Journal of Threatened
Taxa | www.threatenedtaxa.org | 26 May 2025 | 17(5): 26999–27006
ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)
https://doi.org/10.11609/jott.9647.17.5.26999-27006
#9647 | Received 24 January 2025 | Final received 12 May 2025 | Finally
accepted 15 May 2025
Wallowing site observations of
the Philippine Tamaraw Bubalus mindorensis Heude, 1988 (Artiodactyla: Bovidae) using
camera trap data
Jean-Matthew B. Bate 1,
Nikki Heherson A. Dagamac
2, John Carlo Redeña-Santos 3, Emmanuel
Schütz 4 & Fernando Garcia Gil
5
1,2 The Graduate School, University
of Santo Tomas, España, Manila 1015, Philippines.
1,2 Research Center for the Natural
and Applied Sciences, University of Santo Tomas, España,
Manila, 1015, Philippines.
2 Department of Biological
Sciences, College of Science, University of Santo Tomas, España,
Manila, 1015, Philippines.
2,3,4,5 d’Aboville
Foundation and Demo Farm Inc. Makati, 1200, Philippines.
3 WildlifeLink
Research Consultancy Services, Mabitac, 4020,
Philippines.
1 matthewbate22@gmail.com
(corresponding author), 2 nhadagamac@gmail.com, 3 santosjcarlo@gmail.com,
4 manuschutz.ph@gmail.com, 5 fgarciagil@gmail.com
Editor: L.A.K. Singh, Bhubaneswar, Odisha, India.
Date of publication: 26 May 2025 (online & print)
Citation: Bate,
J-M.B., N.H.A. Dagamac, J.C. Redeña-Santos,
E. Schütz & F.G. Gil (2025). Wallowing
site observations of the Philippine Tamaraw Bubalus
mindorensis Heude, 1988
(Artiodactyla: Bovidae)
using camera trap data. Journal of
Threatened Taxa 17(5):
26999–27006. https://doi.org/10.11609/jott.9647.17.5.26999-27006
Copyright: © Bate et al. 2025. Creative Commons Attribution 4.0 International License.
JoTT allows unrestricted use, reproduction, and
distribution of this article in any medium by providing adequate credit to the
author(s) and the source of publication.
Funding: This study is funded by the Department of Science and Technology - Accelerated Science and Technology Human Resource
Development Program (DOST-ASTHRDP).
Competing interests: The authors declare no competing interests.
Author details: Jean-Matthew B. Bate, MSc, The Graduate School, University of Santo Tomas,
España 1015 Manila, Philippines, Research Center for the Natural and Applied Sciences, University of Santo
Tomas, España 1015 Manila, Philippines. Nikki Heherson Dagamac, associate professor, The Graduate School, University of Santo Tomas, España 1015 Manila, Philippines,
Research Center for the Natural and Applied Sciences, University of Santo Tomas, España 1015 Manila, Philippines, Department of Biological Sciences, College of Science, University of Santo Tomas, España 1015 Manila,
Philippines, D’ABOVILLE Foundation and Demo Farm Inc. Makati, 1200, Philippines. John Carlo Redeña-Santos, protection and law enforcement officer, D’ABOVILLE Foundation and Demo Farm Inc. Makati, 1200, Philippines, WildlifeLink Research Consultancy Services, Mabitac, 4020, Philippines. Fernando Garcia Gil, Conservation Research Officer, D’ABOVILLE Foundation and Demo Farm Inc. Makati, 1200, Philippines. Emmanuel Schütz, program director, D’ABOVILLE Foundation and Demo Farm Inc. Makati, 1200, Philippines.
Author contributions: All authors have contributed to data acquisition, data analysis, and manuscript writing.
