International Journal of Primatology

https://doi.org/10.1007/s10764-024-00425-7

Ranging Behavior of Wild Silvery Lutungs (Trachypithecus

cristatus) in the Coastal Forest of West Sumatra, Indonesia

Muhammad Azhari Akbar1,2 · Dyah Perwitasari‑Farajallah1,3 · Rizaldi4 ·

Ani Mardiastuti5 · Muhammad Ikhsan6 · Thoriq Alfath Febriamansyah4 ·
Kanthi Arum Widayati1 · Yamato Tsuji7

Received: 29 August 2023 / Accepted: 22 February 2024

© The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature
2024

Abstract
Ranging behavior, including home range use and daily travel distance, provides
valuable information on the behavioral responses of primates to their habitats. We
evaluated the ranging behavior of wild silvery lutungs (Trachypithecus cristatus)
inhabiting a coastal forest in West Sumatra, Indonesia, for 614 h over 15 months
in 2018–2021, focusing on the relationships between ranging behavior and activity
budgets, dietary composition, and food availability. The annual home range was 8.1
ha (minimum convex polygon) and 11.7 ha (95% kernel), and the mean (± stand-
ard deviation [SD]) daily travel distance was 926 ± 385 m. Daily travel distance
showed a positive correlation with percent resting and negative correlations with
percent feeding and moving. Furthermore, daily travel distance was correlated posi-
tively with percent young leaf feeding and negatively with percent mature leaf feed-
ing, ripe fruit feeding, and dietary diversity. In contrast, home range size did not
vary significantly across the study period, and we found few significant correlations
between home range size and dietary composition or activity budget. The longer
daily travel distances in food scarce season are likely due to the lutungs needing
to travel further to find trees bearing young leaves. We conclude that the silvery
lutungs’ ranging behavior is linked to their high-risk, high-return strategy.

Keywords Daily travel distance · Gunung Padang · Home range size · Silvery
langur · Variation

Handling Editor: Joanna (Jo) M. Setchell

Extended author information available on the last page of the article

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Vol.:(0123456789)
 M. A. Akbar et al.

Introduction

Home range use is the result of complex interactions between animal behavior
and the environment that provides the resources (Dugatkin, 2020). Many primates
adjust their ranging patterns in response to temporal changes in food availabil-
ity (Clutton-Brock, 1975; Buzzard, 2006), water resources (Sigg & Stolba, 1981;
Hill, 1999; Scholz & Kappeler, 2004), and shelter (Harrison, 1983). A primate’s
energy expenditure correlates with their ranging behavior, such as daily travel dis-
tance and home range use, and this influences activity budgets, particularly feed-
ing time (Dunbar, 1992), ultimately influencing reproductive success (Cheney et
al., 1988). Understanding ranging patterns and their determinants also is impor-
tant in primate conservation (Robbins & McNeilege, 2003).
Primates respond to food scarcity in different ways. For example, during the
dry season, when there is less food available, some primate species extend their
home ranges or travel farther (e.g., tufted capuchin Cebus apella (Terborgh,
1983); Indo-Chinese gray langurs Trachypithecus crepusculus (Fan et al., 2015);
François’ langurs T. francoisi (Zhou et al., 2007, Li et al., 2014); Annamese sil-
vered langurs T. margarita (González Monge, 2016); black and gold howler mon-
keys Alouatta caraya (Raño et al., 2016)). Other species, in contrast, reduce their
travel distances (e.g., capped langurs Trachypithecus pileatus (Stanford, 1991);
eastern lesser bamboo lemurs Hapalemur griseus (Overdorff et al., 1999); white-
handed gibbons Hylobates lar (Bartlett, 2007)). This implies that during times
of food scarcity, the former species use a high-risk, high-return strategy, and the
latter use a low-risk, low-return strategy (Strier, 1999). The seasonality of food
availability, however, had no bearing on home range sizes or travel distances in
several primate species (e.g., black-and-white colobus monkeys Colobus ango-
lensis and C. guereza (Bocian, 1997); guenons Cercopithecus lhoesti and C. mitis
doggetti (Kaplin, 2001); red leaf monkeys Presbytis rubicunda (Hanya and Ber-
nard, 2016); Ashy red colobus Piliocolobus tephrosceles (Kibaja et al., 2023),
due to their use of fallback foods (Di Fiore, 2003; Grueter et al., 2009; Hanya &
Chapman, 2013). Even within species, home range size and daily travel distances
differ between sites (Zhou et al., 2007, 2011a).
Asian colobines can subsist on foliage, a relatively abundant and nonpatchy-
distributed resource (Tsuji et al., 2013). Their home range size (2–3500 ha) and
travel distance (<200–1300 m) vary. Several studies of Trachypithecus monkeys
have found that home range increases with group size (Carl, 2009; Fan et al.,
2015; González Monge, 2016), implying that intragroup competition affects home
range size (but see Kirkpatric, 2007). Li & Rogers (2006), in contrast, reported
negative correlations between the daily travel distance and percent moving and
percent feeding, which implied that ranging is also affected by characteristics of
food patches in each season.
Silvery lutungs (Trachypithecus cristatus) are an Asian colobine, which is
common in Sumatra, Borneo, Natuna Archipelago, and the Peninsular Malaysia
(Roos et al., 2008). They have the widest distribution among Trachypithecus spe-
cies and can inhabit coastal forests (Groves, 2001; Meijaard & Shekelle, 2006),

