nr.07 01 07 Lobite

Grassroots Journal of Natural Resources, Vol.7 No.
1 (April 2024)
ISSN: 2581-6853 | CODEN: GJNRA9 | Published by The Grassroots Institute
Website: https://grassrootsjournals.org/gjnr | Main Indexing: Web of Science
ISSN 2581-6853 | 7(1) Apr 2024 M – 00379 | Research Article
Modelling Habitat Suitability and Distribution of the

Endemic Mindanao Horned Frog (Pelobatrachus stejnegeri)
and its Response to Changing Climate
Neil Jun Sala Lobite

Department of Forest Biological Sciences, College of Forestry and Natural Resources,
University of the Philippines – Los Baños, Laguna, Philippines, 4031.
Email: [email protected] ǀ ORCID: https://orcid.org/0000-0002-1767-0997
How to cite this paper: Lobite, N.J.S. (2024). Abstract

Modelling Habitat Suitability and Distribution Climate change is already affecting biodiversity, with special concern to
of the Endemic Mindanao Horned Frog endemics whose range is restricted and limited. This study focuses on the
(Pelobatrachus stejnegeri) and its Response to Mindanao horned frog (Pelobatrachus stejnegeri), an endemic species to the
Changing Climate. Grassroots Journal of
Philippines, susceptible to climate-induced habitat changes. Using MaxEnt
Natural Resources, 7(1): 123-137. Doi:
https://doi.org/10.33002/nr2581.6853.070107 species distribution model (SDM), the current and future (year 2050
projections) habitat suitability and distribution of P. stejnegeri were
modelled. Results showed that annual mean temperature, elevation, and
annual precipitation were the environmental variables having the highest
Received: 12 February 2024 influence on P. stejnegeri's distribution. The model predicts a significant
Reviewed: 17 April 2024 range contraction under representative concentration pathways (RCP) future
Provisionally Accepted: 19 April 2024 scenarios (RCP 2.6 and RCP 8.5), with a more pronounced decrease in
Revised: 24 April 2024 distribution (31.72%) under the high emission scenario (RCP 8.5). These
Finally Accepted: 25 April 2024
findings emphasize the vulnerability of P. stejnegeri to climate change and
Published: 30 April 2024
Copyright © 2024 by author(s) highlight the importance of integrating SDM into conservation and
management strategies to protect endemic species under changing climatic
Publisher’s Note: We stay neutral with conditions.
regard to jurisdictional claims in published Keywords
maps, permissions taken by authors and Mindanao horned frog; Species distribution model; Climate change; Species
institutional affiliations. distribution
This work is licensed under the Creative
Commons Attribution International
License (CC BY 4.0). Introduction
http://creativecommons.org/licenses/by/4.0/
Climate change presents a substantial threat to biodiversity by causing
unprecedented changes in ecosystems (IPBES, 2019; Román-
Palacios and Wiens, 2020). The effects of climate change on
Executive (Chief) Editor biodiversity are substantial and intricate, encompassing changes in
Dr. Hasrat Arjjumend habitat distribution, alterations in species interactions, and a greater
Associate Editors probability of extinction (Bellard et al., 2012; Pecl et al., 2017).
Dr. Usongo Patience
Moreover, changes in climate can lead to a shift in species' habitats
Ms. Areej Sabir
Grassroots Journal of Natural Resources, Vol.7, No.1 (April 2024), p.123-137 | ISSN 2581-6853 | CODEN GJNRA9
Doi: https://doi.org/10.33002/nr2581.6853.070107 Open Access
and distribution (Pecl et al., 2017), which endangers the existence of multiple organisms
(Chen et al., 2011; Trew and Maclean, 2021).
Climate-driven changes in species’ potential distribution can be assessed using species

