Modeling the distribution and movement intensity of Arabian Leopard Panthera pardus nimr in the Arabian Peninsula - Authorea
←
→
Page content transcription
If your browser does not render page correctly, please read the page content below
Modeling the distribution and movement intensity of Arabian
Leopard Panthera pardus nimr in the Arabian Peninsula
M. Zafarul Islam1 , Alexander Gavashelishvili2 , Luka Kokiashvili3 , Ahmed al Boug1 ,
Abdullah as Shehri1 , A. Townsend Peterson4 , and Daniel Jiménez Garcı́a5
1
Prince Saud al Faisal Wildlife Research Center
2
ILIAUNI
3
Ilia State University
4
University of Kansas
5
Benemérita Universidad Autónoma de Puebla
May 5, 2020
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Abstract
Aim: Our research attempts 1) to link the distribution and movement rate of critically endangered Arabian leopard (Panthera
pardus nimr) to environmental variables varying in space and time, and 2) thus to identify environmental constraints and
priority areas for the recovery of Arabian leopard. Location: The Arabian Peninsula. Methods: Generalized linear and additive
models were used to fit leopard presence/absence locations to environmental variables. Movement rates between the modeled
polygons of leopard presence were inferred and mapped using the isolation-by-resistance model, where probability values of the
species distribution model were treated as those of conductance. Results: Currently Arabian leopard prefers to live and move in
terrain that has high NDVI and is difficult for humans to reach. Main conclusions: Our results suggest that 1) Arabian leopard
accumulated genetic and phenotypic differences from its conspecifics at a series of glacial maxima during the last glacial period in
the Yemeni refugium, from where it expanded elsewhere in the Holocene warming following the expansion of suitable landscape
types, (2) humans expanded too, eventually restricting the source populations of Arabian leopard to an area intersecting eastern
Yemen and western Oman today, (3) Most of the survey effort to detect and ensure the survival of the leopard in the peninsula
has taken place outside the polygons identified by our models as core areas for the species. Our models would serve as a
tool for the management of the species in conservation efforts such as an Arabian Leopard and associated species conservation
in western Arabia project. Keywords: Arabian leopard, Panthera pardus nimr, Arabian Peninsula, Distribution, Movement
intensity, Source population, Corridors, the Yemeni refugium.
INTRODUCTION
As mentioned in ‘The Sixth Extinction: An Unnatural History’ the majority of threatened species experi-
encing greatest threats due to human intervention and the way the mass extinction event is reaching across
diverse ecosystems is alarming (Kolbert, 2014; Hilton-Tailor, 2000). Humans have persecuted large preda-
tors for centuries, reducing their distributions and altering niches, species such as Arabian LeopardPanthera
pardus nimr , Arabian Wolf Canis lupus arabs ,Striped Hyena Hyaena hyaena sultana and many others in
Saudi Arabia and rest of the Arabian Peninsula. Around 100 years before Asian Lion Panthera leo persica
and Asian CheetahAcinonyx jubatus venaticus were extinct from the region (Harrison, 1968; Schnitzler &
Hermann, 2019; Harrison, 1983; Harrison and Bates, 1991). The removal of these top predators from much
of the natural world has had diverse direct and indirect effects (Esteset al ., 2011). Some have been extermi-
nated, most others occur as small populations with drastically reduced geographic ranges due to social and
political disruption of carnivore conservation programs.
1
International Union for Conservation of Nature (IUCN) documents that out of nine species of leopards, five
are listed as endangered or critically endangered (Breitenmoser et al ., 2010). The greatest challenge in
today’s world is to restore those species, which are on the verge of extinction including the Arabian Leopard
Panthera pardus nimr (Hemprich and Ehrenberg, 1833), which is one of the Arabia’s flagship predators and
listed as Critically Endangered (Mallon et al ., 2008; Boug et al ., 2009; Islam et al., 2015; 2018). It has
an estimated population of 100–250 across its entire range in the Arabian Peninsula (Spalton et al ., 2006;
Spalton and Al Hikmani, 2014; Breitenmoser et al ., 2006; Islam et al ., 2015, 2018) and is also considered
to be a genetically distinct subspecies (Mallonet al ., 2008). Known locally as Al nimr al-arabi , this leopard
subspecies is small in size, well adapted to desert habitats and endemic to the Arabian Peninsula. It once
occupied the mountainous rim of the Arabian Peninsula, albeit at low densities given the harsh environment
and limited prey base.
Historically, much of the leopard’s range was located within the Kingdom of Saudi Arabia; this has decreased
by about 90% since the nineteenth century (Judas et al ., 2006; Boug et al., 2009, Islam et al ., 2018). In
1982 a live-leopard was seen in Wadi Hiswa in the Asir (Gasperetti et al ., 1985), while Nader (1989) reported
on killings of leopards and the collection of leopard remains in the 1970s and 1980s and who mentioned that
the cat would probably be confined to South-western Highlands in the Kingdom. From field surveys in Asir
region (SW Highlands) Biquand (1990) mentioned the Arabian Leopard probably present although they
made no sightings, while Nader (1996) reported a small population of the leopard still present in the Hijaz
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
(northwestern highlands) and also in South-west, although no live animal was reported. Recently a number
of field surveys were carried in the western highlands of Saudi Arabia (Islam et al ., 2011, 2018) but not
seen. Al Johny (2007) recorded 62 sites where leopard presence was likely in Saudi Arabia. The majority
of these sites were checked between 2010 to 2017 and did not confirm of presence of the species, while one
male was poisoned in February 2014 (Islam et al., 2018).
From 1833 to 2018, there are records of more than 300 (live animals, pugmarks, carcasses, reports) in the
entire region of Arabian Peninsula (Islam et al., 2018) and the population may have declined by over 90%
since the beginning of the 19th century, with the population reaching its presently highly precarious status
at the start of the 21st century.
The population of Arabian Leopard in Oman is believed to be a significant breeding population especially
in Dhofar mountains in the southern part of the country, where prey populations are healthiest (Stuart &
Stuart, 2007). This is one of the important places in the Arabian Peninsula where a sizeable reserve (the
4,500 km2 Jabal Samhan Nature Reserve) provides excellent habitat for the leopard. Leopards were also
sighted in the Musandam Peninsula (Spalton & Al-Hikmani, 2006).
