Characteristics, location, and usage patterns of resting burrows in Chinese pangolins: Insights from radio-telemetry tracking

Abstract

The Chinese pangolin Manis pentadactyla, a burrowing mammal, relies on burrows for foraging, shelter, and rearing its young. Pangolin burrows are typically classified as either resting (or resident) or feeding burrows. Resting burrows, in particular, are crucial for the pangolin’s survival, offering shelter, protection, and stability for the pangolin, especially during colder months and breeding seasons. Consequently, understanding the characteristics and usage patterns of resting burrows is critical for effective in-situ and ex-situ conservation efforts. This study assessed the density of resting burrows using sign surveys along line transects in a fragmented agricultural landscape. Additionally, radio telemetry was employed to specifically locate and characterize the resting burrows of seven Chinese pangolins (1 male and 6 females), as well as to analyze their usage patterns. The results revealed a burrow density of 110.8 per ha, with no significant variation across different environments. However, Chinese pangolins preferred resting burrows in areas with minimal human disturbance. Over tracking periods ranging from 52 to 327 days, female pangolins used between 12 and 32 resting burrows, while a single male pangolin used 48 resting burrows. Pangolins occupied individual burrows for up to 10 days during colder months and breeding seasons. Notably, resting burrows were frequently shared among individuals. Activity patterns showed that pangolins typically exited their burrows between 6:00 PM and 12:00 AM and entered their burrows between 10:00 PM and 4:00 AM. To conclude the findings:1) burrow count, while useful for identifying pangolin presence, is not a suitable quantitative indicator of population density. 2) Most resting burrows had compacted soil at the entrance, which could be mistaken for old or inactive burrows in other studies. 3) Resting burrows function as permanent structures that are reused across seasons instead of seasonal adaptations. 4) Preserving low-disturbance habitats will be critical for ensuring sufficient resting burrow sites, thereby supporting the long-term viability of pangolin populations. These results provide valuable insights into the usage patterns of resting burrows, thus enriching the understanding of pangolin behavior, ecological requirements and conservation strategies for this endangered species.

Full text

Original Research Article
Characteristics, location, and usage patterns of resting burrows in
Chinese pangolins: Insights from radio-telemetry tracking
Nick Ching-Min Sun
a,b,c,*,1
, Jing-Shiun Lin
a,1
, Kurtis Jai‑Chyi Pei
d,**
a
Institute of Wildlife Conservation, National Pingtung University of Science and Technology, Taiwan
b
Pingtung Rescue Center, National Pingtung University of Science and Technology, Taiwan
c
IUCN SSC Pangolin Specialist Group, Zoological Society of London, London, UK
d
Taiwan Wildlife Society, Taiwan
ARTICLE INFO
Keywords:
Burrow density
Fossorial
Habitat selection
Human disturbance
Manis pentadactyla
Taiwan
ABSTRACT
The Chinese pangolin Manis pentadactyla, a burrowing mammal, relies on burrows for foraging,
shelter, and rearing its young. Pangolin burrows are typically classied as either resting (or
resident) or feeding burrows. Resting burrows, in particular, are crucial for the pangolin’s sur-
vival, offering shelter, protection, and stability for the pangolin, especially during colder months
and breeding seasons. Consequently, understanding the characteristics and usage patterns of
resting burrows is critical for effective in-situ and ex-situ conservation efforts. This study assessed
the density of resting burrows using sign surveys along line transects in a fragmented agricultural
landscape. Additionally, radio telemetry was employed to specically locate and characterize the
resting burrows of seven Chinese pangolins (1 male and 6 females), as well as to analyze their
usage patterns. The results revealed a burrow density of 110.8 per ha, with no signicant vari-
ation across different environments. However, Chinese pangolins preferred resting burrows in
areas with minimal human disturbance. Over tracking periods ranging from 52 to 327 days, fe-
male pangolins used between 12 and 32 resting burrows, while a single male pangolin used 48
resting burrows. Pangolins occupied individual burrows for up to 10 days during colder months
and breeding seasons. Notably, resting burrows were frequently shared among individuals. Ac-
tivity patterns showed that pangolins typically exited their burrows between 6:00 PM and 12:00
AM and entered their burrows between 10:00 PM and 4:00 AM. To conclude the ndings:1)
burrow count, while useful for identifying pangolin presence, is not a suitable quantitative in-
dicator of population density. 2) Most resting burrows had compacted soil at the entrance, which
could be mistaken for old or inactive burrows in other studies. 3) Resting burrows function as
permanent structures that are reused across seasons instead of seasonal adaptations. 4) Preserving
low-disturbance habitats will be critical for ensuring sufcient resting burrow sites, thereby
supporting the long-term viability of pangolin populations. These results provide valuable in-
sights into the usage patterns of resting burrows, thus enriching the understanding of pangolin
behavior, ecological requirements and conservation strategies for this endangered species.
