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Socio-ecology of the hoolock gibbon (Hylobates hoolock) in two forests of

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 SECTION IV. FIELD RESEARCH
SOCIO-ECOLOGY OF THE HOOLOCK GIBBON
(HYLOBATES HOOLOCK) IN TWO FORESTS OF BANGLADESH
M. Farid Ahsan1
T he hoolock or white-browed gibbon (Hylobates hoolock) is the
second-largest of the gibbons. Males weigh 6.1–7.9 kg, females
6.0–6.6 kg (Shortridge 1914) with a head and body length of
45.7–63 cm (Roonwal and Mohnot 1977) and have long limbs and
dense, shaggy pelage. The pelagy of males and young females is
black with silver-white eyebrows and adult females are pale-brown
with golden-white eyebrows. Newborns are pale-greyish-white with
yellow tinge but lack hair on part of the face, palms, and soles of
their feet which are black. The female has a long clitoris often mis-
taken for a penis.
The wild hoolock gibbon ranges in the Arunachal Pradesh,
Assam, Meghalaya, Nagaland, Manipur, Tripura, and Mizoram
States of India (Mukherjee 1986); Jamalpur (on the border with
Meghalaya State of India), Sylhet, Chittagong, Chittagong Hill-
Tracts, and Cox’x Bazar Forest Divisions of Bangladesh (Ahsan
1984); Sittwe, Rangoon, and Myitkyina Provinces of Burma, now
Myanmar (Mootnick et al. 1987); and extreme western part of Yun-
nan Province (Yingjiang, Lianghe, Tecgchong, and Baoshan) of
China (e.g., Tan Bangjie 1985).
The main aims of this paper are to describe the ecology and be-
havior of the hoolock gibbon in detail, to fill in the gap in gibbon
studies, to help in the promotion of conservation strategies, and to
compare the results with those of the previous studies of gibbon
species throughout Southeast Asia.
Study Area
The study was conducted in two forested areas of Bangladesh
(20°34' to 26°38' N and 88°01' to 92°41' E): (a) Lawachara of West
Bhanugach Reserve Forest (24°32' N and 91°47' E; altitude 22 m;
total area 2,738 ha) under Sylhet Forest Division in the northeast;
and Chunati (22°55' N and 92°03' E; altitude 15 m; total area 40 ha
[study block]) under Chittagong Forest Division in the southeast
(Fig. 1). The former forest is isolated from the nearest forest by
about 10 km, because of surrounding plantations of tea, coffee, and
rubber, and the cultivation of rice. These two forests are not marked-
ly evergreen, but the majority of small trees are evergreen and most
of the tall trees are deciduous (Ahsan 1994). The latter forest is a
small block isolated from the nearest block by small patch of culti-
vated rice field (of about 500 yards). The detailed description of
these study areas are available elsewhere (e.g., Ahsan 1994) but a
short summary is given here (from Ahsan 1994) to introduce them
to others.
The soils of Lawachara are brown, sandy clay loam to clay
loam of Pliocene origin, and the Chunati are also brown, sandy loam
to sandy clay loam of Pliocene sediments (Hossain et al. 1989). The
forests of Lawachara do not fall markedly under one recognized
type, but as semi- and/or mixed-evergreen, where tall trees are de-
ciduous and the understory evergreen. The forest canopy of Chunati
is open and not markedly of a single type, but it can be categorized
as mixed-evergreen, where tall trees are deciduous and the under-
story evergreen. The former forest is quite rich for both of its faunal
and floral diversities, while the latter is reasonably good. In both
1Department of Zoology, University of Chittagong, Bangladesh
286
areas, weather is hot, humid, and monsoonal in summer, and dry
and mild in winter. Long-term data (1970-1990) on rainfall show a
more uniform fluctuation at Lawachara (Srimangal) than at Chunati
(Chittagong, due to influence of the Bay of Bengal), while temper-
atures and humidity fluctuate less within and between sites.
Data were collected through scan sampling (Altmann 1974) of
10-minute intervals on three groups (two in Lawachara, Sylhet, and
one in Chunati, Chittagong), the first two of every three months
worked at Lawachara and the third month at Chunati (five day al-
ternate blocks and 20 days a month) from January to July 1989.
From August 1989 to July 1990 (then until December 1990 vegeta-
tional and ranging works were done), the first two weeks of every
month was spent at Lawachara of Chunati, following two groups—
alternating for 5 days each. Rest, travel, feed, forage, calling, terri-
toriality, and play were noted as major activities and other behaviors
were scored as notes (Ahsan 1994).
In Lawachara, 114 species of plants (of 70 genera and 37 fam-
ilies) were identified (including many up to generic level but ex-
cluding family level). More than 100 species remained unidentified
(n=1,520) from the phenological transects. While in Chunati 58
species (of 40 genera and 27 families) were identified and six re-
mained unidentified (n=211).
The mean total height of phenological trees were 16.1 m in
Lawachara and 15.6 m in Chunati, which differed significantly be-
tween two sites (M-W U-test, 2-tailed, z=-11.155, p<0.0001). DBH
(diameter at breast height, > 10 cm) was 26.1 cm and 37.2 cm re-
spectively, which varied significantly (M-W U-test, 2-tailed, z=-
8.977, p<0.0001). The former has a more closed canopy than the
latter (Fig. 2). Figure 2 gives a hypothetical representation of the
forest profiles of two sites based on one-meter transects (north-
south and east-west) in the middle of each site.
In Lawachara, the tree density (> 10 cm DBH) was 203
trees/ha and in Chunati 124 trees/ha. The 50 m x 50 m quadrant den-
sity was 93 trees/plot (range 41–182, n=25; i.e., 372 trees/ha) in
Lawachara and 43 trees/plot (range 18–63, n=10; i.e., 171 trees/ha)
in Chunati. These two methods of estimating tree densities varied
within the site (p<0.01).
RESULTS AND DISCUSSION
Social Organization
Group Structure
In the Lawachara study area, eight groups (excluding solitary
females) were located, and each of them comprised 2 to 5 animals
(mean 2.9 at the beginning of the study and 3 at the end) (Table 1).
Of these, most effort was concentrated on 2 groups (L1 and L2). Al-
though, data were also gathered on groups whose home ranges were
overlapped with L1 and L2. In the Chunati area, 7 groups were lo-
cated and each contained 2 to 4 animals (mean 3 at the beginning of
the study and 2.9 at the end) (Table 1). Of these, only group C, who
lived in a small, isolated forest block, was intensively studied. The
group initially had 3 members. The adult female died during the
 IV. Field Research