In Yemen, Arabian leopards used to occur in all mountainous areas especially south-western highlands and
eastern highlands bordering Oman (Al Jumaily et al ., 2006). Sanborn & Hoogstraal (1953) reported that
the species was rare but widespread while Harrison (1968) reports on several specimens of leopard from the
mountains around Aden and Beihan. Obadi (1993) reports the killing of leopards during the late 1970s and
early 1980s in the area of Lodar northeast of Aden. However, most capture records are from the area of Al
Wada’a about 120 km north of the capital Sana where Lagrot & Lagrot (1999) also reported signs of leopard
as well as captures. In subsequent years at least 10 wild caught leopards entered zoos in Sana’a or Ta’iz
(Budd, 2003) and at least nine were reported to have come from the Al Wada’a area (EPAA, 2000; Mellon,
2009).
Nowell & Jackson (1996) reported that there was a small population of 20 in 1970s in Negaev desert in Israel
and 11 individuals were identified in the country, based on genetic analysis of 268 scats collected (Perez et
al., 2006). One leopard was sighted in Sde Boker in 2007 and Granit (2016) reported that in 2010, a leopard
was sighted in the northern Arabah and since then it is considered extinct from Israel (Granit, 2016).
In Jordan, the last confirmed sighting of a leopard was in 1987 (Qarqaz and Baker, 2006).
In the United Emirates, Harrison (1968) reports the presence of the Arabian leopard from Musandam
mountain bordering Oman. Jongbloed (2001) reported that in 1986 one leopard was killed in the mountains
2
while in 1991 a male was caught alive near Masafi and in 1992 one was shot in Wadi Bih. Stuart & Stuart
(1995) mention that around 20 leopards may be there in mountains that were confirmed from the tracks
from field survey in 1995. In Ras alKhaimah in 1999 and 2000 found some signs of leopard but were not
confirmed by camera-traps deployed at the same time (Llewellyn-Smith, 2002).
Depletion of the leopard’s prey base and retributive killing are the greatest threats (Islam et al ., 2011; 2015;
2018; Islam and Boug, 2017). Excessive illegal hunting has greatly depleted key prey populations like the
Nubian ibex (Capra nubiana ), Rock hyrax (Procavia capensis ), Arabian mountain gazelles (Gazella arabica
) and Arabian hare (Lepus capensis arabs ) (Al-Johny, 2007; Islam et al., 2018; Judas et al ., 2006). As
a consequence, the leopard has become increasingly dependent upon domestic livestock for its subsistence,
which in turn leads to violent retaliation by herders who lose animals. Moreover, there are some reports of
the sale of furs and occasionally live animals sold in local markets in parts of the region, where leopard fat
is valued by some local people for its perceived medicinal properties (Judas et al ., 2006).
Habitat modelling for the Arabian Leopard is needed due to fragmented populations in the range states
(Breitenmoser et al ., 2010) and it is one of the important tools for conservation planning (Rougetet al .,
2006; Beier et al ., 2008). The species habitat model can be used to infer movement corridors of the Arabian
Leopard from its significant breeding population in Oman to other suitable areas in Yemen and Saudi Arabia.
These corridors help the leopards, especially sub-adults to move distant areas in search of food, territory
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
and mate and that’s how the species’ gene flow phenomena occur (Haddad et al ., 2003; Dixon et al ., 2006;
Roberts & Angermeir, 2007).
Studies on population connectivity is one of the important ways to understand species movements, and
becomes extremely indispensable to know the niches that also are helpful in maintaining genetic diversity and
demographic exchange in regional populations (Fahrig, 2003; Crooks and Sanjayan, 2006; Cushman, 2006;
Cushman et al., 2013). Studies show that species shift their geographic ranges in response to conservation
threats that include human-wildlife conflicts and also climate change (Heller and Zavaleta, 2009). Species,
especially large carnivores, which require large ranges to meet ecological and energetic needs are especially
prone to to human-wildlife conflicts. Areas such as mountain regions of Arabian Peninsula, where human
populations occupying remote areas, species survival disseminated and habitat is not protected at the extent
that need to be. Maintaining connectivity across a wide landscape would help in expanding the existing
network of protected areas, as suggested (Santini et al., 2016; Bruner et al., 2004; Schmidt-Soltau and
Brockington, 2007). Since the species is surviving in largely unprotected areas, nevertheless, maintaining
corridors across species region might complicate it as majority of the area is privately owned.
Spear et al., (2005) and Cushman et al., (2012) documented that the predicted corridor network for many
taxa depend on three important aspects: (1) the distribution and abundance of the species, (2) the dispersal
and movement ability of the species and (3) the pattern of differential movement cost, or resistance, across
the landscape.
Based on these, we designed our studies with the objective to generate the distribution and movement
intensity models of Arabian leopard.
MATERIAL AND METHODS
Study area is comprised of five major mountain ranges: 1) the south-western highlands in Saudi Arabia
up to Yemen; 2) the Hijaz Mountain to the northwest of Saudi Arabia; 3) Highlands in Jordan, Israel and
Palestine; and 4) Al Hajar mountains in Oman and United Arab Emirates & 5) Hijaz Mountain in Saudi
Arabia. The South-western Highlands consists of well-developed Juniperus forest remains intact above 2,000
m and deciduous woodland, often characterized byAcacia , with many endemic plants (CEPF, 2012) along
with deep valleys (wadis ). The SW Highlands is connected gradually with the NW mountains called Hijaz
(2,100 m) along the Red Sea coast. This range includes Median Mountains in the extreme north-west of
Saudi Arabia. In Israel and Palestine contains the central highlands with Mount Meron 1,208 m. East of
central highlands lies the Jordan Rift Valley contains Jordan River and many lakes provide habitats to many
animals and Arabian Leopard used to persists. Al Hajar Mountains and Musandam Peninsula form one
3
contiguous spread in northeastern Oman and the eastern United Arab Emirates. Jabal Dhofar is in western
par to Oman is continuous mountain range to eastern Yemen where leopard population is still surviving.
Modeling the species distribution: We explored the distribution of Arabian Leopard (Panthera pardus
nimr ) as a function of various environmental variables (Table 1 ). The selection of variables was based on
documented species–habitat associations in west and central Asia (Gavashelishvili and Lukarevskiy 2008).
To model the distribution, presence and absence locations were fitted to the environmental variables. 96
locations where our camera traps failed to detect leopard from 2011 through 2015 were considered absences.
Even though these cameras were deployed in many places of Saudi Arabia, where leopards had been recorded
historically, none were camera-trapped during the study period. We used camera trap locations that were
> 1,000 m from neighboring locations because the coarsest cell size in our analysis was 500 m (Table 1
), and hence to avoid the repeated sampling of environmental variables in a grid of 500 × 500 m cells, the
minimal distance will be SQRT(2 × 5002 ) = 707.1068 m. This spacing would reduce the effects of spatial
autocorrelation as well. Thus, we obtained 44 absence locations. Presences were obtained by generating 44
random locations within the year-round ranges of leopard identified by other studies (Mazzolli, 2009; Spalton
and Al Hikmani, 2014; Fig. 1 ).