Acknowledgements: JMBB and JCRS would like to thank the Department of Science and Technology - Accelerated Science and Technology Human Resource
Development Program (DOST-ASTHRDP) for the scholarship grant. JMBB and NHAD would also like to thank the World Wildlife Foundation
Philippines (WWF-Philippines) for the partnership. NHAD and JCRS acknowledge the support of the Department of Science and Technology – Philippine Council for Agriculture, Aquatic and Natural Resources Research and Development (DOST-PCAARRD) for the Balik Scientist and Project MATAPAT.
Abstract: Due to the combination of
anthropogenic and natural stressors, the endemic Philippine Tamaraw Bubalus mindorensis
was divided into four distinct subpopulations. Currently, the largest
population is found within the protected Mts. Iglit-Baco
Natural Park. Wallowing is one of the most important behavioral adaptations of B.
mindorensis to address heat stress
but it has remained understudied. With the species facing a continuous threat
of extinction, the future conservation plan for the species needs to be
carefully strategized, bringing attention to further understand its wallowing behaviour in the wild. Using camera trap technology, the
wallowing activity of Tamaraws was photo-captured in a wallowing site of Mts. Iglit-Baco Natural Park. The site was continuously
monitored for three years utilizing two sets of cameras capturing opposite
sides for a total of 517 independent wallowing events over 1096 sampling days.
The photographic data reveal a solitary and diurnal behaviour
of Tamaraw wallowing. The correlation of detections with climate variables
reveals temperature as a direct driver of B. mindorensis
wallowing. This study presents the baseline data of the wallowing behavior
of the otherwise data deficient species and brings attention towards its
management and conservation practices.
Keywords: Endemic, ethology, habitat use,
mammalogy, protected area, wildlife monitoring, thermoregulation.
INTRODUCTION
The Philippine Tamaraw Bubalus mindorensis
is the largest endemic land mammal in the Philippines, and it is found only on
the island of Mindoro, Philippines. The species is listed as Critically
Endangered under the Department of Natural Resources (DENR) Administrative
Order 2019–09, the Philippine Red List Committee, and The International Union
for Conservation of Nature (IUCN) and is also listed in the Appendix I of the
Convention on International Trade in Endangered Species (CITES) (Maala 2001). Historically, it was believed to be widespread
on the island; however, the combination of anthropogenic (land conversion,
illegal logging, migration, and hunting) and natural (rinderpest outbreak)
stressors caused a significant decrease in its numbers from an estimated 10,000
individuals in 1900 to 154 individuals by 2000 (Ishihara et al. 2007; Boyles et
al. 2016; Tabaranza et al. 2022; Schütz
et al. 2023). At present, the B. mindorensis
is reported to be confined within four sites (Image 1A) on the island: (1) Aruyan-Malate Tamaraw Reservation Area, (2) Upper Amnay watershed region, (3) Mt. Patrick region, and (4)
Mts. Iglit-Baco National Park (MIBNP) (Gil et al.
2021; Schütz et al. 2023).
Mts. Iglit-Baco
Natural Park (MIBNP) is a protected area proclaimed under the National
Protected Areas System (NIPAS) act in 1992 and was recognized as an ASEAN
heritage site in 2003 due to the rich biodiversity of native and endemic
species inhabiting the mountain (Bonenfant et al.
2023). It is located in the south-central portion of the island, which
coincides with the ancestral domain of the Taobuid
indigenous communities (Schütz et al. 2023). The park
is known for holding over 80% of the entire Philippine Tamaraw population and
has been the focal point of the Tamaraw conservation efforts since its
establishment (Long et al. 2018; Bonenfant et al.