* Corresponding author at: Institute of Wildlife Conservation, National Pingtung University of Science and Technology, Taiwan.
** Corresponding author.
E-mail addresses: pangolin_tw@mail.npust.edu.tw (N.C.-M. Sun), kcjpei@mail.npust.edu.tw (K.J. Pei).
1
These authors contributed equally to this work.
Contents lists available at ScienceDirect
Global Ecology and Conservation
journal homepage: www.elsevier.com/locate/gecco
https://doi.org/10.1016/j.gecco.2025.e03406
Received 30 August 2024; Received in revised form 2 January 2025; Accepted 4 January 2025
Global Ecology and Conservation 57 (2025) e03406
Available online 6 January 2025
2351-9894/© 2025 Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
( http://creativecommons.org/licenses/by-nc-nd/4.0/ ).

1. Introduction
The Chinese pangolin (Manis pentadactyla Linnaeus, 1758) is one of the four extant pangolin species in Asia, distributed across
South Asia, Southeast Asia, and East Asia, including Hainan Island and Taiwan (Corlett, 2007; Wu et al., 2020). As an ecological
engineer, the Chinese pangolin signicantly inuences habitat heterogeneity, impacting the structure and function of forest ecosys-
tems (Sun et al., 2024a). Over recent decades, its population has declined drastically due to poaching for traditional medicines, dietary
supplements, leather products, and decorative items (Challender et al., 2014; Shirley et al., 2023). Human disturbance, climate change,
and habitat fragmentation further endanger Chinese pangolin populations (Gao et al., 2022). Therefore, the Chinese pangolin is
categorized as ‘Critically Endangered’ on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species
(Challender et al., 2019). While Chinese pangolins have been studied more extensively compared to other pangolin species, partic-
ularly in terms of biology, evolution, and captive breeding (Heighton and Gaubert, 2021; Sun et al., 2024b), their life history remains
relatively underexplored, partly due to their solitary, nocturnal, and cryptic behavior (Morin et al., 2020).
Chinese pangolins are fossorial (burrowing) animals, and burrows play a crucial role in their life history by providing shelter,
feeding sites, and protection from extreme weather and predators (Sun et al., 2018, 2021a). Thus, understanding the characteristics
and usage patterns of these burrows is vital for both in-situ and ex-situ conservation efforts, as it can directly inform habitat man-
agement strategies that are critical for the species’ survival. Previous research has examined the physical attributes of pangolin
burrows, such as their depth and structure (Fang and Wang, 1980; Trageser et al., 2017; Wu et al., 2004). Recently, Zhang et al. (2024)
used burrow molding techniques and advanced 3D scanning to precisely reconstruct and analyze burrow structures. As closed
structures, pangolin burrows provide environments with stable temperature and humidity levels throughout the year. During winter,
uctuations in air temperature and humidity inside deep burrows are considerably smaller compared to those observed outside (Bao
et al., 2013). Although most burrows are ground-based (i.e., tunnels excavated into the soil), Chinese pangolins also use hollow tree
trunks and man-made structures, such as cracks in concrete oors, for shelter (Sun et al., 2019a, 2021a). As obligate myrmecophagous
mammals, Chinese pangolins almost exclusively prey on ants and termites, often excavating burrows to access soil-inhabiting prey
(Sun et al., 2020a; Chao et al., 2020). Consequently, pangolin burrows are typically classied as either resting (or resident) or feeding
burrows (Sun et al., 2021b, 2024a; Sabin et al., 2024). However, pangolins may also excavate burrows for multiple reasons. For
instance, they might create a burrow near an ant nest to feed in the burrow before resting (Heath, 1992).