Generalized linear models (GLMs) and generalized additive models (GAMs) were used to fit leopard pre-
sence/absence to environmental variables using the mgcv package (Wood, 2011) in R version 3.5.2. (R Core
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Team, 2018). We used GAMs because they are able to find nonlinear and non-monotonic relationships. GLMs
and GAMs were fitted using a binomial family with a logit link function. For GAMs penalized thin plate
regression splines were used to represent all the smooth terms. The restricted maximum likelihood (REML)
estimation method was implemented to estimate the smoothing parameter because it is the most robust of
the available GAM methods (Wood, 2011).
Model and variable selection were performed by exploring all possible subsets of environmental variables,
where pair-wise correlations between variables were less than 0.9. To get subsets of environmental variables,
variable combinations were generated using the gtools package for R (Warnes et al., 2015). The predictive
power of the models was evaluated through a 10-fold cross-validation. The cross-validation of many models
was handled through R’s parallelization capabilities (Microsoft Corporation and Weston, 2017 & 2018). The
best models were selected by the area under the receiver operating characteristic (ROC) curve (AUC). For
the final decision we also checked concurvity between model terms and between each term and the rest of
the model using the mgcv package.
All gridded environmental layers (raster layers) reflecting the situation in 2018 were resampled to a resolution
of 90m using the nearest neighbor assignment technique of QGIS Desktop 3.4.11-Madeira. We used this cell
size to avoid missing the importance of environmental variables at the finest resolution of ˜90 × 90 m (Table
1 ) in our analysis. The best leopard distribution model was predicted to the raster layers using the raster
package (Hijmans, 2016) in R version 3.5.2. Probabilities of leopard presence were dichotomized into presence
and absence at the cutoff value that maximized the sum of sensitivity and specificity. Then we generated 1000
random points within the inferred presence polygons that represented vast hyper-arid almost barren areas
where leopard populations have never been recorded, and using them as absences, ran the presence/absence
analysis again in order to refine the species distribution model. Thus, we obtained two-step final model
consisting of two models: (1) the first one ignoring the negative impact of hyper-arid low productivity areas
on leopard distribution and (2) the second one considering the negative aridity impact. To map the final
probability and presence/absence of leopard distribution, we multiplied probability maps and dichotomized
maps, respectively.
Also, we inferred and mapped movement rates between the inferred polygons of leopard presence using
the isolation-by-resistance (IBR) model via the Circuitscape 4 software (McRae et al., 2013). IBR is based
on the resistance distance that, as a predictor of movement impedance between populations, is likely to
perform better than Euclidean or least-cost path-based distance measures (McRae, 2006). Unlike Euclidean
and least-cost distances, the IBR algorithm assumes that a disperser does not have complete knowledge
of the landscape being traversing and can use multiple paths to reach a destination. The program models
4
multiple random walk paths between populations across a resistance or conductance grid that is a raster
map, wherein the value of each cell indicates the relative difficulty (or resistance) of moving through that cell.
The program treats the raster map as an electrical circuit, where cells with finite resistances are converted
to nodes, cells with infinite resistance (absolute barriers) are dropped, and adjacent cells with zero resistance
are consolidated into a single node. In this electrical circuit, adjacent nodes are connected by resistors, with
resistances equal to the mean of cell values between a pair of orthogonal neighbors, and the mean resistance
multiplied by the square root of 2 between a pair of diagonal neighbors to reflect the greater distance
between cell centers. By injecting 1 amp of current (by default) into each focal node (population) and using
Kirchoff’s and Ohm’s laws, the program calculates effective resistances (i.e., resistance distances), current,
and voltages that can then be related to ecological processes (e.g., individual movement and migration rate).
The current through nodes or resistors can be ecologically interpreted as expected net movement probabilities
(i.e., movement rates) for random walkers moving through a heterogeneous landscape. We used probability
values of our final distribution model as those of conductance in the IBR algorithm.
Table 1 .—Variables used for modeling the distribution of Arabian Leopard (Panthera pardus nimr ).
Variable Description
presence Leopard present or absent
elev SRTM elevation grid of 90-m cells (Jarvis et al., 2008)
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Slope tan Tangent of slope (°) calculated from a SRTM elevation grid of 90-m cells (Jarvis et al., 2008)
TRI Terrain Ruggedness Index, calculated in a rectangular neighborhood of 1x1 km using QGIS Desktop 3.4.11-Ma
cd Slope-weighted cost distance from human roads and settlements as a proxy for human disturbance, calculated
ndvi min Annual minimum of normalized difference vegetation index of a 16-day interval as a proxy for food, water and
ndvi max Annual maximum of normalized difference vegetation index of a 16-day interval as a proxy for food, water and
ndvi mean Annual mean of normalized difference vegetation index of a 16-day interval as a proxy for food, water and cove
ndvi med Annual median of normalized difference vegetation index of a 16-day interval as a proxy for food, water and co
SNOW Annual sum of snow cover 8-day intervals as a proxy for movement impedance, extracted from 500-m MODIS
*
Obtained from https://search.earthdata.nasa.gov/search. At every presence/absence location, values of
the remotely sensed variables were measured for those date ranges in which the location was monitored.
5Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Fig. 1 .—The locations used for modeling the distribution of Arabian Leopard (Panthera pardus nimr ).
The map is projected to Albers Equal Area Conic; WGS: 1984; central meridian: 47.03187; standard parallel
1: 20.85766; standard parallel 2: 29.12067; latitude of origin: 24.98916.
We validated the leopard presence probability and movement intensity models, using confirmed records of
leopard presence locations dated from 1881 until 2018 (supplementary material: Arabian Leopard points -
Albers.csv). These records included a total of 321 locations, of which we selected those with a location
accuracy of < 100 m – that is, 256 locations. We divided these relatively accurate locations into 10-year
intervals starting with 2008-2018. Then values of our models measured at these locations were compared
across the periods using ANOVA and TukeyHSD (Tukey Honest Significant Differences). The logic behind
this validation procedure was that if our models truly explained leopard distribution and movement intensity,
then their values were to be greater for a period that was closer in time to that of our study. This was expected
because environmental variables – distance from human presence to much greater extent than other ones –
were spatially rather different farther back in time from our study period. For instance, human settlements,
especially vehicular access roads, were much fewer farther in the past. Initial analyses to derive distribution
6models using either all or various subsets of these 256 locations via both presence-only and presence/absence
methods showed much poorer results than those based on our training presence/absence locations. The
initial models were too optimistic about the species distribution in the Arabian Peninsula. This is probably
because the 256 locations were widely spread in time, and included many of those of vagrant and dispersing
leopards whose locations were not really limited by the environmental variables that we used in our analyses.