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Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

tropical rainforests (O’Brien & Kinnaird, 1997; Groves, 2001), secondary forests
(O’Brien & Kinnaird, 1997; Saj & Sicotte, 2007), karst forests (Whitten et al.,
1987), and protected forests, such as national parks (O’Brien & Kinnaird, 1997).
There have been several ecological studies of silvery lutungs (Furuya, 1961;
Bernstein, 1968; Mohd-Daut et al., 2021). Recent studies have shown that the
lutungs spent most of their times resting (56%) (Tahir et al., 2017) and that provi-
sioning affected home range size of the lutungs (Mohd-Daut et al., 2021).
In this study, we evaluated home range size and daily travel distances of wild sil-
very lutungs and examined the relationship between these ranging data and activity
budget, feeding, and food availability.

Methods

Study Site and Subjects

We conducted fieldwork in Gunung Padang (GP), Kampung Seberang Pebayan,

Batang Arau Sub-district, Padang Selatan District, Padang City, West Sumatra,
Indonesia (100° 20′–100° 21′ E and 0° 57′–0° 58′ S). Gunung Padang is a hilly cape
area along the coast of western Sumatra. The western side of GP is directly bordered
by the Indian Ocean, and the eastern side is bordered by the fishermen’s settlement
areas of Kampung Seberang Pebayan, Batang Arau River, and Padang Timur Dis-
trict. The area of GP is 24.3 ha, and the maximum elevation is 115 m above sea level
(Fig. 1). Gunung Padang is a tourist site managed by Padang City Tourism Agency.
Tourists typically use bike tracks on the outskirts of GP. The northern coast of the
study site has steep slopes, and tourists cannot access it. Gunung Padang is 68%
secondary forest dominated by Arecacae (Cocos nucifera and Arenga obstusifolia)
and Moraceae (Ficus spp.) (Ilham et al., 2018). There also are several agricultural
plantations, such as clove (Syzygium aromaticum) and cacao (Theobroma cacao).
During the study, the mean temperature was 27.1 °C (range 20.5–34.6 °C), and the
monthly mean precipitation was 326.8 mm (range 147.3–562.0 mm) in Padang City
(Badan Pusat Statistik, 2022).
Only one group of silvery lutungs inhabits GP (Akbar et al., 2022). During our
study, the group comprised 25 individuals: one adult male (>5 years, largest body
size), five nursing females (adult females carrying infants), ten other adult females
(>5 years, slightly smaller than the adult male, with irregular white patches on the
inside flanks), four juveniles (1.5–5 years, smaller body size than adults), and five
infants (<1.5 years, carried by the mother, bright yellow or orange pelage, which
is common for the genus Trachypithecus until 3–5 months after birth) (Bernstein,
1968; Harding, 2010; Trisilo et al., 2021).

Ranging Data Collection

We followed the lutungs for 3–15 days per month (Table I). We stayed >10 m away
from the lutungs to avoid disturbing them. We collected ranging data from 07:00 h to

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 M. A. Akbar et al.

Gunung
Padang

Fig. 1  Gunung Padang, West Sumatra, Indonesia.

