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Terrestrial Vertebrate Diversity of Caves and Limestone Forests in Southern

Sierra Madre Mountain Range, Luzon Island, Philippines

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Philippine Journal of Science
152 (6A): 2061-2075, December 2023
ISSN 0031 - 7683
Date Received: 20 Jun 2023

Terrestrial Vertebrate Diversity of Caves and Limestone

Forests in Southern Sierra Madre Mountain Range,
Luzon Island, Philippines

1,2 2,3 2,3†

Renz Angelo J. Duco , Anna Pauline O. de Guia , Judeline C. Dimalibot ,
2 2 2 2
Julius A. Parcon , Edison A. Cosico , Charlie R. Malizon , Ammie R. Aguila ,
2,3 2,3
Phillip A. Alviola , and Juan Carlos T. Gonzalez

1
Biodiversity Research Laboratory, Institute of Biology,
University of the Philippines Diliman, Quezon City 1101 Philippines
2
Museum of Natural History, CFNR Quadrangle, Upper Campus,
University of the Philippines Los Baños, College, Laguna 4031 Philippines
3
Institute of Biological Sciences, College of Arts and Sciences,
University of the Philippines Los Baños, College, Laguna 4031 Philippines

The caves and karst forests of southern Sierra Madre are a unique and understudied habitat that
is home to a high number of endemic and globally threatened species. These habitats provide
essential resources for the local terrestrial vertebrate fauna, although increasing anthropogenic
disturbances pose a serious threat to existing wildlife populations. Here, we provide an inventory
of terrestrial vertebrate fauna of caves and limestone forests of three ecotourism areas within the
southern Sierra Madre in CALABARZON Region, Philippines. A combination of various field
survey methods (i.e. line transect surveys, mist netting, opportunistic catching, cage trapping)
was conducted from 2021–2023 to determine the diversity of amphibians, reptiles, non-volant
mammals, bats, and birds in the three sites. A total of 160 species (13 amphibians, 23 reptiles,
98 birds, four non-volant mammals, and 22 bats) were recorded in this study. Endemicity was
high at 46% with 56 Philippine endemics, including 21 species endemic to the Greater Luzon
Island. In addition, we recorded 18 locally and globally threatened species based on the IUCN
Red List and DENR-DAO 2019-09. The high species richness and presence of threatened species
underline the high conservation value of the study areas. However, unmanaged growth of tourism
and habitat fragmentation threaten their biodiversity. The data generated will be used to guide
the classification, proper management, and conservation of these karst areas, caves, and their
resources by the national government and local government units.

Keywords: birds, CALABARZON, ecotourism, herpetofauna, karst forest, mammals

INTRODUCTION reptiles, 115 amphibians, 235 mammals, and 725 birds

(IUCN 2022; Brinkman et al. 2022). Of these, about 62%
The Philippines is considered one of the 17 megadiverse (727 species) are endemic to the Philippines. However,
countries of the world, supporting one of the highest these numbers are still considered an underestimation of
concentrations of endemic species and terrestrial vertebrate the true diversity of terrestrial vertebrates in the country,
life (Brown and Diesmos 2009). The country houses at given the inaccessibility of biodiversity surveys in some
least 1166 species of terrestrial vertebrates – including 326 islands and extreme locations, as well as limited studies
*Corresponding author: [email protected] and ongoing debates among scientists with regard to the
†Deceased: May 2022 taxonomy and delineations of some species.
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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

