Global Ecology and Conservation 37 (2022) e02167

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Global Ecology and Conservation

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

Incorporating satellite remote sensing for improving potential

habitat simulation of Prosopis cineraria (L.) Druce in United
Arab Emirates
Remya Kottarathu Kalarikkal, Youngwook Kim *, Taoufik Ksiksi
Department of Biology, College of Science, United Arab Emirates University, Al Ain, United Arab Emirates

A R T I C L E I N F O A B S T R A C T

Keywords: Predicting the suitable habitat of a species is one of the adequate and enhancing approach in
Habitat suitability biodiversity conservation planning and implementation. Species habitat distribution is closely
Remote sensing linked with environmental factors and bioclimatic variables used as prediction variables in spe­
SDM
cies distribution models (SDMs). Recent application of satellite remote sensing data and the
MaxEnt
NDVI
bioclimatic variables has created an advanced way to improve the SDMs performance. In this
Prosopis cineraria study, MaxEnt was used to predict species habitat distribution. Our objectives are to assess the
Ghaf tree application of satellite remote sensing data in predicting the potential habitat suitability of the
UAE arid plant species. We have selected Prosopis cineraria (L.) Druce (Ghaf), the national tree of the
UAE. We have chosen 33 environmental variables along with 90 species occurrences and for the
final model simulation we have used three modeling scenarios. MaxEnt results showed that the
model simulation with all key variables has substantially improved the potential habitat suit­
ability prediction with a mean Area Under receiver operating Curve (AUC) value of 0.984,
indicating a better predictive accuracy in the integration of satellite remote sensing data, edaphic
variables and topographic parameters. Model results showed that the spatial proportions of the
potential habitat suitability in the UAE consisted of high (2%), medium (6%) and low (9%)
habitat suitability classes. The MaxEnt results revealed that precipitation of the coldest quarter
(32.5%), NDVI (12.1%) and elevation (8%), had significant contribution to the potential habitat
distribution of UAE Ghaf trees. Cold season precipitation is the most significant climate constraint
on the habitat distribution. The most habitat suitability of Ghaf tree in the UAE was within a
certain range of NDVI, LST, elevation, aspect, and precipitation patterns. This paper finding could
be useful for environmental managers for implementing reforestation not only in UAE but also for
the entire middle eastern regions with similar hyper arid conditions.

1. Introduction

Drylands on Earth are critical segment of physical and human environments (Singh and Chudasama, 2021), making up more than
40% of the land surface (Smith et al., 2019). Drylands provide habitat for rare and endemic species, and forage for wildlife (Bes­
telmeyer et al., 2015). Dryland ecosystems are strongly influenced by variation of surface air temperature, drought period, precipi­
tation, and experiencing the increasing vulnerability of these ecosystems to climate change (Ramarao et al., 2019; Yadav and Lal,

* Corresponding author.
E-mail address: youngwook.kim@uaeu.ac.ae (Y. Kim).

https://doi.org/10.1016/j.gecco.2022.e02167
Received 13 January 2022; Received in revised form 12 May 2022; Accepted 18 May 2022
Available online 24 May 2022
2351-9894/© 2022 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/by-nc-nd/4.0/).
R.K. Kalarikkal et al. Global Ecology and Conservation 37 (2022) e02167

