Mammalian Biology 98 (2019) 80–90

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Mammalian Biology
journal homepage: www.elsevier.com/locate/mambio

Original investigation

Dietary habits of wild Javan lutungs (Trachypithecus auratus) in a

secondary-plantation mixed forest: Effects of vegetation composition
and phenology
Yamato Tsuji a,∗ , Masazumi Mitani b , Kanthi Arum Widayati c , Bambang Suryobroto c ,
Kunio Watanabe a
a
Primate Research Institute, Kyoto University, Inuyama, Aichi, 4840094, Japan
b
Institute of Natural and Environmental Sciences, University of Hyogo, Sanda, Hyogo, 6691546, Japan
c
Department of Biology, Bogor Agricultural University, Bogor, West Java, 16680, Indonesia

a r t i c l e i n f o a b s t r a c t

Article history: This study investigated the seasonal dietary habits of wild Javan lutungs (Trachypithecus auratus) inhab-
Received 12 December 2018 iting the secondary-plantation mixed forest of Pangandaran Nature Reserve, West Java, Indonesia, in
Accepted 8 August 2019 relation to vegetation composition and phenology. We conducted behavioral observations of a habitu-
Available online 9 August 2019
ated group for 16 non-consecutive months. The lutungs fed on 164 items (leaf, fruit, flower, etc.) from 85
Handled by Marcus Clauss different plant species. Number of main plant species (>1% usage) was 20, and accumulated percentage of
feeding (all plant parts combined) for the top five, 10, and 20 plant species was 43.4%, 63.8%, and 81.2%,
Keywords: respectively. The percentage of feeding on each plant species was positively correlated with the total
Colobines number of trees and total crown volume. However, selectivity varied; for instance, plant species with a
Diet lower crown volume were preferred. The lutungs mainly fed on young leaves (average over the study
Feeding ecology period, 69.9%), with fruits (both mature and young, 21.2%) contributing more in certain months. The
Indonesia higher percentage of folivory was similar to that found in previous studies for this species and other Tra-
Vegetation chypithecus species. The category-based analyses revealed a significant positive correlation between the
percentage of young fruit and their availability, whereas most other relationships were not significant.
The percentage of feeding on young leaves showed a significant negative correlation with the percentage
of feeding on mature leaves and dietary diversity, whereas the percentage of feeding on mature leaves
had a positive correlation with that of young fruits. Species-based analyses showed that the percentage
of feeding on three main diet items had a significant positive relationship with their availability. Our
results imply that the dietary composition of the lutungs in our study site is determined by vegetation
composition, while the seasonal change in diet is characterized by the availability of main food parts
(young leaves and young fruits).
© 2019 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.

Introduction (especially frugivorous and folivorous) animals must respond to

seasonal fluctuations in the food environment by changing their
Temperature, rainfall, and plant phenology vary seasonally, feeding strategies, including dietary composition and selection of
which in turn, influence the availability of food resources main diets (defined as diets regularly consumed by a population
(Hemingway and Bynum, 2005). Animals must satisfy their nutri- and from which animals obtain a significant proportion of their
tional requirements. For instance, the foraging success of females energy requirements) (e.g., Banack, 1998; Rosalino et al., 2003; de
might ultimately affect population parameters, such as adult mor- Camargo et al., 2014). In primate research, the effects of vegetation
tality (Wrangham, 1981; Cheney et al., 1988), inter-birth interval composition and phenology, proxies of food abundance, on ranging
(Frank, 1986), birth rate (Bulger and Hamilton, 1987), and infant patterns, diet choice and their seasonal changes have been reported
mortality (Borries et al., 1991) via intra-group competition. Thus, frequently (Davies and Bennett, 1998; Agetsuma and Noma, 1995;
Brugiere et al., 2002).
Colobine monkeys (known as “leaf monkeys”) are a large group
∗ Corresponding author. of primates that are widely distributed in Asia and Africa (Fashing,
E-mail address: [email protected] (Y. Tsuji). 2011; Kirkpatrick, 2011). They have large, specialized stomachs

https://doi.org/10.1016/j.mambio.2019.08.001
1616-5047/© 2019 Deutsche Gesellschaft für Säugetierkunde. Published by Elsevier GmbH. All rights reserved.
 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90 81

Fig. 1. Location of Pangandaran Nature Reserve (top) and study area (bottom). Area surrounded by a dashed line represents the forest park (Taman Wisata Alam), which is
located to the north of the nature reserve (Cagar Alam). The location and size of the home range of lutungs (Trachypithecus auratus) (K group) is shown in gray.