Unfortunately, the country’s rich biodiversity and MATERIALS AND METHODS

exceptionally high rate of endemism are continually
threatened mainly by habitat loss and fragmentation due to
Study Sites
various anthropogenic activities such as logging, mining,
Surveys were conducted in three sites within the southern
and accelerated changes in land use (Catibog-Sinha and
Sierra Madre Mountain Range in the CALABARZON
Heaney 2006; Corlett 2019). The Philippines’ forest
Region, Philippines (Figure 1). Study sites generally
ecosystems are considered one of the most ecologically
consisted of lowland forest over limestone characterized by
vulnerable areas in the world owing to rapid tropical forest
abundant karst outcrops and rugged terrain. The climate is
loss, with almost 80% decline in forest habitats in the last
characterized by two pronounced seasons: the dry season
100 years (Turner et al. 2003; Corlett 2019). Currently,
occurring from March–May and the wet season, which is
322 species of terrestrial vertebrates – most of which are
usually marked by heavy rainfall that lasts for the rest of the
endemics – are placed under different categories of threat
year. The following are brief descriptions of each study site:
due to continuous loss and destruction of their natural
habitats, illegal trafficking, overexploitation, which are Cavinti. Sampling was conducted in Cathedral
further exacerbated climate change (Ong et al. 2002a; (14°16'54.48"N, 121°38'9.66"E, 280 masl) and Minalokan
BCSP 2020). (14°16'53.88"N, 121°37'58.44"E, 295 masl) caves located
in Brgy. Paowin, Cavinti, Laguna. These caves are
The Sierra Madre is considered one of the last frontiers in
classified as Class II by the Department of Environment
the Philippines, having one of the largest remaining blocks
and Natural Resources (DENR) and are currently open to
of tropical old-growth rainforest (Poulsen 1995). It is the
tourists. Both caves are long and complex, with several
longest mountain range in the Philippines, spanning most
entrances present, and are traversed by an underground
of the entire eastern Luzon Island from the province of
river. The surrounding vegetation is mainly characterized
Cagayan in the north to the Quezon province in the south.
by secondary-growth forests adjacent to agroforests and
One of its sections – the southern Sierra Madre, which
coconut plantations. Dominant plant families recorded
runs from the southern portion of Mingan Mountains to
in the area were Melastomataceae, Euphorbiaceae, and
Atimonan, Quezon Province – is considered a distinct
Moraceae (Ficus spp.). A paved road from the village
biogeographic region of Luzon Island, considering the
proper to about 250 m from the cave complex was recently
high degree of microendemism observed for herpetofauna
constructed.
and mammals (Devan-Song and Brown 2012; Balete et al.
2013; Vallejo and Telnov 2014). However, southern Sierra Tayabas. The sampling area is located in Sungwan
Madre’s terrestrial forest areas are mostly unexplored, Cave (14°1’29.10”N, 121°39’55.20”E; 215 masl) and
and little is known about the terrestrial biodiversity of surrounding karst forests situated in Brgy. Lawigue,
lowland karst forests and caves present in this protected Tayabas City, Province of Quezon. The cave is also
area. Because of the rapid conversion of forest habitats and classified as Class II and open for tourist visits (DENR-
natural landscapes in the southern Sierra Madre, additional PAWB 2008). The site is fairly remote, taking at least an
efforts to conduct species inventories and monitoring are hour on foot to reach the campsite from Brgy. Lawigue
needed to fill knowledge gaps and capture the full picture proper. The cave is inside a private land, although tourists
of terrestrial biodiversity in this protected area (Haribon often visit the site for recreational purposes and religious
Foundation 2018; Barit et al. 2022). In addition, as caves activities. Sungwan Cave has three identified entrances,
and limestone forests are threatened habitats and home one of which is flooded and connected to a small river.
to a number of known karst and cave obligate species, The surrounding vegetation consisted of secondary
collection and biodiversity documentation are needed to forest and lowland riverine forest mainly dominated by
better understand the usage of karst resources by these Euphorbiaceae and Arecaceae plants. Coconut plantations
unique species toward effective conservation management and agricultural lands are also present about a kilometer
and species-specific intervention (Duco et al. 2021). from the cave.
Here, we present the results of an inventory of terrestrial Pamitinan. Sites were located inside the Pamitinan
vertebrates (amphibians, reptiles, birds, and small volant Protected Landscape (PPL) in Rodriguez, Rizal. Three
and non-volant mammals) in selected caves and karst caves were surveyed during this study: Pamitinan Cave
forest areas in southern Sierra Madre, Luzon Island. (14°43'48.42"N, 121°11'21.18"E; 120 masl), Bat Cave
Important ecological values (i.e. species richness, (14°43'51.78"N, 121°11'23.22"E; 240 masl), and Lobog
diversity, endemicity, and conservation status), as well Cave (14°44'47.28"N, 121°11'42.54"E; 260 masl). The
as threats to terrestrial vertebrate fauna in these areas, are Pamitinan and Bat Caves are both classified as Class II
also provided to guide proper conservation management and located near human settlements, government offices,
of these habitats and their inhabitants. and small markets and business centers of Brgy. San

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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

Figure 1. Location of the sampling areas in CALABARZON region.

Rafael. These caves are also near Wawa Dam, which is such as burrows, rotten logs, and near cave entrances.
frequently visited by tourists. Although these caves are These were left overnight then checked and re-baited
not open for visitors, signs of human disturbance such the following morning. A total of 270 trap nights were
as the presence of man-made structures, water pipes, employed in Cavinti, whereas sampling efforts for Tayabas
vandalism, and speleothem collection are apparent for and Pamitinan were 200 and 60 trap nights, respectively.
both caves. Meanwhile, Lobog cave is situated in Brgy. Signs of the presence of other non-volant mammals (e.g.
Mascap, about 5 km away from the urban areas of Brgy. monkeys, warty pigs, deers, and civets) were also noted.
San Rafael. The cave has a high ceiling (~ 10 m) and is
traversed by a river. Minimal disturbance was observed Bats were captured using mist nets (6 x 2.6 m with 36-
as only a few locals live near the area. mm mesh) set low to the ground near potential flyways
inside the forest area, ridges, as well as near streams and
rivers. These nets were tended from 1800–2000 h and
Field Sampling deployed for a minimum of three nights for each site.
Surveys were carried out from December 2021–February Nets were also established in cave entrances to capture
2023, covering both wet and dry seasons. Amphibians bats emerging from the caves. The total sampling effort
and reptiles were sampled through strip-transect sampling 2
for Cavinti was 23961.60 mist-net hours (m ∙ h), whereas
and opportunistic catching (Delima et al. 2007). For each 2
28454.40 and 11232.00 m ∙ h were spent for Tayabas and
of the three sites, 250-m transect lines were established Pamitinan, respectively. The same mist nets set up for bats
and sampled from 0700–0900 and 2000–2200. Specific were used to capture diurnal and nocturnal bird species.
microhabitats, such as aerial ferns, forest floor litter, tree Mist nets were checked every 30 min to prevent any
holes, rocks, streams, and plant leaves, were searched for uneventful death of captured birds. Mist netting for birds
any herpetofauna occupying them. The total sampling was performed only for Cavinti and Tayabas, with netting
efforts for the three sites were 28 h for Cavinti, 34 h for 2
efforts of 22464.00 and 23961.60 m ∙ h, respectively.
Tayabas, and 10 h for Pamitinan.
Mist netting for birds was also supplemented by transects
Cage traps (28 cm x 18 cm x 12 cm and 56 cm x 36 cm (Duco et al. 2021). At least two transect lines for each site
x 24 cm) were used to capture non-volant mammals in were established, and transect walks were done twice each
the study sites. These were baited with toasted coconut day – from 0600–0800 and 1600–1800. Bird species heard or
and peanut butter, then set up in potential runways