That’s why we used these locations to test rather than develop our models.
RESULTS
GLMs performed better than GAMs. The first step of our analysis suggested that slope-weighted cost
distance from human roads and settlements alone best explained the leopard distribution, while the variables
best explaining the leopard distribution in the second step were annual maximum of normalized difference
vegetation index and terrain ruggedness index (Table 2 ). The response was positive to all three variables
(Fig. 2 and Fig.3 ). The projection of the first model to the entire Arabian Peninsula misclassified large
hyper-arid areas as leopard presence (Fig. 4 ). Rerunning the analysis on absence points generated randomly
within these hyper-arid areas removed those areas (Fig. 5 ). Combining these two models by multiplying
them produced the final model that showed quite a realistic picture of current leopard distribution in the
region (Fig. 6 ). The final model suggested that the least fragmented habitat was in an area intersecting
eastern Yemen and western Oman. This area included top 7 largest neighboring polygons. During our study
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
period all breeding leopards were recorded within or near these polygons. So, we considered these polygons
as source populations and the others as sink populations in our next analysis in order to infer migration
intensity between these populations (Fig. 7 ). Values of our inferred leopard presence probability and
movement intensity at leopard presence records dating from 1881 until 2018 were significantly greater for
the period of 2008-2018 – i.e., the period that contained that of our study (Fig. 8 ).
Table 2 .—Summary of the best generalized Linear models (GLM) of leopard presence/absence fitted to
environmental variables. GLMs are fitted using a binomial family with a logit link function. See
Table 1 for descriptions of variables.
Step n Variables Parameter P-value AUC Cutoff Overall
estimate Accuracy
1 88 cd 0.006965 7.87e-06 0.936 0.712 0.83
Intercept -4.02119 2.74e-06
2 1044 ndvi max 0.01062 6.82e-06 0.996 0.434 0.994
TRI 0.03909 0.0063
Intercept -23.6521 5.16e-07
7Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Fig. 2 .— GLM-fitted relationship of leopard presence with slope-weighted cost distance from human roads
and settlements (cd). The gray band shows the 95% confidence interval. The distribution of the explanatory
variable by leopard presence/absence categories is shown on horizontal axes. The plot is made using the
visreg package (Breheny and Burchett, 2017).
Fig. 3 .— GLM-fitted relationships of leopard presence with annual maximum of normalized difference
vegetation index (ndvi max) and Terrain Ruggedness Index (TRI). The gray bands show the 95% confidence
interval.The distribution of the explanatory variables by leopard presence/absence categories is shown on
horizontal axes. The plots are made using the visreg package (Breheny and Burchett, 2017).
8Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Fig. 4 .— Modeled distribution of Arabian Leopard (Panthera pardus nimr ) constrained by human dis-
turbance. Black points are generated randomly within polygons misclassified as those of leopard presence,
and used to refine the species distribution model. The map is projected to Albers Equal Area Conic; WGS:
1984; central meridian: 47.03187; standard parallel 1: 20.85766; standard parallel 2: 29.12067; latitude of
origin: 24.98916.
9Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Fig. 5 .— Modeled distribution of Arabian Leopard (Panthera pardus nimr ) constrained by terrain pro-
ductivity and ruggedness. The map is projected to Albers Equal Area Conic; WGS: 1984; central meridian:
47.03187; standard parallel 1: 20.85766; standard parallel 2: 29.12067; latitude of origin: 24.98916.
10Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Fig. 6 .— Modeled distribution of Arabian Leopard (Panthera pardus nimr ) constrained by human distur-
bance, terrain productivity and ruggedness. The map is projected to Albers Equal Area Conic; WGS: 1984;
central meridian: 47.03187; standard parallel 1: 20.85766; standard parallel 2: 29.12067; latitude of origin:
24.98916.
Fig. 7 .—Isolation-by-resistance model of cumulative current flow (analogous to probability of gene flow
or migration rate) highlights potential corridors between the assumed source and sink populations, using
the probabilities of the species final distribution model as conductances to leopard movement. Warmer
color predicts higher migration rate normalized to a scale of 0 - 100. Green polygons are assumed source
populations.
Fig. 8 .—Boxplots of values of our inferred leopard presence and movement intensity at leopard presence
records dating from 1881 until 2018.
DISCUSSION
11Leopards progenitors might have started appearing in the Arabian Peninsula from Africa through the Sinai
Peninsula, during the late Pliocene or early Pleistocene (Uphyrkina et al., 2001) and Hedges (2000) mentioned
that the leopard might have followed the same route, approximately at the same time, as humans. During
this period, the Red Sea was already formed (Thompson, 2000). Leopard populations should have spread
all over the Arabian Peninsula, shrinking or expanding their ranges due to natural climate change (Burton,
1995). In the southern half of the Arabian Peninsula only the western part of present-day Yemen harbored
landscape types suitable for the survival of leopard source populations during a series of glacial maxima
(Fig. 9 , Gavashelishvili and Tarkhnishvili, 2016). We hypothesize that the most likely major event in
the geological past that could be a key in triggering the known genetic and phenotypic differences between
Arabian leopard and the other subspecies by reproductively isolating their populations from each other was
a succession of glacial maxima restricting the Arabian population to western Yemen. Our hypothesis about
the Yemeni refugium could also be supported by the higher levels of genetic diversity and unique alleles in
leopards of Yemeni origin, compared to leopards of the current stronghold – that is, the Dhofar mountains
of Oman (Al Hikmani, 2019). In the Holocene warming following the last glaciation suitable landscape
types expanded, and so did leopard populations from the Yemeni refugium to the limits suggested by our
second model (Fig. 5 ). As humans also expanded in and to the peninsula, grew in numbers and started
transforming landscape types through ever-evolving technologies, leopard distribution range has shrunk to
the limits suggested by our final model (Fig. 6 ). According to our analyses the current distribution of
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
leopard in the Arabian Peninsula is explained by human disturbance, terrain productivity and ruggedness,
which is in agreement with the species habitat requirements in west and central Asia (Gavashelishvili and
Lukarevskiy, 2008). Apart from these three explanatory variables snow cover considerably limits the species
distribution in west and central Asia. The reason we did not get a strong response to this variable in this
study is definitely due to the scarcity of snow cover in the Arabian Peninsula.