that contain diverse microorganisms (Lambert, 1998). Initially, the prise non-leaves. Thus, dietary habits of Javan lutungs and their
morphological and physiological characteristics of colobines was seasonal changes are expected to be affected by forest character-
considered as an adaptive trait to increase efficiency in processing istics. We specifically tested two predictions: (1) the composition
fibrous foods, such as foliage (Caton, 1999). However, colobines of the lutungs’ diet is determined by vegetation composition, with
feed on large quantities of other items (such as fruit and seed) lutungs feeding on plant species in proportion to their abundance;
(Sayers, 2013; Tsuji et al., 2013a), and therefore the classical view and (2) the seasonal changes in dietary composition and dietary
of colobines to be leaf specialist must be reconsidered (Sayers, diversity are influenced by the availability of the main diet food
2013). Furthermore, recent studies have shown finer-scale dietary items and species, rather than total food availability.
preferences by colobines, with individuals preferring several plant
species, regardless of tree density (Grueter et al., 2009; Zhou et al.,
Material and methods
2009). Fig fruits (Ficus spp.) and lichen, for example, are used as
“fallback foods” (defined as foods whose consumption correlates
Research protocols
negatively with their availability; Marshall et al., 2009) by red
leaf monkeys (Presbytis rubicunda) and Yunnan snub nosed mon-
Our field research adhered to the legal requirements of
keys (Rhinopithecus bieti) (Grueter et al., 2009; Hanya and Bernard,
Indonesia and Japan.
2012). Thus, the dietary composition of colobines and seasonal vari-
ations in the dietary composition are not simply determined by the
availability of leaves. In addition, the feeding strategy of colobines Study site
should be considered to be equivalent to that of more frugivorous
primate species (such as Cercopithecinae [macaques and baboons] Pangandaran Nature Reserve (PNR) is located on the southern
and great apes), for which the main diet influences their feeding coast of West Java, Indonesia (7◦ 43 S, 108◦ 40 E) (Sumardja and
and ranging behavior (Remis et al., 2001; Tsuji and Takatsuki, 2009; Kartawinata, 1977; Tsuji et al., 2013b, Fig. 1). PNR is divided into
Owens et al., 2015). To understand the feeding strategy of colobines, two sections; namely, a 38-ha forest park (Taman Wisata Alam)
we need to evaluate dietary selection at a finer, plant species-based and a 370-ha nature reserve (Cagar Alam) (Mitani et al., 2009; Tsuji
scale. et al., 2015). PNR is situated at an average elevation of <100 m
In this study, we examine whether vegetation composition and above sea level (Mitani et al., 2009). It has an average annual rain-
phenology of main food items influence the dietary habit, dietary fall (1990–2010) of 2940 mm (Rosleine and Suzuki, 2012), and air
diversity, and seasonal change in dietary composition in wild Javan temperature and humidity range between 25–30 ◦ C and 85–95%,
lutungs (Trachypithecus auratus). Several ecological studies have respectively (Brotoisworo, 1991; Kool, 1993). The main vegeta-
been conducted on this species (Kool, 1993; Beckworth, 1995; Vogt, tion of the forest park is secondary forest; however, there are
2003) and it has been shown that their main diets vary among study also several areas of teak Tectona grandis and mahogany Swietenia
sites with different forest types, and that >40% of their diets com- macrophylla plantation (planted during the 1930s–1950s for wood
82 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90

Table 1
Observation time and results of scanning for a group of Javan lutungs in Pangandaran Nature Reserve, West Java, Indonesia.

Year and Month #Observation day Observation time (hr) #Scan #scanned animals Mean scanned animal

2012
April 10 98.8 539 5661 10.5
October 6 70.0 420 4630 11.0
November 9 86.8 521 4933 9.5
December 4 39.0 234 2104 9.0

2013
June 2 20.3 122 771 6.3
July 5 55.0 330 3678 11.1
August 4 45.0 270 3007 11.1
September 5 55.7 334 4152 12.4
October 5 52.3 314 3713 11.8
November 4 24.7 148 1819 12.3
December 2 19.2 115 130 1.1

2014
January 1 11.0 66 861 13.0
February 1 11.3 68 862 12.7
March 1 10.8 65 761 11.7
April 1 11.3 68 1378 20.3
May 1 11.2 67 934 13.9
Total 61 622.4 3681 39394 10.7