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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

seen were recorded. Photographs were also taken to confirm documented, about 46% (77 species) are endemic to the
identification. Sampling efforts for bird transect were 79 h for Philippines; this includes nine amphibians, 14 reptiles,
Cavinti, 32 h for Tayabas, and 30 h for Pamitinan. 46 birds, one rodent, and seven bat species. Further, 21
species (six amphibians, five reptiles, nine birds, and
All captures were measured and identified following field one bat) are endemic to Greater Luzon Island. Three
guides and books available such as Alcala (1986), Alcala introduced species (Columba livia, Passer montanus, and
and Brown (1998), Brown et al. (1996, 2013), Ingle and Rattus tanezumi) were also recorded in our study areas.
Heaney (1992), Heaney et al. (2016), and Allen (2020).
Information on the species’ residency status, general In terms of herpetofauna diversity, our study areas
population trend, and global and local conservation status exhibited a lower diversity of amphibians and reptiles
were also extracted from the International Union for compared to other surveys conducted in the Sierra Madre
Conservation of Nature’s (IUCN) most recent assessment region. For instance, Brown et al. (1996) reported 52
(2022), DENR-DAO (Department Administrative Order) species in the Central Sierra Madre, whereas Brown
2019-09, and WBCP (2022). et al. (2013) documented 101 species in the northern
Sierra Madre. However, when compared with other areas
Muscle tissue samples and voucher specimens were also surveyed in southern Luzon Island such as Batangas (24
collected for some species, especially those belonging species; Endozo et al. 2017) and Caramoan (22 species;
to species complexes, for molecular verification and Clores et al. 2021), our study sites displayed higher species
identification in combination with the collected taxonomic richness. Birds are also more diverse in our study sites
data. All voucher specimens were deposited at the University compared to nearby protected areas such as Mt. Makiling
of the Philippines Los Baños–Museum of Natural History (25 species; Gonzalez et al. 2020), Mt. Banahaw (30
Zoological Collection. Field sampling was covered by species; Gascon et al. 2013), and Mt. Irid-Angelo (88
Wildlife Gratuitous Permit Nos. R4A-WGP-2021-LAG-004, species; Haribon Foundation 2023). Lastly, mammalian
R4A-WGP-2021-QUE-005, and R4A-WGP-2021-RlZ-010. diversity in the sites surveyed is comparable with those
from other parts of the Sierra Madre region such as those
Data Analysis reported in Mt. Irid (24 species; Balete et al. 2013), the
The adequacy of our sampling effort in the three sites Mingan Mountains (35 species; Balete et al. 2011), as
was assessed using sample size-based rarefaction and well as in the provinces of Cagayan, Quirino, and Quezon
extrapolation sampling curves for species richness using (37 species; Duya et al. 2007) despite the lower sampling
the iNEXT package in R (Hsieh et al. 2016). The curves effort and limited sampling areas in this study.
were computed for each site using abundance data and
Rarefaction and extrapolation curves generated for bird
estimated with 95% confidence intervals based on 100
transect and bats showed an approximation of a plateau
bootstraps, with non-overlapping confidence areas
in species diversity indicating enough sampling effort to
indicating significant differences in species richness
document these taxa in all three sampling areas (Figures
between curves (α = 0.05).
2d and e, respectively); sampling completeness for the
Shannon’s species diversity index (H’) was used as a sampling sites ranged from 73–93% for bat mist netting and
measure of diversity, as it takes into account a community's 82–93% for bird transect. While the species accumulation
species diversity and relative abundance (Magurran 2004). curve for bird mist netting did not reach a plateau (Figure
This index was estimated using the diversity function 2c), all species recorded for this method were documented
from the vegan package in R (Oksanen et al. 2013). during transect sampling. Mist netting has drawbacks that
Diversity values were then compared between the three may result in limited data; factors such as birds and bats
sites using the Kruskal-Wallis test (α = 0.05) using IBM avoiding the nets after multiple attempts, as well as lower
SPSS Statistics for Windows v 20.0. detection rates per unit effort contribute to this (Ralph et
al. 2004). A combination of mist netting and transects may
yield a more complete inventory of birds in an area (Duco
et al. 2021). Further, setting up canopy nets should also be
RESULTS AND DISCUSSION considered to capture birds that are more active in the upper
forest strata (Duco et al. 2021). Alternative techniques such
Terrestrial Vertebrate Community and Sampling as utilizing recorders to detect bird songs and, calls using
Completeness radar or sensors to capture flight, patterns and setting up
A total of 160 species of terrestrial vertebrate species camera traps are also increasingly being used to provide
were recorded from the three sites – comprised of a more comprehensive understanding of the diversity and
13 amphibians, 23 reptiles, 98 birds, four non-volant behavior of local bird communities (Rovero et al. 2010;
mammals, and 22 bats (Appendix I). Of the 160 species Washburn et al. 2022; Budka et al. 2022).