Burrow counts, often used to estimate burrow density, provide insights into habitat selection and preference (Dhami et al., 2023;
Karawita et al., 2018; Sabin et al., 2024; Tamang et al., 2022). Several studies have employed burrow density to estimate population
sizes and usage patterns (Mahmood et al., 2015; Nadeem et al., 2015; Dhami et al., 2023). Some researchers suggest that each active
resting burrow represents an individual Indian pangolin inhabiting the area (Mahmood et al., 2014, 2015; Irshad et al., 2015; Akrim
et al., 2017). However, our current understanding of how pangolins use burrows remains unclear. For instance, feeding and resting
burrows can be further classied as active or inactive. Active burrows are typically identied by the presence of freshly dug soil,
footprints, false walls, or feces near the entrance, while inactive burrows are identied by the presence of dead leaves, spider webs, or
compacted soil near the entrance (Dhami et al., 2023; Sabin et al., 2024; Trageser et al., 2017). Nevertheless, burrow characteristics
that accurately reect pangolin usage remain unvalidated and are difcult to apply (Ichu et al., 2017; Perera et al., 2022). Pangolins
have been observed to use multiple burrows, occasionally staying in a single burrow for several days at a time (Lim and Ng, 2008;
Pietersen et al., 2014; Sun et al., 2018, 2019a, 2021a). However, how usage patterns change over time and across locations (e.g.,
agricultural vs. forested habitats) has yet to be investigated. Similarly, while human-related disturbances are acknowledged as threats,
how these disturbances specically affect burrow usage, daily activity patterns, and overall pangolin behavior is an underexplored
area that warrants deeper investigation.
In Taiwan, an increase in Chinese pangolin populations has been observed in lowland habitats (Sun et al., 2019a, 2023), partic-
ularly in the Coastal Mountain Range in southern Taiwan (Sun et al., 2019a, 2020b). In this study, we used radio telemetry to locate
Chinese pangolin resting burrows and examined trends in burrow usage patterns. Our goals were to: 1) determine burrow density in
the study area, 2) explore the habitat characteristics of resting burrows, 3) examine the physical characteristics of the burrows, 4)
determine how many resting burrows each pangolin used, and 5) investigate burrow usage patterns. To the best of our knowledge, this
is the rst study to exclusively apply ne-scale eld techniques to explore Chinese pangolin burrow usage patterns. Our ndings
provide valuable insights to strengthen conservation strategies for Chinese pangolins by supporting the development of management
practices that mitigate human disturbance, enhance habitat protection, and ensure the long-term viability of pangolin populations.
2. Materials and methods
2.1. Ethical approval
This study monitored pangolin movement patterns from April 2009 to November 2010 using radio telemetry. Ethical approval was
granted by the Laboratory Animal Center of the National Pingtung University of Science and Technology, Taiwan. Pangolin live-
trapping permits (0980129850 and 0991616024) were obtained from the Forestry and Nature Conservation Agency (formerly
Forestry Bureau), Taiwan.
2.2. Study area
The study was conducted in Ruanshan Village, Yanping District, Taitung, Taiwan (22.909, 121.185), located in the southern region
N.C.-M. Sun et al.
Global Ecology and Conservation 57 (2025) e03406
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of the isolated Coastal Mountain Range. This fragmented landscape comprises patches of secondary forests, managed tree plantations
(Acacia confusa, Liquidambar formosana, Fraxinus formosana, Lagerstroemia subcostata, and Cyclobalanopsis glauca), bamboo forests,
grasslands, and agricultural lands (Fig. 1). Primary forests are absent due to human encroachment. The study area covers approxi-
mately 100 ha at elevations between 400 and 600 m above sea level. Ruanshan Village is home to an aboriginal Bunun community and
has a tropical climate, characterized by a hot, wet season from May to October and a cooler, drier season from November to April.
Chinese pangolin density in the study area is estimated at 12.8 pangolins per 100 ha (Pei, 2010).
Based on the classication criteria of the National Land Surveying and Mapping Center, Ministry of the Interior, Taiwan, habitats in
the study area were categorized into medium-disturbance (44 %) and low-disturbance (56 %) habitats based on the degree of human
disturbance (Fig. 1). The low-disturbance habitats, including secondary forests, bamboo forests, and grasslands, experienced minimal
human activity throughout the year. In contrast, the medium-disturbance habitats, which primarily included farmlands for crops and
fruit trees, such as plum trees (Prunus mume), betel nut trees (Areca catechu), and tea oil camellia (Camellia oleifera), saw a marked
increase in human activity during harvesting and pesticide-spraying seasons (March–April). The two habitat types were interspersed
across the landscape.