collection) (Kool, 1993; Mitani et al., 2009; Rosleine and Suzuki, In this study, we defined that plant species (part combined) >1%
2012). usage and plant parts >1% usage over the study period as the main
food plant species and main diet items, respectively.
Behavioral data collection
Vegetation composition
Of the six groups of Javan lutungs that inhabit the forest park,
we selected one group (K group) that had a home range located To assess the abundance of potential resources for the target
at northern part of the forest park. In December 2012, K group lutung group, we used a part of our previous data (Tsuji et al.,
was composed of 27 individuals (one adult male, 11 adult females, 2015) and conducted a follow-up vegetation survey in 2016. We
nine juveniles, and six infants; age class was defined based on divided the home range of K group (11.6 ha, Fig. 1) into 20 m × 20-
Brotoisworo, 1991). We observed their dietary habits for 16 non- m quadrats (n = 291) and recorded the location of all tree (>5 m in
consecutive months (April, October to December 2012, June to height) in all quadrats using a handheld GPS receiver. Shrub and
December 2013, and January to May 2014), with a total observation liana species were not included in this survey. For each tall tree,
time of 622.4 h over 61 days (Table 1). Because the individuals in the we recorded species identity, as well as the tree height (TH) and
groups were habituated to the presence of tourists (Brotoisworo, height of lowest branch (TB) (to 0.1 m) by using a handheld laser
1991), we were able to observe them at a close distance (<10 m). rangefinder (TruPulse200; Laser Technology, Inc., Centennial, CO,
The home range size of the K group was calculated by recording USA). In the case of strangler figs, we measured the size of the host
GPS locations on a handheld device (GPS MapCSx; Garmin, Ltd., trees. We estimated the maximum crown diameter (CD) by eye
Schaffhausen, Switzerland, recorded every 10 min), and was calcu- and categorized it into seven classes: (1) 0–4.9 m, (2) 5.0–9.9 m, (3)
lated as 11.6 ha. The boundaries of the home ranges of neighboring 10.0–14.9 m, (4) 15.0–19.9 m, (5) 20.0–24.9 m, (6) 25.0–29.9 m, and
groups were clear, because each group rarely entered the home (7) ≥30.0 m. We then estimated the crown volume (CV, m3 ) of each
range of the other groups. tall tree by using the following formula:
We recorded the activities of visible animals (except infants) by
using 5-min scan sampling at 10-min intervals (total scans: 3681, CV = (TH–TB)×(CD/2)2 ×
mean number of scanned animals: 10.7). The activities of the ani-
For convenience, we used representative CD values for each
mals were classified into feeding, moving, resting, social grooming,
class: (1) 2.5, (2) 5, (3) 10, (4) 15, (5) 20, (6) 25, and (7) 30 m, and
and other behaviors (such as alarm calling, playing, and copula-
summed the CV values for each species within the home range of
tion). When feeding was observed during the scan, we also recorded
the target group. Because we obtained a significant positive cor-
the plant species and part eaten, which we categorized into bud,
relation between the CV of 685 trees (these trees were marked
bark, leaves (further divided into mature, young, and petiole), fruits
for a phenological survey of trees) and their diameter at breast
(mature and young), seeds (mature and young), flowers (including
height (DBH), known as a proxy for food abundance (Chapman
flower buds), and others (such as insect and soil). When an indi-
et al., 1992)) (rs = 0.770, p < 0.001), our CV appeared to represent
vidual fed on two or more parts during a single scan, we recorded
food abundance.
only the first one. From these data, we obtained: (1) the feeding per-
To evaluate the selectivity of plants by lutungs, we calculated
centage as: number of scanned animals feeding on target species (or
the Jacob’s selection index (D), with values of 0, 1, and −1 corre-
plant parts: R types)/total number of feeding animals × 100; and (2)
sponding to indifference, preference, and avoidance, respectively
dietary diversity (Shannon-Wiener index, H’), given by following
(Jacobs, 1974).
formula:


R D= (r i –pi )/(r i +pi –2r i pi )

H =− [pi × logpi ]
Where, ri and pi represent proportion of annual feeding (across any
i=1
parts) and proportion of crown volume of species i inside the home
Where, pi is the feeding proportion of the plant part i. range, respectively.
 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90 83

Fig. 2. Relationship between the crown volume (m3 ) of plant food species and their Jacobs’s selectivity index (D). Each circle represents each plant species (n = 62). The white
circles represent main food plant species for which annual feeding percentages are >1% (n = 17). Correlation lines (black line: all species, dashed line: only for main food plant
species) and statistics are also shown.