2018). Tree species are considered as major biodiversity supporters and keystone species of the dryland ecosystems (Munzbergova and
Ward, 2002). Arid trees influence their understory vegetation by reducing the extreme temperature (Greenlee and Callaway, 1996),
providing improved soil texture and nutrients (Pugnaire et al., 2004; Moro et al., 1997; Belsky, 1994), and availability of soil moisture
by hydraulic lift (Ludwig et al., 2001). Prosopis species are one of the arid nurse plant canopies which are spatially distributed in Oman,
India, Afghanistan, Pakistan, Iran and Arabian countries (Jinu et al., 2017; El-Keblawy and Abdelfatah, 2014). Prosopis species is
considered as characteristic tree of most of the desert thorn forest, secondary dry deciduous forest and desert dune scrub (Mann et al.,
1977; Bhansali, 2010).
With the good amount of macro-nutrients and organic matter beneath their canopy many Prosopis species cultivate a “resource-
island” in the desert ecosystems (Kaur et al., 2012).
The recent rapid deterioration of biodiversity in the arid regions was reported as a severe environmental challenge in the current
era (Pimm, 2021; Barnosky et al., 2011). Identification, and protection of suitable habitat of significant species within the arid
ecosystem might be a major solution for the rapid biodiversity deterioration (West et al., 2016). Prosopis cineraria (L.) Druce belongs to
Leguminosae family (P. cineraria) plays a significant role of biodiversity in the desert ecosystems (NAS 1979) due to its adaptation to
grow in relatively low fertile soil conditions and moisture availability (Shankarnarayan et al., 1987). These trees are highly compatible
with the agricultural crops due to their nitrogen fixing and soil organic matter recharging ability (Toky and Bisht 1992). In United Arab
Emirates (UAE) P. cineraria is considered as the national tree and known as Ghaf tree (Al Ghais et al., 2020). Entire plant part of Ghaf
tree is versatile, and thus it is also called as the ‘King of Desert’, ‘love tree’ and ‘Pride of the Desert’ (Singh et al., 2021). Common
threats faced by this wild species in the country are mainly urbanization, over grazing, climate change, species invasion and excessive
ground water extraction (Howari et al., 2022; Gallacher and Hill, 2005). In order to conserve this species UAE government has
implemented rules along with massive planting of Ghaf saplings over the country (Aspinall, 2001).
Species distribution models (SDMs) are widely used in the field of ecology for more than two decades (He et al., 2015). SDMs
become a common tool in ecology to map the species habitat distribution (Wang et al., 2020), to find the suitable habitat, to test
ecological theories (Petitpierre et .al., 2012), to identify species invasion risk (Negrini et al., 2020; Truong et al., 2017; West et al.,
2016), to predict the environmental or climate change responses to the species habitat (Elith* et al., 2006; Zimmermann et al., 2010),
and to facilitate implementation and selection of nature reserves for conservation (Engler et al., 2004; Zhang et al., 2012; Murray-­
Smith et al., 2009). SDMs operate statistically based on the species occurrence data and the spatial interaction with its environmental
predictor variables (Morán-Ordóñez et al., 2012). Wide spatial application ranges from global, regional and landscape level often made
SDMs a common tool in ecology (Dakhil et al., 2021; Hao et al., 2019). Even though SDMs provide efficient and effective simulations,
their prediction potential is habitually constrained by lack of predictor variables spatial explicitly and unavailability of species absence
data (He et al., 2015).
Maximum Entropy distribution (MaxEnt; Phillips et al., 2006) model based on an ecological niche machine-learning algorithm is
one of the robust (Guisan et al., 2007) and frequently applied (Almarinez et al., 2021) algorithm among the available numerous SDMs
(Guisan and Zimmermann, 2000; Bolker et al., 2009). MaxEnt works based on presence only data as a source of information along with
the predictor variables, making the model more compatible (Dudov, 2017). MaxEnt is allowed for using both continuous and cate­
gorical predictor variables (Elith et al., 2011). The MaxEnt simulation is stable and consistent with good accuracy even with small
sample size (Pearson et al., 2007). However, the unavailability of continuous fine scale predictor variables turned as a major concern
among the ecological modeling society (Segal et al., 2021). The use of satellite remote sensing data provides a new platform for
ecological modelers to find a solution for addressing this spatial scale issues with SDMs (Randin et al., 2020). Satellite remote sensing
data products available at consistently spatial and temporal resolutions characterize an array of ecologically relevant patterns and
processes (Andrew et al., 2014; Buermann et al., 2008; Girma et al., 2016). Satellite remote sensing data can be used directly or
indirectly in biodiversity monitoring, management and conservation (Turner et al., 2003). Remote sensing data rely on the radiometric
properties and spectral signatures received from the targeted species or community (He et al., 2011; Kokaly et al., 2009). Species
biophysical and biochemical properties together influence the spectral signature of the vegetation (Feilhauer et al., 2012). These
spectral signatures are used as good representation for floristic composition identification in vegetation models (He et al., 2009).
Advanced technology in remote sensing and statistical modeling consistently improves the SDMs prediction efficiency (He et al.,
2015). Hence the objectives of this study are to assess the MaxEnt model performance for predicting the potential habitat suitability of
P. cineraria trees by integrating satellite remote sensing data, to understand how the environment influences the distribution of
P. cineraria trees in the UAE, to find the optimal combination of predictor variables for the best MaxEnt simulation results and to project
the potentially suitable habitat for P. cineraria in the regions where the future conservation strategies can be focused.