2.3. Pangolin burrow density estimation
To estimate pangolin burrow density in different environments, four line transects were established within the home ranges of the
seven tracked pangolins in November 2009 (Fig. 1). Each transect was 5 m wide, covering 1.30 ha (0.69 ha of medium-disturbance
habitat and 0.61 ha of low-disturbance habitat). All pangolin burrows within the transects were identied without categorizing
them by function (e.g., resting or feeding) or determining their activity status (active or inactive), and the environment around each
burrow was documented. Burrow density within each transect was calculated as the number of burrows divided by the area of the
habitat type where they were located (i.e., medium-disturbance or low-disturbance habitats). A chi-square goodness-of-t test was
Fig. 1. Locations of Chinese pangolin resting burrows identied through radio-telemetry tracking in the study area, a fragmented landscape in the
Coastal Mountain Range in southern Taiwan, surveyed from April 2009 to November 2010. Dots represent the resting burrow locations of the seven
tracked pangolins. Gray lines represent transect lines, each measuring 2600 m in length, covering a total area of 1.30 ha. Black contour lines indicate
altitude (m). Red lines mark major roads. Green areas represent low-disturbance habitats, including secondary forests, bamboo forests, and
grasslands. Red-shaded areas represent medium-disturbance habitats, comprising plum farms, betel nut farms, and tree plantations.
N.C.-M. Sun et al.
Global Ecology and Conservation 57 (2025) e03406
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performed to assess pangolin habitat preferences for burrow excavation.
2.4. Radio telemetry and resting burrow location
In addition to the line transect survey, radio telemetry was employed to track pangolins and record their resting burrow locations
within an approximate 100-ha study area. This method allowed direct and continuous monitoring of resting burrowing sites. A total of
12 adult pangolins were live-trapped and tracked (See Supplementary Materials 1 for detailed information on the tracked pangolins).
Of these pangolins, one male and six female pangolins were tracked for at least 50 days. Pangolins were tted with either a Telonics
MOD-125 transmitter (53 g) or an ATS R2030 transmitter (24 g), both with active-mode tags that operated in cycles of 16 and 8 h,
respectively. These transmitters accounted for less than 2 % of the pangolins’ body weight. Transmitters were securely attached to the
pangolins’ tails using screws and bolts. Sun et al. (2019b) contains the detailed improved method for the transmitter attachment.
Lighter pangolins received the 24 g transmitters, which have been shown to have no clear impact on mating, nursing, and foraging
behaviors (Sun et al., 2018, 2021), with multiple individuals successfully tracked for over 2 years before the tracker’s battery ran out.
(Sun et al., 2019b). Radio signals were received using a telemetry receiver (TR4; Telonics) and a directional H-antenna (RA-2AK or
RA-23K; Telonics). The pangolins were tracked for sessions lasting 10–20 days to determine the locations of resting burrows. The
location of each pangolin was triangulated every 30 min, starting at 5:00 PM and continuing until each pangolin entered the resting
burrow in the morning. Burrow entry was conrmed by a marked reduction in the transmitter signal strength, followed by discreet
conrmation of the burrow location in the morning.
2.5. Resting burrow characteristics
After a pangolin vacated a burrow, data on habitat type, burrow slope, inner diameter of the burrow entrance (calculated as the
average of horizontal and vertical widths), burrow angularity, and burrow depth were collected. Burrow depth was measured using an
LED light-equipped webcam, which was connected to a computer via a 550-cm-long exible metal wire (refer to Supplementary
Materials 2 and 3 for details on the webcam setup and burrow footage).
2.6. Burrow usage
Seasonal variations in burrow usage were analyzed by comparing: (1) the circadian rhythm of pangolins’ exit and entry times, and
(2) the number of consecutive days for which individuals returned to the same resting burrow. Because insufcient data were retrieved
on pangolin LF6 during the wet season and pangolin LF1 during the dry season, these two pangolins were excluded from the analysis.
Additionally, nursing females (LF3 and LF4) were excluded from the dry season analysis, as they typically use the same burrow for
extended periods during the dry (nursing) season (Sun et al., 2021a). All statistical analyses were performed using PAST software
version 3.14 (Hammer et al., 2001), with the level of signicance set at
α
=0.05.