Fig. 9 . The inferred biomes at a series of glacial maxima during the last glacial period (Gavashelishvili
and Tarkhnishvili 2016). The Yemeni refugium – i.e. woodlands and grasslands surrounded by vast desert
— must have played a crucial role in survival and evolution of Arabian leopard.
Barring human disturbance (Fig. 5 ), our model predicts the leopard has potential areas in central highlands
in Palestine, Israel and northwest Saudi Arabia, where its historical presence was confirmed (Hemprich and
Ehrenberg, 1833; Lady Anne Blunt, 1881; Doughty, 1888; Carruthers, 1909) mainly along the pilgrimage
routes to Makkah in the Hail region and around Madayn Saleh. In comparison with current modeled
distribution (Fig. 6 ) the leopard presence in hilly Hijaz region confirms, although historically scares
(Gasperetti et al., 1985; Nader, 1989; Harrison and Bates, 1991; Gasperetti & Jackson, 1990).
Based on outputs, the habitat is larger and less fragmented westward in Oman and eastward in Yemen.
However, geographically limited studies in Jabal Dhofar in western Oman seems to support a good nucleus
that is probably due to enforced protection. Our findings show that eastern Yemen should have the species
stronghold too and research resources must be concentrated there as well.
Conservation issues: There is significant distributional decline of the leopard in range countries due to
various factors i.e., habitat degradation and fragmentation due to urbanization, road development, mineral
development, agricultural expansion, overgrazing and related activities; unrestricted trapping, poisoning
and hunting of leopards, often resulting from livestock depredation or other cause of human-wildlife conflict
(Islam et al., 2015; Islam and Boug, 2017, 2018;Fig. 4 ); depletion of natural prey base, particularly declines
in Arabian gazelle and ibex populations (Islam et al., 2011; Islam et al., 2015; Islam et al., 2017); insufficient
protection despite its full protected status under Saudi Arabia’s laws; and (5) lack of awareness among the
general public of the need for protection and conservation of the country’s wildlife, including the endangered
Arabian leopard. To restore the species in the wild, range countries need to step up to address these key
threats.
Habitat degradation and fragmentation:
Habitat fragmentation has long been recognized as a major factor responsible for threatened species loss and
12extinction through two main mechanisms; (a) the decrease of habitat size that directly leads to a decrease of
population size; and (b) the isolation of natural habitats which reduces the ability of individuals to disperse
between range nuclei (Wilcox & Murphy, 1985; Wilcove et al.,1986; Saunders et al., 1991; Wiens, 1994). It
has been documented that it would favor genetic destitution to demographic reductions as documented in
subspeciesP.p. orientalis, P.p. kotiya (Uphyrkina et al., 2001).
Our model predicts limited habitat left for the leopard, where degradation and fragmentation are important
issues to be considered by the range countries to save the species (Nowell & Jackson, 1996; Fahrig, 2003).
Although the leopards appear to be tolerant of humans, when they are in close contact with humans, they
get killed (Islam et al., 2018). Populations in the species range are too small to maintain stable growth, while
protected habitat exist in low percentage and the majority of leopards live outside these protected lands.
For example, the mountain of Musandam are marginal habitat for leopard, where prey-base have been
exterminated and over-browsing-grazing by livestock, changing agricultural practices and the rapid expansion
of roads and house constructions have all contributed to habitat degradation.
The model clearly shows very little habitat remains in Jordan, where historically people used traps made of
stone to catch leopards in 1999 and the species has been poached and even commercialised alive in part of
its current range (Qarqaz & Abu Baker, 2006).
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Saudi Arabia has not been spared from the habitat deterioration and loss for many species including the
Arabian Leopard. Habitat loss probably started many centuries ago with tree cutting to answer wood re-
quirements of growing iron and copper industries (Collenette, 1999). This was already largely developed in
the 16th century. Tree cutting and deforestation was still important in the southern Hijaz Mountains for
charcoal production till late 1990s (Biquand et al. , 1990) and continued by 2007, before it was banned by
that year to cut trees for charcoal or other purposes. In the last twenty years, governmental policy encour-
aged pastoralism extension and agriculture intensification to obtain national self-sufficiency in production
of meat and agricultural products (Judas et al. 2004). The Habitat of leopard is greatly affected by con-
struction of new tracks and roads in the escarpment that gave way to encroachment on once remote areas.
Considerable development of highway construction occurred between 1985 and 1990 (Gasperetti & Jackson,
1990) and continued till date and the majority of western highlands are well connected by tracks or roads,
even with a few houses in remote mountains. Along these new access roads, anarchic urbanization was and
is still implemented without any Environmental Impact Assessment and/or management plan. Rehabilita-
tion of terraces and water supplies with tanks allowed otherwise unsustainable development and intensified
encroachment on natural habitat (Judas et al., 2004; Islam et al., 2018).
Killing/Poisoning of carnivores: The importance of the output of our model increases if we want to
look for the remaining population of the Arabian Leopard in the region and protect them from killing,
especially by shepherds. Beside habitat deterioration for the carnivores and prey in the region, the Arabian
Leopard faces tremendous threats from people, when they started occupying the same area. Although legally
protected, current law enforcement is ineffective, a total of 52 killing of Arabian leopards have been recorded
in Saudi Arabia itself (Islam et al., 2018; Al-Johany, 2007; Judas et al., 2006; Jackson et al., 2010) and in
other countries similar reports are there too and killing of predators and hanging them on trees or posts is
a normal practice in remote areas of the region.
Depletion of prey-base: Relation predator-prey is also virtually unknown in Saudi Arabia. Although the
subject has been highly debated for different big predator besides leopards, Arabian wolf, Striped hyenas
etc (Erlinge et al., 1984; Wright et al., 1994), leopards may select and control wildlife populations. Arabian
Leopard and prey populations respond dynamically to one another. Many prey species are facing equal
conservation challenges and population declined dramatically over the decades. Certain species such as
Hamadrayas Baboon adapted themselves to humans, which created commensalism issues and they carry
lethal diseases (Olarinmoye et al., 2016 & 2019). Over time, the two populations cycle up and down in
number.
Although the diet of the Arabian Leopard is poorly known, and from studies of African leopards, they are
13considered opportunistic, and have a wide range of prey species within a large panel of size (Schaller, 1972;
Bertram, 1982; Hayward et al., 2006; Macaskill, 2009). They are able to kill a cow, but will also eat insects.