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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

Insufficient sampling for herpetofauna (Figures 2a and Among the three study sites surveyed, Cavinti was the
b) suggests a higher likelihood of finding more species. most species-rich (109 species), followed by Tayabas
Although the sampling completeness of herpetofauna (106 species), whereas Pamitinan had a significantly
for the Tayabas site was high (79% for amphibians and lower number of species recorded (72 species). Shannon’s
89% for reptiles), sites in Cavinti and Pamitinan yielded diversity index, meanwhile, showed that the Tayabas site
low results (< 61%). This is expected as more effort was is relatively more diverse (H’= 1.901) compared to Cavinti
provided for herpetofauna sampling in Tayabas (34 man- (H’= 1.771) and Pamitinan (H’= 0.921), although this
hours) than in either Cavinti (26 man-hours) or Pamitinan difference is not significant based on the Kruskal-Wallis
(10 man-hours). Employing additional techniques like test (H= 4.067, p = 0.131). The presence of a few intact
pitfall traps, drift fences, and bioacoustics may be needed old-growth forests in Cavinti and Tayabas and a relatively
for a more comprehensive herpetofauna survey in the lesser degree of anthropogenic disturbance experienced
study areas (Bury and Corn 1987; Bolitho et al. 2023). in these sites may have contributed to the observed

Figure 2. Sample completeness rarefaction (solid line segment) and extrapolation (dotted line segments) sampling
curves for species richness with 95% confidence intervals (shaded areas) using abundance data from [a]
amphibians, [b] reptiles, [c] birds mist netting, [d] birds transect, [e] bats mist netting, and [f] non-volant
mammal trapping.

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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

high diversity and richness of terrestrial vertebrate study area poses a threat, as their proliferation and spread
species. These sites are adequately managed by the local could result in the displacement of native fauna (Pili et al.
government units (LGUs) to ensure that the forested 2019). Identifying species undergoing population decline
areas are conserved and protected. Further, these areas or those that could potentially threaten local wildlife is of
are fairly remote and are still relatively underdeveloped, utmost importance for conservation prioritization, as this
so anthropogenic disturbance is still considered minimal. will be an important guide for policymakers in ensuring
In contrast, although the site in Pamitinan is inside a that resources and efforts are directed toward the most
protected area, human encroachment and deforestation vulnerable species.
are apparent. Human settlements, small business centers,
and markets were established a few meters away from Conservation assessment for most of the species
the Pamitinan Cave entrance. Further, plastic pipes that documented in this study is already outdated. For instance,
supply water to nearby communities were installed inside 112 out of the 160 species had their last conservation
the cave. These factors may have contributed to the low assessment done more than five years ago, including one
diversity of terrestrial vertebrates in the site, as observed species (Polypedates leucomystax) that was last assessed
in other studies (Seltmann et al. 2017; Quibod et al. 2019). in 2004. Meanwhile, the local conservation assessment
of threatened species in the Philippines was updated in
2019 through DENR DAO 2019-09. To properly manage
Conservation Status and Assessment and conserve these species, an updated conservation
Our survey also yielded a record of six Threatened and six assessment and population status is needed.
Near-Threatened species of terrestrial vertebrates based
on IUCN (2022), which includes four species (Varanus The presence of threatened species in the sites surveyed
olivaceus, Bubo philippensis, Buceros hydrocorax, and highlights the significance of the southern Sierra Madre as
Desmalopex leucopterus) under Vulnerable category an important habitat for species of conservation concern,
and two Endangered species (Cuora amboinensis and particularly in the CALABARZON region. This region in
Macaca fascicularis) (Appendix I). In addition, three southern Luzon is rapidly developing into a metropolitan
endemic species (Gonocephalus sophiae, Malayotyphlops area with an increasing human population. Despite having
ruficaudus, and Miniopterus eschscholtzii) are either 23 key biodiversity areas recognized as priority conservation
categorized as Data Deficient or Not Assessed. Meanwhile, areas, only a few are adequately protected by law (Ong et al.
a total of 17 locally threatened species were recorded in the 2002b). Thus, utmost attention to protecting the remaining
study areas based on DENR-DAO 2019-09. This includes lowland forests of southern Sierra Madre is important to
four Vulnerable species (Varanus olivaceus, Copsychus provide a suitable habitat for these threatened species.
luzoniensis, Penelopides manillae, and Desmalopex Further, the CALABARZON region is believed to have
leucopterus), two Endangered (Bubo philippensis and a significant number of caves, many of which are still
Buceros hydrocorax), three Critically Endangered bird unknown and awaiting discovery. Currently, 79 caves have
species (Ceyx cyanopectus, Loriculus philippensis, and been identified, 45 of which are open for tourists (DENR-
Phapitreron amethystinus), and eight species under Other CALABARZON Regional Cave Committee, pers.
Threatened Species category. comm.). However, comprehensive data on the terrestrial
Based on the recent IUCN assessment (2022), a significant vertebrates of these caves is yet to be generated. Thus,
proportion of the species recorded in this study are additional surveys are needed to generate data on these
experiencing a decrease in population size (39.4% or 63 caves to guide proper classification and the management
out of 160 species recorded), whereas about 38.1% (61 and conservation of these biologically important habitats.
species) still have a stable population. Meanwhile, ten
species have increasing population size based on the latest Threats to Terrestrial Vertebrates and
IUCN assessment, most of which are introduced invasive Recommendations for Management of Caves and
species (i.e. Rhinella marina, Rattus tanezumi) as well Karst Areas
as migratory bird species that have a wide distribution Since our study areas are frequented by tourists, tourism-
range. Surprisingly, the population trend for the remaining related visits and activities may have a great impact on the
26 species (16%) is as unknown due to limited data for cave population of resident cave fauna. In this study, some
assessment. Given that a significant proportion of species bird species (i.e. Aerodramus amelis, Collocalia spp.) and
within our study area is experiencing population decline, most of the bat species we recorded are cave-dependent,
it underscores the significance of these sites as important utilizing caves as nesting and roosting sites. Caves provide
habitats for these species. The presence of migratory specific and stable microclimatic conditions (temperature
species further highlights the global importance of these and relative humidity) and physical structures that many
areas as resting stops during migration (Ouano et al. bat populations require for protection, roosting, and feeding
2022). In contrast, the presence of invasive species in the