3. Results
3.1. Pangolin burrow density
A total of 144 pangolin burrows were located across the study area based on the line transect survey. The overall burrow density
was 110.8 per ha, with 98.6 per ha in medium-disturbance habitats and 124.6 per ha in low-disturbance habitats. Statistical analysis
revealed no signicant difference in burrow density between habitat types was not signicant (goodness-of-t
χ
2
=3.0629,
p=0.0814, df =1).
Table 1
Information on the seven tracked Chinese pangolins and their resting burrow usage.
ID Sex
(M =Male,
F =Female)
Body
mass
(kg)
Tracking
duration
(days)
Total number of
resting burrows
located
Total
instances of
resting
burrow use
Number of
resting
burrows shared
with other
tracked
pangolins
Resting burrow use by other tracked pangolins
(Number of burrows shared)
LF1 F 4.4 91 29 37 13 (44.8 %) LM1 (2), LF3 (7), LF4 (3), LF5 (2), LF6 (1)
LF2 F 5.1 52 16 23 1 (6.0 %) LM1(1)
LF3 F 5.6 239 23 52 15 (65.2 %) LM1 (3), LF1 (7), LF4 (7), LF5 (2), LF6 (1)
LF4 F 5.4 327 32 83 16 (50.0 %) LM1 (8), LF1 (3), LF3 (7)
LF5 F 5.3 204 20 41 8 (40.0 %) LM1 (3), LF1 (2), LF3 (2); LF6 (5)
LF6 F 3.4 95 12 17 6 (50.0 %) LM1 (3), LF1 (1), LF3 (1); LF5 (5)
LM1 M 4.8 249 48 71 14 (29.2 %) LF1 (2), LF2 (1), LF3 (3), LF4 (8), LF5 (3), LF6 (3)
N.C.-M. Sun et al.
Global Ecology and Conservation 57 (2025) e03406
4

3.2. Number and locations of resting burrows
Each pangolin was tracked for 52–327 days (Table 1). A total of 137 distinct resting burrows were used 324 times by the tracked
pangolins (Table 1). Female pangolins used between 12 and 32 distinct resting burrows (mean =22 ±7.6), whereas the male pangolin
(LM1) used 48 distinct resting burrows. Pangolins exhibited some preference for specic burrows, which they returned to frequently,
and multiple pangolins occasionally used the same burrows. The proportion of resting burrows shared with other tracked pangolins
ranged from 6 % to 65 % (Table 1). All female pangolins, except LF2, shared burrows with at least two other females, while the male
pangolin shared his burrows with all six females. However, simultaneous use of a burrow by different pangolins was rare, occurring
only ve times—once with a male and a female, and four times with pairs of females.
As illustrated in Fig. 1, the resting burrows were scattered across the study area. Most of the burrows were ground burrows (85.4 %,
n=117), with smaller numbers located in tree root hollows (n=11), under large rocks (n=7), beneath concrete oors (n=2), and in
a leaf pile (n=1) (Fig. 2). Notably, only four resting burrows were located in freshly dug soil, indicating recent excavation or
Fig. 2. Representative images of different types of resting burrows used by Chinese pangolins. (a) Resting burrow with freshly dug soil near the
entrance. (b–e) Resting burrows with compacted soil near the entrance. (e) Resting burrow under a large rock. (f) Resting burrow inside a cracked
rock. (g) Resting burrow under a tree root hollow. (h) Resting burrow under a concrete oor. The white arrow indicates the entrance of each burrow.
N.C.-M. Sun et al.
Global Ecology and Conservation 57 (2025) e03406
5

renovation. Of the 137 distinct resting burrows, habitat types were determined for 111, with 96 burrows (86.5 %) found in low-
disturbance habitats (Table 2), indicating a clear preference for these areas (goodness-of-t
χ
2
=23.193, p<0.001, df =1).
3.3. Burrow characteristics
Due to variations in on-site factors, not all resting burrows were fully measured, resulting in discrepancies in sample size. Of the 93
burrow depth measurements taken, 18 were incomplete because the deepest point could not be reached due to the burrow’s curved
structure. Based on the remaining measurements, the average burrow depth was 201.6 ±94.8 cm (n=75, range =60–505 cm). Most
resting burrows (n=50, 66.7 %) had a depth of 100–250 cm (Fig. 3a), and the average slope of the burrows was 32.2◦±15.6◦
(n=97, range =0◦to 75◦). The majority of the burrows (n=67, 69.1 %) had a slope between 10◦and 40◦, with only 10 % having
slopes exceeding 50◦(Fig. 3b). Moreover, most burrows (n=71, 74.8 %) had an entrance angle ranging between 0◦and −40◦
(Fig. 3c). The average inner diameter of the burrow entrance was 17.1 ±3.2 cm (n=87, range =10.5–27 cm), with the majority of
burrows (n=52, 59.8 %) having an inner burrow entrance diameter of 15–20 cm (Fig. 3d).