distribution model (SDM) (Zurell et al., 2023). SDM can be used to predict suitability
of habitats for different species under different environmental conditions making it an
extremely significant tool for conservation planning and decision-making (Rahman et
al., 2019). This modeling approach is especially beneficial for species that are vulnerable
to climate change, such as the Mindanao horned frog. The predictive capability of SDM
enables the targeted allocation of conservation areas, highlighting the significance of
focusing conservation efforts in areas of high conservation value, especially for endemic
species with restricted ranges (Guisan et al., 2013).
Amphibians, due to their distinct physiological requirements and life cycles, are
exceptionally susceptible to the consequences of climate change, making them one of
the most endangered animal groups (Hof et al., 2011; Li, Cohen and Rohr, 2013; Luedtke
et al., 2023). Fluctuations in temperature, changes in rainfall patterns, and an increased
occurrence of severe weather events greatly affect the ability of amphibians to survive,
reproduce, and spread (Li, Cohen and Rohr, 2013; Campbell, Miller and Muths, 2020;
Rollins-Smith and Le Sage, 2023). These impacts are intensified for endemic species
whose distribution is restricted and limited (Manes et al., 2021) urging the call for
additional studies for this vulnerable group (Luedtke et al., 2023).
The Mindanao horned frog (Pelobatrachus stejnegeri) is an endemic amphibian species

found exclusively on the island of Mindanao in the Philippines. It is known for its unique
ecological role and its importance as a bio-indicator (IUCN SSC Amphibian Specialist
Group, 2020). This terrestrial frog is commonly found in both main and secondary
montane and lowland rainforests having mountain streams as the primary breeding
grounds. Being endemic, these species are extremely susceptible to habitat alterations
resulting from climate change, mostly due to their specific biological requirements and
restricted geographic range (Bickford et al., 2010; Manes et al., 2021). However, no
studies have been conducted yet, exploring the habitat suitability and distribution of the
Mindanao horn frog and assessing its response under changing climates.
This study aims to model the distribution of the endemic Mindanao horned frog (P.
stejnegeri) under current and future climatic conditions. Additionally, it aims to measure
the impact of climate change on this species' geographic distribution through
determination of regions where the habitat may become more or less suitable based on
future climate scenarios.
Methodology
Study Area and Occurrence Records
The study was carried out in the island of Mindanao in the Philippines, where P.
stejnegeri is endemic (Figure 1). Mindanao is located in the southern part of the
Philippines and is one of the three island groups of the country, covering an approximate
land area of 95,468.17 square kilometers (PhilAtlas, 2022). The occurrence points of P.
stejnegeri were collected from the Global Biodiversity Information Facility
Neil Jun Sala Lobite 124

(https://www.gbif.org/) and survey data conducted in 2012–2023 by the author.

Duplicate geographic coordinates and coordinates falling outside the study area
boundary were removed from the data points to clean them up. Using SDMtoolbox
(Brown, Bennett and French, 2017), the distribution data were subjected to spatial
rarefaction at a 5 km distance in order to eliminate sampling bias and spatial
autocorrelation.
Figure 1: Location of the study area with observations of P. stejnegeri (red dots).
Environmental Variables
This study utilized 21 environmental layers with a spatial resolution of 30 seconds to

create a distribution model of P. stejnegeri. These include topographical factor
(elevation), vegetation factor (enhanced vegetation index), and bioclimatic factors (bio
1–19). The bioclimatic factors were the following: Bio1: Annual Mean Temperature;
Bio2: Mean Diurnal Range; Bio3: Isothermality; Bio4: Temperature Seasonality; Bio5:
Max Temperature of Warmest Month; Bio6: Minimum Temperature of Coldest Month;
Bio7: Temperature Annual Range; Bio8: Mean Temperature of Wettest Quarter; Bio9:
Mean Temperature of Driest Quarter; Bio10: Mean Temperature of Warmest Quarter;
Bio11: Mean Temperature of Coldest Quarter; Bio12: Annual Precipitation; Bio13:
Precipitation of the Wettest Month; Bio14: Precipitation of the Driest Month; Bio15:
Precipitation Seasonality; Bio16: Precipitation of the Wettest Quarter; Bio17:
Precipitation of the Driest Quarter; Bio18: Precipitation of the Warmest Quarter; and
Bio19: Precipitation of the Coldest Quarter. Elevation data were obtained using the
digital elevation model (DEM) of the Shuttle Radar Topography Mission (SRTM),
which has a 90 m spatial resolution. The Enhanced Vegetation Index (EVI) map was
obtained from Moderate-Resolution Imaging Spectroradiometer (MODIS) data

