To achieve
a model precision of SE 50.05, based on the weighted mean
w50.56 and weighted mean p50.05 (Table 1), 368 camera-
trap stations would be required in the study area.
Visitation rates ranged from a minimum of 0.01 visits
per day in Area 1to a maximum of 0.05 visits per day in
Area 3and the rates in Areas 2,4and 5were 0.02 visits per
day. Visitation rates were not correlated with the numbers
of individual leopards camera-trapped in the areas (r
2
5
0.43,F
1,3
52.31,P50.226).
Discussion
Our results indicate there are seven leopards in Bamu
National Park. In the late 1970s their number was estimated
to be 15–20 (Kiabi et al., 2002). Whether these figures
indicate a population decline cannot be ascertained as the
two studies used different methodologies. Our estimates
show that camera-trapping over 150 km
2
for 400 trap-nights
that obtains seven photographs of leopards gives the same
unbiased estimate of leopard density as does a survey cov-
ering all of Bamu (Fig. 2). We did not find the thresholds
or curve asymptotes that would indicate a stabilization of
leopard densities in relation to increase in study area, sam-
pling effort and sampling efficiency. Although this could
indicate an insufficiently large study area and overestimation
of density (Maffei & Noss, 2008), lack of stabilization in this
case is most likely caused by differences in leopard numbers
photo-captured in each area, which inevitably affects area-
specific densities in a small population.
PLATE 2 Examples of leopard photo-captures in Bamu National
Park: (a) adult female, (b) adult male, (c) adult female and (d)
subadult female. Photos: Plan for the Land Society.
Leopard in Bamu National Park, Iran 555
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Low detection probability (Table 1) brings about a high
rate of non-detections in the areas of actual presence (false
absence) that, if ignored, underestimates leopard occu-
pancy by 47%. Thus the area inhabited by leopards in this
protected area is much larger than that predicted from
photographic capture sites alone; a pattern commonly
found in rare and elusive species (MacKenzie et al., 2006;
Linkie et al., 2007).
At 1.87 –SE 0.07 per 100 km
2
the leopard density in
Bamu is higher than elsewhere in Iran and than in two
other areas where it was estimated by camera-trapping:
Jabal Samhan Nature Reserve in Oman (0.4individuals
per 100 km
2
; Spalton et al., 2006) and the Russian Far East
(1.1–1.2individuals per 100 km
2
;Kostyriaetal.,2003). In-
tensive year-round use of territorial markers such as scrapes is
further evidence of high leopard density in Bamu (Ghoddousi
et al., 2008a). This density is, however, lower than in an equa-
torial rainforest in Gabon (2.7–12.1individuals per 100 km
2
)
where the same photographic capture-recapture technique
was employed (P. Henschel, pers. comm.).
Poaching and habitat fragmentation are threats to the
existence of leopards in Bamu (Ghoddousi et al., 2008b).
Although this National Park is well-protected, with nu-
merous and capable game wardens (46 covering the 356.1
km
2
), occasional cases of poaching still occur. Rapid in-
dustrial and agricultural development beyond its boundaries
makes Bamu an isolated island surrounded by the Isfahan–
Shiraz highway and a refinery to the west, Shiraz city and its
suburbs to the south, and agricultural lands to the north and
east (Fig. 1; Ghoddousi et al., 2008b). Habitats in Bamu are
affected by illegal grazing in the north-east and unregulated
local tourism along the Park edge. Such intensive fragmen-
tation and encroachment limits space and dispersal routes
for leopards in Bamu (Ghoddousi et al., 2008b).
We detected spatial segregation of individual leopards in
relation to human factors. The subadult male was photo-
captured only in south-western Bamu, which is the part of
Bamu most fragmented by industrial barriers. The subadult
female and an adult female were photo-captured in the
south-east close to agricultural lands. The adult male and
most of the adult females shared the central part of Bamu,
least affected by human pressures (Area 3).
The relatively high leopard density in Bamu could be
a result of a connection with other areas of Fars Province by
corridors such as along the Kor river from the easternmost
part of Bamu to Bakhtegan National Park and Wildlife
Refuge, where the presence of leopards has been confirmed
(Darvishsefat, 2006). Leopard conservation measures in
Bamu, partly already underway, need to focus on mitiga-
tion of the effects of habitat fragmentation and degradation,
and anti-poaching activities and awareness-raising.
The Persian leopard project in Bamu is ongoing and is
now focused on capacity building and educational pro-
grammes for villagers and farmers around the National
Park. In spring 2009, with the collaboration of governmen-
tal organizations and international funders, 1,400 students
in 14 villages around Bamu were educated on the impor-
tance of the leopard and the National Park. Research
priorities in Bamu are a detailed study of the species’
spatial distribution and a radio telemetry study of possible
connections to other populations.
Acknowledgements
We thank the personnel and volunteers of the Plan for the
Land Society and the Fars office of the Department of
Environment, especially H. Zohrabi (Head of the Biodiver-
sity Bureau), for their continued support of this project.
FIG. 2 Curvilinear relationships between leopard density and (a)
sampling effort, (b) area size and (c) number of independent
pictures in Bamu National Park (Fig. 1).
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We thank B.H. Kiabi, B.F. Dareshouri and P. Henschel for
provision of information, D. MacKenzie for assistance in
using PRESENCE, and B. Gruber for his DoubleTrack
workbook. Financial support for this project was generously
provided by individual Iranian donors.
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Biographical sketches
ARASH GHODDOUSI is a member of Plan for the Land Society and
coordinator of the Persian leopard project in Iran. Since 2005 he has
been carrying out research on threatened mammals and their
conservation and is a member of the IUCN/SSC Cat Specialist group.
AMIRHOSSEIN KHALEGHI HAMIDI is a research associate of the
Persian leopard project. He is also involved in wildlife management
and community-based conservation of large carnivores in Iran.
TAHER GHADIRIAN is a wildlife specialist involved in several research
projects on the Asiatic cheetah and Persian leopard. DELARAM
ASHAYERI is a field zoologist and manager of a community-based
Asiatic cheetah conservation project, and she also participates in the
Persian leopard and other wildlife projects. IGOR KHOROZYAN
carries out research on the Persian leopard and its conservation in
Armenia. He cooperates with the Plan for the Land Society and
prepared the Persian leopard assessment for the 2008 IUCN Red
List.
Leopard in Bamu National Park, Iran 557
ª2010 Fauna & Flora International,
Oryx
, 44(4), 551–557