3.4. Burrow usage
The pangolins typically exited their burrows before midnight (92.9 % of observation, 299/322; Fig. 4a). The most common exit
time was between 6:00 PM and 12:00 AM (97.2 % of observation, 313/322; Fig. 4a). The entrance time was between 10:00 PM and
4:00 AM (92.3 % of observation, 287/311; Fig. 4a). No signicant difference was observed in pangolin exit time between the dry
(n=168) and wet seasons (n=154;
χ
2
=15.596, p=0.076, df =9), though two peaks were observed in the dry season at 7:00 PM and
10:00 PM (Fig. 4b). In contrast, the pangolins entered their resting burrows signicantly later during the wet season (n=169) than
during the dry season (n=142;
χ
2
=35.741, p<0.001, df =10; Fig. 4c).
Overall, the pangolins (n=6) spent an average of 1.47 ±0.3 days in a burrow during the wet season and 2.97 ±0.5 days during
the dry season (n=3; t (7) =5.7282, p<0.001). Resting burrow usage was shortest from May to August (1–3 days for over 90 % of
usage events; Fig. 5). However, from November to February, the pangolins were more likely to return to the same burrow on the same
evening (about 40 % of usage events). Longer usage patterns, exceeding 10 days, were recorded in the periods from November to
December (28 %) and March to April (42 %; Fig. 5).
Many resting burrows (39.1–89.1 %) were used only once throughout the study period (Fig. 6), even by pangolins monitored for
extended durations (e.g., pangolins LF3, LF4, LF5, and LM1). However, some pangolins exhibited a clear preference for specic
burrows, with certain burrows being used seven times or more during the study period, particularly by pangolins LF3, LF4, and LF5.
4. Discussion
4.1. Habitat selection for resting burrows
Overall, the density of Chinese pangolin resting burrows in the study area (110.8 burrows per ha) is higher than that reported in a
previous survey in northern Taiwan (56.7 burrows per ha) (Fan, 2005). Notably, both densities are considerably higher than those
reported in other countries. For example, studies in Nepal reported signicantly lower burrow densities: only 8 burrows per ha in Kavre
(Suwal, 2011), 1.04 active burrows per ha in Gorkha and Palungtar (Dhami et al., 2023), and 0.83 active burrows per ha in Shivapuri
Nagarjun National Park (Bhandari and Chalise, 2014). The higher burrow density observed in Taiwan compared to other countries
may be attributable to the larger pangolin population in Taiwan, consistent with their current population status worldwide (Sun et al.,
2019a). Nevertheless, these discrepancies should be interpreted qualitatively, as the number of burrows identied within an area does
not necessarily correlate with habitat usage or population density, which remains unconrmed. For instance, the number of feeding
burrows may be inuenced more by the availability and seasonality of underground food resources than by the population density of
pangolins. Therefore, burrow count, while useful for identifying pangolin presence, is not a suitable quantitative indicator of popu-
lation density.
Our ndings indicate that Chinese pangolins do not exhibit a strong environment preference when digging burrows as the overall
burrow density between habitat types was no signicant difference, implying that medium-disturbance habitats also play an important
role. However, as most resting burrows were located in low-disturbance habitat over the 100-ha study area, suggesting most burrows in
the medium-disturbance habitat were were likely feeding burrows. Chinese pangolins are known to effectively use agricultural elds
and tolerate moderate disturbance levels (Sun, 2023; Suwal et al., 2020). Agricultural elds, with their increased abundance of
Table 2
Number and proportion of Chinese pangolin resting burrows (n=111) across different habitat types.
Medium-disturbance habitats (44 % of the study area) Low-disturbance habitats (56 % of the study area)
Plum farm Betel nut farm Tree plantation Bamboo forest Secondary forest Grassland
Number of
resting burrows
8 6 1 36 57 3
Proportion 7.2 % 5.4 % 0.9 % 32.4 % 51.4 % 2.7 %
Sum 13.5 % 86.5 %
N.C.-M. Sun et al.