18:00 h but stopped following if the lutungs moved to the northern coast. Therefore,
the size of the home range might be underestimated. We used a handheld Garmin
GPSMAP 64s to record the group’s location (approximate center of the group) every
10 min. From this data, we calculated home range sizes (minimum convex polygon
[MCP]) and 50% and 95% kernel (Börger et al., 2006). We defined the 50% kernel
as the “core area” and the 95% kernel as the “home range.” We used the open-source
Quantum Geographic Information System (QGIS) 3.16 Hannover (QGIS Develop-
ment Team, 2020) and “adehabitatHR” package in R ver 4.3.0 (R Development Core
Team, 2023) to calculate the home ranges. We determined the daily travel distances
(m) by summing the distances between location points recorded on a given day. We
excluded data collected on days with <30 location points or when we did not cover
the entire day.

Behavioral Data Collection

We recorded the behavior of the lutungs was at 10-min intervals using scan sam-
pling (Martin & Bateson, 1993). In each scan, we spent up to 10 s observing each
visible individual to record its activity. We classified their activities into feed-
ing, moving, resting, social grooming, and others (playing, mating, aggression,
and mothering). We observed each individual once per scan. We calculated the

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Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

Table I  Home range sizes and daily travel distances of Trachypithecus cristatus at Gunung Padang, West
Sumatra, Indonesia from August 2018–July 2019 and September–November 2021
Month, year # Sam- # Location points Home range size (ha) Daily travel
pling days distance (m)

Fixed kernel Minimum con- Mean SD

vex polygon
50% 95%

August 2018 7 185 0.7 2.9 3.1 651 458

September 2018 3 99 0.6 2.8 1.8 751 -
October 2018 4 54 0.7 2.9 5.0 NA NA
November 2018 5 97 1.6 9.0 6.9 433 -
December 2018 4 73 3.1 13.7 5.9 592 -
January 2019 7 188 3.3 11.8 6.2 1080 359
February 2019 5 194 3.3 11.7 6.7 793 364
March 2019 13 436 2.2 10.6 7.5 1057 384
April 2019 15 647 1.2 8.0 7.0 1119 456
May 2019 10 430 1.7 9.4 6.7 1269 718
June 2019 10 313 2.1 10.6 7.5 899 239
July 2019 10 303 1.9 10.0 7.8 919 137
September 2021 5 146 1.3 6.8 6.2 748 623
October 2021 5 213 1.5 7.4 5.6 1150 865
November 2021 7 356 1.8 9.0 6.1 1517 782
Annual range 7.3 239.9 5.6 11.7 8.1 926 385

monthly proportion of each behavior by dividing the number of scans in which

we recorded that behaviour for any animal by the total number of scan samples
for all animals in that month.
When the lutungs were feeding, we recorded the food item. Based on a previ-
ous study of the diets of Asian colobines (Tsuji et al., 2013), we classified the
dietary components into mature leaves, young leaves, ripe fruit and seeds, unripe
fruit and seeds, flowers, and others, including bark, stem, pith, and underground
storage. We calculated the monthly proportion of each food by dividing the num-
ber of scans in which we recorded that food for any animal by the total number of
feeding scans for all animals in that month. Therefore, the calculation was based
on the feeding effort of the animals, rather than the actual food intake.
We used the Shannon-Wiener index (H’) to examine food diversity in a given
month, using the formula:
r
∑
�
H =− [pi × logpi ]
i=1

where pi is proportion of plant part i, and r is the total number of plant parts
observed in the dietary composition. We excluded data for October 2018, in which
we observed only <300 min from analyses.

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 M. A. Akbar et al.

Plant Phenology

In August 2018, we selected 204 forest trees along a hiking path (ca. 1 km in length
and ca. 5 m in width) for a phenological survey. The lutungs eat 88 of these 204
trees (14 species) (Akbar et al., 2019). To examine food availability, we evaluated
the phenology of these 88 food plant trees monthly by checking for the presence or
absence of (1) mature leaves, (2) young leaves, (3) flowers, (4) ripe fruits, and (5)
unripe fruits. We used the percentage of trees in which the target plant parts were
present as the phenology index for that plant part in any given month (Tsuji et al.,
2019).