Plant phenology correlation tests between the phenology index of the plant part
eaten and monthly percentage of feeding on a target plant part and
Before this study, we had established monitoring trees along a dietary diversity index (H’). Second, for the species-based analy-
pre-existing forest path (ca. 2.5 km in length and ca. 5 m in width) ses, we conducted corresponding analyses between the phenology
inside the forest park (N = 675) (Tsuji et al., 2015). Furthermore, in index of the main diet items and monthly percentage of feeding on
July 2013 we added 10 Ficus spp. trees inside the home range of K the target items. If no trees were monitored of a given plant species,
group. The phenology of the labeled trees was evaluated by exam- we excluded this species from the analyses. For these analyses, we
ining each plant for the presence or absence of: (1) mature leaves, again omitted data for months with shorter observation periods (<3
(2) young leaves, (3) flowers, (4) mature fruits, and (5) young fruits days).
twice a month. We calculated two different phenology indices: the We conducted all statistical analyses in R version 3.2.3 (R
one across species and another for specific plant species. The pro- Development Core Team, 2015). The significance level was set to
portion of trees on which the respective plant part was present 0.05.
represented the phenology index for that plant part at a given
time. For our analyses, we averaged the phenology index for each Results
month. For Cynometra ramiflora, Vitex pubescens, Sterculia coccinea,
and Ficus sumatrana, for which there was only a small number of Diet composition
monitored trees inside the home range of K group, we added several
trees outside the home range (and inside the forest park). Over the course of the study, lutungs fed on 85 plant species, of
which most were trees (77), followed by shrubs (4) and lianas (4)
Data analysis (Table 2). However, number of main plant species was 19 (Table 2).
Among them, five species, namely, Vitex pubescens (11.2%), Ficus
To test the relationship between plant abundance and feeding by benjamina (10.2%), Tectona grandis (8.2%), Cynometra ramiflora
lutungs, we conducted Spearman’s rank correlation tests between (8.0%), Pterospermum javanicum (5.9%), and Swietenia macrophylla
the percent of annual feeding (devoted to the consumption of any (5.1%) showed >5% annual feeding (Table 2). The accumulated
parts of target species) of the K group and (1) tree number and (2) percentage of feeding for top five, 10, and 20 plant species was
crown volume of the target plant species within the home range. 43.4%, 63.8%, and 81.2%, respectively. Notably, lutungs fed on two
We also conducted correlation tests between the CV and Jacobs’s plantation species: young leaves and petioles of T. grandis and S.
selectivity index (D) to examine whether vegetation composition macrophylla (Table 2). The diet of lutungs encompassed 164 differ-
affects diet selectivity. Finally, we conducted other correlation tests ent diet items, most of which were leaves and their parts (young
to examine how the percent of feeding of monkeys for specific leaves of 78 species, mature leaves of 10 species, petioles of five
plant parts affect the percent of feeding on other food types or species, and buds of one species) (Table 2). Finally, number of the
dietary diversity. For this analysis, we omitted data for months with main diet items was 22 (Table 2).
shorter observation periods (<3 days) to ensure the reliability of the
analysis. Relationship between plant abundance and feeding
To examine how lutungs respond to fluctuations in the availabil-
ity of food resources, we conducted two types of analyses. First, for Significant positive correlations between the annual feeding
the plant-part category-based analyses, we conducted Spearman’s proportion of woody plants and their crown volume (rs = 0.477,
84 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90

Table 2
Annual dietary composition of a groups of Javan lutungs (Apr 2012 - May 2014) in Pangandaran Nature Reserve, West Java, Indonesia, and vegetation within home ranges of
the subject group.

# Family Species Local name Plant Part Annual Tree Crown Crown Jacob’s
type eaten feeding number volume volume index (D)
percentage (m3 ) (%)
(%)

1 Flacourtiaceae Hydnocarpus heterophylla BUNTUT LUTUNG Tree Ylf 0.44 61 3539 0.36 0.09
(Bl.) Sloot.
2 Anacardiaceae Buchanania arborescens POHPOHAN Tree Fr 0.03 65 2994 0.31 −0.07
(Bl.) Bl.
Mlf 0.12
Yfr 0.01
Ylf 0.11
3 Dracontomelon mangiferum DAHU Tree Ylf 0.24 22 2051 0.21 0.06
Bl.
4 Mangifera indica L. MANGGA Tree Ylf 0.04
5 Mangifera sp. MANGGA PALI Tree Yfr 0.21 19 8069 0.83 −0.12
Ylf 0.44
6 Spondias pinnata (L.F.) Kurz KEDONDONG HUTAN Tree Ylf 0.15 1 1272 0.13 0.06
7 Annonaceae Cananga odorata Hook F. et KANANGA Tree Fl 0.01 23 20086 2.07 −0.89
Thoms.
Pt 0.01
Ylf 0.11
8 Stelechocarpus burahol BRAHOL Tree Ylf 0.02 8 291 0.03 −0.19
Hook. F. et Thomson
9 Arecaceae Arenga obtusifolia Bl. Ex. LANGKAP Tree Yfr 0.05 6 179 0.02 0.44
(=Palmae) Mant.
10 Bignoniaceae Oroxylum indicum (L.) PONPORANG Tree Ylf 0.01 1 628 0.06 −0.81
Benth. Ex Kurz
11 Radermachera gigantea (Bl.) PADALI Tree Ylf 0.03
Miq
12 Cannabaceae Celtis philippensis Blanco KIPEPETEK Tree Ylf 0.47
13 Clusiaceae Garcinia celebica L. MANGIS HUTAN Tree Ylf 0.10 1 35 0.00 0.93
14 Garcinia dioica Bl. CEURI Tree Ylf 0.03
15 Combretaceae Terminalia bellirica JAHA Tree Ylf 0.12 1 6440 0.66 −0.69
(Gaerth.) Roxb.
16 Terminalia catappa L. KETAPANG Tree Ylf 1.50 4 1228 0.13 0.85
17 Connaraceae Agelaea macrophylla (Zoll.) SUSUMUNDING Liana Ylf 0.11
Leenh.
18 Cucurbitaceae Mukia maderaspatana (L.) BARAKATIDA Liana Mlf 0.01
M.Roem.
Ylf 0.05
19 Dilleniaceae Tetracera scandens (L.) KIASAHAN Liana Fl 0.18
Ylf 0.21
20 Euphorbiaceae Alchornea rugosa (Lour.) BURUTU Tree Ylf 0.03 3 58 0.01 0.67
Muell.Arg.
21 Antidesma bunius (L.) HUNI Tree Mlf 0.07 4 798 0.08 0.22
Spreng.
Ylf 0.05
22 Aporosa sphaeridophora KIENDOG Tree Fl 0.01 1 21 0.00 0.72
Merr.
23 Excoecaria agallocha L. MATABUTA Tree Ylf 0.01 1 51 0.01 0.13
24 Mallotus philippensis (Lam.) PARENGPENG Tree Fl 0.06 97 6508 0.67 0.60
Muell.Arg.
Fr 0.04
Ylf 2.54
25 Mallotus ricinoides (Pers.) KIBAJING Tree Fl 0.05 33 1727 0.18 −0.27
Mull. Arg.
Fr 0.05
26 Suregada glomerulata (Bl.) KIBENTEUR Tree Ylf 0.09
Baill.
27 Fabaceae Acacia auriculiformis A. ACACIA Tree Fl 0.89 15 3273 0.34 0.02
Cunn. Ex Benth.
Sd 0.95
Ylf 0.59
28 Albizia lebbeck (L.) Benth. KITOKE Tree Ylf 0.43 8 2019 0.21 0.35
29 Bauhinia purpurea L. KIKUKUPU Liana Ylf 0.83
30 Cynometra ramiflora L. KATENG-KATENG Tree Bd 1.74 12 3358 0.35 0.92
Fl 0.78
Fr 0.06
Mlf 0.02
Yfr 0.03
Ylf 5.40
31 Dalbergia latifolia Roxb. SONOKELING Tree Ylf 0.93 14 3853 0.40 0.41
32 Desmodium umbellatum (L.) KIBALANAK Tree Ylf 0.02 1 25 0.00 0.77
Benth.
33 Tamarindus indica L. ASEM Tree Ylf 0.22 2 379 0.04 0.01
 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90 85