Global Ecology and Conservation 57 (2025) e03406
6

termites and ants (Richer et al., 1997), can provide valuable foraging resources. Therefore, maintaining agricultural landscapes like
organic orchards and eco-friendly tea gardens could support pangolins by offering alternative foraging habitats (Tseng et al., 2021).
For instance, Gurung (1996) observed that farmers encountered Chinese pangolins more often in agricultural elds during July and
August, at a time when prey was abundant. This suggests that seasonality and human disturbances may also inuence the distribution
of feeding burrows. Given that resting burrows are typically found in low-disturbance habitats (Table 2), preserving secondary and
bamboo forests adjacent to agricultural elds would ensure the availability of safe habitats for resting. However, our results are based
on a small study area and a limited sample size, which may not adequately capture patterns across the species’ broader range.
4.2. Physical characteristics of resting burrows
Resting burrows exhibited a variety of physical characteristics. Most had compacted soil at the entrance (Fig. 2), which could have
been mistaken for old or inactive burrows in other studies (Dhami et al., 2023; Sabin et al., 2024). In this study, only four resting
burrows showed signs of freshly dug soil at their entrance. This suggests that many of the “active” burrows identied in previous
research are likely feeding burrows rather than resting ones. Therefore, burrow count should not be used as a quantitative indicator of
population density. Moreover, we observed signicant variation in the depth and diameter of resting burrows. In some cases, adult
pangolins used burrows with shallow depths (e.g., 62 cm) and narrow diameters (e.g., 10.5 cm). This challenges previous assumptions
that burrow size could reliably differentiate between feeding and resting burrows (Wu et al., 2003; Mahmood et al., 2013). While
previous studies categorized resting burrows into seasonal types—summer and winter—with winter burrows being signicantly longer
(70–964 cm vs. 22–89 cm) (Fang and Wang, 1980; Wu et al., 2002, 2004), our ndings reveal that resting burrows function as per-
manent structures reused across seasons rather than seasonal adaptations. Additionally, the slopes of the resting burrows observed in
the present study (32.2◦±15.6◦) align with values reported in previous studies (Wu et al., 2003; Suwal et al., 2020; Dhami et al.,
Fig. 3. Resting burrow characteristics of Chinese pangolins. (a) Depths, (b) slopes, (c) entrance angles, and (d) inner entrance diameters.
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Fig. 4. Circadian rhythm of resting burrow usage in Chinese pangolins. (a) Entry and exit times across the whole year. (b, c) Entry and exit times
during each season.
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2023). These ndings suggest that resting burrows are more complex and variable than previously assumed, and highlight the need for
a more nuanced approach in both identifying burrows and developing conservation strategies for Chinese pangolins
4.3. Number of resting burrows used by each pangolin
Our data indicate that each pangolin used multiple resting burrows, with the number of newly found burrows decreasing over time,
suggesting a limited availability of burrows in the study area. The approximate number of resting burrows used by each pangolin was
likely underestimated, as additional resting burrows were discovered toward the end of the study period. Pangolin LF4, which was
tracked over an extended period, exhibited a high frequency of burrow usage (Table 1), making it a reliable representative of burrow
usage in female pangolins. Sun et al. (2021a) reported that the distribution area of resting burrows for Chinese pangolins aligns closely
with their home range. In our study, the home range of the male pangolin LM1 was three to four times larger than that of a female
pangolin (Lin, 2011). Also, most of LM1’s resting burrows were newly found during the study period, indicating that the actual number
of resting burrows used by the male pangolin LM1 is likely much higher than the 48 recorded.
Chinese pangolins are polygynous (Sun et al., 2020b), which could explain why the male pangolin LM1 shared multiple resting
burrows with all six females. However, this study only provides burrow usage data from one male pangolin, limiting our ability to draw
conclusions about burrow usage patterns in male pangolins. For instance, burrow usage patterns of roving males may differ from those
of territorial males, which could inuence the number of resting burrows. The availability of suitable habitats is a key factor that may
limit the number of resting burrows, ultimately affecting pangolin density in the area. Additionally, as resting burrows are typically
used by female pangolins for nursing during the breeding season (Sun et al., 2021a), preserving low-disturbance habitats will be
critical for ensuring sufcient resting burrow sites, thereby supporting the long-term viability of pangolin populations.
