Science of the Total Environment 777 (2021) 146031

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Science of the Total Environment

journal homepage: www.elsevier.com/locate/scitotenv

Landscape-level habitat management plan through geometric reserve

design for critically endangered Hangul (Cervus hanglu hanglu)
Tanoy Mukherjee a,1, Vandana Sharma b,1, Lalit Kumar Sharma a,⁎, Mukesh Thakur a, Bheem Dutt Joshi a,
Amira Sharief a, Avantika Thapa a, Ritam Dutta a, Stanzin Dolker a, Basudev Tripathy a, Kailash Chandra a
a
Zoological Survey of India, Kolkata 700053, India
b
Indian Institute of Remote Sensing, Dehradun 248001, India

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• Only 9% area of the Dachigam landscape

found suitable for Hangul in Kashmir
Valley.
• Conservation Reserves adjacent to
Dachigam NP are vital for the viability
of Hangul.
• Geometric reserve designing is benefi-
cial and cost-effective management
strategy.
• Creation of Greater Dachigam NP will be
vital for the critically endangered
Hangul.

a r t i c l e i n f o a b s t r a c t

Article history: Hangul (Cervus hanglu hanglu), the only red deer subspecies surviving in the Indian subcontinent, is of top con-
Received 20 December 2020 servation priority with global importance. Unfortunately, it has lost much of its historical distribution range, and
Received in revised form 17 February 2021 it is now confined to Dachigam landscape within the Kashmir valley of India. The Government of India initiated a
Accepted 17 February 2021
recovery plan in 2008 to augment their numbers through ex-situ conservation programs. However, it was neces-
Available online 22 February 2021
sary to identify potential hangul habitats in Kashmir valley for adopting landscape-level conservation planning
Editor: Fernando A.L. Pacheco for the species. Based on geometric aspects of reserve design, we modeled hangul habitat using an ensemble ap-
proach to identify hangul habitats. The present model indicates that the conifer and broadleaf mixed forests were
Keywords: the most suitable habitats. Only 9% of the total study landscape was found suitable for the species. We identified
Hangul corridors among the suitable habitat blocks, which may be vital for the species' long-term genetic viability. We
Dachigam landscape suggest reorganizing the existing management of Dachigam National Park (NP) following the landscape level
Ensemble model and habitat block-level management planning based on the core principles of geometric reserve design. We rec-
Landscape structure ommend that the identified patch (PID-6) in the southern region of the landscape to be converted into a Conser-
Landscape connectivity
vation Reserve or merged with the Overa-Aru Wildlife Sanctuary. This habitat patch PID-6 may be a stepping
Reserve design
stone habitat and vital for maintaining the species landscape connectivity and metapopulation dynamics.
© 2021 Elsevier B.V. All rights reserved.

1. Introduction

Hangul or Kashmir Stag (Cervus hanglu hanglu), the only surviving

⁎ Corresponding author at: O/C Wildlife and GIS Section, Zoological Survey of India,
Kolkata, India.
red deer in the Indian subcontinent, is one of the four easternmost dis-
E-mail address: lalitganga@gmail.com (L.K. Sharma). tributed subspecies of red deer (Gee, 1965; Grzimek, 1990; Kurt, 1978;
1
These Authors Contributed Equally. Schaller, 1969). It occupies temperate coniferous forests in the Western