Data Analysis

To evaluate the ranging patterns of lutungs, we used Spearman’s correlation tests

between ranging (monthly home range size and daily travel distance) and 1) activ-
ity budget (percentages of feeding, moving, resting, grooming, and other activi-
ties), 2) diet (percentages of mature leaves, young leaves, flowers, ripe fruits, and
unripe fruits) and its diversity (H’), and 3) availability of food resources (phenol-
ogy index for mature leaves, young leaves, flowers, ripe fruits, and unripe fruit). We
have reported the activity budgets, feeding percentages, and food availability previ-
ously (Akbar et al., 2021; 2022). We omitted data for September to December 2018
from analyses because we had <100 location points for these months (Table I). We
conducted statistical analyses by using R version 4.3.0 (R Development Core Team,
2023) and set α at 0.05.

Ethical Note

We collected all data from habituated, wild animals without interfering with their
activities. The research complied with the laws of Indonesia, and the protocols were
approved by Indonesian Foreign Research Permit Division, Ministry of Research.
The authors declare that they have no conflict of interest.

Data Availability The dataset on location points of the lutungs is available as Elec-
tronic Supplementary Materials. The dataset on activity budgets, feeding percent-
ages, and food availability is available elsewhere (Akbar et al., 2021; 2022).

Results

We monitored the lutungs on 110 days (614 h) and recorded 3684 location points
(Table I). The mean number of location points per day was 33.5 (range 4–67). We
lost the monkeys on 54 days, but in 46 cases we found them again on the same day.
The cumulative area used curves reached an asymptote after 1000 location points
(Fig. 2), suggesting that our home range estimates are correct. The annual MCP and

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Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

Fig. 2  Cumulative areas used by 12

95%Kernel
lutungs at Gunung Padang, West
Sumatra, Indonesia from August 10
2018–July 2019 and Septem-

Home range area (ha)

ber–November 2021. MCP = 8 MCP
minimum convex polygon.
6
50%Kernel

0
0 500 1000 1500 2000 2500 3000 3500 4000

Number of location points

95% kernel covered approximately 33.7% and 48.1% of GP. The core area of the
lutungs’ home range was located in the northern part of GP (Fig. 3).
We found no significant correlations between home range size and monthly activ-
ity data (Table II). In contrast, we found a significant positive correlation between
daily travel distance and percent resting (Fig. 4a) and significant negative cor-
relations between daily travel distance and percent feeding and percent moving
(Figs. 4b–c; Table II).
The home range size and the composition of their diet were not significantly cor-
related (Table III). In contrast, we found a significant positive correlation between
daily travel distance and the percent young leaf feeding (Fig. 4d) and significant
negative correlations between daily travel distance and the percent mature leaf feed-
ing, percent ripe fruit feeding, and dietary diversity (Figs. 4e–g; Table III).
Finally, we found no significant correlations between ranging and phenology
indices (Table IV), except for negative relationships between home range size and
the phenology index of unripe fruits (Table IV).

Discussion

We found strong negative correlations between daily travel distances and 1) percent
mature leaf feeding, 2) percent ripe fruit feeding (especially Ficus variegata, their
main food source, Akbar et al., 2022), and 3) the diversity index. These negative
correlations suggest that when young leaves are less available, the lutungs, like other
primate species, rely on food items whose distribution is relatively even (Newton,
1992; Grueter et al., 2009; Lambert et al., 2004; Akbar et al., 2022; Hanya & Chap-
man, 2013), and their travel distance became shorter. Since the cumulative area used
curves reached an asymptote after 1000 locations, our sample size was adequate to
evaluate ranging behavior. According to Li and Rogers (2006), the negative correla-
tions between the daily travel distance and percent moving and percent feeding is
attributed to the lutungs’ dependence on alternative foods: they tend to remain in
fewer feeding patches for longer periods of time. Given that digestion of leaves takes

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 M. A. Akbar et al.

August 2018 September 2018 October 2018

November 2018 December 2018 January 2019

February 2019 March 2019 April 2019

Fig. 3  Home ranges (95% kernel) and core areas (50% kernel) of a group of silvery lutungs at Gunung
Padang, West Sumatra, Indonesia from August 2018–July 2019 and September–November 2021.