4.4. Duration and frequency of resting burrow usage
Chinese pangolins typically exit their burrows before midnight, and in our study area, human activity was relatively low,
particularly at night. However, in Taiwan, Chinese pangolins are sometimes found in rural areas with higher human activity, such as in
villages with extensive road networks (Sun et al., 2019a). Therefore, further research is required to determine how the degree of
disturbance in different habitats affects pangolin activity patterns. Chinese pangolins are generally more active outside their burrows
during the wet season or summer, when high temperatures and humidity drive increased activity among ants and termites (Levings and
Windsor, 1984; Nondillo et al., 2014). These ndings suggest that Chinese pangolins may spend more time foraging than resting during
these periods.
Our ndings revealed notable seasonal variation in burrow usage patterns. During the wet season or summer (May to October),
pangolins changed their resting burrows almost every day. In contrast, during the dry season (November to April), they often remained
in the same burrow for an extended period, sometimes exceeding 10 days. Given the decrease in available food resources during the
winter months (Sun et al., 2020a), this behavioral adaptation—where pangolins remain in the same burrow for extended
Fig. 5. Burrow usage periods. Data were exclusively collected from nonbreeding pangolins. Percentages represent the number of consecutive days
for which a resting burrow was used.
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periods—may help minimize energy expenditure, similar to patterns observed in larger myrmecophagous mammals (McNab, 1984).
Chinese pangolins exhibit reduced metabolic rates and lower activity levels in winter (Khatri-Chhetri et al., 2015; Lin, 2011), sup-
porting the hypothesis that they adjust their behavior to conserve energy during this period of decreased food availability.
Matthews et al. (2023) suggested that focusing camera traps on specic burrows can enhance the detectability of giant pangolins
Fig. 6. Frequency of resting burrow usage in Chinese pangolins (Female LF1–LF6, Male LM1).
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(Smutsia gigantea). In our study, we further observed variability in burrow usage frequency, which highlights the need for careful
verication of detection probabilities in camera trap surveys of resting burrows (Willcox et al., 2019). We also hypothesize that factors
such as competition, attraction, or avoidance between males and females inuence the frequency with which burrows are used,
potentially causing different individuals to use the same burrow with varying frequency. Therefore, resting burrows that are infre-
quently used by one pangolin may be frequented more often by others. To better understand these dynamics, future research should
focus on monitoring individual burrows to track usage by all residents. Additionally, as the pangolins tracked in our study were all
adults with established home ranges (Lin, 2011), it is possible that certain resting burrows served as potential nursing sites. Investi-
gating how subadult pangolins use these burrows could yield valuable insights into the species’ broader ecological and behavioral
patterns. For animal translocation, our ndings can also provide an important reference for burrow usage monitoring after release.
5. Conclusions
Feeding and resting burrows of Chinese pangolins are often difcult to distinguish through direct observation, even by experienced
researchers or hunters. This study reveals distinct patterns in the characteristics, number, and usage of resting burrows by Chinese
pangolins in a fragmented landscape. Specically, it validates the usage patterns of resting burrows and highlights how these patterns
may change over time and vary by location. However, this study has certain limitations. The relatively small study area (approximately
100 ha) may not be representative of broader landscapes, and the limited number of tracked individuals and tracking duration might
restrict the generalizability of the ndings. Furthermore, the data collection methods employed in this study may have introduced
potential biases of single burrow usage frequency, such as competition, attraction, or avoidance between individuals. To draw more
comprehensive conclusions, future research should focus on exploring the differences in structural characteristics and distribution
patterns of feeding and resting burrows, the number of resting burrows used by each pangolin, and individual burrow usage by all
residents. Additionally, incorporating more study sites would provide more robust insights into pangolin burrow usage and contribute
to more effective conservation strategies.
Declaration of Competing Interest
The authors declare no competing interests.
Acknowledgments
The authors thank Jia-Hui Lin, Man-Rong Yu, Xiang-Hua Hu, and Ya-Wen Leu for their invaluable support throughout the extensive
eldwork. They are also grateful to the Pangolin Conservation Association, Taitung County Government, Taitung County Police Bu-
reau (Ruanshan Station), and the Taitung Forest District Ofce for their assistance with logistics. Finally, the authors thank the Forestry
and Nature Conservation Agency, Taitung Branch, for their nancial support (grant no. 104-737-1).
Appendix A. Supporting information
Supplementary data associated with this article can be found in the online version at doi:10.1016/j.gecco.2025.e03406.
Data availability
No data was used for the research described in the article.
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