However, even when large prey is present, leopards will feed in a large part upon small and easy to catch
prey, such as small birds, rodents (rock rats and girds) or insects. They often include a large proportion
of other smaller carnivore species (Genet, Mongoose, foxes, dogs, and Hyena’s offspring). Individual choice
and specialization are a common trait including antelopes, deer, pigs, and primates (Kingdon, 1997).
Scat analyses in Oman (Wright, 1999) showed that the main prey species were, by order of importance, the
Arabian Gazelle (Gazella gazella ), Nubian Ibex (Capra ibex ), Cape Hare (Lepus capensis ), Rock Hyrax
(Procavia capensis ), bird species, Porcupine (Hystrix indica ), Ethiopian Hedgehog (Paraechinus aethiopicus
), small rodents and insects while 14 mammal species were recorded by the camera traps at Jabal Samhan
NR that include (Spalton et al., 2006). Human interference, through depletion of prey base and killing by
hunters and shepherds, has been suggested as the main cause of decline (Spalton & Al Hikmani, 2006).
In Israel/Palestine, the diet of the leopard was recorded mainly comprised of Hyrax and Ibex (90%) and 5%
of porcupines (Ilany, 1990).
In Saudi Arabia, the diet of the leopard could be mainly constituted of Hyrax, Arabian Mountain gazelle
and similar size prey species. Nubian Ibex and gazelles could have been an important part of the diet of
leopards in the past, but the range of these ungulates has been significantly reduced; leopards should have
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
had no choice to survive to shift their diet according to prey availability (Judas et al., 2004; Islam et al.,
2018). If leopard preys on baboons, it happens at night when they sleep among rocks (Shortridge, 1934 in
Kingdon, 1977). Since leopards are known to be opportunistic predators, we can presume that they will also
prey on species such as partridges in mountain regions (Ammoperdix heyi, Alectoris melanocephala and A.
phiibyi), porcupines, hares and even fish, frogs and turtles that are still abundant in some places (Judas et
al., 2004; Islam et al., 2011).
Predation upon goat, sheep, young camels and feral ass has been reported (Islam et al., 2018; Biquand, 1989;
Biquand & Boug, 1992). As known from Africa, leopards are attracted by domestic dogs that they captured
at night around or even inside camps (Haltenorth & Miller, 1985) or South America for Jaguar (Panthera
onca, Rabinowitz, 1986).
Insufficient protection despite its full protected status under laws: Predators such as the Arabian
Leopard is critical for healthy ecosystems, ensuring that a greater variety of species survive and thrive by
keeping prey populations in check. As we know, the re-introduction of wolves into Yellow Stone National
Park in the United States of America caused trophic cascading effects on flora and fauna of the Park (Smith
et al., 2013). If we are proactive, conservation programs are capable of ensuring a positive future for these
and other species.
Our model shows that Jabal Dhofar (i.e. western parts of Dhofar mountains) in Oman and eastern parts
of Dhofar mountains in Yemen are the source. Moreover, our model did not indicate the known eastern
population in central Oman as large as its western one, and this might have to do with better protection
enforcement (Spalton et al., 2006; Mellon, 2009).
Arabian leopard is considered a flagship species throughout its range, and every range country provides
protection but according to our model the majority of the population occurred outside the protected areas.
Conservation efforts need to be focused towards leopard populations outside the current network of protected
areas as well, as the majority of available leopard habitats presently receive no legal protection (Jacobson et
al., 2016).
In Saudi Arabia, there is no specific reserve for the Arabian Leopard conservation, while hunting is restricted
under the National Hunting Law of 1978 #457 under the Wildlife Protected Areas System (Seddon, 1996).
All forms of hunting are officially prohibited in all Protected Areas managed by the SWA, but the presence
of leopards has been attested from only two of them, which are Ar Raydah Reserve (9 sqkm, established
in 1989) in Abha and Jabal As Shada Reserve (50 sqkm established in 2002) in Al Mikhwa states exists in
14South-western Highlands. Raydah and As Shada reserves are very small to shelter viable Leopard populations
(Judas et al., 2004). So, it was highly recommended to protect more areas, where the possibilities of leopard
are high or where the leopard persisted in the past (Islam et al., 2014; 2018).
Hilly areas with wadis in Yemen is continuous up to Jabal Dhofar in Oman provide a good habitat for the
Arabian leopard and 12 are important locations for the leopard within this strip, where leopard persistence
is likely and need to be surveyed properly, which are also highlighted in our movement intensity model.
In Yemen although there are 10 declared reserves and among them are Jabal Bura valley forest and Dhamar
Montane Plains Mahjur Traditional Reserve but leopards have not been recorded in these reserves, while
at Wada’a, in Amran district, where the majority of the leopards have been captured before (Stanton et
al., 2008) classifies as very poor by our model for the species presence. In 2008 no leopard was recorded
(Stanton et al., 2008) and in 2010 intensive camera-trapping in Hawf Protected Area in eastern Yemen was
carried out, even using attractive scents but could not detect any leopards (Pittet, 2011). The area where
camera-traps were deployed, were outside of our modeled polygons of the inferred source populations. This
also emphasizes that the survey effort should be more focused following our findings.
Lack of awareness among the general public of the need for protection and conservation of the
country’s wildlife, including the endangered Arabian leopard: Public support towards conservation
of wildlife is extremely necessary that majority of the wildlife persists outside of the protected areas. It is
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
highly suggested that beside intensive monitoring in modeled areas, range countries should initiate public
awareness programs to get support for long-term conservation of the species.
Conservation measures proposed: Our models would be beneficial not only to the Arabian Leopard but
other biodiversity in general through linkage of science to policy and decision processes. This model would
be helpful in future landscape-based planning on known and projected corridors and prioritize policies, and
articulate the range of solutions. That is how our model would bring together scientists, public stakeholders
and policy makers, and are used as an adaptive management tool to understand complex landscapes that
are undergoing short- and long-term change.
It is highly recommended to intensify the field search and camera trapping in all the important areas such
as western mountains in Oman e.g., Jabal Samhan NR, Jabal Qamar, Jabal al Qara in Dhofar (Spalton
& Al Himani 2014), eastern mountain of Yemen: Hawf, and Jadib (Pittet 2011), and locations at western
highlands in Saudi Arabia e.g., Wadi Tarj, Wadi Lajab; Jabal Uthrub, Wadi Nauman, remote mountains of
Tanomah, Billasmar, Billahmar, Bani Saad) and probably Fiqrah mountain (Islam et al . 2014). We also
recommend to carry out field search for leopard in modelled areas and especially corridors.
