proceedings
Proceeding Paper
Variability in Philippine Coffea liberica Provides Insights into
Development Amidst a Changing Climate †
Ma. Carmen Ablan Lagman
Citation: Ablan Lagman, M.C.
Variability in Philippine Coffea liberica
Provides Insights into Development
Amidst a Changing Climate.
Proceedings 2023, 89, 27. https://
doi.org/10.3390/ICC2023-14852
Academic Editor: Dirk W.
Lachenmeier
Published: 21 August 2023
Copyright: © 2023 by the author.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Proceedings 2023, 89, 27. https://doi.org/10.3390/ICC2023-14852
Practical Genomics Laboratory, Center for Natural Science and Environment Research, De La Salle University,
Manila 1004, Philippines; ma.carmen.lagman@dlsu.edu.ph
† Presented at the International Coffee Convention 2023, Mannheim, Germany, 30 September–3 October 2023.
Abstract: The Philippines is one of only a few countries where Coffea liberica is commercially produced.
Two infra-species, C. liberica liberica (also locally known as ‘liberica’) and C. liberica var dewevrei (also
known as ‘excelsa’) comprise 1.1% and 5.7% of the national coffee production, respectively. These rare
varieties are produced widely in the provinces of Batangas and Cavite mainly because of historical
affinity, a local market, and more recently the renewed interest from importers. Aside from its
unique flavor profile, renewed interest comes from the potential of the larger beans and deeper root
systems of the species to thrive on a warming planet. In this paper, we present work that has been
performed in evaluating bean morphology, chlorogenic acid, caffeine content and genetic variability
of C. liberica varieties from different areas of the country. The initial efforts to predict areas for
increased production were based on a maximum entropy model. Combining these data provides
insights into development areas for increased production in the Philippines.
Keywords: Philippine coffee; climate change; liberica
1. Introduction
Coffea ‘liberica’ is often referred to as the rare and even elusive variety of commercially
grown coffee. This is because only 2% of the global coffee production is C. liberica [1]. The
species is grown commercially in five countries—Indonesia, Malaysia, Vietnam, Ethiopia,
and the Philippines—mainly for local consumption, although some have exported to
the emerging specialty coffee industry. Surprisingly, C. liberica comprises 90% of coffee
cultivated in Malaysia [2].
C. liberica is locally well known to the countries that produce them; however, C. liberica
remains a gem to be discovered for the rest of the world. C. liberica is referred to as a
‘heirloom species’ in the Western world, where it is sold. In Australia, it is called a ‘heritage
variety’ (Adams and Russell, 2018). Two varieties of this species are traded: ‘liberica’ and
‘excelsa’. C. liberica trees are taller with larger leaves, making harvesting more difficult.
The beans are much bigger than C. arabica or C. canephora resulting in less production per
tree [3].
C. liberica cups may have a very complex full-bodied flavor profile, distinct from the
other commercial species. Not many have experimented roasting the species given the
limited availability of this variety. C. liberica is characterized as having a “fruity, woody
taste similar to jackfruit, with bitter top-notes”. This is very different from the varied
floral and fruity, sweet, and slightly acidic profile of C. arabica. The bitter earthy character
of C. canephora and C. liberica has also found its way into the instant or soluble coffee
market. C. liberica is slowly gaining attention globally and is being sought from farms in
the Philippines [4].
The World Checklist of Vascular Plants recognizes three species of coffee. Coffea arabica,
a species originating from the natural hybridization of C. eugenioides and C. canephora. Coffea
canephora, another familiar species, and ‘robusta’, which is the widely cultivated variety;
https://www.mdpi.com/journal/proceedings
Proceedings 2023, 89, 27 2 of 7

thus, the term used for commercial purposes. Two infra-species of C. liberica are noted.
These are Coffea liberica var liberica and Coffea liberica var dewevrei [5]. These varieties are
commercially referred to as ‘liberica’ and ‘excelsa’.
This paper discusses the C. liberica in the Philippines and the work in progress on the
two commercial varieties: ‘liberica’ and ‘excelsa’. Initial results of the science the industry
needs for development of these varieties are presented in the country. Although it is country
specific; the hope is to contribute to the growing research to understand C. liberica from
the perspective of a country that has grown the species, and has also been growing the
other commercial varieties: Coffea arabica and Coffea canephora. This paper chooses to focus
on the research performed in the Philippines in support of the coffee industry roadmap
2021–2025 [6] and acknowledging the distinction in C. liberica.