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Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

May 2019 June 2021 July 2021

September 2021 October 2021 November 2021

Annual range Legend

Fixed Kernel 50%

Fixed Kernel 95%

Fig. 3  (continued)

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Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

Table II  Results of Spearman’s rank correlation tests between ranging behavior (home range size and
daily travel distance) and activity budgets of Trachypithecus cristatus at Gunung Padang, West Sumatra,
Indonesia from August 2018–July 2019 and September–November 2021
Ranging Percent activity
Feeding Moving Resting Grooming Others

Home range size

50% kernel rs 0.027 0.118 0.045 -0.482 0.455
p 0.946 0.734 0.903 0.137 0.163
95% kernel rs −0.073 0.036 0.155 −0.355 0.236
p 0.839 0.924 0.654 0.286 0.486
Minimum convex poly- rs −0.236 −0.155 0.155 0.200 −0.209
gon (MCP)
p 0.486 0.654 0.654 0.558 0.539
Daily travel distance rs −0.736 −0.691 0.782 −0.327 −0.082
p 0.013** 0.023* 0.007** 0.327 0.818

*p < 0.05; **p < 0.01. Others includes playing, aggression, mating, and mothering

1.6
1.4
a) b) c)
1.2
1.0
0.8
0.6
0.4
0.2
0.0
20 30 40 50 60 4 6 8 10 12 14 16 25 30 35 40 45 50
% Resting % Feeding % Moving
Daily travel distance (km)

1.6
1.4
d) e) f)
1.2
1.0
0.8
0.6
0.4
0.2
0.0
50 60 70 80 90 0 5 10 15 0 2 4 6 8 10
% Young leaves % Mature leaves % Ripe fruits

1.6 g)
1.4
1.2
1.0
0.8
0.6
0.4
0.2
0.0
0.0 0.2 0.4 0.6
H'

Fig. 4  Significant correlations between daily travel distances (km) and a) % resting, b) % feeding, c) %
moving, d) % young leaves, e) % mature leaves, f) %ripe fruits, and g) dietary diversity (H’) of a group
of silvery lutungs at Gunung Padang, West Sumatra, Indonesia from August 2018–July 2019 and Sep-
tember–November 2021.

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 M. A. Akbar et al.

Table III  Results of Spearman’s rank correlation tests between ranging behavior (home range size and
daily travel distance), monthly feeding percentages of Trachypithecus cristatus at Gunung Padang, West
Sumatra, Indonesia from August 2018–July 2019 and September–November 2021
Ranging Percent feeding H’
(0.0–0.5)
Mature leaves Young leaves Ripe fruits Unripe fruits Flowers
(0.0–13.8) (58.6–100.0) (0.0–8.1) (0.0–30.0) (0.0–3.2)

Home range
size
50% kernel rs 0.200 −0.009 −0.052 −0.245 0.220 −0.227
p 0.558 0.979 0.881 0.468 0.515 0.503
95% kernel rs 0.182 −0.004 −0.126 −0.136 0.211 −0.273
p 0.595 0.989 0.713 0.694 0.533 0.418
Minimum con- rs 0.355 −0.405 0.056 0.345 0.349 0.164
vex polygon
p 0.286 0.216 0.871 0.299 0.293 0.634
Daily travel rs −0.745 0.769 −0.786 −0.200 −0.147 −0.827
distance
p 0.012* 0.006** 0.004** 0.558 0.667 0.003**

* p < 0.05, ** p < 0.01

Table IV  Results of Spearman’s rank correlation tests between ranging behavior and monthly phenology
indices for Trachypithecus cristatus at Gunung Padang, West Sumatra, Indonesia from August 2018–July
2019 and September–November 2021
Ranging Phenology index (range)
Mature leaves Young leaves Ripe fruits Unripe fruits Flowers
(37.7–38.0) (13.8–21.3) (0.7–4.0) (1.6–7.4) (2.5–8.7)

Home range size

50% kernel rs 0.300 0.173 −0.164 −0.536 −0.255
p 0.371 0.614 0.634 0.094 0.451
95% kernel rs 0.300 0.435 −0.055 −0.636 −0.455
p 0.371 0.264 0.882 0.040* 0.163
Minimum convex polygon rs 0.136 0.300 −0.327 −0.827 −0.491
p 0.694 0.371 0.327 0.003** 0.129
Daily travel distance rs 0.564 −0.109 −0.182 −0.109 −0.445
p 0.076 0.755 0.595 0.755 0.173

*p < 0.05; **p < 0.01

longer time than fruits, the positive correlation between the daily travel distance and
percent resting is thought to be a secondary effect of dependence on leaves (Arse-
neau-Robar et al., 2021). From these results, we conclude that the lutungs’ daily
travel is linked to their high-risk, high-return strategy. A similar strategy has been
reported for proboscis monkeys in Sabah, Malaysia (Matsuda et al., 2013) and may
be true for other lutungs.