https://doi.org/10.1016/j.scitotenv.2021.146031
0048-9697/© 2021 Elsevier B.V. All rights reserved.
T. Mukherjee, V. Sharma, L.K. Sharma et al.
Himalayas in the Jammu and Kashmir (J&K) Union Territory (UT) of
India (Gee, 1965; Schaller, 1969). Our previous studies on the molecular
genetic diversity, population demography, viability, and phylogenetic
status of hangul provided the first phylogeographic status of hangul
among the 22 subspecies of red deer (Mukesh et al., 2015). Lorenzini
and Garofalo's (2015) insights into the evolutionary history of Cervus
corroborate our findings on the genetic distinctness of the Tarim clade
of red deer and support our ideas regarding raising the status of hangul
to a species level. Historically, the hangul distribution formed an arc
covering the majority of the north-eastern mountains of the Jhelum
and Chenab Valleys of J&K UT and some parts of the Chamba Valley of
the Himachal Pradesh State in India (Schaller, 1969; Holloway, 1971).
Regrettably, hangul has lost much of its distribution range due to a vari-
ety of factors and is now confined to a cluster of protected areas (PAs)
collectively known as the Dachigam Landscape (DL), ca. 1000 km2
(Schaller, 1969; Ahmad et al., 2009). The overall population size of han-
gul shrank dramatically from 3000–5,000 in the 1900s to approximately
148 individuals in 2015 (Brook et al., 2017; Gee, 1965; Sofi and Ahmad,
2009; Bacha, 2015). The factors behind this unfortunate population de-
cline during the last few decades are largely unknown. However, previ-
ous studies have attributed population decline with habitat loss at the
landscape level (Ahmad et al., 2009; Charoo et al., 2010; Gee, 1965;
Schaller, 1969). A short survey commissioned by the Department of
Wildlife Protection documented the existence of hangul outside the
DL as small, isolated and fragmented populations in forested areas of
Kupwara, Bandipora, Ganderbal, Pulwama and Anantnag (Sofi and
Ahmad, 2009). Considering the population status and conservation
threats, the hangul has been placed under Schedule-I of the Indian
Wildlife (Protection) Act of 1972 and the J&K Wildlife Protection Act
of 1978. Additionally, the Convention on International Trade in Endan-
gered Species of Wild Fauna and Flora (CITES) forbids the trade of han-
gul and lists it among the Appendix-I species. However, based on our
latest studies, the International Union for Conservation of Nature
(IUCN) has reassessed the conservation status of the species and sepa-
rated it from other subspecies, listing the species as ‘critically endan-
gered’ (Brook et al., 2017). Previously, hangul was lumped with the
red deer complex and listed as ‘least concern’ (Mukesh et al., 2013;
Mukesh et al., 2015). The scientific information on hangul is scant, and
no systematic study has been conducted to model its current distribu-
tion range or scrutinize the limiting factors determining its distribution
in the DL. However, several short-term studies and surveys have been
carried out to assess its distribution, habitat use, feeding habits and pop-
ulation status (Gee, 1965; Schaller, 1969; Holloway, 1971; Sofi and
Ahmad, 2009; Kurt, 1978; Inayatullah, 1985; Ahmad et al., 2013).
Recognizing the IUCN World Conservation Union initiative, the
Government of India developed a recovery plan for hangul in 2008
(IUCN, 1987; MoEF, 2012). The recovery plan is implemented by the
Jammu and Kashmir Wildlife Protection Department with an overarch-
ing goal of augmenting the existing population and reclaiming its
historical distribution range. There is a need to identify sites and
corridors that are suitable for the long-term viability of the introduced
population after captive breeding (MoEF, 2012).
Hence, we developed a model for landscape-level reintroduction
using remote sensing and geographic information system. The recent
advancements in remote sensing techniques allow for modeling the po-
tential habitats of a species reliably and are imperative for establishing
species and habitat associations (Raxworthy et al., 2003; Meier et al.,
2010; Wisz et al., 2013; Franklin, 2013; Kalle et al., 2013). GIS-based
species prediction models use a combination of habitat factors to predict
potential habitat and have been applied to a wide range of taxonomic
groups for conservation planning and management. Moreover, species
distribution models have been used to understand the impact of climate
change on the distribution range of a species (Bakkenes et al., 2002;
Porfirio et al., 2014). A variety of species distribution prediction models
are available, and the most commonly used are presence-only models
such as maximum entropy (MaxEnt) (Phillips et al., 2006), the genetic
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Science of the Total Environment 777 (2021) 146031
algorithm for rule-set production (GARP) (Stockwell, 1999), ecological
niche factor analysis (ENFA) (Stockwell, 1999), and Mahalanobis factor
analysis (Calenge et al., 2008). The ENFA model has been applied for
hangul (Kaul et al., 2018) to identify suitable sites for the species. How-
ever, it is dependent on single-model algorithms, and lacks information
on habitat connectivity and the role of existing conservation reserves in
the landscape. As it does not provide a definitive management-friendly
strategies, DL demands a re-evaluation using a more robust and
multimodel approach with a definitive phase wise action plan for
hangul.
Thus, we adopted an ensemble modeling approach in which multi-
ple modeling algorithms such as boosted regression trees (BRT), a gen-
eralized linear model (GLM), multivariate adaptive regression splines
(MARS), and random forest (RF) to develop an ensemble model for
the species. The ensemble modeling approach is one of the most robust
techniques due to its predictability and usefulness in identifying
species-habitat relationships (Elith and Leathwick, 2009; Guisan et al.,
2007a, 2007b; Miller, 2010; Mukherjee et al., 2020).
The traditional approaches to increasing the effectiveness and sus-
tainability of a protected area network are based upon the theories of
biogeography and metapopulation dynamics (Hanski and Gilpin,
1991). However, landscape-level conservation and management strate-
gies based on the classic idea of reserve design, which encourage an in-
creasing number of habitat patches along with enhanced connectivity
and improved habitat quality, are considered useful for threatened spe-
cies (Diamond, 1975; Donaldson et al., 2017). In the earlier approach,
there was a significant trade-off between habitat quality and the geo-
metric principles of reserve design, mainly in the size, number, and
quality of the patches and the connectivity between suitable habitat
patches (Diamond, 1975). However, in recent decades, conservation ac-
tion plans and initiatives have evolved from a reserve-based approach
to a landscape-level approach (Opdam and Wascher, 2004; Watts and
Handley, 2010). This remarkable change in policy planning could be
due to the accelerated threat of climate change and habitat fragmenta-
tion, which have led to an increasingly dynamic pattern of colonization
and extinction (Heller and Zavaleta, 2009).
Hence, in the present study, we created a landscape-level multiscale
conservation plan for improving habitat networks by identifying and
evaluating connectivity within the landscape, the integrity of suitable
patches, and the importance of conservation reserves (CRs) in develop-
ing a protected area network. We identified potential habitat patches in
line with the core principle of the classic ideas of reserve design to sus-
tain the long-term viability of hangul in the Kashmir Valley.
2. Materials and methods
2.1. Study area
The study area comprises of Zabarwan mountain ranges and the ad-
joining areas of Kashmir Himalayas in the State of Jammu and Kashmir,
India, where hangul was reported to be present historically (Fig. 1). The
intensive study area comprised of DL, which is a mosaic of Protected
Areas (PA) along with other forest areas classified as Reserves Forest
under Territorial Forest Divisions. The DL is composed of two National
Parks, i.e., Dachigam National Park & City Forest National Park, two
Wildlife Sanctuary (Overa-aru Wildlife Sanctuary, Thajwas Wildlife
Sanctuary), and 10 Conservation Reserves. The landscape
biogeographically can be classified as North-West Himalaya 2A zones
(Rodgers and Panwar, 1988), where vegetation is typically Himalayan
Moist Temperate with Broadleaf Mixed Forest in valley areas and
Conifer Forests in middle elevation slopes. The higher altitudes are rep-
resented by Subalpine and Alpine meadows (Champion and Seth,
1968). The altitudinal gradients range from 1600 m to 5000 m and
majority of the study area is composed of mountainous terrain with
moderate to precipitous mountains and Valley regions. The climate is
the sub-Mediterranean type with bixeric regime having two dry spells
T. Mukherjee, V. Sharma, L.K. Sharma et al.
Fig. 1. Study area representing the hangul presence location along with the different LULC classes of the study landscape.
during June–July and September–November (Singh and Kachroo,
1978).
2.2. Study design and data collection
The locations used to model the potential hangul habitats in the
Kashmir Valley were obtained from previous studies (Mukesh et al.,
2013, 2015; Sharma et al., 2010; Charoo et al., 2010; Mukesh et al.,
2015) and published government reports (Charoo et al., 2011; Sofi
and Ahmad, 2009). The locations used in this study cover the current
distribution range of hangul and relic sites. Spatiotemporal autocorrela-
tion testing of the locations was performed using SDMtoolbox 2.0
(Brown et al., 2017) to remove the clustered location of hangul. The
minimum distance between the location points was set at 1 km for