Table 2 (Continued)

# Family Species Local name Plant Part Annual Tree Crown Crown Jacob’s
type eaten feeding number volume volume index (D)
percentage (m3 ) (%)
(%)

34 Trachylobium verrucosum KISAPI Tree Fr 0.03

Hayne
35 Hernandiaceae Hernandia peltata Meism. BOROGONDORO Tree Fl 0.01 39 13590 1.40 −0.31
Fr 0.41
Ylf 0.32
36 Lamiaceae Tectona grandis L. JATI Tree Pt 7.94 483 365453 37.58 −0.74
Ylf 0.21
37 Vitex pubescens Vahl. LABAN Tree Fl 0.24 129 46138 4.74 0.43
Fr 0.47
Mlf 0.01
Yfr 0.01
Ylf 10.48
38 Lauraceae Neolitsea cassia (L.) HURU Tree Ylf 0.05 13 569 0.06 −0.11
Kosterm.
39 Lecythidaceae Barringtonia spicata Bl. PUTAT Tree Fl 0.55 3 2630 0.27 0.44
Fr 0.01
Pt 0.14
40 Lythraceae Lagerstroemia flasreginae BUNGUR Tree Ylf 0.13 32 4795 0.49 −0.59
Retz.
41 Lagerstroemia speciosa (L.) BENGER Tree Ylf 0.01 24 5466 0.56 −0.01
Pers.
42 Malvaceae Grewia paniculata Roxb. DERAWAK Tree Ylf 0.01 8 579 0.06 −0.63
43 Heritiera littoralis Aiton DUNGUN Tree Ylf 0.86 1 261 0.03 0.94
44 Hibiscus similis Bl. WARU Tree Fl 0.01 28 3854 0.40 0.83
Ylf 4.17
45 Hibiscus tiliaceus L. WARU LAUT Tree Fl 0.03
Ylf 1.51
46 Kleinhovia hospita L. TANGKOLO Tree Pt 0.40 3 1009 0.10 0.59
47 Pterocymbium javanicum R. TONGTOLOK Tree Fl 0.18 1 393 0.04 0.75
Br.
Ylf 0.11
48 Pterospermum divirsifolium CERLANG Tree Fl 0.14 23 3863 0.40 0.39
Bl.
Ylf 0.76
49 Pterospermum javanicum CARUY Tree Fl 1.99 327 225869 23.22 −0.66
Jungh
Fr 0.01
Sd 0.07
Yfr 3.82
Ylf 0.02
50 Schoutenia ovata Korth. WARIKUKUN Tree Fl 0.85 3 291 0.03 0.94
Mlf 0.03
Ylf 0.09
51 Sterculia coccinea Jack HANTAP HEULANG Tree Fl 0.06 78 42648 4.39 −0.14
Mlf 0.03
Sd 0.39
Ylf 2.86
52 Meliaceae Dysoxylum caulostachyum KOKOSAN MONYET Tree Fl 0.41 374 32221 3.31 −0.57
Miq.
Fr 0.20
Sd 0.30
Ylf 0.01
53 Swietenia macrophylla MAHONI Tree Fl 0.14 157 117656 12.10 −0.44
King
Pt 4.36
Ylf 0.57
54 Moraceae Ficus annulata Bl. KIARA KONENG Hemi- Fr 0.73 2 772 0.08 0.81
epiphyte
Yfr 0.01
55 Ficus benjamina L. BERINGIN Hemi- Fr 6.54 3 406 0.04 0.99
epiphyte
Mlf 0.01
Yfr 1.03
Ylf 2.65
56 Ficus elastica Roxb. KARET MUNDING Tree Ylf 0.03
57 Ficus microcarpa L.f. PERENG Hemi- Ylf 0.04
epiphyte
58 Ficus obscura Bl. HAMPELASAN Hemi- Ylf 0.14
epiphyte
59 Ficus pubinervis Bl. KOPENG Tree Yfr 0.24 2 285 0.03 0.98
Ylf 2.69
60 Ficus septica Burm. KICIAT Tree Fr 0.01 103 5707 0.59
86 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90