2. Data and methods

2.1. Study area and species occurrence

The study domain is situated in the UAE, between 22.40–26◦ N: 51–56◦ E. The country consists of seven states (emirates), including
Abu Dhabi, Dubai, Sharjah (SHJ), Ajman, Um Al-Quwain (UAQ), Ras al-Khaimah (RAK) and Fujairah. The UAE is a subtropical desert
with hot and humid climate along the shoreline and comparatively hot and dry climate in the interiors (Özçevik et al., 2015). Average
minimum surface air temperature of UAE is 14 ◦ C and the average maximum surface air temperature is 45 ◦ C (Aldababseh and Temimi,
2017). The warmest months are June, July & August with temperature approximately 50 ◦ C and the coolest months are December,
January & February with a temperature approximately 10 ◦ C (Alqasemi et al., 2021). Mean annual rainfall of UAE is 78 mm, varying
from the southern desert region (40 mm) to the northeastern mountains (160 mm) (Ouarda et al., 2014). The primary habitat of the

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UAE includes sand dunes, sabkhas, coastal mangroves, mountains, and wetlands (Tourenq and Launay, 2008). Even though the UAE
represents a simple ecosystem with a basic food chain, the species occupied in the area has unique and remarkable adaptations to the
harsh climate (Tourenq and Launay, 2008). The occurrence of P. cineraria was recorded by systematic random floristic field survey
conducted during 2018–2019 with the help of potable GPS (Ksiksi et al., 2019). We have selected 90 field observations mainly from Ras
al-Khaimah, Um al-Quwain, Ajman and Abu Dhabi emirates. All these occurrence data were converted to the input file format of
MaxEnt model (Kamyo and Asanok, 2020).

2.2. Satellite remote sensing data

Remote sensing data are widely used to improve SDM performance (Girma et al., 2016; He et al., 2019). Incorporating remote
sensing data in SDM provide more robust predictors with refined habitat distribution maps (Morán-Ordóñez et al., 2012; Peterson and
Nakazawa, 2008; Prates-Clark et al., 2008). Spectral responses to different environmental conditions provide significant information
on vegetation status, land use cover and soil moisture (Franklin and Wulder, 2002) at different spatial and temporal scale which has
important role on explaining species fundamental and realized niche (Feilhauer et al., 2012). We used three satellite remote sensing
data records (Table S1) derived from the Moderate Resolution Imaging Spectroradiometer (MODIS) sensor during 2000–2019 from
Application for Extracting and Exploring Analysis Ready Samples (AppEEARS Team, 2021). Monthly normalized difference vegetation
index (NDVI) with 1 km spatial resolution (MOD13A3; Didan, 2015) was used as optical measure of canopy greenness.
Satellite-derived NDVI is widely used in studying composition and structure of a leaf and canopy (Feilhauer et al., 2012; Huete, 2012;
Kim et al., 2014). Several studies have applied NDVI to assess vegetation responses to climate variability and environmental change
(Fensholt et al., 2012; Kim et al., 2012). The NDVI is sensitive to canopy photosynthesis (Kim et al., 2010) and has been widely used to
analyze regional patterns and temporal variability in vegetation growth (Huete, 2012; Kim et al., 2014).
The MODIS day and night land surface temperature (LST) with 1 km spatial and 8-day temporal resolution ((MOD11A2; Wan et al.,
2021. 4.) were used as the temperature constraint on vegetation growth. The LST represents the radiation temperature measured
between the top of surface materials (plant canopy, water) and the atmospheric interface (Alqasemi et al., 2021; Stroppiana et al.,
2014). LST is one of the significant input variables in most of the soil vegetation modeling (Vlassova et al., 2014) since it is correlated
with the soil moisture and canopy evapotranspiration (Wang and Liang, 2008). The 500 m MODIS yearly total potential evapo­
transpiration (PET) (Gilani et al., 2020; Li et al., 2014) data records (MOD16A3; Running et al., 2017) were used as moisture constraint
on vegetation growth, since the atmospheric evaporative demand has major role in structuring the vegetation communities for a water
limited dry ecosystem (Dakhil et al., 2021; Li et al., 2013a, 2013b; Mod et al., 2016). A 1-km MODIS plant functional type classification
data for 2019 (MCD12Q1; Friedl and Sulla-Menashe, 2019) was used to classify land cover type in the UAE.