the rarefication process. Finally the spatially independent (n = 38) loca-
tions were used for model building.
2.3. Predictor variable selection and description
The habitat predictors that define potential hangul habitats were
chosen based on the biological relevance and data availability
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Science of the Total Environment 777 (2021) 146031
(Ahmad, 2006; Kurt, 1978; Sharma et al., 2010). The predictor variables
comprised the habitat types relevant to hangul, topography, landscape
configuration and anthropogenic variables (Figs. S1–S3). The selection
of these key environmental variables is crucial in determining the hab-
itat niche to model the potential distribution habitat (Guisan and
Zimmermann, 2000; Manly et al., 2002; Porfirio et al., 2014). The pri-
mary focus was on producing the best fit model for the data with the
combination of the possible variables (Johnson et al., 2006).
2.3.1. Habitat and vegetation predictors
The Landsat satellite imagery (Landsat-8 OLI/TIRS) with spatial res-
olution of 30 m was downloaded from https://earthexplorer.usgs.gov
(Path: 149; Row; 36; Date of accusation: 2017-08-17; Cloud cover
>10%) and classified following Anderson et al. (1976) using ERDAS
Image version 9.0 (ERDAS®, Inc., Atlanta, Georgia). The overall accuracy
was assessed using Kappa Statistics (Landis and Koch, 1977). We classi-
fied the study area into nine land use and land cover (LULC) classes, and
four classes, viz., conifer forest, broadleaf mixed forest, temperate grass-
lands and scrubland, and alpine meadows, were used for further analy-
sis since these classes were more appropriate for defining hangul
habitat (Fig. 1). The major land cover types identified in the final
T. Mukherjee, V. Sharma, L.K. Sharma et al.
classified image were separated into different vector files and the
Euclidian distance function for each land cover type in the study area
was determined using ArcGIS 10.6 software.
2.3.2. Topographic variables
Topographic predictors such as elevation zones, slope, and aspect
categories were used to model potential habitats, considering their rel-
evance in hangul life history (Kurt, 1978; Sharma et al., 2010). Schaller
(1977) and Sharma et al. (2010) reported the differential use of land-
scape topography to enhance adaptability in extreme weather condi-
tions and minimize predation risk. The slope, elevation and aspect
maps were generated from the digital elevation model (DEM) layer
using the surface analysis tool in the Spatial Analyst toolkit of ArcGIS
10.6 (ESRI®). After extracting the water bodies (streams) from the
DEM, the distances (meters) of the water bodies from the hangul loca-
tion were calculated using a log-Euclidean distance function in ArcGIS
10.6 (ESRI®).
2.3.3. Anthropogenic variables
Unlike other red deer subspecies, hangul is sensitive to anthropo-
genic disturbances and is reported to avoid areas with moderate-to-
high levels of anthropogenic disturbance (Charoo et al., 2010). Several
studies have used the distance to roads and human use traits as a
proxy for disturbance in habitat modeling (Huijser and Bergers, 2000;
Kramer-Schadt et al., 2004; Torres et al., 2012). The distance of the han-
gul location to the nearest road (human use trails and permanent roads)
was calculated using a log-Euclidean distance function in ArcGIS 10.6
(ESRI®).
2.3.4. Bioclimatic predictors
Bioclimatic predictors are important in describing the climatic iso-
therm and can influence the distribution of the target species (Paudel
et al., 2015). A preliminary set (n = 19) of bioclimatic predictors was
downloaded from the WorldClim database (https://www.worldclim.
org/).
The potential predictor raster data were resampled at 30-m spatial
resolution using the Spatial-Analyst Extension of ArcGIS 10.6. The spa-
tial multicollinearity among the variables was tested using the SAHM
package in the VisTrails environment, and predictors with Pearson cor-
relation (r) >0.8 were omitted from the modeling environment
(Warren et al., 2010).
2.4. Model building and validation
We used an ensemble modeling approach to evaluate the habitat vi-
ability of hangul in the Kashmir Valley. Four different modeling ap-
proaches were used to develop the ensemble, namely, boosted
regression trees (BRT), the generalized linear model (GLM), multivari-
ate adaptive regression splines (MARS), and random forest (RF). The
modeling procedure was implemented through the VisTrails pipeline
with the SAHM (Software for Assisted Habitat Modeling) package
(Morisette et al., 2013; Talbert and Talbert, 2012). The final output of
the individual models resulted in a probability surface ranging from 0,
which indicates unsuitable habitat, to 1, which indicates the highest
score of habitat suitability for the species. In addition to generation of
the respective probability surfaces for the respective models, individual
Boolean maps were generated using the minimum training presence as
a threshold (Hayes et al., 2015).
To build the ensemble model, the AUC threshold was set to >0.75.
After the final model selection and ensemble model building, the en-
semble probability (continuous) and ensemble count maps were gener-
ated based on the model argument (MA). The final ensemble count
maps ranged from 0, indicating that the null of the participating models
agreed on the suitability in the respective pixel, to the highest MA of 4,
indicating the agreement of all participating models on the suitability of
the species (Mukherjee et al., 2020). The SAHM package for VisTrails
4
Science of the Total Environment 777 (2021) 146031
was used for model fitting and evaluation of the model performance
metrics (Morisette et al., 2013). The appropriateness of the model com-
plexity was examined by taking the difference between the AUC (train)
and AUC (CV), represented as ΔAUC (Jarnevich et al., 2018). The scores
of the ΔAUC from the respective models provide information on how
sensitive the models are to the fitting data (Jarnevich et al., 2018;
Mukherjee et al., 2020). In addition to the AUC-based evaluation, the
Cohen's kappa (Cohen, 1968), true skill statistic (TSS) (Allouche et al.,
2006), specificity and sensitivity were evaluated for training and
cross-validation (n = 10) using SAHM Package (Jiménez-Valverde
et al., 2013; Phillips and Elith, 2010). The variable importance of predic-
tors was evaluated using the increase in the AUC after fitting the final
model with and without each predictor variable (Grenouillet et al.,
2011; Mukherjee et al., 2020). The combined importance of the predic-
tors was evaluated by the mean AUC (μ AUC) scores obtained for each
predictor from the four models (Jarnevich et al., 2018; Jiménez-
Valverde et al., 2013).
2.5. Habitat connectivity and patch-level habitat evaluation for hangul
The biological connectivity was assessed using the circuit model,
which is a widely used approach for animal corridor design (Ruiz-
González et al., 2014; Sharma et al., 2019; Wang et al., 2014). We simu-
lated the possible biological corridor for hangul using the Circuitscape
toolbox in ArcGIS Ver. 10.6. (McRae et al., 2008; McRae and Shah,
2009; Wang et al., 2014). The final ensemble habitat suitability model
for hangul was used as the conductance surface in the circuit model
(Sharma et al., 2019). To eliminate the complexity in the final circuit
model, we used only the confirmed locations of hangul in the study
landscape. The extent of the proposed connectivity surface was de-
signed to cover the past and present distribution ranges for hangul,
starting from Bandipur (past range) in the far north-western region of
the landscape to the southernmost suitable habitat node, i.e., the Desu
range (past range). Qualitative assessment of the habitat ranges was
performed by computing the zonal statistics across the protected area
(PA) boundary, including the national park (NP), wildlife sanctuary
(WLS) and adjacent conservation reserves (CR) in the DL. In addition,
a zonal evaluation was performed for the past distribution range bound-
ary to understand the present habitat conditions in those areas and in-
vestigate the possibility of reintroduction.
2.6. Habitat management strategy through geometric reserve design
Considering the logistic and socioecological limitations of the entire
landscape, we evaluated possible strategies for landscape-level hangul
habitat management planning in the study landscape. We evaluated
the integrity and connectedness of the suitable patches in the DL to sug-
gest habitat block-level planning in the landscape at a finer scale. Habi-
tat block-level planning in the landscape can be considered a
compromise that originates from the trade-off between the funds and
resources available for managing suitable patch-level integrity and the
connectedness among the respective patches. In the present study, we
followed the suggested approaches of geometric reserve design and
landscape-level conservation strategies suggested by Diamond (1975)
and Donaldson et al. (2017). For comparative evaluation of the habitat
quality and geometric integrity at the habitat block and landscape
levels, we used FRAGSTATS 4.2 and ArcGIS 10.6.
2.6.1. Landscape-level planning for the long-term viability of hangul
To develop a landscape-level conservation plan for the long-term vi-
ability of hangul, three habitat blocks were selected, i.e., greater
Dachigam, Overa-Aru-Desu and Bandipur-Wangat. The selection of the
habitat blocks was based on the final ensemble habitat model, to in-
crease the habitat connectedness and on expert opinion. The habitat
configuration within the habitat block was evaluated using the final en-
semble model designed based on the magnitude of model arguments
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