It is important to maintain the integrity of small and no longer viable subpopulations or local nuclei, and
the connection between neighboring populations needs to be restored (CMS, 2017). Sometimes, animals
move from the nuclei population to other areas and such individuals need adequate migration corridors for
dispersing animals. Our corridor study put forward to facilitate landscape-level, large-scale trans-boundary
conservation, and the Arabian leopard conservation will be benefitted.
These models provide the base for further investigation to conserve the species and we still need to collect
detailed information, especially in predicted corridors across the range (Judas et al., 2006; Islam et al.,
2011; 2018): (1) population assessment - some studies carried out at Jabal Dhofar in Oman (Spalton and
Al Hikmani, 2014); (2) number of individuals per sub-population; (3) range use pattern and home range
size; (4) Activity pattern; (5) basic knowledge of habitat requirements; (6) food requirement (prey species);
(7) prey availability, dynamics and requirements; (8) relation predator-prey (such as seasonal movements of
Ibex, gazelles); (9) competition with other predators (Caracal, Hyena, Wolf); (10) quantification of conflict
with man. To assure long term survival of the wild populations, population viability analysis has to be
developed to project population trends with regularly updated data. This would require: (1) to understand
population dynamics; (2) to study dispersal rates and quantify movements between populations; (3) to
establish long-term monitoring’ with regular periodic surveys (camera trapping); (4) to assess genetic pool
(determine genetic identity from blood and scats, DNA microsatellite mapping.
15Since the Arabian Leopard population is so low and many individuals were poisoned, killed, and trapped, the
range countries need to have conservation support programs on local as well as regional scale as suggested
in the regional and national strategies (Islam et al., 2011, 2014; Breitenmoser et al., 2010) to protect the
Arabian Leopard. It requires constant population monitoring; implementation of regional and national
conservation strategies; protecting important locations, where the Arabian Leopard still persists and carry
out intensive monitoring using camera trapping or ‘track impression pads’ (patches of sand or soft earth
placed along movement corridors) based on our findings; our model suggests that the habitat is larger and
less fragmented westward in Oman and eastward in Yemen. However, based on geographically limited studies
areas in western Oman seems to support a good nucleus that is probably due to enforced protection. Our
findings show that eastern Yemen should have the species stronghold too and research resources must be
concentrated there as well and areas must be protected; and the last suggestion is to Ex-situ conservation-
captive-breeding to safeguard the species in captivity in case it locally extinct.
DATA ACCESSIBILITY
It is declared that we will deposit data in a public repository, which is used in the modeling and analysis in
this paper and the link of data is already been shared with the Journal through email.
REFERENCES
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Al Hikmani, H. (2019). Evolutionary Genetics and Conservation of the Critically Endangered Arabian
Leopard (Panthera pardus nimr ). Doctor of Philosophy (PhD) thesis, University of Kent., United Kingdom.
Al Jumaily, M., Mallon, D. P., Nasher, A .K., & Thowabeh, N. (2006). Status report on Arabian leopard in
Yemen. CAT NEWS Special Issue, 1: 20-25.
Al-Johany, A. M. H. (2007). Distribution and conservation of the Arabian leopards Panthera pardus nimr
in Saudi Arabia. Journal of Arid Environments , 68, 29-23.
Beier, P., Majka, D.R., & Spencer, W.D. (2008). Forks in the road: Choices in procedures for designing
wildland linkages. Conserv Biol, 22(4), 836–851.
Bertram, B.C.R. (1982). Leopard ecology as studied by radio-trackingin Telemetric Studies of Vertebrates .
Cheeseman, C.L. and Mitson R.B. Eds. Symp. Zoo1. Soc. Lond. No. 49:341-352.
Biquand, S. (1990). The Arabian Leopard in Saudi Arabia. Unpublished annual report. NWRC, Taif, p.
27. Saudi Arabia.
Biquand S., Boug, A., & Gaucher, P. (1990). Field reconnaissance survey of southern Hijaz. Unpublished
report, NWRC, Taif, 37 pp. Saudi Arabia.
Biquand, S., & Boug, A. (1989). Protection of the Arabian Leopard in Saudi Arabia. Unpublished report,
NWRC, Taif, Saudi Arabia.
Biquand, S., & Boug, A. (1992). An update of leopard status in AI Fiqrah and recommendations for
immediate action. Unpublished report, NWRC, Taif. 4 pp., Saudi Arabia.
Blunt, L.A., (1881). A Pilgrimage to Nejd : the Cradle of the Arab Race. A visit to the court of the Arab
Emir, and our “Persian Campaign”. Vol 1 & 2, London, Murray, Albemarle Street, UK.
Boug, A., Islam, M. Z., & Shehri, A. (2009). Captive-breeding of Arabian Leopard Panthera pardus nimr in
the Kingdom of Saudi Arabia. Wildlife Middle East 4 (3): 2.
Breheny, P., & Burchett, W. (2017). Visualization of Regression Models Using visreg. The R Journal, 9,
56-71.
Breitenmoser, U., Breitenmoser C., Mallon D. & Edmonds A. J. (Eds). (2006). Status and Conservation
of the Leopard on the Arabian Peninsula. Cat News Special Issue No 1 , IUCN/SSC Cat Specialist Group
and Breeding Centre for Endangered Arabian Wildlife. 47 pp.
16Breitenmoser, U., Breitenmoser-Wursten, C., Mallon, D., & Edmonds, J.A. (2010). Strategy for the Conser-
vation of the Leopard in the Arabian Peninsula . Sharjah: IUCN/SSC Cat Specialist Group, Environment
& Protected Areas Authority.
Bruner, A.G., Gullison, R.E., & Balmford, A. (2004). Financial costs and shortfalls of managing and
expanding protected-areas systems in developing countries. Biosci . 54, 1119–1126.
Budd, K. (2003). Arabian Leopard Regional Studbook. Breeding Centre for Endangered Arabian Wildlife,
Sharjah.
Burton J.F. (1995). Birds and climate change . Christopher Helm Eds. London, 376 pp.
Carruthers D. (1909). Big game of Syria, Palestine and Sinai . The field, London 114 (2973):1135-1136.
CEPF. (2012). Ecosystem Profile, Eastern Afromontane Biodiversity Hotspot. BirdLife International.
http://legacy. cepf.net/Documents/Eastern Afromontane Ecosystem Profile FINAL.pdf
CMS. (2017). Proposal for the inclusion of the leopard (Panthera pardus ) on Appendix II of the Convention.
12th Meeting of the Conference of the Parties, Manila, Philippines, 23 - 28 October 2017 Agenda Item 25.1.
UNEP/CMS/COP12/Doc.25.1.4.