2.3. Bioclimatic and environmental data

Bioclimatic variables are used in the SDM studies since these variables describe the eco-physiological tolerance of species (Zhu
et al., 2020; Graham and Hijmans, 2006; Murienne et al., 2009). In this study, we have extracted nineteen bioclimatic variables
(Table S1) with 1-km spatial resolution from the WorldClim-global climate database version 2.1 (Fick and Hijmans, 2017) for the latest
climate condition (1970–2000). Nineteen bioclimatic variables represent annual trends, seasonality and climate constraints derived
from temperature and precipitation. We have also combined seven edaphic variables, including soil quality (Table S1) from the
harmonized world soil database (v 1.2) with 1 km spatial resolution (Fischer et al., 2008). We have extracted the four topographic
parameters, including elevation, slope, aspect, and hill shade from the SRTM (shuttle radar topography mission) with 90 m spatial
resolution (Table S1). All extracted from satellite data records, bioclimatic, and edaphic variables, and topographic parameters are
considered as predictor variables for the model simulation (Table S1).

2.4. Predictor variable selection

All 33 predictor variables were resampled to 1 km spatial resolution by applying nearest neighborhood method from arc toolbox of
ArcMap (10.8.1). All these variables were spatially projected with World Geodetic System (WGS84) and converted to asci files using
ArcMap (10.8.1). The 30-year mean bioclimatic variables were obtained from the WorldClim data base (worldclim.org). We have
derived long term mean (2000–2019) and annual means of NDVI, LST and PET by using ArcMap tool before executing the variable
selection. Selecting the set of predictor variables to simulate the model is one of the crucial phases in SDM, influencing the model
performance (Braunisch et al., 2013). Eliminating the multicollinearity among these predictor variables are significant unless this will
hamper the species-variable relationship when simulating potential habitat suitability (Thapa et al., 2018). Pearson correlation
analysis is one of the major multicollinearity elimination methods widely used in variable selection by assessing the sign and strength
of relationships between variables (Li et al., 2020; Ji et al., 2020; Heikkinen et al., 2006).
Pearson correlation matrix table was used to check multicollinearity among the 33 variables using the SPSS statistics software (IBM
SPSS 20). We eliminated the variables shown r-value ≥ 0.8 with two or more other variables (Table S2). The 0.8 of r-value was used in
the previous study (Fan et al., 2020). After the elimination of colinear variables we run the model to finalize the key variables by
enabling the Jackknife method available within the model. To run the model, we have used the long term means of bioclimatic and
remote sensing variables along with the topographic and edaphic variables. For the final model simulation, we excluded the variables
which resulted ≤ 0.1% of variable contribution based on the Jackknife variable selection method (Hundessa et al., 2018). Jackknife
test identifies the predictor variable with highest gain in the model simulation when applied in isolation and the gain represent the

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average probability of the point localities (Cotrina Sánchez et al., 2021; Moratelli et al., 2011; Phillips and Dudík, 2008).