(MA), ranging from 0 MA, indicating the lowest level of habitat suitabil- Table 1
ity, to the highest of 4 MA, indicating the highest level of suitability Evaluation metrics for each of participating distribution models. Boosted Regression Trees
(BRT), Generalized Linear Model (GLM), Multivariate Adaptive Regression Splines (MARS)
(Mukherjee et al., 2020). The PLAND (percentage of suitable land), NP and Random Forest (RF). Model evaluation metrics included area under the receiver oper-
(number of suitable patches), and PD (patch density of suitable patches) ator curve (AUC), Proportion Correctly Classified (PCC), True Skill Statistic (TSS), Cohen's
were calculated to quantify suitable areas within the habitat blocks. The kappa, sensitivity, and specificity. Model evaluation was assessed on the training data used
shape complexity of the habitat patches within the selected habitat to the model as well as the withheld CV (Cross Validation) data used for model evaluation
for hangul.
block was assessed using the LPI (largest patch index of the suitable
patches), ED (edge density of the suitable patch) and LSI (landscape Model Dataset AUC Δ AUC PCC TSS KAPPA SPECIFICITY SENSITIVITY
shape index). In addition, the fragmentation within the respective hab- BRT TRAIN 0.96 0.07 89.3 0.78 0.77 0.89 0.89
itat blocks was measured using the IJI (interspersion and juxtaposition CV 0.89 83 0.66 0.65 0.81 0.85
index) and AI (aggregation index) of the suitable patches (Midha and GLM TRAIN 0.84 0.02 78.9 0.57 0.56 0.78 0.79
Mathur, 2010; McGarigal and McComb, 1995). CV 0.82 73.9 0.47 0.45 0.75 0.72
MARS TRAIN 0.92 0.03 85.2 0.70 0.69 0.85 0.85
CV 0.89 76.5 0.51 0.50 0.81 0.7
2.6.2. Patch susceptibility and role of conservation reserves (CR) in the RF TRAIN 0.83 0 74.6 0.49 0.48 0.74 0.75
Dachigam landscape CV 0.83 73.2 0.45 0.45 0.75 0.69
The habitat patches susceptible to fragmentation were selected
using the mean suitability of the highest model argument (4 MA) within
the total study area. The structural complexity and level of patch vulner-
ability were evaluated using the patch-level configuration metrics in surpassed the standard AUC threshold of 0.75, all models were selected
FRAGSTATS 4.1. The compactness of the selected patches was quantified to build the ensemble model. The comparative scores of the PCC, TSS,
using the PERIM (patch perimeter), SHAPE (shape index), CIRCLE (re- Kappa, sensitivity, and specificity were also in useful ranges for both
lated circumscribing circle) and FRAC (fractal dimension index) indices. the test and CV data (Table 1). The ΔAUC was computed to understand
The perimeter-area ratio (PARA) and radius of gyration (GYRATE) were the efficacy of model fitting, and suggests excellent fitting of the training
calculated for the area and extent evaluation. The connectedness among data with the CV split; the RF model had the best ΔAUC score, with a
the patches was evaluated using the CONTIG (Contiguity index), in perfect zero, followed by GLM with a score of 0.02 (Table 1,
which the lower limit of 0 indicates a single pixel and the value in- Figs. S11, S12).
creases as the patch contiguity and connectedness increase (LaGro, Among the variables, climatic predictors play the most significant
1991). role in determining the potential hangul habitat in Kashmir Valley.
To understand the importance of conservation reserves (CRs) in The annual temperature range (bio_7) is the most important variable
hangul conservation and management planning, we selected (n = 8) affecting the distribution of the species, with a μAUC of 0.24 (Table 2).
CRs located in the DL, these PA forest areas are closely associated with The contribution of precipitation in the warmest quarter (bio_18) is
suitable habitats and the existed PAs in the landscape. The zonal statis- the next most important factor, with a μAUC of 0.198. Among the land
tics for each CR was computed to understand their relative importance cover type predictors, the distance from the coniferous forest (ConFor)
for long-term hangul habitat viability within the study area. and distance from broadleaf forest contribute the most, with μAUC
scores of 0.245 and 0.223, respectively.
3. Results The mean suitability for hangul is the highest for Dachigam NP,
0.72 ± 0.12, suggesting the habitability of the park (Table 3). However,
3.1. LULC mapping the Overa-Aru WLS lies adjacent to Dachigam and has lower mean
values but the highest sum suitability due to its proportionate area.
An area of ca. 9669 km2 was delineated for the distribution predic- The suitable regions in the Overa-Aru WLS are located in the southern
tion model, considering the historical distribution of hangul in the Kash- region of the Dachigam NP; however, suitable habitat patches are also
mir Valley. Forest cover accounts for 67.7% (6553 km2), followed by distributed outside the PA network of the study landscape. The compar-
human habitation, ca. 10.3% (999.2 km2), and orchard and cropland, col- ative analysis of the patch distribution sites suggests that the Desu and
lectively ca. 9.4% (909.4 km2). Among the forest types, coniferous and Sindh Valley forest ranges are still habitable for the species, with mean
broadleaf mixed forest are the dominant forest types, accounting for scores of 0.49 ± 0.18 and 0.45 ± 0.16, respectively (Table 3). However,
ca. 40% (3904 km2) and 7.5% (728 km2), respectively (Table S1). The the Lidar Valley, which lies between the Dachigam NP and Overa-Aru
non-forest LULC includes temperate grassland and scrubland, human WLS, shows comparatively lower ranges of suitability. Out of the
settlements, croplands (horticulture, agriculture crops), rocky clefts, 9669 km2 of study landscape, only 950.28 km2 (9.82%) is highly suitable
water bodies and snow-covered areas. The overall classification accu- for hangul, supported by an MA of four, and 911.06 km2 had an MA of
racy was 83.33% and the overall kappa's statistic was 0.812, indicating three. The unsuitable area for hangul (0 MA) is 5940 km2 (61.43%)
relatively good classification (Table S2). (Fig. 2).