Table 2 (Continued)

# Family Species Local name Plant Part Annual Tree Crown Crown Jacob’s
type eaten feeding number volume volume index (D)
percentage (m3 ) (%)
(%)

Ylf 0.01
61 Ficus subcordata Bl. KIARA KEBO Tree Fr 0.46 2 1343 0.14 0.66
Yfr 0.20
Ylf 0.01
62 Ficus sumatrana (Miq.) Miq. KIARA BEAS Hemi- Fr 0.03
epiphyte
Mlf 0.04
Yfr 0.08
Ylf 1.22
63 Ficus variegata Bl. KONDANG Tree Fr 1.00 2 2857 0.29 0.74
Yfr 0.87
Ylf 0.07
64 Ficus sp. KIARA SP. Tree Fr 0.40 8 19941 2.05 −0.24
Yfr 0.05
Ylf 0.83
65 Myristicaceae Myristica guatterifolia DC. KIMOKLA Tree Fr 0.19 7 234 0.02 0.93
(M. quercifolia?)
Ylf 0.47
66 Myrtaceae Decaspermum fruiticosum IPIS KELIT Tree Yfr 0.09 11 782 0.08
J.R. et G. Foster
Ylf 0.45
67 Eugenia polyantha Wight. SALAM Tree Fl 0.01 192 26229 2.70 −0.45
Sd 0.82
Yfr 0.03
Ylf 0.19
68 Rhodamnia cinerea Jack KIBESI Tree Fr 0.45 12 451 0.05 0.81
69 Syzygium antisepticum (Bl.) KIPANCAR Tree Sd 0.15 179 8422 0.87 −0.65
Merr.& L.M.Perry
Ylf 0.03
70 Syzygium aqueum Alston JAMBU Tree Fl 0.03 6 655 0.07 0.31
Fr 0.09
Ylf 0.01
71 Syzygium racemosum (Bl.) KOPO Tree Fl 0.06 11 914 0.09 0.91
DC.
Ylf 1.87
72 Phyllanthaceae Baccaurea racemosa MENTENG Tree Ylf 0.28
(Reinw. Ex Bl.) Mull.Arg.
73 Glochidion molle Bl. KIHUUT Tree Ylf 0.12 1 8 0.00 0.99
74 Sauropus rhamnoides Bl. KEBOJARU Shrub Ylf 0.01
75 Rhizophoraceae Carallia brachiata (Lour.) KIKUKURAN Tree Ylf 0.01
Merr
76 Rubiaceae Hypobathrum frutescens Bl. KIHAPIT Tree Fr 0.04 12 271 0.03 0.19
77 Ixora paludosa (Bl.) Kurz SOKA Shrub Fl 0.01 1 652 0.07 0.17
Yfr 0.02
Ylf 0.07
78 Nauclea orientalis (L.) L. KELEPU Tree Fl 0.41 11 10493 1.08 0.34
Fr 1.11
Yfr 0.38
Ylf 0.26
79 Neonauclea excelsa (Bl.) CANGCARATANG Tree Fl 0.03 7 1706 0.18 −0.68
Merr.
80 Psychotria viridiflora Reinw. KIKORES Shrub Fr 0.14
Ex Bl.
Ylf 0.01
81 Rutaceae Clausena excavata Burm. KIBACATA Tree Fr 0.03 2 41 0.00 0.88
Ylf 0.03
82 Sapindaceae Erioglossum rubiginosum KILALAYU Tree Fr 2.07 101 4960 0.51 0.81
(Roxb.) Bl.
Mlf 0.41
Ylf 2.23
83 Mischocarpus sundaicus Bl. KIHOE Tree Ylf 0.02
84 Schleichera oleosa Merr. KESAMBI Tree Ylf 0.90 71 5238 0.54 0.25
85 Urticaceae Villebrunea rubescens (Bl.) NANGSI Shrub Ylf 0.01
Bl.
- - - LIANA Liana ? 0.08
- - Liana Bd 0.01
- - Liana Ylf 0.43
- - - Unknown ? ? 0.20
- - ? Ylf 0.02
- - - PAKIS Fern Ylf 0.01
- - - SOIL - - 0.08

Bd: buds, Fl: flower, Fr: mature fruits, Mlf: mature leaves, Pt: petiole, Sd: seeds, Ylf: young leaves, Yfr: young fruits,?: unknown.
The species and parts eaten in bold letters represent main plant species (annual feeding percentage >1%, part combined) and main diet items (annual feeding percentage
>1%), respectively.
 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90 87

Fig. 3. Monthly changes (Aug 2012–May 2014) in the dietary composition of the K group lutungs (Trachypithecus auratus) in Pangandaran Nature Reserve, West Java,
Indonesia. Young leaves include the petiole and buds, and others include soil.