2.5. MaxEnt model simulation

The habitat distribution for P. cineraria was simulated with the selected key environmental variables using MaxEnt model version
3.4.1. (Phillips et al., 2006). MaxEnt model provides reasonable predictions when compare with other environmental niche models
(Wakie et al., 2020; Kumar et al., 2016). The principle behind the model is maximum entropy, which makes minimal assumptions
about the unknown information and consider the information generated by species presence data (Phillips et al., 2006). The MaxEnt
model has many advantages over other SDMs, including high accuracy and consistent area prediction even with spatially restricted
organisms (Giovanelli et al., 2010).
MaxEnt is more effective for prioritizing areas for invasive species management (Underwood et al., 2004). This model is an
appropriate tool for finding new populations of narrowly endemic and rare organisms (Rhoden et al., 2017). To get better environ­
mental envelope for P. cineraria in the UAE, we selected three scenarios for MaxEnt model simulation.
The first scenario (scenario A) was using all key variables from bioclimate, edaphic, topographic and satellite remote sensing data.
In this scenario we used annual means of satellite data as continuous variable to predict the habitat suitability for the years starting
from 2001 to 2019. The remaining variables, including bioclimate, edaphic and topographic data, were constant for model simulation
in scenario A. We produced habitat suitability map for 2000 as the training year and for 2001–2019 were projected with the scenario A.
Second scenario (scenario B) used bioclimatic and edaphic variables, and topographic parameters, excluding satellite remote sensing
data. In the third scenario (Scenario C), only bioclimatic variables were used. For all these scenarios, same model simulation criteria
was applied to find which scenario provides better performance. From the species record, 75% was used for training the model and
remaining 25% was used for testing the model (Li et al., 2019). Ten model replications were used with cross validation method and
with the clog log output format. Maximum 5000 iteration was selected with a clamping option enabled and the regularization
multiplier was kept as inbuilt.
Model performance was evaluated using the area under the receiver operator characteristic (ROC) curve (AUC) (Gebrewahid et al.,
2020; Pratumchart et al., 2019; Pérez Chaves et al., 2018; Phillips et al., 2006). ROC is a conceptually simple visualizing method which
explain the performance of a classifier and the output of which depends on a threshold parameter (Elith et al., 2011). ROC is a
two-dimensional interpretation and in order to compare the classifiers normally use the single scalar value and the common method is
called AUC (Fawcett, 2006). AUC is widely used for measurement of Maxent performance (Cotrina Sánchez et al., 2021; Du et al., 2021;
Biddle et al., 2021; Phillips and Dudík, 2008). It plots the sensitivity against the fractional predicted area for species (Rodgers et al.,
2019; Allouche et al., 2006). AUC value ranges from 0 to 1, indicating the higher value is a perfect fit (Chaiyos et al., 2018). Jackknife
in-built tool in MaxEnt model was enabled to assess the variable importance to the model simulation (Wei et al., 2018). Jackknife
variable importance test help to identify the variables with significant individual influence on the species habitat distribution
(Promnikorn et al., 2019; Elith et al., 2011). The final model output was reclassified into four habitat suitability classes for 2000–2019.
Finally, we developed a mean habitat suitability distribution map by spatially averaging all the simulated habitat distribution for 20
years. We further classified our potential habitat suitability into four suitability classes based on Wakie et al. (2020): not suitable
habitat (0–0.08; NS), low suitability (0.08–0.2; L), medium suitability (0.2–0.6; M), and high suitability (0.6–1.0; H).

Table 1
All key variables used in the simulation of potential habitat suitability of P. cineraria and their percentage of contribution in the UAE.
Variables Abbreviation Percent Permutation Source
contribution importance

bio19 Precipitation of coldest quarter 32.5 14.3 https://www.worldclim.org/

NDVI Normalized vegetation index 12.1 20.6 https://lpdaac.usgs.gov/products/mod13a3v006/
bio3 Isothermality 8.7 1.6 https://www.worldclim.org/
ELEV Elevation 8 20.9 Generated from SRTM DEM (https://srtm.csi.cgiar.org/)
bio18 Precipitation of warmest 7.3 6 https://www.worldclim.org/
quarter
bio9 Mean temperature of driest 6 3.4 https://www.worldclim.org/
quarter
Slope Slope 5.7 2.4 Generated from SRTM DEM (https://srtm.csi.cgiar.org/)
Aspect Aspect 4.7 3 Generated from SRTM DEM (https://srtm.csi.cgiar.org/)
bio4 Temperature Seasonality 3.6 2.5 https://www.worldclim.org/
bio7 Temperature annual range 3.4 6.5 https://www.worldclim.org/
hillshade Hill shade 2.8 1.2 Generated from SRTM DEM (https://srtm.csi.cgiar.org/)
Sq5 Soil excess salt 1.7 1.7 http://www.fao.org/soils-portal/data-hub/soil-maps-and-
databases/harmonized-world-soil-database-v12/en/
Bio1 Annual mean temperature 1.2 10.1 https://www.worldclim.org/
PET Potential Evapotranspiration 1.2 0.2 https://lpdaac.usgs.gov/products/mod16a2v006/
LST Land Surface Temperature 0.7 0.7 https://lpdaac.usgs.gov/products/mod11a2v006/
Bio 6 Minimum temperature of 0.3 4.5 https://www.worldclim.org/
coldest month
Sq7 Soil workability 0.1 0.4 http://www.fao.org/soils-portal/data-hub/soil-maps-and-
databases/harmonized-world-soil-database-v12/en/