2. Variation and Role of Nurture vs. Nature in Philippine C. liberica

The local market in the Philippines refers to a coffee branded as ‘barako’, the colloquial
name for ‘stud’ and the Spanish term for ‘wild boar’, i.e., varraco. The term is associated with
characteristics of fearlessness, masculinity, and toughness which are evidently associated
with C. liberica and loosely associated with coffee produced in the provinces of Cavite and
Batangas, which are known as the historical coffee centers in the Philippines [7]. ‘Liberica’
was apparently the primary variety grown in these provinces prior to ‘branding’, which
could be confusing for people because ‘Liberica’ may refer to any coffee grown in these
traditionally ‘Liberica’ growing areas, regardless of species.

2.1. Variety vs. Environment in Caffeine, CGA and Trigonelline in Philippine C. liberica
Sensorial coffee quality is affected by the of volatile and non-volatile compounds.
Accumulation of non-volatile compounds, such as caffeine, chlorogenic acid, trigonelline,
and sucrose in green beans is affected by environmental factors, such as altitude, shade
and moisture. It is also affected by processing and variety [8]. Compared to C. arabica,
C. liberica liberica and C. liberica dewevrei have a stronger taste and a more pungent aroma.
The characteristic differences in flavor among coffee varieties is hypothesized to be related
to the flavor profiles among coffee varieties. Bitterness and astringency in coffee are partly
due to polyphenols. Polyphenols are also recognized for their antioxidant properties
which are beneficial against several oxidative stress, aging, and related diseases, such
as cardiovascular disease and cancer [9]. Caffeine, chlorogenic acid, diterpenes, and
trigonelline are some of the more prominent polyphenols in coffee.
Caffeine is a naturally occurring stimulant commonly sourced from coffee, tea, or
cacao. Chlorogenic acid and trigonelline are important alkaloids alongside caffeine in
coffee. Chlorogenic acid or CGA strongly influences the taste of coffee, such as astringent,
sweet, and sour tastes. This changes with concentration. Trigonelline is also considered to
be the origin of the unpleasant–complex taste found after prolonged brewing. Trigonelline
breaks down to pyrroles, nicotinic acid, and pyridine to produce desirable aroma attributes,
such as toasted, nutty, and roasted flavors [8].
An initial study on Philippine coffee compared variation in trigonelline in all four-
coffee variety sites for the same variety, and among varieties also gained insight into the
contribution of the environmental factors. This study showed that the highest trigonelline
content was observed in both infra-species of C. liberica, followed by C. canephora and then
by C. arabica. The same study also compared trigonelline content across different sites for
the same variety of coffee (Figure 1). The results indicate that significant differences in
trigonelline were observed across C. arabica cultivated in different environments [10].
Proceedings 2023, 24, x 3 of 7

also presented no significant differences in trigonelline, caffeine, and CGA content among
Proceedings 2023, 89, 27 beans of the same infra-species of C. liberica liberica and C. liberica dewevrei harvested from
3 of 7
farms from different elevations and growing environments.