3.2. Model performance and temporal change dynamics of suitable 3.3. Biological connectivity and landscape configuration analysis of suitable
hangul areas hangul habitat blocks

After eliminating the correlated locations through spatial autocorre-
lation testing, n = 38 presence locations were used for modeling. After
elimination of the multicollinearity among the primary predictor vari-
ables using the Pearson's correlation (r < 0.8), eight variables were se-
lected for model building (Figs. S1–S3). The high AUC of all four
models ranges from 0.83 to 0.96 for the training data, and from 0.83 to
0.89 for the (n = 10) fold cross-validation (CV) split (Table 1, Fig. S4),
indicating excellent performance. Among the models, the BRT had the
highest AUC score of 0.96 for training and 0.89 for CV (Table 1,
Figs. S4–S6), followed by MARS, with an AUC of 0.92 for training and
0.89 for CV (Table 1, Figs. S4, S9, S10). As all the participating models
We evaluated the biological connectivity of the past and present dis-
tribution sites of the species in Bandipur, Sindh Valley, Dachigam, Lidder
Valley and Desu Forest. The Lidder Valley habitat block contains an over-
lapping region with the Dachigam NP and Overa-Aru WLS protected re-
serves (Fig. 3). The habitat connectivity is the highest for greater
Dachigam, with a mean score of 0.055, followed by the Bandipur-
Wangat and Overa-Aru-Desu habitat blocks, with mean connectivity
scores of 0.035 and 0.023 respectively (Table 4). We found a high level
of connectivity between the Dachigam NP and Wangat region, which
covers the eastern part of the Sindh Valley in central Kashmir. However,
the habitat connectivity between the Dachigam NP and Sindh Valley did

5
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

Table 2
Variable importance using the increase in Area under the Curve (AUC) when each predictor variable is permuted using four different modeling environments to model habitat suitability of
hangul.

Predictor Code GLM MARS BRT RF μ (mean)

Temperature Annual Range (BIO5-BIO6) bio_7 0.328 0.295 0.245 0.134 0.2415
Precipitation of Warmest Quarter bio_18 0.224 0.251 0.124 0.195 0.1985
Euclidian distance to road road 0.131 0.033 0.065 0.085 0.0785
Precipitation of Wettest Quarter bio_16 0.05 0.058 0.087 0.065 0.065
Euclidian distance to Broad leaf mixed forest BrolFor 0.186 0.145 0.276 0.286 0.223
Euclidian distance to Conifer forest ConFor 0.251 0.164 0.298 0.267 0.245
Slope Slope 0.182 0.145 0.098 0.270 0.1737
Euclidian distance to Temperate grassland and scrubland TemFor 0.055 0.03 0.014 0.08 0.044

Table 3 of the Dachigam NP has high connectivity with Shikargaha-Tral and par-
Zonal statistics of the past and present distributional ranges of hangul in Kashmir valley. tial connectivity with the historic distribution sites of the Desu forest
Name Area km2 Min Max Range Mean Std Sum range.
Based on the number of model arguments and the mean patch size
Desu 32.68 0.04 0.84 0.80 0.49 0.18 20,249
Sindh Valley 107.48 0.07 0.79 0.72 0.45 0.16 60,475 of the most suitable patches, three habitat blocks were selected for
Lidder Valley 157.39 0.07 0.84 0.78 0.43 0.17 85,278 landscape-level planning for hangul: greater Dachigam, Overa-Aru-
Bandipur 266.18 0.01 0.87 0.86 0.24 0.25 79,902 Desu and Bandipur-Wangat (Fig. 4). With the highest (4 MA) model ar-
Overa -Aru WLS 429.55 0.01 0.87 0.87 0.31 0.22 166,667 gument and lowest edge density (ED) (12.08), the greater Dachigam
Dachigam NP 168.36 0.25 0.91 0.66 0.72 0.12 152,373
(n = 228 patches) habitat block is the most favourable for the long-
term viability of hangul (Table 4). Moreover, the large patch index
(LPI) score of greater Dachigam was much higher than that of the rest
not radiate from the central region of the landscape. There are bottle- of the habitat blocks, with a score of 42.49, indicating that the area is
neck areas, and one bottleneck in the landscape was identified in the composed of continuous larger suitable habitat patches. The Overa-
Akhal region. A considerable level of habitat connectivity was observed Aru-Desu suitable habitat block has a higher number of suitable patches
from the Wangat region to the Bandipur region (historical site). The two with higher patch density (PD) than the Bandipur-Wangat habitat
adjacent protected areas, i.e., the Dachigam NP and Overa-Aru WLS, are block. However, the higher LSI and ED values of the Overa-Aru-Desu
connected to the southern region through the lower region of the Lidder habitat block suggest a higher shape complexity and disaggregation of
Valley (Fig. 3). In addition, the model suggests that the southern region patches.

Fig. 2. Ensemble probability and Ensemble count map for hangul within the study landscape. A. Represents the ensemble probability surface for hangul, where the float values ranged from
0 (Unsuitable) to 1 (Highest suitable). B. represents the categorical demarcation of the suitability, resulted from the number of model argument (MA), where 0 MA is lowest argument
(represents areas found unsuitable by all the participating models) to 4 MA at highest model argument (represents found suitable by all the four participating models).

6
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

Fig. 3. Biological connectivity of hangul within the Kashmir valley. The map represents the cumulative current flow within the study area, the potential habitat blocks identified for
landscape level conservation planning (Bandipur-Wangat, Greater Dachigam and Overa_Aru-Desu) was demarcated by different colour. Total of eight Conservation Reserves (CR) was
found to be associated with the protected area network within the area, demarcated in different colour code. The past distributional sites (Bangipur, Desu, Lidder Valley and Sindh
Valley) of the hangul were obtained from Ahmad (2006). The circular dots represent the important connecting location within the landscape. (For interpretation of the references to
colour in this figure legend, the reader is referred to the web version of this article.)