Table 3
Summary of Spearman’s rank correlation tests between monthly feeding percentages and dietary diversity (H’) of Javan lutungs in Pangandaran Nature Reserve, West Java,
Indonesia.

Feeding percentage Feeding percentage Dietary diversity (H’)

(range) Young leaves Mature fruits Young fruits Flowers (0.07–0.53)

Mature leaves rs = −0.305 rs = −0.085 rs = 0.707 rs = −0.017 rs = 0.424

(0.0–6.5) p = 0.425 p = 0.828 p = 0.03* p = 0.966 p = 0.256
Young leaves rs = −0.717 rs = −0.271 rs = −0.350 rs = −0.883
(33.2–82.3) p = 0.037* p = 0.480 p = 0.359 p = 0.003**
Mature fruits rs = −0.271 rs = −0.083 rs = 0.483
(0.0–34.7) p = 0.480 p = 0.843 p = 0.194
Young fruits rs = 0.000 rs = 0.593
(0.0–40.8) p = 1.000 p = 0.092+
Flowers rs = 0.233
(0.0–27.0) p = 0.552
+
p < 0.1.
*
p < 0.05.
**
p < 0.001.

p < 0.001) and tree number (rs = 0.379, p = 0.002) were detected. and September 2013), mature fruits/seeds (April 2012 and April
However, on closer inspection, the relationship between abun- 2014), and young fruits/seeds (January 2014). Finally, percentages
dance and consumption was more complicated. Plant species that of the mature leaves were constantly low. The percentage of young
had a lower crown volume were more likely to be selected (in terms fruits showed a significant positive correlation with the percent-
of Jacob’s index, D) as foods than those with higher crown volume age of mature leaves (rs = 0.707, p = 0.033), and the percentage
(rs = −0.565, p < 0.001). This pattern was maintained even when of mature fruits and dietary diversity exhibited significant neg-
the analyses were limited to main food plant species (rs = −0.885, ative correlations with the percentage of young leaves (mature
p < 0.001) (Fig. 2). fruits: rs = −0.717, p = 0.037, dietary diversity: rs = −0.883, p = 0.003)
(Table 3).
Seasonal change in diet and relationship with plant phenology We found a significant positive relationship for the phenology of
young fruits with their percentage in the diet (rs = 0.848, p = 0.004).
With respect to the food item category, lutungs mainly fed on However, most of the category-based relationships were not sig-
young leaves (averaged over the study period, 69.9%), followed nificant (Table 4).
by fruits (both young and mature, 21.2%). Mature leaves were With respect to species, three out of 17 main diet items for which
the least consumed (0.8%). Fig. 3 shows monthly changes in the we could conduct the correlation tests (young leaves of C. ramiflora,
dietary composition of lutungs. Young leaves formed the main com- young leaves of S. macrophylla, and young fruits of P. javanicum)
ponent of the diet throughout the study period, but other items had significant positive correlations with their availability (Table 5),
sometimes had a higher percentage, including flowers (April 2012 partly supporting our second prediction.
88 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90

Table 4
Summary of Spearman’s rank correlation tests between monthly phenology index and monthly feeding percentages and dietary diversity (H’) of Javan lutungs in Pangandaran
Nature Reserve, West Java, Indonesia.

Food types Feeding percentage (range) Dietary diversity (H’)

(phenology range) Mature leaves(0.0–6.5) Young leaves(33.2–82.3) Mature fruits(0.0–34.7) Young fruits(0.0–40.8) Flowers(0.0–27.0) (0.07–0.53)

Mature leaves rs = −0.300 rs = −0.300 rs = 0.167 rs = −0.136 rs = 0.216 rs = 0.033

(87.3–99.8) p = 0.509 p = 0.437 p = 0.678 p = 0.728 p = 0.581 p = 0.948
Young leaves rs = 0.700 rs = 0.367 rs = −0.550 rs = 0.373 rs = −0.067 rs = -0.133
(2.5–69.5) p = 0.053+ p = 0.336 p = 0.133 p = 0.323 p = 0.880 p = 0.744
Mature fruits rs = 0.170 rs = 0.083 rs = −0.083 rs = −0.186 rs = -0.050 rs = −0.300
(7.5–19.0) p = 0.663 p = 0.843 p = 0.843 p = 0.631 p = 0.912 p = 0.437
Young fruits rs = 0.610 rs = -0.150 rs = −0.333 rs = 0.848 rs = 0.100 rs = 0.533
(2.2–15.1) p = 0.081+ p = 0.708 p = 0.385 p = 0.004** p = 0.810 p = 0.148
Flowers rs = 0.220 rs = 0.050 rs = −0.100 rs = 0.034 rs = −0.317 rs = 0.133
(3.8–20.8) p = 0.569 p = 0.912 p = 0.810 p = 0.931 p = 0.410 p = 0.744
+
p < 0.1.
**
p < 0.01.