Figure 2. Hypothetical cross section of the forest profile trees of two study
sites (top–Lawachara; bottom–Chunati).
Figure 1. Forested areas of Bangladesh (after Rashid 1977) and location of
study sites. Shaded areas: more or less continuous forest cover. Stippled
areas: scattered forest patches. (a) Moist deciduous forest, (b) Semi/mixed-
evergreen forest. (c) Mangrove forest. (1) West Bhanugach and (2) Chunati.

Table 1
Group Size and Composition of Hoolock Gibbon in Two Study Sites
(initial data in parentheses)

Location Group Adult Young Total

Male Female Juvenile-II Juvenile-I Infant-II Infant-I

Lawachara L1 1 (1) 1 (2) - 1M (1M) - 1 (-) 4 (4)

L2 1 (1) 1 (1) 1 F (1F) 1 F (1 F) 1 (-) - (1) 5 (5)
L3 1 (1) 1 (1) - - 1 (-) - (1) 3 (3)
L4 1 (1) 1 (1) - 1 (-) - (1) - 3 (3)
L5 1 (1) 1 (1) - - - - 2 (2)

RLB 1 (1) 1 (1) - - - 1 (-) 3 (2)

SR 1 (1) 1 (1) - - - - 2 (2)
VA 1 (1) 1 (1) - - - - 2 (2)

Chunati C 1 (1) - (1) - 1 F (1 F) - - 2 (3)

CE1 1 (1) 1 (1) - 1 (1) - - 3 (3)
CE2 1 (1) - - 1 F (1F) - - 2 (2)
CE3 1 (1) 1 (1) - 1 (1) - - 3 (3)
CW1 1 (1) 1 (1) - 1 (1) - - 3 (3)
CW2 1 (1) 1 (1) 1 (1) 1 (1) - - 4 (4)
CW3 1 (1) 1 (1) - - - 1 (1) 3 (3)

Composition (%) 34.1 (34.9) 29.5 (34.1) 25.0 (23.2) 11.4 (7.0)