2023). One instance is the established agreement between the wildlife managers
and the indigenous communities that prevents the hunting of Tamaraws,
harvesting of resources, or establishment of settlements within a 16 km2
area inside the Park which they refer as the “2016 IP No Hunting Agreement
Zone” which became the Strict Protection Zone when the Protected Area
Management Plan was formulated, deemed vital for conserving the species in the
mountain (Bonenfant et al. 2023; Schütz
et al. 2023). Although protection against killing by poachers and traditional
hunting from indigenous communities was enhanced, the confinement of this
endemic mammal has led to several problems; one of which is the increase in
hunting pressure, especially at the border of the strict protection zone where
residing indigenous communities are allowed to set traditional traps (Bonenfant et al. 2023). Another challenge for the confined
species is the overgrowth in population, resulting in crowding, slowing the
population growth, as well as forcing a source-sink dynamic due to the limited
habitat range that may be detrimental to future conservation efforts (Bonenfant et al. 2023).
Wallowing is a behavioural adaptation displayed by several mammals including
bisons, pigs, and buffaloes which is done by
submerging their bodies in mud/water puddle to cover a thick coat of mud on
their body (Coopedge & Shaw 2000; Bracke et al. 2011). This behavior is mainly done to
alleviate heat stress, improving the overall well-being in buffaloes. In fact,
wallowing was shown to be the most effective cooling strategy of buffaloes,
significantly increasing skin temperature, milk production, and overall
productivity under heat stress (Aggarwal & Singh 2008; Petrocchi
et al. 2023). Despite its ecological significance to the species, wallowing
behavior remains understudied since most of B. mindorensis
studies have been focused on population estimates, distribution and occurrence
(Ishihara & Kanai 2010; Ishihara et al. 2015; Gil et al. 2021; Bonenfant et al. 2020, 2023), while behavioral studies
remain outdated, and limited (Custodio et al. 1996; Cebrian et al. 2014; Tabaranza et
al. 2022). The only study of the species’ wallowing is primarily described from
a small population of captive Tamaraws (Momongan
& Walde 1993). Little is known about the Tamaraw
wallowing in the wild besides the earlier descriptions (Custodio
et al. 1996; Cebrian et al. 2014; Tabaranza
et al. 2022).
Given these challenges,
understanding behavioral ecology in response to the rapid shift in climate is
critical in preserving natural behaviours and
ensuring the species’ survival. Especially in MIBNP where intense drought
remains a recurring threat to the bovine (Perez et al. 2022). Hence, we report
the wallowing observations from comprehensively monitoring a wallowing site in
the wild through a camera-trap survey conducted in Mts. Iglit-Baco
Natural Park from 2016–2018. The objective of the study is to (i) determine the B. mindorensis
daily and monthly wallowing patterns and (ii) correlate climatological
variables with the wallowing patterns observed. The results of the study will
pioneer a descriptive study for the species’ wallowing behavior in the wild, as
well as highlight its importance to their natural ecology.
MATERIALS AND METHODS
Study site
Here, we report the observed
activities of B. mindorensis from a three-year
comprehensive camera trapping survey (2016–2018) by monitoring a
wallowing site (849 m) within the safe zone of the MIBNP (Images 1B, 1C). Here the study area is the
wallowing site found at the edge of the grasslands and forest within the 16 km²
of SPZ, which contains two mud puddles and is surrounded by shrubs and tall
grasses (Image 2). The camera trap survey utilized two cameras, capturing
opposite sides of the site.
Camera-trap survey
The initial camera-trap survey
was conducted from 04 March 2016 to 24 October 2018 by The World
Wide Fund for Nature (WWF-Philippines) in collaboration with the
Hubbs-SeaWorld Research Institute (HSWRI), the MIBNP- Protected Area Management
Office (MIBNP-PAMO), and Tamaraw Conservation Program (TCP) under the
Department of Natural Resources (DENR), and the Far Eastern University (FEU).
Due to the critical status and limited distribution of the species, the team
opted for targeted sampling in a forest within the no-hunting zone of MIBNP to
understand its behavior.