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(Murray and Kunz 2005). Human visits are known to proper management of these caves and karst areas,
influence and cause abrupt changes in the microclimate following existing environmental laws in the Philippines
(i.e. fluctuations in temperature, relative humidity, and (i.e. RA 9072 or the National Caves and Cave Resources
carbon dioxide concentrations) inside the cave, which Management and Protection Act and RA 9147 or the
could negatively affect cave-dwelling wildlife populations Wildlife Resources Conservation and Protection Act).
(Rakovec 2020; Debevec and Rakovec 2021; Alviola et al. Limited and closely guided visits of small groups in these
2022). The anthropogenic disturbance caused by tourism areas are recommended to ensure minimal disturbance of
activities has also been known to result in abandonment wildlife populations – specifically, avoiding visitations
and a decrease in the abundance and richness of cave- during mating and maternity seasons of bats and breeding
dwelling bats (Luo et al. 2013; Alcazar et al. 2020), birds and nesting of swiftlets, ensuring local guides are always
(Soldatini et al. 2015; Watson et al. 2021), and herpetofauna present during visits, and reducing light and noise
(Nuñeza and Galorio 2015; Ayu et al. 2020). Furthermore, disturbance inside the cave (Ramji et al. 2013; Medellin
disturbance during sensitive periods of pregnancy, lactation, et al. 2017; Torres-Flores and Santos-Moreno 2017).
and weaning in cave bats can lower the survival rate of pups,
whereas higher mortality rates in nests exposed to human
recreational disturbance were observed for cave-nesting
seabirds (Watson et al. 2021). Garbage and graffiti in the CONCLUSIONS
cave walls probably caused by the influx of unguided tourist
This study provides an inventory of terrestrial vertebrate
visits were also observed during our survey, indicating signs
species present in lowland karst forests and caves in the
of negative visitor impact. Future studies should prioritize
southern Sierra Madre. Given the inadequacy of sampling
monitoring and examining the impacts of the identified
for some taxa in the sites, as well as the increasing number
disturbances on the cave fauna and establish effective cave
of newly discovered and unclassified caves and unexplored
management strategies.
karst areas in the region, it is possible that more species
Land-use change resulting from the development of these can be documented in the future. Thus, intensive surveys
tourism areas is also a major threat to terrestrial vertebrate and long-term studies of caves and karst areas of southern
diversity in our study areas. For instance, a road going to Sierra Madre are steps forward and we expect a significant
Cathedral Cave in Cavinti, Laguna was paved recently, increase in the current species list and possible discovery
resulting in an influx of tourists and the construction of more of novel species, as more surveys utilizing a variety of
human settlements and business centers due to easier access. sampling methodologies are conducted.
The construction of the road has also caused damage and
The presence of island endemics, as well as species of
fragmentation to the surrounding karst landscape.
high conservation concern, underlines the potential of the
Other threats to wildlife population in the caves and caves and the surrounding karst landscapes as important
surrounding karst areas observed during this survey areas to support viable terrestrial vertebrate communities
include hunting, illegal logging, agricultural expansion, in the southern Sierra Madre. Further, the sites surveyed
speleothem and bird-nest collection, guano mining, and are also important habitats for both cave- and forest-
the presence of infrastructures such as cemented stairs, dependent species, providing suitable nesting and roosting
water pipes, and religious grottos inside the cave. Such resources despite the apparent increase in human visitation
human activities and alterations in the cave and karst and activities and the ongoing developments in the area.
habitat can have detrimental effects on the wildlife Hence, raising community awareness, developing of
population and the natural features of the karst landscape. conservation management plan, strict law enforcement,
and long-term monitoring are needed to protect these
Ecotourism has contributed significantly to economic important habitats and the wildlife population, as well as
growth by providing income for local communities make the ecotourism industry more sustainable.
and supporting biodiversity conservation initiatives
by augmenting funds for the management of protected
areas (Clements et al. 2006; Tolentino et al. 2020). If
unregulated, ecotourism activities inside the caves, as well ACKNOWLEDGMENTS
as the anthropogenic pressures present in the surrounding
karst landscape, may result in long-term negative impacts We acknowledge the support given by DOST Philippines
on wildlife populations (e.g. low species richness and (Department of Science and Technology) through the
diversity, abandonment of roosting sites) and destruction NICER-CAVES (Center for Cave Ecosystems Research)
of various karst features (Perry 2013). Given the presence Program of the UPLB (University of the Philippines Los
of endemic and species of conservation concern in the Baños) Museum of Natural History, under the research
sites visited, this study recommends protection and management of the DOST-PCAARRD (Philippine Council