(https://modis.gsfc.nasa.gov/). Bioclimate factors were obtained from the WorldClim

database (Fick and Hijmans, 2017; http://www.worldclim.org). All layers were
resampled based on a bilinear interpolation in order to match the resolution.
In order to prevent overfitting and identify the most appropriate variables, a correlation
analysis was performed using Pearson's correlation coefficient (r) to examine the
relationship between the 21 environmental variables. A Pearson's correlation value
greater than 0.7 signifies a strong correlation between the two variables. All but one of
any given environmental layers with a high correlation with other layers were removed.
For future climate, bioclimatic variables using the MIROC-ESM climate model for the
period 2041–2060 were downloaded from the WorldClim database
(http://www.worldclim.org). Two representative concentration pathways (RCP), namely
RCP 2.6 and RCP 8.5, were used as scenarios to predict future climate change. The
RCP2.6 and RCP8.5 scenarios reflect low and high greenhouse gas emissions in the
future, respectively.
Species Distribution Modelling
The species distribution models were created using MaxEnt v. 3.4.4 (Phillips, Anderson
and Schapire, 2006). MaxEnt modeling was conducted by allocating 25% of the
occurrence data for testing purposes and 75% of the occurrence data for training
purposes. In order to identify the primary environmental factors that influence the
distribution of P. stejnegeri habitat, a jackknife permutation analysis was performed to
rank the environmental factors based on their percentage contribution. In order to assess
the effectiveness of the MaxEnt model, the area under the receiver's operating curve
(AUC) was employed as a measure of the model's predictive accuracy. A model's
prediction accuracy increases as the Area Under the Curve (AUC) value increases.
Evaluation of model performance were based on three specific criteria: poor, defined as
an AUC value below 0.8; good, defined as an AUC value between 0.90 and 0.95; and
excellent, defined as an AUC value above 0.95. Once the model computations were
finished, the habitat distribution of P. stejnegeri was displayed on ArcMap.
The continuous habitat suitability maps of P. stejnegeri for both current and future
climate conditions (RCP 2.6 and RCP 8.5) were transformed into presence/absence
binary maps using a 10 percentile training presence clog threshold. In order to evaluate
the changes in the distribution of P. stejnegeri due to changing climate, binary maps
representing the current climate conditions were overlaid onto the projected habitats and
compared using the SDMtoolbox in ArcMap. Afterwards, the results were categorized
into three distinct groups: (1) range expansion, (2) no occupancy (absent in both), (3) no
change (present in both), and (4) range contraction.
Results
Variables Correlation and Selection
The correlation analysis of all 21 predictor variables showed a high correlation for some
variables, as shown in figure 2. The variables with the highest correlation are Bio6, Bio9,
Bio10, and Bio11 in relation to Bio1, which has a correlation coefficient above 90%

positive correlation. Nine variables were selected out of 21 variables after the variable
selection process, namely Bio1 (Annual Mean Temperature), Bio2 (Mean Diurnal
Range), Bio3 (Isothermality), Bio4 (Temperature Seasonality), Bio12 (Annual
Precipitation), Bio15 (Precipitation Seasonality), Bio18 (Precipitation of Warmest
Quarter), Bio19 (Precipitation of Coldest Quarter), and elevation.
a. b.
Figure 2. Correlation analysis of 21 predictor variables. a.) Heat map showing the
correlation between variables, b.) Dendrogram showing the grouping of highly
correlated variables with correlation coefficient threshold of 0.7.
Key Environmental Factors Influencing the Habitat Distribution
The importance and contribution of each environmental variable to the prediction model
were analyzed by the jackknife method. The results showed that the contributions of
Bio1 (Annual Mean Temperature), Elevation, and Bio12 (Annual Precipitation) were
24.39%, 18.65%, and 17.21%, respectively (Table 1). The environmental variable with
the highest gain when used in isolation is Bio1 (Figure 3), which, therefore, appears to
have the most useful information on the distribution of P. stejnegeri. The environmental
variable that decreases the gain the most when it is omitted is Bio4, which appears to
have the most information that is not present in the other variables.
Table 1: Estimates of relative contributions of the environmental variables to the MaxEnt