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3. Results and discussion

3.1. Predictor variables envelop and model performance

After reducing multi-collinearity among the predictor variables, eighteen predictor variables were selected. The result of Jackknife
test revealed that mean temperature of the wettest quarter (Bio 8) had zero contribution. In the final model simulation, Bio 8 was
excluded as well. Seventeen variables were ultimately selected in this study and are summarized in Table 1. The AUC values resulted
from the three model simulation scenarios were analyzed. The result found that the model simulation with all key variables (scenario
A) showed the highest AUC value (Fig. 1).
The mean ROC curve generated from the P. cineraria with all key variable (scenario A) explains a good model fit with a mean AUC of
0.984 (Fig. 2). The ROC generally appears at the lower right triangle considered as worst model fit than the random (Fawcett, 2006)
and for a precise model fit the ROC appears at the top left (Chitale and Behera, 2012). Thus, we selected the model simulation with all
key variables as the final model for explaining potential habitat suitability distribution of P. cineraria in UAE.

3.2. Habitat suitability

Fig. 3 depicts spatial distribution of potentially not suitable, low suitable, medium suitable, and high suitable habitats for
P. cineraria in the UAE. The areal proportion of NS consists of 83% of the UAE. L habitat suitability class for P. cineraria in UAE
contributes 9% of the total area. The M and H habitat suitability classes cover 6% and 2% of the total area, respectively. The habitat
suitability distribution of P. cineraria revealed that H potential habitat suitability distribution is towards the northeastern regions of the
country, particularly north portion of Rasal Khima (N.RAK), Al Ain, Umal quaine, Dubai and some parts of Abu Dhabi (Fig. 3). This
could be due to the relatively higher water availability in the northeastern part of the country compared to the southern part and the
topography of the northeastern part of UAE. Whereas the southern part of the country generally found to be potentially L habitat
suitable and NS, except some southeastern regions. Major potential habitat suitability of this species spatially distributed in two soil
types hapalocalcids and torripsamments which are found in UAE (ESM, 2016). Generally P. cineraria develop an isolated wood habitats
and separated by large distance (Brown, 1991).

3.3. Variable contribution and response

The Jackknife variable contribution test revealed that ELEV, NDVI and precipitation of coldest quarter were the major influencing
variables (Fig. 4). These three variables together contributed 52.6% of total variable contribution. Bio3 (8.7%), bio18 (7.3%), bio9
(6%), slope (5.7%) and aspect (4.7%) also had considerable influence on the potential habitat suitability distribution of P. cineraria in
UAE. Cumulative contribution of these five variables were 32.4% of the total variable contribution. Remaining variables showed
14.9% cumulative contribution for the habitat suitability in UAE (Table 1). The environmental variables which observed with major
variable contribution also showed high AUC values when they used for isolation. Among the satellite remote sensing data, the NDVI is
the highest contributing variable for estimating P. cineraria habitat suitability distribution, implying potential NDVI capability for
monitoring spatial distribution patterns of P. cineraria. A study from Indian arid zones suggests that annual rainfall has direct influence
on the density of P. cineraria (Sharma and Joshi, 1982). In a similar way, model simulation study of P. juliflora from the arid regions of
India revealed that precipitation seasonality significantly affects the distribution (Singh et al., 2021). P. juliflora is an invasive species
and recognized as major threat to indigenous biodiversity (Singh et al., 2021). P. juliflora was introduced in UAE to combat deserti­
fication in the country and its present aggressive distribution could be a major threat to the native P. cineraria by means of genetic
contamination (Gallacher and Hill, 2005). P. juliflora can sustain in the harsh soil and climatic conditions like P. cineraria and the
diversified propagation method of P. juliflora aids the species to establish easily to new areas and helps to over crown the native flora
(Dakhil et al., 2021).