287
The Apes: Challenges for the 21st Century
middle of the study, and later on in the study the juvenile-II female
replaced the adult female.
Gibbon group size is not usually more than 6 in the wild
(Ahsan 1994), but the composition varies slightly due to habitat in-
fluence and population dynamics.
Group Cohesion
Empirical data shows that hoolock groups are highly cohesive
during active hours of the day. Of three groups (L1, L2, and C), all
members were found together in 91.8% of the scans (n=4,838) and
in 4.1% of the scans (n=217) one member was missing, giving a
total of 95.6% (n=5,272). Overall the minimum percentage of sight-
ings with all group members present was 85.8% during resting and
a maximum of 96.1% during feeding. The minimum percentage of
all group members observed was 67.3% during foraging for Group
C and a maximum of 100% during calling and playing for all
groups. No significance difference was found in sighting all group
members during scan samples between groups (c2=1.97, df=2,
p>0.05).
There is no doubt that a small family group of gibbons is co-
hesive. Overall, minimal group spread showed in 91.8% of cases
observed all members present in scan samples and sometimes in
huddles during resting, which signifies the hoolocks’ cohesiveness.
Group Dispersion
The mean group dispersion of three groups during diurnal ac-
tivities was 12.5 m (median 10 m, range 0-350 m, n=3,978), which
differed significantly among groups controlling for activities (K-W
1-way ANOVA H=791.868, p<0.0001). The overall group disper-
sion also varied significantly between groups of the two study sites,
Lawachara and Chunati (M-W U-test, 2-tailed, 2=-27.196,
p<0.0001) and even between groups of the same site, Lawachara
(M-W U-test, 2-tailed, z=-7.836, p<0.0001). The probable cause of
difference between groups may be the absence of conspecific terri-
torial conflict in C Group and some sort of coordination loss be-
tween siblings of L1. The pattern of dispersal of two study sites is
different, however it is similar between groups of the Lawachara.
On average, the minimum group dispersal was 8.3 m during calling
and a maximum of 28.9 m during traveling. Group dispersal during
territorial behavior was not recorded for Group C, because they live
in a small patch of forest where no other gibbon group occurs.
Interindividual Spacing
Interindividual spacing was estimated for the two nearest indi-
viduals to the center of the group (i.e., adult female). The overall
mean distance of the closest individual to the center of the group
was 4.3 m (median 3 m, range 1-45 m, n=488) and for the second
closest, it was 9 m (median 6 m, range 1-45 m, n=588). These dis-
tances varied highly significantly between three groups (K-W 1-
way ANOVA–1st closest: H=82.027, p<0.0001; 2nd closest:
H=82.776, p<0.0001); it was also varied between two study sites
(M-W U-test, 2-tailed–1st closest: z=-8.334, p<0.0001; 2nd closest:
z=-3.627, p<0.0003); and between the two groups at Lawachara (M-
W U-test, 2-tailed–1st closest: z=-4.135, p<0.0001; 2nd closest: z=-
8.627, p<0.0001). The probable cause of difference between groups
may be the distribution of standing crop size and the lack of coordi-
nation between siblings of L1 (Ahsan 1994 for detail), which may
result in higher spacing between group members. On average, the
minimum distance of the first individual from adult female was 1 m
during calling (but sometimes individuals huddled during resting or
were in contact during grooming, when the distance is 0) and a max-
imum of 16.4 m was observed during traveling (see Ahsan op cit).
288
Generally, the adult male was the closest individual to the adult
female (1 m) during calling and ranged from 3.9 to 8.6 m). The Ju-
venile-I was the second closest to her during calling bouts. The dis-
tance between the adult male and juvenile-I, as first and next nearest
individuals did not vary significantly (M-W U-test, 2-tailed – 1st
closest: z=-0.365, p=0.715; 2nd closest: z=-0.541, p=0.588). The
subadult and juvenile-II were furthest from other members of the
group both during the day and a night. The mean distance between
individuals was 9 m (median 7 m) during feeding, 17 m (median 15
m) during foraging, 13 m (median 10 m) during resting, and 29 m
(median 20 m) during traveling.
The overall mean distance to the closest individual in the cen-
ter was lowest (4.3 m) in hoolock than in larger-bodied gibbon
species (siamang 8.9 m: Chivers 1974), but its mean range and other
activities are higher than other gibbon species (Ahsan 1994). The
subadult is always the farthest from the center in all gibbon species,
including hoolock.
Intergroup Spacing
On average, the hoolock groups of Lawachara maintained an in-
tergroup spacing of 34.2 m (median 30 m, range 5-200, n=112) only
when the second and/or third group was observed during scan sam-
ples. Intergroup spacing of Chunati was not determined because
Group C lived alone in a small forest block. The natural spacing of
gibbon groups may need to be determined by locating populations
that have lived free from human interference for a number of gener-
ations. The hoolock study areas were heavily affected by human in-
terference, so it could not be compared with other gibbon species.
Travel Order Sequence and Group Leading
The adult female is most likely to be first during travel, and the
juvenile last. Overall, the adult female led the group in 61% of
cases, followed by the adult male (33%) and the juvenile (6%). The
Group C adult female led only 43% and the adult male 48%. After
the female’s death, the male led 67% cases and the juvenile female
led 39%.
The adult female of a siamang group led 65% of the travel
bouts (Chivers 1974). An agile gibbon adult female led travel 36%,
and the adult male 52% (Gittins 1979). The adult female hoolock,
who in this case led travel bouts, is close in size to its larger-bodied
relative, the siamang. The animal who initiates group movement
may not be the leader of the group. It follows the indicator—decider
model given by Kummer (1968) for group movement in hamadryas
baboons. In hoolock gibbons, a young and/or infant-II initiated by
traveling in one direction, however the group did not follow unless
an older decider (i.e., adult male/female) followed in that direction.
Similar instances were observed for the young and infant of agile
gibbon (Gittins 1979). When an adult hoolock traveled in one di-
rection, he or she would look back to see if the other partner was fol-
lowing the same route. This appears to be a mutual agreement
between them.
Grooming Behavior
Grooming was most frequent between adult pairs during social
interactions within group. It helps strengthen the social bond, re-
duces tension, and keeps the pelage clean. It is mostly performed by
the adult male and typically lasts for short periods of 5 sec to 14 min
(mean 5.9 min, median 3 min, n=83). Two peaks of grooming were
observed during warm hours, one at 1100 h (between 1100 and
1200 h) and another at 1400 h. The monthly distribution of groom-
ing shows marked increases between February and May, which may
result from the warm weather after the winter months. Grooming