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Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

for Agriculture, Aquatic, and Natural Resources Research eastern region of the Sierra Madre Mountain Range,
and Development). We also want to thank the LGU of Philippines. iForest 15: 63–70.
Cavinti, Tayabas City ENRO (Environment and Natural
[BCSP] Biodiversity Conservation Society of the Phil-
Resources Officer), and PAMB of PPL for providing
ippines. 2020. Philippine Red List of Threatened
logistical assistance and local guides. We acknowledge
Wild Fauna. Department of Environment and Natural
Wilson Bulalacao and Benjie Gurobat for their assistance in
Resources Biodiversity Management Bureau (DEN-
data gathering and for helping us throughout the duration of
RBMB), Quezon City, Philippines. 120p.
fieldwork. We thank DENR-CALABARZON for granting
the necessary permits to conduct the research. BOLITHO L, NEWELL D, HINES H. 2023. Bioacoustic
Monitoring Reveals the Calling Activity of an En-
dangered Mountaintop Frog (Philoria kundagungan)
in Response to Environmental Conditions. Diversi-
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Appendix I. List of terrestrial vertebrates recorded including residence status, endemicity, conservation status based on IUCN (2022) and
DENR DAO 2019, and distribution.
Conservation status Recording Study area recorded
Taxa Residency status
IUCN (2022) DENR DAO 2019 method Cavinti Tayabas Pamitinan
Amphibians
Kaloula conjuncta Philippine endemic Least Concern Not listed Transect +
Kaloula kalingensis Luzon endemic Least Concern Not listed Transect + +
Other Threatened
Limnonectes macrocephalus Luzon endemic Near Threatened Transect + + +
Species
Limnonectes woodworthi Luzon endemic Least Concern Not listed Transect + +
Occidozyga laevis Resident Least Concern Not listed Transect + + +
Platymantis corrugatus Philippine endemic Least Concern Not listed Transect + +
Platymantis dorsalis Philippine endemic Least Concern Not listed Transect + +
Platymantis luzonensis Luzon endemic Near Threatened Not listed Transect + +
Platymantis mimulus Luzon endemic Least Concern Not listed Transect + +
Platymantis pygmaeus Luzon endemic Least Concern Not listed Transect +
Polypedates leucomystax Resident Least Concern Not listed Transect + + +
Rhacophorus pardalis Resident Least Concern Not listed Opportunistic +
Rhinella marina Resident Least Concern Not listed Transect +
Reptiles
Ahaetulla prasina Resident Least Concern Not listed Transect +
Aplopeltura boa Resident Least Concern Not listed Transect +
Other Threatened
+
Boiga cynodon Resident Least Concern Species Transect
Other Threatened
+ +
Bronchocela cristatella Resident Least Concern Species Transect
Calamaria gervaisi Philippine endemic Least Concern Not listed Opportunistic +
Other Threatened
+
Cuora amboinensis Resident Endangered Species Opportunistic
Cyclocorus lineatus Luzon endemic Least Concern Not listed Transect +
Cyrtodactylus philippinicus Philippine endemic Least Concern Not listed Transect + +
Dendrelaphis luzonensis Luzon endemic Least Concern Not listed Transect + +
Dendrelaphis marenae Resident Least Concern Not listed Transect +
Draco spilopterus Philippine endemic Least Concern Not listed Transect +
Eutropis multicarinata Resident Least Concern Not listed Transect + +
Other Threatened
+ +
Gekko gecko Resident Least Concern Species Transect
Other Threatened
+
Gonocephalus sophiae Philippine endemic Data Deficient Species Transect
Hemibungarus calligaster Philippine endemic Least Concern Not listed Opportunistic +
Lamprolepis smaragdina Resident Least Concern Not listed Transect +
Lipinia pulchella Philippine endemic Least Concern Not listed Transect +
Malayotyphlops ruficaudus Luzon endemic Data Deficient Not listed Transect +
Pinoyscincus coxi Philippine endemic Least Concern Not listed Transect +
Other Threatened
+
Ptyas luzonensis Philippine endemic Least Concern Species Transect
Rhabdophis spilogaster Luzon endemic Least Concern Not listed Transect +