Figure 1. Principal component analysis of environment across different farms where C. arabica (green),
Figure 1. Principal component analysis of environment across different farms where C. arabica
C. liberica liberica (blue), and C. liberica dewvrei (orange) are harvested.
(green), C. liberica liberica (blue), and C. liberica dewvrei (orange) are harvested.
The same cannot be said for C. liberica liberica and C. liberica dewevrei varieties. No
2.2. Morphological
significant Characterization
differences of C. liberica
in trigonelline content vs. were
C. canephora
observed. and C.
Syarabica
et al. [11] also found
2.2.1. Leaves and Tree Morphology
significant differences in protein, DNA, caffeine, and CGA content between C. arabica and
either of C. liberica
Variation libericaand
in the leaves andtreeC. liberica
structure dewevrei, whilebecause
are distinct the twoC.infra-species
liberica libericawere
leavesnot
different
are wider at in the
content
apex.ofThethe length
same biologically
of the leaves important
of Coffeamolecules. Interestingly,
liberica of variety libericathis
may study
be
also presented
twice as long as no C. significant differences
arabica. C. liberica in trigonelline,
dewevrei caffeine,
leaves, on the and CGA
other hand, content among
are smaller. Note
beans of the same infra-species of C. liberica liberica and C. liberica dewevrei
that in farms, the variations between the infra-species of C. liberica increase in range, and harvested from
farms from different elevations and growing environments.
a greater overlap in sizes may occur. The range of leaf sizes within a tree also differs.
Distinction between leaf morphology among the four varieties was completed.
2.2. Morphological Characterization of C. liberica vs. C. canephora and C. arabica
Hence, C. liberica is distinct from C. canephora and C. arabica and between the latter two
2.2.1. Leaves and Tree Morphology
species. Furthermore, there is a wider overlap in the leaf lamina morphology between the
Variation in There
two infra-species. the leaves and tree
are distinct structure are
differences distinct
between C. liberica C. liberica liberica
becauseinfra-species leaves
in laminar
are wider at the apex. The length of the leaves of Coffea liberica
size, domestic distribution, major secondary attachment, and intercoastal tertiary veinsbe
of variety liberica may
twice as
among leaves as C.This
long[12]. arabica.
studyC. did
liberica
not dewevrei
decide toleaves,
changeon the
the taxonomic
other hand,rank are smaller. Note
of C. liberica
that in farms,
dewevrei without thefurther
variations betweenstudies.
taxonomic the infra-species of C. liberica
A more detailed increase
variation in range,
within and a
C. liberica
greater overlap in sizes may occur. The range
infra-species has to be investigated with farmers in Batangas. of leaf sizes within a tree also differs.
InDistinction
both Cavite between leaf morphology
and Batangas provinces,amongfarmers theattest
four varieties was completed.
to the resilience of the treesHence,
to
C. liberica is distinct from C. canephora and C. arabica and between
long term exposure to the sun and erratic periods of drought. They believe that, although the latter two species.
Furthermore,
the leaf lamina there is a wider
is wider, the C. overlap in the leaf lamina
liberica infra-species morphology
have reduced between
the loss the two
of moisture
infra-species. There are distinct differences between C. liberica infra-species in laminar size,
through transpiration because of the upward orientation of the leaves and branches, with
domestic distribution, major secondary attachment, and intercoastal tertiary veins among
leaves facing up more in comparison to the orientation of the leaves in C. canephora and C.
leaves [12]. This study did not decide to change the taxonomic rank of C. liberica dewevrei
arabica (Figure 2). The upward facing leaves may also reduce the leaves’ surface area ex-
without further taxonomic studies. A more detailed variation within C. liberica infra-species
posure to sunlight as well as water loss through transpiration, as initial data seem to indi-
has to be investigated with farmers in Batangas.
cate. Root systems for C. liberica are also longer, almost twice as long as C. canephora root
In both Cavite and Batangas provinces, farmers attest to the resilience of the trees to
systems, as seedlings provide them with a deeper access to water reservoirs.
long term exposure to the sun and erratic periods of drought. They believe that, although
the leaf lamina is wider, the C. liberica infra-species have reduced the loss of moisture
through transpiration because of the upward orientation of the leaves and branches, with
leaves facing up more in comparison to the orientation of the leaves in C. canephora and
Proceedings 2023, 89, 27 4 of 7

C. arabica (Figure 2). The upward facing leaves may also reduce the leaves’ surface area
exposure to sunlight as well as water loss through transpiration, as initial data seem to
Proceedings 2023, 24, x 4 of 7
indicate. Root systems for C. liberica are also longer, almost twice as long as C. canephora
root systems, as seedlings provide them with a deeper access to water reservoirs.

Figure 2.
Figure 2. From
From left
left to
to right—young
right—young C.
C. canephora,
canephora, C.
C. liberica
liberica liberica,
liberica, C.
C. arabica,
arabica, and
and C.
C. liberica
liberica dewevrei
dewevrei
trees presenting
trees presenting distinct
distinctleaf
leafmorphologies
morphologiesand
andtree
treebranch
branchstructure.
structure.