The survey used two camera models
at two different angles: the Bushnell camera capturing one side of the site,
which recorded the date and time from 12 March 2016 to 30 July 2017. The Reconyx Ultrafire XR6 camera
captured photos from 01 August 2017 to 24 October 2018, which recorded the time
stamp, date, temperature, and moon phase on the opposite side of the site. The
camera was strategically placed in tree trunks, 1.5 m above ground, ideal for
capturing large-sized mammals. Image retrieval and maintenance were conducted
once every month by the WWF-Philippines staff, the TCP & the MIBNP-Park
Area Management Office (PAMO) rangers. The cameras were set on high
sensitivity, which captures three photographs per trigger and a one-second
interval per picture. Photographs were recorded with an image size of 5
megapixels and a 640x480 image resolution.
Image processing
The camera trap data was
organized per year, with subfolders arranged by months to determine the
seasonality of the observations. The photos were tagged in the metadata using DigiKam ver. 8.4.0. To minimize potential errors in
identification, data on the age class & sex were excluded from the image
tagging, and only activity, species wallowing behaviour
& number of individuals per wallowing observations were tagged. The
detection of the images was defined at a 30-minute interval, which is estimated
based on the time spent for each Tamaraw activity before moving away from the
detection radius of the camera. This ensures that independent events are
captured, and it also minimizes potential duplicate counts of individuals and
behavior recorded. The statistical analysis was done through R software version
4.4.1 (Kumar & Tiyagi 2024). The function “recordTable“ of
camtrapR package (Niedballa
et al. 2016) was used to extract the metadata from the images into a CSV file.
The metadata includes the timestamp, the tags, the minDeltaTime, and the camera station.
Climate data
The nearest available
meteorological station of the Philippine Atmospheric, Geophysical, and
Astronomical Services Administration (PAGASA) to MIBNP is located in San Jose
City, south of the island. The elevation, geographic, and topographic
difference between the available station and the study site may reflect
different conditions, potentially affecting the outcome of the analysis. The
climatological data of San Jose, Mindoro station, spanning from 01 January 2016
to 31 December 2018, was requested from The Climate and Agrometeorological Data
Section (CADS) of PAGASA. The three-year data include the minimum, maximum, and
mean temperature, rainfall, humidity, and wind speed.
Statistical analysis
The statistical analysis and
visualization were done through the R software using the reshape2, ggplot2, and
corrplot packages. A correlation heatmap was used to
determine the correlation of wallowing observations with the available climate
data which includes rainfall, relative humidity, max temperature, minimum
temperature, mean temperature, wind speed, and wind direction.
RESULT AND DISCUSSION
Tamaraw ethogram
The survey lasted for 1,096 days,
capturing a total of 9,560 photographs. Using 30-minute intervals, a total of
517 independent wallowing detections were observed. The image tagging revealed
18 different activities exhibited at the wallowing site (Table 1). The
activities were further classified into six distinct categories including (1)
feeding & drinking, (2) movement & navigation, (3) social interactions,
(4) rest & relaxation, (5) wallowing transition, and (6) hygiene
maintenance.
This preliminary ethogram
provides the basic activities that are observed from B. mindorensis
at wallowing sites which should be considered when strategizing conservation
plans for the species. For example, the observed wallowing behavior denotes the
importance of mud holes for their natural ecology; therefore, such landscape
features must be present in their potential expansion ranges to ensure their
survival and overall well-being. Furthermore, the ethogram reports behaviors
that have never been described for the species such as sparring and mock
aggression, which are commonly observed between juveniles, and bulls with their
juveniles.
Wallowing site activity patterns
Photo analysis revealed a
distinct diurnal pattern (Figure 1). Tamaraw wallowing was primarily observed
at 0700–1800 h, with two peak periods at mid-day, at 1000–1200 h, and another at 1500–1700
h. No wallowing was observed at night, contrasting the observations of Momongan & Walde (1993) on
captive B. mindorensis individuals, which
reported wallowing during midnight. This suggests that thermoregulation is the
primary purpose of wallowing and not ectoparasite protection.