sessions for all groups were more or less similar except for Group C
due to the of the death of adult female of the group.
In the siamang, the adult male initiates grooming and a mean
grooming bout lasted 12.9 min at Kuala Lompat and 11.5 min at Ulu
Sempam (Chivers 1974). The adult males of both siamang and lar
groups groomed their adult females more than other group females
(Palombit 1992). In polygynous primates such as rhesus and pig-
tailed macaques, Hanuman langurs (Ahsan), and olive baboon (C.
Drews, personal communication), males are groomed more by fe-
males than they groom the females. In monogamous gibbons,
females are groomed more by the males and females do not recip-
rocate. The grooming asymmetry between males and females of
monogamous and polygynous primates is unique.
Sexual Behavior
Hoolock gibbons reach sexual maturity at about 7 years of age
(Mathews 1946). The observed sexual behavior of hoolocks was
classified (modified from Chivers 1974) as (1) attempted copula-
tion; (2) possible copulation, when observation conditions obscured
the behavior; (3) brief copulation, when mounting was either for
very short time (less than 5 sec) and/or intromission was possibly
not accomplished (mostly observed during ventro-ventral attempts
of copulation between adult males and juveniles); and (4) complet-
ed copulation, when mounting was achieved and pelvic thrusts ob-
served. Mating occurred in all months throughout the year except
August and September, and the mean duration of complete copula-
tion between adult pairs was 36.6 sec (median 30 sec, range 5-120
sec, n=39). Results suggest that the birth peak of hoolock gibbons is
between mid-September and January (see Ahsan 1994). The gesta-
tion period appears to be 6 to 8 months (180-240 days) which is in-
termediate among gibbon species (Geissmann 1991) and the
inter-birth interval is assumed to be 3 to 4 years (Ahsan 1994).
Group Formation
The following methods of new group formation observed in
hoolock gibbons included: brother-sister pair formation, father-
daughter pair formation, group formation through mate leaving and
remating, and dispersed subadult pair formation.
RANGING BEHAVIOR
Day range length (DRL): The overall total distance traveled by
three hoolock groups was 1367 m (median 1404, range 278-3,375,
n=56). The total DRL varied significantly among groups (K-W 1-
way ANOVA, H=20.496, df=2/55, p=0.0001) and groups within the
same site (M-W test, U=230, z=-2.667, p=0.0076).
The general trend in each group for DRL was to be higher dur-
ing the early hours of the day (i.e., before 0900 h) and to decrease
gradually thereafter, but Group L2 traveled more during 1200–1500
h than the 0900–1200 h period of the day. This is because Group L2
had to cover the large part of its home range area.
It is obvious that, after the long overnight fast, animals have to
eat more, find sufficient quantity and better quality of food, espe-
cially sugary fruits, which are a favorite of gibbons and sparsely dis-
tributed in the forest. Hence, they have to travel more during the
early hours of the day.
One of the most striking aspects of the animals’ ranging be-
havior is the regularity with which they move around their range.
Not just one factor totally accelerates ranging variation in gibbons’
behavior, but a group of interrelated factors regulates it, although
foods, especially the distribution of figs, generally restrict to some
extent the ranging of gibbons.
289
IV. Field Research
Night position shift (NPS): The overall NPS (between consecutive
nights) was 352 m (median 28 m, range 0-1,229 m, n=56), which
varied significantly among groups (K-W 1-way ANOVA,
H=23.507, df=2/55, p=0.0001), and between groups at Lawachara
(M-W test, U=272, z=-4.068, p=0.0001).
Night position distance (NPD): NPD (distance between sleeping
trees of a group used in one night) mean for three groups was 59 m
(median 37 m, range 10-486 m, n=38), which differed significantly
among groups (K-W 1-way ANOVA,H=13.47, df=2/36, p=0.0012),
and also varied between L Groups (M-W test, U=-3.524, p=0.0004).
Maximum radius (MR): The overall mean MR of the day range
covered was 478 m (median 411 m, range 95-1,229, n=56), which
varied significantly among groups (K-W 1-way ANOVA,
H=32.828, df=2/55, p=0.0001), and also differed between L Groups
(M-W test, U=267.7, z=-3.922, p=0.0001).
Maximum radius time (MRT): The average time taken to cover
maximum radius area of the day was 376 min (median 392 min,
range 80-667, n=56), which did not vary significantly among groups
(K-W 1-way ANOVA, H=1.107, df=2/55, p=0.6014), or
between groups in the same area (M-W test, U=174, z=-0.8, p=
0.4237). It is reasonable to predict that each group covered propor-
tionately the same amount of area of its home range during the same
length of time.
Activity period (AP): The mean AP was 543 min (median 541 min,