2.2.2.
2.2.2. Seeds Morphology within C.
Morphology within C. liberica
liberica and against C.
and against C. arabica and C.
arabica and C. canephora
canephora
Coffea
Coffea liberica
liberica of variety liberica
of variety liberica has very large fruits, fruits, and compared to Coffea arabica,
the
the fruit
fruit color
color is is greatly
greatly varied,
varied, ranging
ranging from
from light
light yellow
yellow to to deep
deep purple.
purple. The
The bean
bean shape
shape
varies
varies in the the ratio
ratioofoflength
lengthtotowidth
width and
and from
from oblong
oblong to teardrop
to teardrop shapes.
shapes. WithinWithin the
the Phil-
Philippines,
ippines, more more than
than 15 15 different
different ‘kinds’
‘kinds’ ofofC.C.liberica
libericaliberica
libericawere
werefound
found just
just by observing
observing
the
the fruit
fruit from
from the the trees.
trees.
C. liberica dewevrei,
C. liberica dewevrei, although
although moremore available,
available, is is less
less variable.
variable. It It ranges
ranges inin color
color from
from
light red and yellow to bright and deep red. Green beans are smaller than C. liberica liberica
light red and yellow to bright and deep red. Green beans are smaller than C. liberica liberica
with
with aa less
less elongated
elongated shape.
shape. Previous
Previous research
research has has demonstrated
demonstrated that that the
the length
length ofof the
the
minor
minor radial
radial axisaxis in
in the
the bean
bean isis the
the characteristic
characteristic which differentiates C.
which differentiates C. liberica
liberica dewevrei,
dewevrei,
C.
C. liberica
liberica liberica,
liberica, C.
C. arabica andC.
arabicaand C.canephora.
canephora.
With
With the limitation of only using the
the limitation of only using the green
green coffee
coffee bean,
bean, the
the morphometrics
morphometrics of of height,
height,
length,
length, and depth at the length of the minor radial axis were compared among beans from
and depth at the length of the minor radial axis were compared among beans from
various
various farms
farms for for beans
beans that
that farmers
farmers identified
identified as as C.
C. liberica
liberica liberica.
liberica. The
The elevation
elevation ofof the
the
farms
farms waswas variable.
variable. The The farm
farm trees
trees were
were simultaneously
simultaneously planted planted in in the
the farms;
farms; thus,
thus, the
the
trees
trees were
were of of aa very
very similar
similar‘kind’.
‘kind’.
The
The data
data indicates
indicates aa wide
wide overlap
overlap andand aa high
high variability
variability of of measurements
measurements (Figure(Figure 3).
3).
Although highly overlapping, the data from the two farms marked
Although highly overlapping, the data from the two farms marked as purple and yellow as purple and yellow
hints at a higher similarity within farms. Farmers are observed to have farms that focus on
hints at a higher similarity within farms. Farmers are observed to have farms that focus
a specific ‘kind’ of tree, as the elevation was the same for both locations. The only difference
on a specific ‘kind’ of tree, as the elevation was the same for both locations. The only dif-
was that the dominant ‘kind’ of tree was planted, rather than focusing on the environment
ference was that the dominant ‘kind’ of tree was planted, rather than focusing on the en-
in which it was grown. This highlights the relative importance of the nature of the coffee
vironment in which it was grown. This highlights the relative importance of the nature of
plant as well as its nature.
the coffee plant as well as its nature.

2.2.3. Genetic Characterization

Genetic characterization has received greater attention from other researchers. There
is a very clear differentiation of the three species of C. arabica, C. canephora, and C. liberica
through sequences from the matK of the rbcL gene or through simple sequence repeats
[13]. Differentiation of C. liberica dewevrei and C. liberica liberica has been less straight for-
ward using standard barcoding genes (i.e., matK, rbcL and trnl-psba); however, natural
reproductive barriers exist between these infra-species, as evidenced by weak pollen via-
bility of F1 hybrids [14]. Our current work involves genetic identification of mother trees
and seedlings for the development of nurseries. In addition, work is on-going on the
Proceedings 2023, 24, x 5 of 7

Proceedings 2023, 89, 27 5 of 7

caffeine-related gene HCT in C. liberica, as the aim is to develop C. liberica disease-resistant
varieties of coffee with the desired CGA and caffeine content.

Figure 3.
Figure 3. PCA
PCAononcoffee
coffeebean
beanmorphometrics
morphometrics of of
C. C.
liberica grown
liberica in six
grown in different farms
six different in theinPhil-
farms the
ippines. The purple and yellow circles are drawn around the most different clusters. These
Philippines. The purple and yellow circles are drawn around the most different clusters. These two
two
‘varieties’ are more different from each other than from others.
‘varieties’ are more different from each other than from others.