Fig. 1. AUC values of model replications with three scenarios: (A) with all key variables (17 variables), including bioclimate, edaphic, topographic
and satellite remote sensing data, (B) with bioclimatic and edaphic variables, and topographic parameters (14 variables), excluding satellite remote
sensing variables and (C) with bioclimatic variables (7 variables).

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Fig. 2. Mean area under receiver operating characteristic (ROC) curve (AUC) generated from MaxEnt model.

Fig. 3. Mean potential habitat suitability distribution of P. cineraria in the UAE for 2000–2019. H is high habitat suitability, M is medium habitat
suitability, L is low habitat suitability and NS is not suitable habitat.

Compared to the native P. cineraria species, the high rate of intrinsic water use efficiency of P. juliflora support their faster invasion
in UAE (Hussain et al., 2019). Dormant seed germination is often induced by increased soil water availability rather than temperature
change (Vidiella and Armesto, 1989). Several studies also agreed that water availability has significant role in warmer regions with dry
soil conditions by affecting the plant height, photosynthesis, number of branches (Zhang et al., 2018; Baskin and Baskin, 1998; Wang
et al., 2003). Our results also confirms that precipitation of the coldest quarter (bio19) of the UAE has significant influence on the
potential habitat suitability distribution of P. cineraria.
Variable response curves indicate the relationship between the probability of P. cineraria occurrence and environmental variables,
and define how the logistic prediction changes with alteration in these variables (Li et al., 2020a, 2020b). The variable response curves
generated from the model explains that the NDVI ranging from 0.1 to 0.2 are the most suitable values for UAE P. cineraria habitat
distribution. The probability of maximum occurrence of P. cineraria was high in an elevation range up to 390 m, but the higher al­
titudes showed least suitability. The result showed the relatively higher probability when precipitation of coldest quarter (Bio19)
varied from 50 to 65 mm. Around 85% of probability of presence was simulated when the precipitation of warmest quarter (Bio18) was

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Fig. 4. All key variable importance test using Jackknife method.

approximately 10 mm (Fig. 5).

P. cineraria showed a high suitability with an annual mean temperature (Bio1) range of 27–28.5 ◦ C and temperature of the driest
quarter (Bio9) of approximately 36 ◦ C. The most suitable isothermality (Bio3) range was 42–47% and the large isothermality explains
the use of relatively high temperature for photosynthesis during daytime and low temperature at nighttime which helps to reduce the
energy consumption of respiration (Hong et al., 2017). The response curve of aspect indicated a high suitability with no downslope and
a higher possibility in the southwest (202–245◦ ), west southwest (247.5–292.5◦ ) and west-northwest (292.5–337.5◦ ) aspects. Many
studies also have mentioned the significance of geomorphological heterogeneity, particularly slope and aspect in vegetation and soil
development at arid and semi-arid climate regimes (Monger and Bestelmeyer, 2006; Cantón et al., 2004; Carson and Kirkby, 1972;
Hutchins et al., 1976; Nichols et al., 1998).