range 374-745, n=154), which varied significantly among groups
(K-W 1-way ANOVA, H=7.924, df=2/55, p=0.019), but did not dif-
fer between L Groups (M-W test, U=205, z=-1.833, p=0.0668) pos-
sibly due to the same habitat.
Number of 0.25 ha quadrant entered: A hoolock group entered 8-
85 (mean 32.9, median 32, n=56) 0.25 ha quadrants during a whole
day period, which varied highly significantly among groups (K-W
1-way ANOVA, H=244, p=0.0004), and very significant between
two groups at L site (M-W test, U=231, z=-2.701, p=0.0069). The
maximum use of any quadrant depended on the distribution and
density of most available and favored food trees.
Home range (HR) and pathway: Home range was delimited indi-
vidually by drawing a line around the cumulative day ranges for the
entire study period (Figs. 3-5). The overall home range was 50.8 ha
(Table 2), which varied significantly among groups (c2=38.8, df=2,
p<0.001). The mean area covered by each group during full-day fol-
lows was 31.6 ha i.e., 634% of the total actual home range (Table 2),
which also varied significantly among groups (c2=20.909, df=2,
p<0.001).
It is evident that hoolocks used some parts of their home range
more than others, and a number of lacunae are present. Most of the
pathways that exit in Lawachara forest seem to be suitable for trav-
eling through, from my observation points on the ground. There is
one point where the members of Group L1 have to jump about three
meters between trees to cross the railway lines, to use some food
sources in that part of the home range. Although there is an easier al-
ternative path to cross the railway lines, the group only used the for-
mer path during my study period, perhaps because food sources there
are close to it. For Group C, there are several places where the group
did not visit during my study, possibly due to lack of canopy links
between trees. In some places, the group used the suitable alternative
arboreal route, but in other places individuals came down to a mini-
mum height (about 3 m) to visit some parts of the home range.
The Apes: Challenges for the 21st Century
Figure 3.Pathways of group L1 during full-day follows (home range
area–dotted line; railway tracks–crossed line).
Figure 4.Pathways of group L2 during full-day follows (home range
area–dotted line; railway tracks–crossed line).
Core area (CA): The mean CA of a hoolock group was 27.8 ha
(Table 2), which varied significantly among groups (c2=26.612,
df=2, p<0.001).
Territory size (TS): The mean TS of L1 and L2 was 51.3 ha (Table
2), which varied significantly among groups (c2=18.613, df=1,
p<0.001). A small portion of the shared home range between neigh-
bors was controlled by one of the group as its own territory.
Night sleeping trees: Hoolocks spend more than 60% of their life in
sleeping trees. Chosen night trees were to some extent different in
appearance in that the majority were relatively taller than neighbor-
ing trees. Most of them were situated on slopes or near a hilltop.
When the group size of a hoolock family was more than two, they
used more than one sleeping tree for the same night (except one—
when L1 [3-member] used two sites in the same tree). Usually the
youngest member, infant/juvenile-I, slept with the mother, the adult
male either slept separately or together with the juvenile-I, if present.
In Lawachara, 47 sleeping trees (19 species) were located for
L1, and 52 (13 species) for L2, with 24 trees (11 species) for C in
Chunati. Some other sleeping trees could not be located due to the
gibbons’ fast and quiet movement in the different directions during
290
Figure 5. Pathways of group C during full-day follows (home range
area–dotted line).
the eleventh hour of active period. The distribution of sleeping trees
was not evenly dispersed within the home ranges.
Maintenance of territory: Hoolocks defend or maintain their ter-
ritory by:
1. loud morning song bouts;
2. visiting all parts of the boundary at regular intervals;
3. aggression through disputes with neighboring groups, calling
and/or chasing towards solitary intruders; and
4. gradual exclusion of maturing offspring from the territory.
TIME BUDGET AND ACTIVITY PATTERN
O ne important aspect of a species’ ecology is the proportion of
time that animals spend in different activities and the distribu-
tion of these activities throughout the day. Activity profiles are one
way to approach the problem of how animals budget their time and
may suggest how a species uses resources and adapts to its environ-
ment.
Start of activity period: It was not possible to determine at what
time a hoolock gibbon awoke, but the time at which it left the sleep-
ing site and/or tree was recorded. After leaving a sleeping site and/or
tree, animals urinated and defecated. All members of a group tend-
 IV. Field Research