model
Variable Percent contribution Permutation importance
Bio1 24.39 28.31
Elevation 18.65 2.026
Bio12 17.21 11.42
Bio4 14.17 18.61
Bio2 9.47 18.19
Bio18 8.2286 19.5545
Bio19 7.0256 0.1321
Bio3 0.4861 0.1612
Bio15 0.3655 1.5926

Figure 3: Jackknife test of variable importance on P. stejnegeri distribution
Modelled Distribution of P. stejnegeri under Current Conditions
The Maxent model predicts the distribution of P. stejnegeri across the entire island, as
shown in figure 4. The species' current modeled distribution closely matches the region's
published distribution data, including the range map produced by the IUCN
(International Union for Conservation of Nature, 2024). The presence/absence binary
map showed that the entire area where P. stejnegeri is predicted to be present is
38,550.18 km2. The model AUC value of 0.917 indicates that the model's performance
is superior to random prediction, hence confirming its validity.
a. b.
c.
Figure 4: Species distribution model of P. stejnegeri under current conditions.

a.) MaxEnt model of P. stejnegeri showing probability of occurrence;
b.) Presence/absence map of P. stejnegeri; c.) Receiver operating characteristic (ROC)
curve for evaluating the model performance of MaxEnt model.

Modelled Distribution of P. stejnegeri under Future Climate Scenarios
Using the MIROC-ES model, the distribution of P. stejnegeri is projected under future
climatic scenarios. Both models under RCP 2.6 and RCP 8.5 indicated a decrease in the
species' range, as shown in figure 5, using the 10th percentile threshold cutoff. The
predicted extent of P. stejnegeri's present under RCP 2.6 is 17,890.26 km2, but in RCP
8.5 it is 8,452.72 km2. The findings indicated a decrease in the extent of the area occupied
under projected future climate conditions.
a. c.
b. d.
Figure 5: Species distribution model of P. stejnegeri under future conditions (2050). a.)
MaxEnt model showing probability of occurrence under RCP 2.6; b.) MaxEnt model
showing probability of occurrence under RCP 8.5; c.) Presence/absence map under
RCP 2.6; d.) Presence/absence map under RCP 8.5.
Figure 6 illustrates the geographical areas where the distribution of P. stejnegeri will
expand and contract in response to changing climate conditions, as determined by
different RCPs. According to the RCP 2.6 climatic scenario for the year 2050, the
suitability for P. stejnegeri would decrease, resulting in a total range contraction area of
23,164.77 km2 (Table 2). The majority of the contractions are located in the northern
region of Mindanao. On the other hand, under the extreme climate scenario (RCP 8.5),
the area of suitable habitat is predicted to be greatly reduced, with a total contraction
range of 30,200.16 km2, or 31.72% contracted in reference to the current model. The
areas where no change occurs, i.e., species that are predicted to be present in both current
and future climatic scenarios, are primarily concentrated in the high-elevation parts of
Mindanao.
a. b.
Figure 6: Distribution changes of P. stejnegeri from the present climate to the future
year (2050) under climate scenarios: (a) RCP 2.6; (b) RCP 8.5.
Table 2: Relative changes in P. stejnegeri distribution area under climate scenarios RCP
2.6 and RCP 8.5 for the year 2050.
Distribution Total area of change (km2) Percentage of change
Change RCP 2.6 RCP 8.5 RCP 2.6 RCP 8.5
Range 2,519.68 101.99 2.65 0.11
expansion
No occupancy 54,116.55 56,534.24 56.84 59.38
No change 15,400.04 83,64.65 16.18 8.79
Range 23,164.77 30,200.16 24.33 31.72
contraction
Discussion
The present distribution modeling of P. stejnegeri is crucial for understanding the