Collenette, S. (1999). Wildflowers of Saudi Arabia . NCWCD. 799 pp., Riyadh, Saudi Arabia.
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Compton, B.W., McGarigal, K., Cushman, S.A., & Gamble, L.R. (2007). A resistant-kernel model of
connectivity for amphibians that breed in vernal pools. Conserv Biol 21:788–799.
Crooks, K.R., & Sanjayan, M. (2006). Connectivity Conservation . Cambridge, UK: Cambridge University
Press.
Cushman, S.A. (2006). Effects of habitat loss and fragmentation on amphibians: a review and prospectus.
Biol Conserv 128, 231–240.
Cushman, S.A., Landguth, E.L., & Flather, C.H. (2012). Evaluating the sufficiency of protected lands
for maintaining wildlife population connectivity in the US northern Rocky Mountains. Divers Distrib18,
873–884.
Cushman, S.A.; McKelvey, K.S.; & Schwartz, M.K. (2009). Use of empirically derived source-destination
models to map regional conservation corridors. Conserv Biol 23, 368–376.
Cushman, S.A., McRae, B., Adriaensen, F., Beier, P., Shirley, M., & Zeller, K. (2013). Biological corridors
and connectivity. Key Topics inConservation Biology 2, 384-404.
Dixon, J.D.; Oli, M.K.; Wooten, M.C.; Eason, T.H.; McCown, J.W. et al. (2006). Effectiveness of a regional
corridor in connecting two Florida black bear populations. Conserv Biol 20(1), 155–162.
Doughty C. M., (1888). Travels in Arabia Deserta . 2 Vo1. Cambridge University Press., UK
EPAA (2000). CAMP for Arabian Carnivores & PHVA for the Arabian Leopard and Tahr. Final Report.
Breeding Centre for Endangered Arabian Wildlife, Sharjah.
Erlinge, S., Goransson, G., Hogstedt, G., Jansson, G., Liberg, O., Loman, J., Nilsson, L.N., Von Schantz,
T. & Sylven, M. (1984). Can vertebrate predators regulate their prey? Am. Nat . 123(1), 125- 133.
Estes, J.A., Terborgh, J., Brashares, J.S., Power, M.E., Berger, J., Bond,W.J. et al. (2011). Trophic
downgrading of planet earth.Science 333: 301–306.
Fahrig, F. (2003). Effects of habitat fragmentation on biodiversity.Annual Review of Ecology, Evolution, and
Systematics 34, 487–515.
Gasperetti, J., & Jackson, P. (1990). Preliminary Report of the Status of the Arabian Leopard. Unpublished
report.15 pp. NWRC, Saudi Arabia.
17Gasperetti, J., Harrison, D.L., & Biittiker, W. (1985). The Carnivora of Arabia. Fauna of Saudi Arabia 7,
397-461.
Gavashelishvili, A. & Lukarevskiy, V. (2008). Modelling the habitat requirements of leopard Panthera pardus
in west and central Asia.Journal of Applied Ecology 45, 579–588.
Gavashelishvili, A. & Tarkhnishvili, D. (2016). Biomes and human distribution during the last ice age.
Global Ecology and Biogeography 25, 563-574.
Granit, B. (2016). Once there were Leopards. Israel Birding Portal.
Haddad, N.M., Bowne, D.R., Cunningham, A., Danielson, B.J., Levey, D.J., et al. (2003). Corridor use by
diverse taxa. Ecology 84(3), 609–615.
Haltenorth, T., & Miller, H. (1985). Les mammiferes d’ Afrique et de Madagascar. Delachaux et Niestle,
397 pp.
Harrison, D. L. & Bates, P.J.J. (1991). The Mammals of Arabia . Second edition. Harrison Zoological
History Museum Publication, England. ISBN: 0951731300.
Harrison, D. L. (1983). The mammal fauna of Oman, with special reference to conservation and the flora
and fauna surveys. Journal of Oman Studies 6(2): 329-339.
Posted on Authorea 1 Feb 2020 — CC BY 4.0 — https://doi.org/10.22541/au.158057220.09724014 — This a preprint and has not been peer reviewed. Data may be preliminary.
Harrison, D.L. (1968). The mammals of Arabia . Vol. 2. Carnivora, Artiodactyla, Hyracoidea. Ernest Benn
Ltd., London, XIV: 193-381.
Hayward, M., Henschel, P., O’Brien, J., Hofmeyr, M., Balme, G., & Kerley, G. (2006). Prey Preferences of
the Leopard (Panthera pardus ). Journal of Zoology , 270/2: 298-313.
Hedges S.B. (2000). Human evolution. A start for population genomics.Nature 408:652653.
Heller, N.E., & Zavaleta E.S., (2009). Biodiversity management in the face of climate change: a review of
22 years of recommendations.Biological conservation 142, 14–32.
Hemprich, F.W., & Ehrenberg, C.G. (1828-1833). Symbolae physicae seu icones et descriptiones mammal-
ium, 1 and 2. Berlin.
Hijmans, R. J., (2016). Raster: geographic data analysis and modeling. R package version 2.5-8.
https://CRAN.R-project.org/package=raster.
Hilton-Taylor, C. (2000). 2000 IUCN Red List of Threatened Species. IUCN, Gland, Switzerland and
Cambridge, UK.
Ilany, G. (1990). Leopard Panthera pardus in Israel, Cat News, Gland, 12:4-5. In Gasperetti & Jackson,
1990.
Islam, M. Z. & Boug, A. (2017). National Strategy and Action Plan for Arabian Leopard in the Kingdom
of Saudi Arabia. CAT NEWS 66, 14-17.
Islam, M. Z., Boug, A., & Shehri, A. (2011, 2012, 2013). Status of Arabian Leopard and camera trapping in
Western Highlands of Saudi Arabia. PSFWRC internal reports. Saudi Arabia.
Islam, M. Z., Hashim, K., & Boug, A. (submitted 2019). Pharmaceutical drug cause mortalities of some
species of raptors in Saudi Arabia: an overview. Journal of Threatened Taxa .
Islam, M. Z.; Boug, A.; Judas, J.; and As-Shehri, A. (2018). Conservation challenges for the Ara-
bian Leopard (Panthera pardus nimr ) in Western Highlands of Arabia. BIODIVERSITY , DOI:
10.1080/14888386.2018.1507008.
Islam, M. Z.; Boug, A.; Shehri, A.; & Jaid, M. (2015). Poisoning of endangered Arabian Leopard in
Saudi Arabia and its conservation efforts.CAT NEWS 60, 16-17.
18You can also read