Table 2
Home Range (HR), Full-day Follows Range Area, Territory Size and Core Area (in ha)

Group HR Home Range* Territory Size** Core Area

Area % of HR Area % of HR Area % of HR

L1 40.7 25.5 62.7 29.4 72.2 22.0 54.1

L2 86.0 52.0 60.5 73.1 85.0 49.3 57.3
C 25.7 17.2 66.9 - - 12.0 46.7
All 50.8 31.6 63.4 51.3 80.9 27.8 52.7

*From full-day follows

**Excluded group C, because the lack of home range overlapping with conspecifics

ed to become active within a few minutes of each other, and the gib-
bons moved away from the sleeping area together.
Overall, hoolock groups left sleeping sites –104 to +30 min
(mean 1 min, median 8 min, n=276) before/after sunrise and sleep-
ing trees –105 to +30 min (mean –2 min, median 7 min, n=276) be-
fore/after sunrise. Dispersal from sleeping sites and trees was highly
correlated with dawn and sunrise (rs=0.97, P<0.001).
The active period of Hoolocks was delayed until late dawn dur-
ing winter (January) due to heavy fog, and began early during warm
weather (June) due to early dawn.
Siamangs started activity 12 min before sunrise (Chivers
1974); while agile (Gittins 1979) and Kloss (Whitten 1980) gibbons
sing before dawn and start morning activity respectively 7 min and
2 min (male only) after dawn. The hoolock starts 2 min before sun-
rise, which is intermediate between larger (siamang) and smaller
(agile and Kloss) gibbons.

End of activity period: Overall hoolocks ceased their diurnal ac-

tivities 37–303 min (mean 153, median 148 min, n=330) before sun-
set and either instantly or a few minutes later entered into the
sleeping site. End of daily activity was highly correlated with sun-
set (rs=0.876, p<0.001).
Siamangs ceased diurnal activities and entered into sleeping
trees 120 min (at Kuala Lompat) and 126 min (at Ulu Sempam) be-
fore sunset, and into sleeping position 7 min thereafter (Chivers
1974). Kloss gibbons ended activities 85 min before sunset (Whit-
ten 1980), while hoolocks entered sleeping trees 153 min before
sunset and 5 min into sleeping sites, which is longer than any gib-
bon species.
Why do hoolocks retire so early in contrast to many other di-
urnal forest animals? The probable answer is that gibbons retire so
early to avoid overlap in feeding time with sympatric monkeys with
when their diets overlap (but whom they rarely meet)—having fed
enough by that time of the day by avoiding prolonged midday sies- Figure 6. Time spent (%) in different activities by different groups.
tas, characteristic of monkeys.

Time spent in different activities: Hoolocks spent most of their di-

urnal time (66.6%) eating foods (30.8% feeding and 25.8% forag-
ing) then resting (27.5%) (Fig. 6), which varied significantly among
groups and sites (c2-tests, p<0.001), but within site it varied only for
rest and play (c2-tests, p<0.001).
Age/sex classes variation in activity budget: Gross activity budg-
ets of different age/sex classes varied widely among three groups,
between two groups at Lawachara site, and all groups together (c2-
tests, p<0.05 to 0.0001).
Monthly activity pattern: For each group, each activity varied sig-
nificantly between months (p<0.0001) and there was also significant
variation between different activities for each group in each month
(p<0.001 to 0.0001).