The monthly observations reveal
higher counts of wallowing during the longer wet season compared to the dry
season (Figure 2). Wallowing was observed to peak in May, when the dry season
transitions to the wet season, which was further confirmed by the high
correlation between minimum temperatures, and wallowing detections (Figure 3).
The observation suggests that perhaps B. mindorensis
wallowing is induced by the availability of mud puddles, which are limited
when these mud puddles are sometimes dry during the hotter months of January to
early May. Low wallow observations during the dry season may indicate that
these bovines search for other wallowing site instead or rather seek shade in
forest during hotter hours when the mud holes dry up, which are normally
observed behavior of buffaloes (Katwal et al. 2024)
The correlation of climatological
variables detections reveals temperature (minimum temperature, maximum
temperature, and mean temperature) as the primary driver of wallowing. This can
simply validate the importance of wallowing as a thermoregulatory mechanism in B.
mindorensis, inferring that increase in ambient
temperature induces the wallowing behavior as long as the mud puddles are
readily available in their habitat.
Further analysis of individuals
during the Tamaraw wallowing (Figure 4) events reveals a preference for
solitary wallowing. This is in line with the solitary nature of Tamaraws, where
both adult bulls and cows are observed to be lone individuals (Custodio et al. 1996). Occurrences of wallowing with
multiple (3–5) individuals were rare, which may also be attributed to the
agonistic behavior between bulls, resulting in the competition for mud puddles.
However, this behavior was only associated with breeding season where bull
fights are common in order to assert dominance in a herd and displace the
losing individual (Custodio et al. 1996)
CONCLUSION
The study provides a preliminary
descriptive information regarding the wallowing pattern of B. mindorensis on a single wallowing site in MIBNP. It is
highly recommended for future that Philippine Tamaraw behavioral studies be
conducted on multiple sites to extrapolate its general wallowing patterns in
the wild. Nevertheless, the present results highlight the importance of wallowing
on the behavioral ecology of Bubalus mindorensis, particularly wallowing as their primary
thermoregulation during heat stress as suggested by its diurnal pattern. The
seasonal patterns show the importance of mud puddle availability to induce the
behavior, which should be immediately identified and preserved by the park
managers. The preservation of these landscape features should be considered
when expanding their range as well as translocating to ensure their
survivability. The correlation of climate variables further validate
temperature as the primary driver that induces wallowing behaviour
and is a critical consideration for future conservation planning of B. mindorensis.
Table 1. A
constructed ethogram of Bubalus mindorensis observed activities.
|
Activity |
Description |
|
Feeding and
drinking |
|
|
Foraging |
The head is lowered, chewing
food |
|
Drinking |
The head is lowered in front of
a body of water, consuming water |
|
Suckling |
Juvenile feeding from adult
mammary glands |
|
Movement and
navigation |
|
|
Traversing |
The head is straightforward,
traveling across the landscape |
|
Investigating
environment |
Standing, the head is held up
high, either moving left or right |
|
Running |
Traversing rapidly |
|
Social Interactions |
|
|
Mounting |
A male, positioned on top of a
female |
|
Nose-to-nose
touching |
Standing, physical contact
between the noses of two individuals |
|
Sparring |
Two individuals, heads are
lowered, physical contact of horns |
|
Mock aggression |
One or two individuals, heads
are lowered, no physical contact of horns |
|
Rest and relaxation |
|
|
Idling |
Individual/s standing,
stationary |
|
Resting |
Individual/s lying down on the
grass, stationary |
|
Wallowing |
Individual/s submerged/standing
on top of a mud puddle, |
|
Stretching |
Extending limbs or entire body |
|
Wallowing
transitions |
|
|
Enter wallow |
Moving into a mud puddle |
|
Exit wallow |
Moving out of a mud puddle |
|
Hygiene and
Maintenance |
|
|
Scratching |
Rubbing or scraping the body
against a surface or its feet |
|
Urinating |
Releasing urine, usually in a
different stance |
For
figures & images - - click here for full PDF
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