3.4. Spatial distribution of habitat suitability and environmental variables

The areal percentage of potential habitat suitability of P. cineraria revealed that 60% of H habitat suitability area and 80% of the M
suitability area occupied in an elevation range between 95 and 195 m (Fig. 6a), whereas about 60% of L suitability was observed in an
elevation range of 195 m. Mean NDVI from 2000 to 2019 showed that above 80% of the H, M and L habitat suitability classes occupied
in the area where NDVI value ranges from 0.05 to 0.1 (Fig. 6b). The remaining 20% of the habitat suitability distributed in 0.1–0.2
NDVI range. In addition, a significant correlation between the mean annual NDVI and the mean potential habitat suitability of
P. cineraria was observed. The LST ranging from 31 ◦ C to 34 ◦ C showed spatial and temporal suitability in the entire country (Fig. 6c).
Annual mean NDVI and LST generally have a negative correlation, indicating higher LST reduces vegetation activity in the desert
ecosystems (Dall’Olmo and Karnieli, 2002). Most of the desert areas with heat attenuation have observed in the presence of vegetation
(Lazzarini et al., 2013). Spatial distribution of NDVI from 0.1 to 0.2 mostly exists in the barren area (Fig. 6d) which includes 83% of the
H potential habitat suitability for P. cineraria in the UAE. Shrub land exhibited 11.4% H potential habitat suitability for P. cineraria and
also 4% of H potential habitat suitability was predicted in the urban & build-up land of UAE, remaining land cover types were with
minimum suitability.

3.5. Limitations of the study

Use of different time scale variables for model simulation because of the unavailability of finer resolution bioclimatic data; though,
application of same temporal scale data may further improve the model prediction. Also, this study has utilized single model approach;
however, the use of multiple model combinations may enhance the prediction.

4. Conclusion

This study represents a primary attempt to include satellite remote sensing data to improve the habitat suitability prediction model
of P. cineraria in the UAE. Our results suggest that the integration of satellite remote sensing data, edaphic variables and topographic
parameters has improved the model accuracy with a mean AUC value of 0.984. In addition, the results showed that 17% of the UAE is

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R.K. Kalarikkal et al. Global Ecology and Conservation 37 (2022) e02167

Fig. 5. Variable response curves for major contributing bioclimatic and remote sensing variables for P. cineraria. The curves indicate the correlation
between the variables used for the probability of the presence of P. cineraria.

suitable for P. cineraria and the remaining area is potentially unsuitable for this species under recent environmental condition. This
study also emphasizes that when implementing afforestation in the UAE with desert species, such as P. cineraria (Ghaf tree), the
propagation should be in its naturally or potentially distributed habitats in order to get an optimal growth. The previous studies

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Fig. 6. Areal percentage of mean potential habitat suitability to P. cineraria in the UAE binned by (a) Elevation, (b) Mean NDVI (2000–2019), (c)
Mean LST (2000–2019), d) Land cover classification derived from MCD12Q1 for 2019.

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R.K. Kalarikkal et al. Global Ecology and Conservation 37 (2022) e02167

Fig. 6. (continued).

(El-Keblawy and Abdelfatah, 2014; El-Keblawy and Al-Rawai, 2007) have reported that P. juliflora (invasive species) is growing
aggressively and crowds out the native P. cineraria canopy in the UAE. We recommend that this study could be used to identify the
potential suitable habitat of P. cineraria in UAE and eradicate the invasive P. juliflora from the same habitat in order to control the
overcrowding. Understanding the potential habit suitable area of P. cineraria might help to reduce the future risk of invasion of other

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species like P.Juliflora by providing special species management practices at their suitable habitats. Mangers can also prioritize the
spatial zones for implementing species eradications and ground truthing for P.Juliflora based on the three simulated habitat suitability
classes (H, M & L) from this study. We recommend that the environmental envelope used for the final simulation of P. cineraria could be
applied in other arid regions of the world particularly in the middle eastern regions to understand the habitat distribution of this
species. Understanding the potential habitat suitability of species might help to manage the invasive species. For the future appli­
cations in the arid regions, in conclusion we propose the use of finer resolution bioclimatic variables which can be derived by using real
time satellite remote sensing sources to enable smaller scale conservation prioritization in the drylands.

Declaration of Competing Interest

The authors declare the following financial interests/personal relationships which may be considered as potential competing in­
terests: Youngwook Kim reports article publishing charges was provided by United Arab Emirates University. Youngwook Kim reports
a relationship with United Arab Emirates University that includes: employment.

Acknowledgement

This work was conducted at United Arab Emirates University, UAE under Start-up and 2020 ASPIRE-AARE grants.

Appendix A. Supporting information

Supplementary data associated with this article can be found in the online version at doi:10.1016/j.gecco.2022.e02167.

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