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 Table V  Ranging profiles in the genus Trachypithecus. Taxonomy of the species in this table is based on IUCN (2023)
Species Study site Group size Observation Forest type Home range size (ha) Daily travel dis- References
days (hours) tance (m)
50% kernel 95% kernel Minimum
convex
polygon

T. auratus Pangandaran, 14.5 65 (NA) Secondary forest 0.5–1.0 2.2–3.2 5.8–8.8 442–530 Kool (1989)
Indonesia
T. crepusulus Wuliangshan, >80 89 (1738) Deciduous- 176–207 453 446 1011 ± 406 Fan et al. (2015)
China evergreen mixed
forest
T. cristatus Gunung Padang, 25 110 (614) Coastal forest 5.6 11.7 8.2 926 ± 385 This study
Indonesia
T. cristatus Kuala Selangor, 20–51 NA (1302) Coastal forest NA NA 20 350 Bernstein (1968)
Malaysia
T. francoisi Fusui, China 23 74 (NA) Limestone forest NA NA 19 438 ± 127 Zhou et al. (2007)
Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

T. francoisi Nonggang, China 10–12 83 (739) Limestone forest NA NA 69 541 ± 146 Zhou et al. (2011a)
T. johnii Nilgiri Mountains, 16–18 54 (270) Deciduous- 2.3–3.9 5.0–7.1 5.6–8.3 NA Horwich (1972)
India evergreen mixed
forest
T. leucocephalus Fusui, China 6–11 110 (444) Limestone forest NA NA 23.8–33.8 215–970 Zhou et al. (2011b)
T. margarita Veun Sai Siem 6–62 49 (NA) Deciduous- NA NA 221 NA González Monge
Pang, Cambodia evergreen mixed (2016)
forest
T. obscuras Penang Island, 14 157 (NA) Secondary forest NA NA 12.9 NA Leen et al. (2019)
Malaysia
T. phayrei Phu Khieo, Thai- 9–31 877 (NA) Evergreen forest NA 58.0–107.1 NA 940–1107 Carl (2009)
land
T. pileatus Madhupur, India 10–13 70 (NA) Moist deciduous NA NA 21.6 325 ± 91 Stanford (1991)
forest

13
 Table V  (continued)
Species Study site Group size Observation Forest type Home range size (ha) Daily travel dis- References
days (hours) tance (m)
50% kernel 95% kernel Minimum

13
convex
polygon

T. poliocephalus Cat Ba Island, Viet 17–20 180 (549) Limestone forest NA NA 22–50 NA Hendershott et al.
Nam (2018)
T. shortridgei Dulongjiang Val- 7–9 69 (NA) Evergreen forest NA NA 38–88 NA Li et al. (2015)
ley, China
M. A. Akbar et al.
Ranging Behavior of Wild Silvery Lutungs (Trachypithecus…

Silvery lutungs at GP had a smaller home range than other Trachypithecus spe-
cies (Table V). The smaller home range size may be attributed to relatively richer
food productivity in GP or the small group size (25 animals). Several studies of Tra-
chypithecus monkeys have found that home range increases with group size (Carl,
2009; Fan et al., 2015; González Monge, 2016), implying that intragroup competi-
tion affects home range size. The home range size of the lutungs at GP was more sta-
ble than the daily travel distance and showed few correlations with activity, percent
feeding, and food availability. However, we found negative correlations between
home range size and the availability of unripe fruits (mainly Ficus spp., known as a
fallback food in GP (Akbar et al., 2022)). This implies that food resources only con-
strain ranging behaviour during the food scarce season. The home range expansion
in the food scarce season matches that reported in other lutungs (Zhou et al., 2007;
Li et al., 2014; González Monge, 2016), and might be a general trend in lutungs.
Another possible reason for the lutungs’ small home and stable home range use is
the effects of tourism in GP; when tourists came, the lutungs often stopped feeding
and ran into the forest (Akbar, personal observation). The lutungs may be forced to
stay in the northern part of the peninsula where tourists rarely visit.