environmental factors that influence its distribution. The MaxEnt model precisely
delineates the habitat range of P. stejnegeri, exhibiting a close correspondence with
established distributions and conservation evaluations, such as those undertaken by the
IUCN. The high AUC score validates the model's accuracy in identifying the significant
environmental factors that influence the distribution of P. stejnegeri. The importance of
the average annual temperature, altitude, and annual rainfall in the model emphasizes
the species' reliance on these biological parameters. This is consistent with overarching
ecological theories that credit temperature and precipitation as the primary factors that

influence patterns of species distribution (Bickford et al., 2010; Guisan and Thuiller,
2005; Mi et al., 2022). The mean annual temperature significantly influences the
distribution of amphibians, as these organisms, being ectothermic, depend on external
heat sources to regulate their body temperature. The metabolic rates, reproductive cycles,
and survival of frogs are intricately linked to the prevailing ambient temperature
conditions (Rollins-Smith and Le Sage, 2023). Wake and Vredenburg (2008) have
provided evidence of the significant impact that changes in temperature patterns can
have on amphibian populations. The vulnerability of P. stejnegeri to the annual mean
temperature underscores the species' sensitivity to climate change, hence reflecting
broader concerns regarding the impact of global warming on amphibian populations.
Additionally, annual precipitation is another important factor that indicates the essential
requirement for moisture by amphibians. Due to their permeable skin, these organisms
rely on a moist environment to facilitate gas exchange (Harvey Pough, 2007). This
unique physiological trait makes them highly susceptible to fluctuations in moisture
levels within their habitat (Harvey Pough, 2007; Lips et al., 2003; Stuart et al., 2004).
The study conducted by Mi et al. (2022) clearly illustrates the correlation between
precipitation patterns and the availability of suitable habitats for amphibians, showing
that the average annual precipitation had the greatest impact on the spatial distribution
pattern. Based on the model, the reliance of P. stejnegeri on the annual precipitation
factor further demonstrates the complex interplay between amphibians and their
hydrological habitats. Lastly, elevation was found to be a significant factor for P.
stejnegeri's distribution. It has been proposed that elevation affects the local weather
conditions, which, in turn, affects amphibian population diversity and dispersal. McCain
and Grytnes (2010) demonstrate how elevation gradients influence biodiversity patterns,
such as those of amphibians, by generating diverse ecological niches at different
altitudes. The importance of elevation in P. stejnegeri's distribution model emphasizes
how changes in altitude can define the boundaries of acceptable habitats for amphibians,
affecting their distributional range and population dynamics (Fu et al., 2006; Supsup et
al., 2022). Overall, the model's ability to accurately predict the current distribution of P.
stejnegeri suggests that the species is highly influenced by a range of climatic and
topographic parameters, specifically temperature, precipitation, and elevation. This
highlights the crucial significance of these environmental conditions in the conservation
of this species.
According to future climate projections, it is predicted that the range of P. stejnegeri

would see substantial changes, with a large reduction in appropriate habitats, especially
under the high-emission scenario (RCP 8.5). This estimate is consistent with the global
consensus on the anticipated effects of climate change on biodiversity suggesting that
several species would have reduced habitats because changing climatic conditions
(Bellard et al., 2012; Blaustein et al., 2010; Chen et al., 2011). The model forecasts a
significant decrease in the spatial range of P. stejnegeri under both RCP 2.6 and RCP
8.5 scenarios, highlighting the severe threats that climate change poses to its habitat
suitability. The range contractions in the northern part of Mindanao are particularly
noteworthy, suggesting that these areas may no longer be suitable for the species due to
shifts in climatic conditions. The anticipated decrease in habitat in Mindanao could have
significant implications for the population of P. stejnegeri, given that this species is
endemic to the island. The projected changes in the distribution of P. stejnegeri
emphasize the broader challenges that endemic species in biodiversity hotspots are
facing because of climate change (Trew and Maclean, 2021). In light of these concerns,