3.4. Patch susceptibility and the role of conservation reserves for the conser-
vation of Hangul
Based on the mean suitability of the highest model argument and
highest patch area, seven patches were selected for the comparative
patch-level shape complexity analysis (Fig. 4). Higher values of patch
disintegration and other shape complexity indices, viz., PARA, SHAPE,
FRAC and CIRCLE, were observed for Patch-2 (PID-2) (Table 5). How-
ever, considering the relatively large size of Patch-5 (PID-5) and its
lower PARA and higher CONTIG values, PID-5 has higher relative
contiguity and connectedness among the suitable patches. PID-5 en-
compasses Dachigam NP and Dara/Kimber CR and continues to the
north-western section of the landscape. Patch-6 (PID-6) is one of
the largest patches habitable for hangul and has structural similarity
to PID-5 (Fig. 4, Table 5). However, the high PARA value for PID-6 indi-
cates a high susceptibility for fragmentation in the future.
To understand the role and imperatives of CRs in hangul conserva-
tion within the study landscape, we evaluated the zonal statistics for
hangul within the CR from the final ensemble probability surface. Our
results suggest that all the CRs in the surrounding areas of Dachigam
NP, viz., Dara/Kimber, Brein, Nishat, Khonmoh, Khrew, and Khangund,
representing greater Dachigam, possess suitable habitat patches for
hangul. Among them, the Dara/Kimber CR has the most suitable
patches, with a mean score of 0.72, followed by the Khangund CR with

Table 4
Habitat configuration estimation and zonal connectivity within landscape level habitat block.

Habitat block Class PLAND NP PD LPI ED LSI IJI AI Connectivity score

Greater Dachigam 1 MA 12.42 465.00 0.81 2.06 12.37 22.19 51.86 92.89 (Mean-0.055)
2 MA 9.57 887.00 1.54 1.64 19.49 38.54 59.23 85.67 (Sum score 39,964)
3 MA 16.11 807.00 1.40 3.13 22.28 33.77 55.96 90.36
4 MA 43.91 228.00 0.40 42.49 12.08 11.08 11.05 98.21
Overa_Aru-Desu 1 MA 25.01 947.00 1.49 15.32 18.48 24.35 50.30 94.76 (Mean-0.023)
2 MA 15.54 1909.00 3.00 3.23 35.82 57.71 55.70 83.85 (Sum score-18,378)
3 MA 22.48 1614.00 2.54 6.61 38.11 51.14 54.89 88.14
4 MA 27.19 684.00 1.08 16.43 16.47 20.07 11.38 95.90
Bandipur-Wangat 1 MA 23.92 1061.00 1.63 10.81 19.72 26.71 49.15 94.17 (Mean-0.035)
2 MA 24.46 1266.00 1.94 7.67 31.49 40.89 57.11 91.06 (Sum score-29,395)
3 MA 17.05 1847.00 2.84 1.95 33.85 52.73 59.48 86.10
4 MA 28.21 355.00 0.55 12.78 13.50 16.28 16.84 96.81

Note: Class represents suitable habitat of hangul as per number of model argument, where (1 MA) represents suitable habitat for hangul by one model to a highest (4 MA), represents
suitable habitat as per the all participating model.

7
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

Fig. 4. Represents the identified most suitable patches for hangul (PID-1-PID-7) where the colour code represents the respective size of the suitable patch. Potential habitat blocks
identified for landscape level conservation planning where A. represents the total landscape B. represents Bandipur-Wangat, C. represents Overa_Aru-Desu and D. represents Greater
Dachigam and was demarcated by different colour. Past distributional ranges (Bangipur, Desu, Lidder Valley and Sindh Valley) of the hangul obtained from Ahmad (2006). Circular
dots represents the important connecting location within the landscape. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of
this article.)

Table 5
Shape complexity evaluation of the selected suitable patches for hangul in Kashmir valley.

Patch-ID Percentage of area PERIM GYRATE PARA SHAPE FRAC CIRCLE CONTIG

PID -1 14.51 160,769.07 4225.20 16.11 4.02 1.15 0.60 0.99

PID -2 8.82 258,110.82 4184.80 42.53 8.27 1.24 0.76 0.97
PID -3 7.87 154,392.48 3341.53 28.53 5.24 1.19 0.60 0.98
PID -4 12.08 229,105.80 5413.95 27.57 6.28 1.20 0.75 0.98
PID -5 35.81 471,359.79 7343.79 19.14 7.50 1.21 0.72 0.99
PID -6 15.33 289,824.48 4708.52 27.50 7.05 1.21 0.64 0.98
PID -7 5.58 128,096.10 2766.30 33.39 5.16 1.19 0.74 0.97