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Other Threatened
+ +
Trimeresurus flavomaculatus Philippine endemic Least Concern Species Transect
Varanus olivaceus Luzon endemic Vulnerable Vulnerable Transect +
Birds
Transect/ Mist-
Accipiter virgatus Resident Least Concern Not listed +
net
Transect/ Mist-
Actenoides lindsayi Philippine endemic Least Concern Not listed + +
net
Actitis hypoleucos Migratory Least Concern Not listed Transect +
Transect/ Mist-
Aerodramus amelis Philippine endemic Least Concern Not listed + + +
net
Aethopyga flagrans Philippine endemic Least Concern Not listed Transect +
Alcedo atthis Migratory Least Concern Not listed Transect +
Aplonis panayensis Resident Least Concern Not listed Transect +
Resident/
Ardea intermedia Least Concern Not listed Transect + +
Migratory
Ardea purpurea Resident Least Concern Not listed Transect +
Artamus leucorynchus Resident Least Concern Not listed Transect + +
Bolbopsittacus lunulatus Philippine endemic Least Concern Not listed Transect + +
Bubo philippensis Philippine endemic Vulnerable Endangered Transect +
Resident/
Bubulcus coromandus Least Concern Not listed Transect + +
Migratory
Buceros hydrocorax Luzon endemic Vulnerable Endangered Transect + + +
Centropus bengalensis Resident Least Concern Not listed Transect + + +
Centropus viridis Philippine endemic Least Concern Not listed Transect + + +
Transect/ Mist-
Ceyx cyanopectus Philippine endemic Least Concern Critically Endangered + +
net
Transect/ Mist-
Chalcophaps indica Resident Least Concern Not listed + + +
net
Chlidonias hybrida Migratory Least Concern Not listed Transect +
Chrysocolaptes haematribon Luzon endemic Least Concern Not listed Transect +
Cincloramphus timoriensis Resident Least Concern Not listed Transect +
Cinnyris jugularis Resident Least Concern Not listed Transect + + +
Collocalia isonata Philippine endemic Least Concern Not listed Transect + +
Collocalia marginata Philippine endemic Least Concern Not listed Transect + +
Transect/ Mist-
Collocalia troglodytes Philippine endemic Least Concern Not listed + + +
net
Columba livia Introduced Least Concern Not listed Transect +
Transect/ Mist-
Copsychus luzoniensis Philippine endemic Least Concern Vulnerable + +
net
Coracina striata Resident Least Concern Not listed Transect +
Corvus macrorhynchos Resident Least Concern Not listed Transect + + +
Cypsiurus balasiensis Resident Least Concern Not listed Transect + + +
Dasylophus cumingi Luzon endemic Least Concern Not listed Transect +
Dasylophus superciliosus Luzon endemic Least Concern Not listed Transect + +
Transect/ Mist-
Dicaeum australe Philippine endemic Least Concern Not listed + + +
net
Dicaeum bicolor Philippine endemic Least Concern Not listed Transect +

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Dicaeum hypoleucum Philippine endemic Least Concern Not listed Transect + +

Dicaeum pygmaeum Philippine endemic Least Concern Not listed Transect + + +
Transect/ Mist-
Dicrurus balicassius Philippine endemic Least Concern Not listed + + +
net
Resident/
Egretta garzetta Least Concern Not listed Transect + +
Migratory
Elanus caeruleus Resident Least Concern Not listed Transect +
Erythropitta erythrogaster Philippine endemic Least Concern Not listed Transect +
Eudynamys scolopaceus Resident Least Concern Not listed Transect + +
Falco severus Resident Least Concern Not listed Transect +
Gallus gallus Resident Least Concern Not listed Transect +
Geopelia striata Resident Least Concern Not listed Transect + + +
Gerygone sulphurea Resident Least Concern Not listed Transect + +
Halcyon gularis Philippine endemic Least Concern Not listed Transect + + +
Haliastur indus Resident Least Concern Not listed Transect + + +
Hemiprocne comata Resident Least Concern Not listed Transect +
Hirundo rustica Migratory Least Concern Not listed Transect + +
Hirundo tahitica Resident Least Concern Not listed Transect + + +
Hypothymis azurea Resident Least Concern Not listed Transect + +
Transect/ Mist-
Hypsipetes philippinus Philippine endemic Least Concern Not listed + + +
net
Irena cyanogastra Philippine endemic Near Threatened Not listed Transect +
Ixobrychus cinnamomeus Resident Least Concern Not listed Transect +
Lalage nigra Resident Least Concern Not listed Transect + +
Lanius cristatus Migratory Least Concern Not listed Transect + + +
Leptocoma sperata Philippine endemic Least Concern Not listed Transect + + +
Lonchura atricapilla Resident Least Concern Not listed Transect +
Lonchura punctulata Resident Least Concern Not listed Transect +
Loriculus philippensis Philippine endemic Least Concern Critically Endangered Transect + + +
Megalurus palustris Resident Least Concern Not listed Transect +
Merops americanus Resident Least Concern Not listed Transect + +
Merops philippinus Resident Least Concern Not listed Transect +
Resident/
Monticola solitarius Least Concern Not listed Transect +
Migratory
Motacilla cinerea Migratory Least Concern Not listed Transect + + +
Mulleripicus funebris Luzon endemic Near Threatened Not listed Transect +
Ninox philippensis Philippine endemic Least Concern Not listed Transect +
Oriolus chinensis Resident Least Concern Not listed Transect + + +
Orthotomus derbianus Luzon endemic Least Concern Not listed Transect + + +
Otus megalotis Luzon endemic Least Concern Not listed Transect + +
Transect/ Mist-
Pachycephala philippinensis Philippine endemic Least Concern Not listed +
net
Transect/ Mist-
Pardaliparus elegans Philippine endemic Least Concern Not listed + + +
net
Passer montanus Introduced Least Concern Not listed Transect + + +
Penelopides manillae Luzon endemic Least Concern Vulnerable Transect + + +
Pericrocotus divaricatus Migratory Least Concern Not listed Transect + +