Conclusions

The silvery lutungs’ ranging behavior is linked to a high-risk, high return strategy,
in which they travel further in food scarce season to find trees bearing young leaves.
In the future, we need to examine the effects of both intrinsic (e.g., group size) and
extrinsic factors (e.g., temperature, food productivity, and characteristics of food
patches) on ranging behavior of lutungs, using both long-term data from GP and
comparative data for other species or other populations, to test the generality of our
findings.
Supplementary Information The online version contains supplementary material available at https://​doi.​
org/​10.​1007/​s10764-​024-​00425-7.

Acknowledgments We thank the local people of Gunung Padang for their cooperation and for providing
accommodation during the fieldwork, the staff of Herbarium ANDA, Andalas University for their assis-
tance with plant identification, Try Surya Harapan and Alitha Mas Juanes for their assistance with rang-
ing data collection at the study site, Vovi Ratna Qori for helping with field data recap from the study site,
Mr. K. Imai for helping with home-range analyses, and a handling editor and two anonymous review-
ers for giving constructive comments and suggestions to draft. This study was financially supported by
the Ministry of Education, Culture, Research, and Technology of Indonesia (PMDSU – Kemdikbudris-
tek), the Cooperative Research Fund of Primate Research Institute, Kyoto University, KAKENHI (Nos.
15H05242 and 16K18619), JSPS Bilateral Open Partnership Joint Research, Future Development Fund-
ing Program of Kyoto University Research Coordination Alliance, and JSPS Core-to-Core Program (A.
Advanced Research Networks).

Author Contributions MAA performed the fieldwork, analysed the data, and wrote the MS. YT devel-
oped methodology, analysed the data, and provided advises. Other authors (AF, MI, KAW, DPF, R, and
AM) provided editorial advises.

Funding Ministry of Education,Culture, Research, and Technology of Indonesia, The Coop-

erative Research Fund of Primate Research Institute, Kyoto University, Japan Society for the

13
 M. A. Akbar et al.

Promotion of Science,15H05242, Yamato Tsuji, 16K18619, Yamato Tsuji, Bilateral Open Partnership
Joint Research,Yamato Tsuji, Future Development Funding Program of Kyoto University Research Coor-
dination Alliance, Yamato Tsuji, Core-to-Core Program (A. Advanced Research Networks), Yamato Tsuji

Declarations
Conflicts of Interest The authors declare that they have no conflict of interest.

Inclusion and Diversity Statement While citing references scientifically relevant for this work, we actively
worked to promote gender balance in our reference list. The author list includes contributors from the loca-
tion where the research was conducted, who participated in study conception, study design, data collection,
analysis, and/or interpretation of the findings.

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Authors and Affiliations

Muhammad Azhari Akbar1,2 · Dyah Perwitasari‑Farajallah1,3 · Rizaldi4 ·

Ani Mardiastuti5 · Muhammad Ikhsan6 · Thoriq Alfath Febriamansyah4 ·
Kanthi Arum Widayati1 · Yamato Tsuji7

* Yamato Tsuji
ytsuji1002@gmail.com
1
Biology Department, Faculty of Mathematics and Natural Sciences, IPB University, Bogor,
West Java, Indonesia
2
Biology Department, Faculty of Forestry and Sciences, Universitas Lancang Kuning,
Pekanbaru, Riau, Indonesia
3
Primate Research Centre, IPB University, Bogor, West Java, Indonesia
4
Biology Department, Faculty of Mathematics and Natural Sciences, Andalas University,
Padang, West Sumatra, Indonesia
5
Conservation of Forest Resources and Ecotourism Department, Faculty of Forestry, IPB
University, Bogor, West Java, Indonesia
6
Yayasan Ekosistem Lestari, Medan, North Sumatra, Indonesia
7
Department of Science and Engineering, Faculty of Biosciences, Ishinomaki Senshu University,
Ishinomaki, Miyagi, Japan

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