291
The Apes: Challenges for the 21st Century
Overall traveling peak was in May 1989 (22.9%), feeding peak
during winter months (December-January [51.2% to 57.4%], little
variation among groups), foraging peak in June 1990 (41.7%), rest-
ing peak in July 1990 (35.4%), calling peak in February 1989
(10.5%) (i.e., end of winter, varied between groups), territorial peak
by L1 and L2 in August 1989 (7.7%), and playing peak was in July
1990 (9.2%).
Daily activity pattern: Overall time spent in different activities
during each hour of the day differed significantly (c2=27 to 437,
df=6, p=0.0001) except 1700 h (c2=10.545, p=0.1035). For each
group, each activity varied significantly between different hours of
the day (p=0.001). It also varied significantly between different ac-
tivities for each group in each hour of the active period (p=0.0001).
Two peaks of traveling pattern were found, highest (21.7%)
during the last hour (1600 h) and second-highest (19.1%) in the first
hour (0500 h). The prediction is that during the first hour, they trav-
eled more to have the first morning meal after the overnight fast.
Again, during the last hour they traveled more because they had to
find food to prepare for the overnight fast, and travel towards a suit-
able sleeping area.
A feeding peak occured in the morning at 0700 h (37.5%), and
a foraging peak in early afternoon at 1300 h (33.8%). A resting peak
was in the afternoon at 1500 h (34.4%). Hoolocks did not call after
1200 h (1200-1250 h) but called mostly in the morning. Territorial
behavior had a similar pattern to calling. Play occurred during rest-
ing times of the day and was mostly restricted to the young individ-
uals with a peak at 1000 h (5.3%).
When time budget of major activities in gibbon species are
compared (Ahsan 1994), it is found that feeding (feeding and for-
aging) resembles that of larger species (e.g., siamang) and resting
that of smaller species (e.g., agile). It is very difficult to draw a gen-
eral conclusion as to whether the hoolock resembles its larger or
smaller relatives.
DIET AND FEEDING
Food source: Hoolock gibbons eat a variety of plant foods (leaf,
liana, flower, nectar, and fruit, etc.) and animal matters of which
fruits are important and figs are dominant. Hoolocks used 101
species (48 genera and 32 families) of plants as food sources at
Lawachara (of which 10 were fig species), and 80 species (47 gen-
era and 34 families) at Chunati (of which 13 were fig species).
The actual number of food plants may be higher, especially at
Lawachara, because some of the plant species were not in the fruit-
ing stage during the study period. Also, some species could not be
identified. The number of plant taxa used by a gibbon species de-
pends on the diversity and density of food plant species. Hoolocks
of Assam (India) used 43 species (Tilson 1979), and of Tripura
(India) 13 taxa (Gupta pers. comm.). Ahsan (1994) compiled the
number of plant taxa used by gibbon species in the wild and found
a wide variation within the same species.
Feeding and foraging heights: The mean feeding heights were
14.3 m (median 14 m, range 4-37 m, n=4,882) at Lawachara, and
11.5 m (median 11 m, range 2-28 m, n=1,102)at Chunati; while the
total heights of plants were, respectively, 18.9 m (median 18 m,
range 6-40) and 17.6 m (median 17.6, range 4.8-45). Although food
height depends on various factors, both feeding and total heights of
two sites varied significantly (M-W U-test, 2-tailed, p<0.0001). On
the other hand, foraging heights were 12.8 m (median 12m, 4.2-
26.2, n=527 for L Groups), and 12.4 m (median 12 m, range 2-22,
292
n=205 for Group C), when the total heights were 17.3 m (median 18
m, range7-33.2) and 17.6 m (17.2 m, range 7.5-30) respectively for
those two sites, which did not vary significantly (M-W U-test, 2-
tailed, p>0.05). Actually, feeding and foraging heights depend on
the height of the food trees and as well as the height of the food
sources.
Feeding time: Of the total amount of time feeding (66.6%, 30.8%
feeding and 25.8% foraging), hoolocks spent 72.6% of time eating
fruit (41.4% figs, 34% other fruits, and 1.2% fruit exudates), 11.7%
on leaves, and 11.7% on other foods. The difference of fruit feeding
(including figs) among sites is not significant—80.7% at Lawachara
and 74.6% at Chunati, while the difference between the two groups
at Lawachara is negligible (L1 - 80.5% and L2 – 81%). On the other
hand, the difference of foraging time for fruit among groups is no-
ticeable (L1–48.1% and L2–57.8%).
The possible explanation for this variation during feeding and
foraging are that plant species’ richness and density of figs and other
fruits were higher at Lawachara (site for L1 and L2 Groups) than
Chunati (site for C group). L Groups had more opportunities to use
more of these types than Group C. Group L2 spent more time for-
aging for figs and other fruit because the larger group had to main-
tain a larger home range. This large group patrolled the area to
maintain possession of fruiting trees as their home range overlapped
with three groups.
The combined feeding and foraging time for different food
types can be compared (Fig. 7). Different age-sex classes did not
spend the same amount of time feeding and foraging on food types.
Figs and other fruits provided the main portion (more than 75%) of
the diet for all classes.
Proportion of time spent feeding and foraging is commonly
used in primate field studies, and it is a most useful technique in as-
sessing species diet importance. Figs and other fruit are frequently
eaten by hoolocks during feeding, leaves during foraging. Hoolocks
also forage on figs and other fruit, but their feeding bouts on leaves
are always shorter.
Monthly variation in food choice: Monthly diet of all hoolock
groups was dominated by figs and other fruit, both during feeding
and foraging. Groups L1 and L2 had a somewhat similar trend of
monthly fluctuations of different items during feeding, while Group
C did not. A similar trend was also found during foraging. When
feeding and foraging are combined, the variations among groups are
also noticeable, even within the same site (L1 and L2). If the two
sites are compared, the same trend of variations is found.
The relative proportion of dietary items (both phenology and
diet) throughout the year at both sites were highly significant (rs-
test, p=0.002 to 0.0001) except for mature fruit at Chunati
(p=0.0623). Based on the Shannon-Weiner index, there was highly
negative correlation between the diversity of mature fruits eaten and
the number of trees with mature fruits each month (rs=-0.867,
p=0.0001).
Diurnal variation in food choice: When hoolocks became active in
the morning after urinating and defecating, they almost invariably
traveled to a rich food tree and have a long feeding bout. In the later
few hours of the day, group members spread and foraged on small-
er food sources. Before retiring to the night tree(s), they always had
a long feeding bout as in the morning.
During full-day follows (i.e., without losing them between one
sleeping tree and the next), hoolock groups on average exploited 7.9
 IV. Field Research