3. Defining
2.2.3. GeneticAreas for Cultivation of Coffea liberica
Characterization
Different
Genetic climatic factorshas
characterization andreceived
environmental
greater requirements
attention fromgreatly vary among There
other researchers. Coffea
species.
is a very Some of these factorsofinclude
clear differentiation the three thespecies
lowland and
of C. highland
arabica, locations,and
C. canephora, theC.different
liberica
elevationsequences
through of coffee plantation,
from the matK optimum annual
of the rbcL rainfall
gene range,simple
or through optimum mean annual
sequence repeats tem-
[13].
perature range,ofairC.humidity,
Differentiation and wind
liberica dewevrei andstress. In addition,
C. liberica thebeen
liberica has effectslessofstraight
shade have also
forward
been proven
using standardtobarcoding
have great interaction
genes with
(i.e., matK, theand
rbcL biochemical
trnl-psba);composition
however, naturalof coffee, most
reproduc-
especially
tive barrierswith green
exist coffee
between beans.
these These contribute
infra-species, to changes
as evidenced in thepollen
by weak different properties
viability of F1
hybrids
of coffee.[14]. Ourenvironment
Thus, current workcan involves genetic identification
be classified as a contributingof mother
factor.trees
Dataandon seedlings
the envi-
for the development
ronment of nurseries.
can also be used to predict In suitable
addition,areas
workfor is developing
on-going onC.the RNASeq
liberica and QTL
plantations.
mapping of disease
A species resistancemodel
distribution genes from
(SDM) C. was
liberica (e.g., SH3)using
developed and the thecaffeine-related
maximum entropy gene
HCT C. liberica,
modelin(Maxent) as the
with theaim is to develop
presence of data onlyC. liberica disease-resistant
[15]. They predict possible varieties of coffee
suitable areas
with the of
for each desired CGA
the three and caffeine
species of coffeecontent.
based on the data from existing locations. The models
were based on nineteen bioclimatic variables as well as altitude data on the current con-
3. Defining
dition, Areas
covering forfrom
data Cultivation of Coffea
1950 to 2000 from liberica
the WorldClim database (http://www.world-
Different climatic
clim.org/current). The factors and environmental
downloaded requirements
files have a spatial greatly
resolution of 30vary among Coffea
arc seconds (1 km
species.
grid). AsSome of these factors
a preliminary output,include
the MaxEnt the lowland
performance and highland
was goodlocations, the different
for the calculated AUC
elevation of coffeetest.
and the jackknife plantation,
The initialoptimum
models for annual rainfall
the three range,are
varieties optimum
presented mean
belowannual
(Fig-
temperature
ure 4). range, air humidity, and wind stress. In addition, the effects of shade have
also been proven to have great interaction with the biochemical composition of coffee,
most especially with green coffee beans. These contribute to changes in the different
properties of coffee. Thus, environment can be classified as a contributing factor. Data
on the environment can also be used to predict suitable areas for developing C. liberica
plantations.
A species distribution model (SDM) was developed using the maximum entropy
model (Maxent) with the presence of data only [15]. They predict possible suitable areas
for each of the three species of coffee based on the data from existing locations. The
models were based on nineteen bioclimatic variables as well as altitude data on the current
condition, covering data from 1950 to 2000 from the WorldClim database (http://www.
worldclim.org/current). The downloaded files have a spatial resolution of 30 arc seconds
(1 km grid). As a preliminary output, the MaxEnt performance was good for the calculated
Proceedings 2023, 89, 27 6 of 7

Proceedings 2023, 24, x 6 of 7

AUC and the jackknife test. The initial models for the three varieties are presented below
(Figure 4).

Figure 4.4. MaxEnt

Figure MaxEnt model
model outputs
outputs for (a) C.
for (a) C. arabica; (b)C.
arabica; (b) C.canephora;
canephora; and (c)C.
and(c) C.liberica.
liberica. The
The limited
limited
distribution
distributionininC.
C.liberica
libericaisisshown.
shown.