Table 5
Summary of Spearman’s rank correlation tests between phenology index and monthly feeding percentage of the lutungs on the main diet items. N in parenthesis represents
number of monitoring trees within the home range.

Species (phenology range) Family Part eaten Annual feeding (%) rs p-value

Vitex pubescensa (N = 21) Verbenaceae Ylf 10.48 −0.092 0.814 n.s.

(2.4–100.0)
+
Tectona grandis (N = 20) Verbenaceae Pet 8.15 0.661 0.053
(0.0–100.0)
Cynometra ramifloraa (N = 4) Fabaceae Ylf, Bd 7.14 0.692 0.039 *

(0.0–100.0)
Ficus benjamina (N = 2) Moraceae Fr 6.54 −0.632 0.252 n.s.
(25.0–100.0)
**
Swietenia macrophylla (N = 17) Meliaceae Ylf, Pet 4.93 0.810 0.008
(0.0–67.6)
*
Pterospermum javanicum (N = 20) Sterculiaceae Yfr 3.82 0.764 0.017
(0.0–62.5)
Sterculia coccineaa (N = 5) Sterculiaceae Ylf 2.86 −0.176 0.650 n.s.
(10.0–100.0)
Ficus pubinervis (N = 2) Moraceae Ylf 2.69 0.632 0.252 n.s.
(50.0–100.0)
Ficus benjamina (N = 2) Moraceae Ylf 2.65 −0.081 0.897 n.s.
(50.0–100.0)
Mallotus philippensis (N = 2) Euphorbiaceae Ylf 2.54 −0.185 0.635 n.s.
(0.0–100.0)
Erioglossum rubiginosum (N = 2) Sapindaceae Ylf 2.23 0.475 0.197 n.s.
(0.0–100.0)
Erioglossum rubiginosum (N = 2) Sapindaceae Fr 2.07 −0.188 0.629 n.s.
(0.0–25.0)
Pterospermum javanicum (N = 20) Sterculiaceae Fl 1.99 0.380 0.314 n.s.
(0.0–65.0)
Ficus sumatranaa (N = 5) Moraceae Ylf 1.22 0.667 0.219 n.s.
(30.0–100.0)
Nauclea orientalis (N = 2) Rubiaceae Fr 1.11 −0.125 0.749 n.s.
(0.0–30.0)
Ficus benjamina (N = 2) Moraceae Yfr 1.03 – – –
(0.0–100.0)
Ficus variegata (N = 1) Moraceae Fr 1.00 0.000 1.000 n.s.
(0.0–100.0)

n.s.: not significant.

Bd: buds, Fl: flowers, Fr: mature fruits, Pet: petiole, Yfr: young fruits, Ylf: young leaves.
We could not perform the analysis for young fruits of F. benjamina due to limited number of months when the lutung fed on it.
**
p < 0.01.
*
p < 0.05.
+
p < 0.10.
a
phenology index was obtained from all monitoring trees within the forest park.

Discussion West Java: 62.1% of diet was foliage; Beckworth, 1995; Bali West
National Park, Bali: 33.6–41.3% of diet was foliage; Vogt, 2003). This
The Javan lutungs in PNR mainly fed on young leaves; how- phenomenon seems to be a common feature of this species. Overall,
ever, the percentage of non-leaves, such as fruits (both mature and 85 plant species were fed on in PNR; however, feeding was mainly
young) was also considerable (Fig. 3). This foliage-dominant dietary focused on a limited number of plant species. Similarly, small num-
composition was similar to that reported by Brotoisworo (1991) bers of plant species contributing to the dietary composition have
and Kool (1993), who studied the dietary habits of Javan lutungs been reported for other Trachypithecus species. For instance, six,
in PNR in the 1970–1980s (annual feeding percentage: 55–80%), eight, and 11 species contributed to >75% of the diet for T. francoisi
as well as studies conducted at other sites in Indonesia (Cibodas, (Zhou et al., 2006), T. delacouri (Workman, 2010), and T. pileatus,
 Y. Tsuji et al. / Mammalian Biology 98 (2019) 80–90 89

respectively (Solanki et al., 2008). Thus, it is important to evalu- Clauss, Michael A Huffman, and an anonymous reviewer for their
ate the key plant species that affect the activity, range, and inter- constructive comments.
and intra-specific competition to elucidate the feeding strategy of
Trachypithecus monkeys. We found that these main diet species
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