potential strategies could involve habitat restoration, the creation of biological corridors,
and the safeguarding of climate refuges that may remain suitable amidst changing
conditions (Rahman et al., 2019; IUCN SSC Amphibian Specialist Group, 2020). The
establishment of corridors will be useful in linking habitats and refuges of P. stejnegeri
that are expected to remain viable under future climate scenarios. This will facilitate the
movement of the species to more suitable environments and is crucial for its adaptation
to environmental changes (Beier, 2012).
Conclusion
This study models the habitat suitability and distribution of P. stejnegeri under both
current and projected future climate scenarios using MaxEnt. The study determined that
annual mean temperature, elevation, and yearly precipitation are the main factors
affecting P. stejnegeri's distribution. Future projections demonstrate the vulnerability of
P. stejnegeri to climate change, highlighting a substantial decrease in its geographical
range under both mild (RCP 2.6) and severe (RCP 8.5) future climate conditions. The
reduction in range, specifically in the high-emission RCP 8.5 scenario, emphasizes the
pressing risk that climate change poses to the species. Given the species' sensitivity to
environmental factors such as temperature and precipitation, together with the projected
decrease in geographical range due to climate change, it is important for conservation
efforts to prioritize the adaptability and quality of P. stejnegeri's habitats.
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Grassroots Journal of Natural Resources, Vol.7 No.

ISSN 2581-6853 | 7(1) Apr 2024 M – 00379 | Research Article

Modelling Habitat Suitability and Distribution of the

Neil Jun Sala Lobite

How to cite this paper: Lobite, N.J.S. (2024). Abstract

Climate-driven changes in species’ potential distribution can be assessed using species

The Mindanao horned frog (Pelobatrachus stejnegeri) is an endemic amphibian species

Study Area and Occurrence Records

Neil Jun Sala Lobite 124

(https://www.gbif.org/) and survey data conducted in 2012–2023 by the author.

This study utilized 21 environmental layers with a spatial resolution of 30 seconds to

Neil Jun Sala Lobite 125

(https://modis.gsfc.nasa.gov/). Bioclimate factors were obtained from the WorldClim

Species Distribution Modelling

Variables Correlation and Selection

Neil Jun Sala Lobite 126

Key Environmental Factors Influencing the Habitat Distribution

Table 1: Estimates of relative contributions of the environmental variables to the MaxEnt

Neil Jun Sala Lobite 127

Figure 3: Jackknife test of variable importance on P. stejnegeri distribution

Modelled Distribution of P. stejnegeri under Current Conditions

Figure 4: Species distribution model of P. stejnegeri under current conditions.

Neil Jun Sala Lobite 128

Modelled Distribution of P. stejnegeri under Future Climate Scenarios

The present distribution modeling of P. stejnegeri is crucial for understanding the

Neil Jun Sala Lobite 130

According to future climate projections, it is predicted that the range of P. stejnegeri

Neil Jun Sala Lobite 131

Neil Jun Sala Lobite 132

Neil Jun Sala Lobite 133

Neil Jun Sala Lobite 134

Academy of Sciences, 117(8): 4211–4217. DOI:

Neil Jun Sala Lobite 135

Authors’ Declarations and Essential Ethical Compliances

Authors’ Contributions (in accordance with ICMJE criteria for authorship)

Research involving human bodies or organs or tissues (Helsinki Declaration)

Research involving animals (ARRIVE Checklist)

Research on Indigenous Peoples and/or Traditional Knowledge

Research involving Plants

Research Involving Local Community Participants (Non-Indigenous) or Children

Neil Jun Sala Lobite 136

Competing Interests/Conflict of Interest

Attribution and Representation

Rights and Permissions

Neil Jun Sala Lobite 137

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