The
The Maxent
Maxent model for C.
model for C. coffea
coffea in
in the
the Philippines
Philippines generally
generally reflects
reflects the
the current
current areas
areas
of
of cultivation
cultivation of C. arabica
of C. arabica and
and C.C. canephora
canephoraindicating
indicatingthe
therobustness
robustnessofofthethemode.
mode.ForFor
C.
C. liberica,
liberica, since
since cultivation
cultivation data
data is is only
only from
from a small
a small area,
area, the
the suitable
suitable areas
areas forfor cultivation
cultivation is
is limited [16], The maps may be further expanded with data on areas where the species
limited [16], The maps may be further expanded with data on areas where the species
have established culture.
have established culture.
4. Conclusions
4. Conclusions
This paper presents some new insights on the diversity of C. liberica and how it
This paper
compares presents
to C. arabica and some new insights
C. canephora. on the
It also diversity
presents of C. libericaamong
the differences and how theitinfra-
com-
pares toC.C.liberica
species arabicaliberica
and C.and canephora. It also
C. liberica presents
dewevrei. Thethe aimdifferences
is to add among the infra-species
to the limited literature
C. liberica liberica and C. liberica dewevrei. The aim is to add
on the species from the data acquired in the Philippines. This study highlights to the limited literature on the
its potential
species from the data acquired
for cultivation amidst a changing climate. in the Philippines. This study highlights its potential for
cultivation amidst a changing climate.
Both C. liberica infra-species were found to be different from C. canephora and C. arabica
in leafBoth
and C.beanliberica infra-species
morphology, were found
branching to be different
architecture, from C.pattern.
and the rooting canephora andspecies
These C. ara-
bica in
have leaf and
bigger front bean
facingmorphology, branching
leaves, deeper roots, architecture,
and larger beans and the rootingstorage
for water pattern. andThese
are
species have bigger front facing leaves, deeper roots, and larger
more adapted to greater variation in climatic conditions, temperature stress, and drought. beans for water storage
and Initial
are more data adapted to greater
also indicates thatvariation in climatic
for infra-species of conditions,
C. liberica thetemperature
nature, or thestress, and
genetic
drought.
variety of the plant, the environment has a lesser influence on the phenotype in the species,
Initial data
particularly in the also indicates that
polyphenols, e.g.,for infra-species
trigonelline, of C. liberica
chlorogenic acidthe nature,
(CGA), andor caffeine.
the genetic
variety of the
Finally, theplant, the environment
approach of predicting has a lesser
areas influence
suitable on the varieties
for planting phenotype of in the spe-
coffee can
cies,performed
was particularly in the
using polyphenols,
a species distributione.g., model
trigonelline,
based on chlorogenic
maximumacid (CGA),
entropy. and caf-
Parameters
feine.WorldClim from 1950 to 2000 produced reasonable maps for C. arabica and C. canephora
from
whenFinally,
the currentthe approach
data on where of predicting areas suitable
they are placed for planting
in the Philippines varieties
were given.ofThe coffee can
results
wasC.performed
for using a species
liberica showed limited distribution
area given that modelthebased
current onareas
maximum
whereentropy.
they areParame-
grown.
ters from
When WorldClim
validated, these from
models 1950
may to be2000
veryproduced
useful for reasonable
future work mapson for C. arabica
Philippine and C.
coffee.
canephora when the current data on where they are placed in the Philippines were given.
Funding:
The results Thisforresearch was showed
C. liberica funded by the United
a limited States
area given Agency for International
that the current areasDevelopment
where they
(USAID),
are grown. through
When RTIvalidated,
International, under
these modelsUSAID STRIDE
may grantuseful
be very agreement No. 0213997-G-2015-002-
for future work on Philip-
00, and
pine internal support by De La Salle University using the Practical Genomics Laboratory.
coffee.
Institutional Review Board Statement: Not applicable.
Funding: This research was funded by the United States Agency for International Development
Informed
(USAID), Consent Statement:
through RTI Not applicable.
International, under USAID STRIDE grant agreement No. 0213997-G-2015-
002-00, and internal support by De La Salle University using the Practical Genomics Laboratory.
Institutional Review Board Statement: Not applicable.
Proceedings 2023, 89, 27 7 of 7

Data Availability Statement: Data used in this paper are from the original research and thesis found
at https://animorepository.dlsu.edu.ph/.
Acknowledgments: The researcher would like to acknowledge the contributions of Aldrin Mallari
and Victor Fuentebella for the work on the species distribution model. The contributions of Janlyn
Santos, Jewel Macato, Joshua Kyle Sy, and Aize Villegas are also acknowledged.
Conflicts of Interest: The author declares no conflict of interest. The funders had no role in the design
of the study; in the collection; analyses; or interpretation of data; in the writing of the manuscript; or
in the decision to publish the results.

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