8
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

a mean score of 0.61 (Table 6). However, the Shikargah and Khiram CRs prevent them from scorching heat from the sun (Ahmad, 2006;
situated in the southern region of the landscape did not represent signif- Holloway, 1971; Sharma et al., 2010; Staines, 1976;). Previous studies
icantly useful patches, except for a few small and fragmented patches also suggest that in the winter when the forage availability in Dachigam
(Table 6; Figs. 2 and 4). landscape dropped to its lowest, most of the forages were restricted in
the conifer and broadleaf habitats in the form of browse such as fallen
4. Discussion leaves, acorns, ferns, grasses and forbs, which remained green through-
out the winter. Further, the close vicinity of forested habitats with the
The LULC analysis showed there are still a considerable amount of water sources makes them suitable for the species (Staines, 1976).
forest habitats in the landscape, indicating a healthy future for wildlife Apart from the contribution of the conifer forest and broadleaf
conservation in the Kashmir Valley (Table S1). The landcover classifica- mixed forest, substantial contribution of temperate grassland and
tion results suggest that coniferous forest constitutes approximately shrubland (TemFor) was also observed (Table 2), which can be attrib-
40% of the landscape, followed by alpine meadows (15.4%), and the uted to the fact that these habitats offered abundant fodder (Ahmad,
human settlements cover 10.30%. A rapid change in the LULC due to 2006). The grassland habitats are rich in platable grasses and used by
the conversion of forested habitats to other land types may negatively the species predominately in the spring season because of high nutri-
impact wildlife and severely hamper ecosystem services in the Kashmir tional values (Segelquist et al., 1972; Short, 1975). Some grasses and
Valley. Sharma (2012) documented low-to-medium levels of fragmen- herbs remain green in these areas even during the winter and become
tation in the DL due to unplanned developmental and anthropogenic ac- available to hangul when show covered the ground (Ahmad, 2006).
tivities. Visual interpretation and landscape complexity indices such as The distance to roads was a negative predictor, suggesting that areas
interspersion & juxtaposition (IJI), the contiguity index, and cohesion away from roads are relatively more suitable for the species (Supple-
index show poor connectivity and sparse spatial distribution of forest mentary Figs. S5, S7, S9 and S11). Red deer are reported to avoid
patches, suggesting fragmentation in the DL (Sharma, 2012). good-quality habitats that are close to roads and can resolve the
trade-off between the use of high-quality resources and risk avoidance
by modifying their habitat use pattern in days and nights (Guisan and
4.1. Suitable hangul habitats and landscape ecology
Zimmermann, 2000). Studies conducted on other red deer subspecies
have documented that deer behaviour and habitat use can be signifi-
The present ensemble habitat model of hangul habitat provides a
cantly influenced by human activities and roads (Torres et al., 2012;
clear and easily interpretable map of potential hangul habitat in the
Bonnot et al., 2013). The connectivity among the forest patches medi-
Kashmir Valley (Fig. 2). It also identifies the key habitat features that
ates dispersion and maintains gene flow, which is imperative for the
are significant predictors of hangul habitat and their relative importance
species' long-term viability (Manel et al., 2003). The habitat patch struc-
(Table 3). The final ensemble model indicates the combined influence of
ture and composition substantially impact the landscape ecology of
bioclimatic and landcover predictors in shaping the suitable habitat for
mammals (Bowers and Matter, 1997). Thus, it is vital to assess the
hangul in the study area. Among the bioclimatic predictors, the annual
level of habitat fragmentation for long-term sustenance of the focal spe-
temperature range (bio 7) was found to be most significant, suggesting
cies (Jaeger et al., 2005).
that the correlation with hangul and its distribution is affected by the
Our results from the circuit model suggests high mean connectivity
range of extreme temperatures in the region. Moreover, recent climate
between Dachigam NP and its associated CRs, which can collaborate
change studies have indicated that the Himalayan region is warming
with a higher level of legal protection in these regions (Ahmad, 2006;
faster than the global average rates (Liu and Chen, 2000; Shrestha
Charoo et al., 2010). However, lower mean connectivity values in the
et al., 1999), which increases the threat of geographic isotherm contrac-
Overa_Aru-Desu was observed due to the lack of suitable habitats and
tion in the study region for the entire biota. A number of studies indi-
the lack of CRs in the adjacent ranges (Ahmad, 2006). Moreover, the
cated that the species are vulnerable to climate change in mountain
southern region of the Overa_Aru-Desu was also known to be predom-
ecosystems (Negi et al., 2012; Liu and Chen, 2000). Among the
inantly influenced by excessive grazing by livestock. Furthermore, the
landcover predictors, the distance from coniferous and broadleaf
region between the DNP and Bandipur-Wangat is also under the influ-
mixed habitats significantly positively influenced hangul habitat.
ence of livestock grazing, lopping and firewood collection by the com-
These habitat types have high connectivity, and patches are spatially
munities located on both the banks of the Sind River of the Sind Forest
close in proximity (Supplementary Figs. S5, S7, S9 and S11), which can
Division (Ahmad, 2006).
be corroborated by the habitat ecology findings for hangul (Ahmad,
While at the finer scale visualization of the prediction map, the fine
2006; Sharma et al., 2010). The conifer habitats in Dachigam provides
patchwork of the probability (dark colour) area indicates regions with
preferred food items, shelter and escaping hides from predators;
relatively high importance for hangul (Fig. 2). The most suitable habitats
hence were found to be utilized more by hangul (Ahmad, 2006).
are distributed in medium elevation zones with flat to moderately
Other studies indicate that the Red deer occupy dense canopy forests
steeper slopes and available food and water resources (Ahmad, 2006;
during summer and winter. The dense canopy forests provide shelter
Sharma et al., 2010). However, seasonal changes in the use of elevation
to Red deer in winter and prevent them from wind-chill, whereas, in
zones have been reported for hangul as a strategy to utilize resources
summer, these areas were used as day resting sites as a strategy to
depending on availability and also as an adaptation to extreme cold dur-
ing the winter (Kurt, 1978; Sharma et al., 2010; Ahmad, 2006). Highly
suitable hangul areas are mostly distributed in the DL and north Kash-
Table 6
Zonal Statistics of the Conservation Reserve (CR) residing in close proximity of Dachigam
mir region, where hangul was historically reported to be distributed
National park and Overa-Aru Wildlife Sanctuary in Kashmir valley. (Gee, 1965) (Figs. 1 and 2). Our results also suggest that hangul has
lost much of its historic range, and the magnitude of loss is the highest
CR name Area km2 Min Max Range Mean Std Sum
for the Bandipora Forest range. Among the past and present distribution
Dara/Kimber 1.22 0.33 0.88 0.55 0.72 0.13 1110.04 of hangul, the Dachigam landscape was found to be the most habitable
Brein 0.47 0.15 0.74 0.59 0.32 0.15 191.07
region, which has been reported by other studies (Ahmad et al., 2009;
Nishat 0.46 0.23 0.88 0.65 0.45 0.23 259.91
Khonmoh 1.19 0.06 0.91 0.84 0.44 0.31 653.19 Kaul et al., 2018). In addition, our findings suggest the relative unsuit-
Khrew 1.01 0.04 0.90 0.87 0.34 0.29 440.72 ability of much of the Overa-Aru WLS, especially in the northern and
Khangund 1.07 0.27 0.89 0.62 0.61 0.17 824.00 central parts of the WLS. This contradicts the findings of Ahmad
Shikargah 1.06 0.03 0.83 0.80 0.26 0.23 350.79 (2006), which suggest the potential of the Overa-Aru WLS as a relic
Khiram 1.07 0.04 0.83 0.78 0.24 0.20 325.73
site for breeding programs.