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Resident/
Pernis ptilorhynchus Least Concern Not listed Transect +
Migratory
Pernis steerei Philippine endemic Least Concern Not listed Transect +
Phapitreron amethystinus Philippine endemic Least Concern Critically Endangered Transect + + +
Transect/ Mist-
Phapitreron leucotis Philippine endemic Least Concern Not listed + + +
net
Phylloscopus borealis Migratory Least Concern Not listed Transect +
Transect/ Mist-
Phylloscopus cebuensis Philippine endemic Least Concern Not listed +
net
Pitta sordida Resident Least Concern Not listed Transect +
Transect/ Mist-
Poliolophus urostictus Philippine endemic Least Concern Not listed + + +
net
Psilopogon haemacephalus Resident Least Concern Not listed Transect + +
Ptilinopus leclancheri Near Endemic Least Concern Not listed Transect + + +
Pycnonotus goiavier Resident Least Concern Not listed Transect + + +
Transect/ Mist-
Rhipidura cyaniceps Luzon endemic Least Concern Not listed + +
net
Rhipidura nigritorquis Philippine endemic Least Concern Not listed Transect +
Sarcops calvus Near Endemic Least Concern Not listed Transect +
Saxicola caprata Resident Least Concern Not listed Transect +
Sitta oenochlamys Philippine endemic Least Concern Not listed Transect +
Spilopelia chinensis Resident Least Concern Not listed Transect + +
Transect/ Mist-
Spilornis holospilus Philippine endemic Least Concern Not listed + + +
net
Terpsiphone cinnamomea Philippine endemic Least Concern Not listed Transect +
Todiramphus chloris Resident Least Concern Not listed Transect + +
Treron vernans Resident Least Concern Not listed Transect + +
Yungipicus maculatus Philippine endemic Least Concern Not listed Transect + + +
Zosterops meyeni Philippine endemic Least Concern Not listed Transect + +
Non-volant mammals
Rattus everetti Philippine endemic Least Concern Not listed Cage trapping + +
Rattus tanezumi Introduced Least Concern Not listed Cage trapping + +
General
Macaca fascicularis Resident Endangered Not listed + +
observation
General
Paradoxurus hermaphroditus Resident Least Concern Not listed +
observation
Bats
Cynopterus brachyotis Resident Least Concern Not listed Mist-net + +
Desmalopex leucopterus Philippine endemic Vulnerable Vulnerable Mist-net +
Eonycteris spelaea Resident Least Concern Not listed Mist-net +
Haplonycteris fischeri Philippine endemic Least Concern Not listed Mist-net +
Macroglossus minimus Resident Least Concern Not listed Mist-net + +
Otopteropus cartilagonodus Luzon endemic Least Concern Not listed Mist-net +
Ptenochirus jagori Philippine endemic Least Concern Not listed Mist-net + +
Rousettus amplexicaudatus Resident Least Concern Not listed Mist-net + + +
Mist-net/
Hipposideros antricola Resident Least Concern Not listed +
Acoustic

2074
 Philippine Journal of Science Duco et al.: Terrestrial Vertebrate Diversity
Vol. 152 No. 6A, December 2023 of Southern Sierra Madre

Mist-net/
Hipposideros bicolor Resident Least Concern Not listed
Acoustic
Mist-net/
Hipposideros diadema Resident Least Concern Not listed +
Acoustic
Hipposideros lekaguli Resident Near Threatened Not listed Mist-net + +
Mist-net/
Hipposideros pygmaeus Philippine endemic Least Concern Not listed +
Acoustic
Megaderma spasma Resident Least Concern Not listed Mist-net +
Mist-net/
Miniopterus eschscholtzii Philippine endemic Not Assessed Not listed + +
Acoustic
Mist-net/
Miniopterus paululus Resident Least Concern Not listed + + +
Acoustic
Mist-net/
Myotis horsfieldii Resident Least Concern Not listed + + +
Acoustic
Mist-net/
Rhinolophus arcuatus Resident Least Concern Not listed + + +
Acoustic
Mist-net/
Rhinolophus macrotis Resident Least Concern Not listed +
Acoustic
Mist-net/
Rhinolophus philippinensis Resident Least Concern Not listed + +
Acoustic
Mist-net/
Rhinolophus rufus Philippine endemic Near Threatened Not listed + +
Acoustic
Tylonycteris pachypus Resident Least Concern Not listed Mist-net +

Appendix II. Some of the endemic, threatened, and other notable species of terrestrial vertebrates observed from the karst areas in southern
Sierra Madre Mountain Range: [a] giant Philippine frog (Limnonectes macrocephalus), [b] Gray’s monitor (Varanus olivaceus),
[c] white-browed shama (Copsychus luzoniensis), [d] mottle-winged flying fox (Desmalopex leucopterus), [e] large Asian
leaf-nosed bat (Hipposideros lekaguli), [f] Philippine long-tailed macaque (Macaca fascicularis), [g] Philippine hanging parrot
(Loriculus philippensis).

2075

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