9
T. Mukherjee, V. Sharma, L.K. Sharma et al. Science of the Total Environment 777 (2021) 146031

4.2. Landscape-level strategy for long term viability
Funding and resources are considered limitations for the implemen-
tation of habitat improvement strategies in the entire suitable habitat of
the species; hence, the present study focused on habitat block-level
planning. Our results suggest that, within the present and past distribu-
tion ranges of hangul, Dachigam NP is the most suitable for the species.
Moreover, the forested ranges of the Sindh Valley Forest Division and
Desu regions have a substantial level of suitable habitat for the species
(Fig. 2). Enhanced protection in the remaining suitable forest patches
of the Sindh Valley region may be crucial for hangul since these patches
act as a bridge between the greater Dachigam and Bandipur-Wangat
habitat blocks. However, the Desu forest range is one of the last habitat
nodes in the south-eastern range of the species (Fig. 2).
We created a rating system for conservation prioritization of the
areas based on the geometric approach of reserve planning (Diamond,
1975) (Table 7). Our reserve planning strategy is based on the habitat
quality, habitat integrity, connectedness, size and number of habitable
patches. The greater Dachigam landscape is the most suitable for species
conservation and habitat management planning based on the rating
system of the geometric approach (Table 7). Moreover, greater
Dachigam has secure perimeters comprised of several protected areas
(NP, WLS, CR), which makes the area appropriate as it is cost-effective
and follows the principles of landscape ecology. The comparative evalu-
ation of the CR habitat configuration at the patch level suggests that ex-
pansion of the Dara/Kimber CR should occur in the north-western
direction to cover the remaining habitat patch (PID-5) under phase I
landscape-level conservation planning.
The conservation prospects for hangul in the remaining habitat
blocks, i.e., Bandipur-Wangat and Overa-Aru-Desu, can be considered
for habitat planning in phase II. Based on the comparative evaluation,
Table 7
Evaluation of habitat blocks through geometric reserve design principle, derived from
Diamond (1975). For all classes (Size, Number, Proximity, Connectivity and Shape),
species extinction rate is expected to be lower for Better than the Worst category. Color
codes represents ranking of habitat blocks for prioritization.
Design Better Worst Habitat block priority ranking
for conservation planning
Principal
Better Greater Dachigam
Size
Medium Overa_Aru-Desu
Smaller
Worst Bandipur-Wangat
Better
Number Greater Dachigam
Medium
Bandipur-Wangat
Single Several
Worst
Overa_Aru-Desu
Better
Proximity Greater Dachigam
the Bandipur-Wangat habitat block is more suitable than the
Overa_Aru-Desu habitat block. However, the PID-6 in the southern re-
gion of the Overa_Aru WLS can't be ignored, as the patch-level integrity
results suggest recent disintegration and increasing fragmentation of
the habitat. PID-6 can potentially act as a stepping stone habitat for han-
gul to establish landscape-level connectivity from the Desu region to the
greater Dachigam habitat block. Thus, we propose including the suitable
region of PID-6 in the Overa_Aru WLS or establishing a new CR for the
conservation of hangul.
5. Conclusion
The present approach to identifying the conservation priority areas
through geometric reserve designing may provide a simpler and cost-
effective way for the park managers to make conservation decisions.
Our finding suggests that protection of remaining habitat for hangul
should be prioritized because it has lost a significant area of its historic
range in the recent past. In recent decades, conservation action plans
and initiatives have evolved from a reserve-based approach to a
landscape-level approach due to the accelerated threat of climate
change and habitat fragmentation. Such threats led to an increasingly
dynamic pattern of colonization and extinction. Our proposed approach
of habitat block-level planning in the landscape can be considered a
compromise that originates from the trade-off between the funds and
resources available for managing suitable patch-level integrity and the
connectedness among the respective patches. The government of
India and the Department of Wildlife Protection, Jammu and Kashmir
have initiated a hangul recovery plan. However, much of the efforts
are focused on captive breeding for restocking wildlife populations.
But the mapping and delineating areas where hangul can be
reintroduced are pivotal for the success of hangul recovery plans. The
potential hangul habitats identified in the present study should be con-
sidered for the future release/reintroduction program of captive-bred
individuals. Finally, we recommend creating Greater Dachigam National
Park (GDNP) based on the habitat block identified in the present analy-
sis. The creation of GDNP through joining the small and isolated PAs in
the Dachigam landscape may be critical for maintaining the long-term
viability of this critically endangered species.
Supported Furthermore, there is a need to invest in habitat enhancement activ-
ities in identified corridor areas and to stop nomadic grazing in the en-
Indices
tire landscape. Additionally, efforts should be made to restore hangul
LPI habitat while considering the habitat predictors and requirements
with respect to food resources, especially the restoration of coniferous
PLAND
and broadleaf mixed forests. Hence, we suggest reorganization of the
existing management regime of Dachigam NP to support landscape-
NP level and habitat block-level management planning. We also recom-
mend notification of PID-6 in the southern region of the landscape as
PD
Conservation Reserve or may be brought into Overa-Aru WLS, as this
habitat patch may act as a stepping stone habitat. PID-6 may be vital
for maintaining the landscape connectivity and metapopulation dy-
AI namics of the species.

Medium
Bandipur-Wangat
IJI
Data availability statement
Worst
Aggregated Widely Overa_Aru-Desu
Raster data pertaining to land use, climatic data, topography and an-
Better
Greater Dachigam Mean
thropogenic variables were used for model building using open source
Connectivity
data. The presence locations of the species can be made available upon
current
Medium
Bandipur-Wangat reasonable request.
flow
Connected Isolated Worst
Overa_Aru-Desu
CRediT authorship contribution statement
Better
Shape Greater Dachigam ED
LKS and VS conceptualized and designed the study. LKS, VS, TM, AS,
Medium
Bandipur-Wangat LSI AT, RD and MT participated in data generation, quality check and pri-
Simple Complex Worst
Overa_Aru-Desu
mary data analysis. TM, VS, LKS, AT, BDJ, RD, SD, MT and BT and wrote
the primary draft of the manuscript. LKS, TM, VS, BDJ, AS, AT, RD, SD,

10
T. Mukherjee, V. Sharma, L.K. Sharma et al.
MT, BT and KC finalized the manuscript. LKS supervised the overall ac-
tivities and KC raised funds for the study.
Declaration of competing interest
The authors do not have any conflict of interest with respect to pub-
lication of this manuscript.
Acknowledgements
Authors are thankful to Director and Head of the Office, Zoological
Survey of India, Kolkata for encouragement and support. The authors
thank Dr. Hitendra Padalia, Scientist-SF, Indian Institute of Remote
Sensing, Dehradun for their help and guidance during the P.G Diploma
project work of the third author. We acknowledge the National Mission
for Himalayan Studies, Ministry of Environment, Forest and Climate
Change (MoEF&CC), Government of India for the funding support under
the Grant No. NMHS/2017-18/LG09/02.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.scitotenv.2021.146031.
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