Figure 7. Time spent (%) feeding, foraging, and combined feeding and foraging on different food types in two study sites, Lawachara (L), Chunati (C): (a) L
feeding; (b) C feeding; (c) L+C feeding; (d) L foraging; (e) C foraging; (f) L+C foraging; (g) L feeding and foraging; (h) C feeding and foraging; (i) L+C
feeding and foraging.

species of plants (median 8, range 2-17, n=81) per day, which did
not vary among groups (K-W test, H=0.802, p=0.67).
Generally, figs and other fruit are clearly dominant in the diet
during feeding, while animal matter is dominant over figs and other
fruit during foraging. If feeding and foraging are combined together,
the general trends remain the same. When compared between two
sites, the trends are: (1) figs and other fruit are dominant diets; (2) an-
imal matter and fruit exudate occupy a significant portion in the diet
of L Groups than the small amount in C Group; and (3) nectar occu-
pies a noticeable portion of the diet for group C, while in L groups it
is negligible. If the data for all groups are combined to produce an
overall diurnal diet of hoolocks, the following conclusion can be
made: (1) figs (41.4%) and other fruit (34%) are the most dominant
items in the diet of hoolock gibbon; (2) figs are highest in the first
(58.3%) and last (52%) hours, while other fruit are less (29.9% and
29.6% respectively) in these two hours; (3) the proportion of leaf ma-
terials increases through the day and is proportionately high between
the very late morning and afternoon; (4) the proportion of animal
matter is relatively high from morning (not early morning) until af-
ternoon; and (5) fruit exudate and nectar are variable.
Fecal analysis: Most of the time it is extremely difficult to collect
entire fecal samples from wild gibbons. Heavy seeds tend to sepa-
rate from the stools as soon as they appear and the remainder fre-
quently shatter on the twigs, branches, and leaves when falling, at
best being scattered over the forest floor and at worst never reach-

293
The Apes: Challenges for the 21st Century

ing the ground. However, it was possible to collect 33 fresh samples Squeal: a fear note of subordinate member; and (11) Grunt: territo-
of feces for three groups during different hours of the day. More- rial dispute note by adult male.
over, 180 samples of feces (including those 33) were partially or
fully analyzed to identify the plant materials, especially seeds, Variation in the units of song bout (Fig. 8): Although a song bout
which were passed through the guts of the hoolocks. Forty species can be divided into a collection of three major types (song/call
(22 families) of plant taxa (including all fig species considered as units)—great call, organizing sequence, and alarm call—but to
one) were identified from seeds and undigested food in the feces. compare intergroup individual variation others are also included.
The actual number of plant species will be more as seeds swallowed
1. Female small tuunk—1 to 5 notes (mean 2.9, n=23) in one song
by the hoolocks could not be recovered.
bout;
Food passage rate: Food passage rates of hoolocks were measured 2. Male tuunk—94 to 594 notes (mean 269.5, n=30) in one song
for 30 different species (20 families) of food plants. It seems to be bout;
that the passage rates do not vary markedly between age/sex class-
es except in two cases. This variation may be possibly due to the 3. Great call—3 to 21 units (mean 10.5, n=10) in one song bout;
physiology of the individual animal and at times undigested food 4. Organizing sequence—7 to 33 units (mean 18.3, n=10) in one
particles are retained in the gut, which delays the passage rate. Out song bout; and
of four selected species, the food passage rate between human and
hoolocks varied significantly for only Sapium baccatum (Family 5. Alarm call—0 to 27 units (mean 5.3, n=10) in one song bout.
Euphorbiaceae) (c2=3.919, df=1, p<0.05).
Distribution of song bouts: Groups L1 and L2 delivered the high-
Germination trial: The germination success ranged 10 to 100% est number of song bouts between 0800 and 1000 hours in the
and was higher in the garden trials than in the forest. This variation morning, which comprised 62% and 70% of the total bouts, respec-
occurs simply due to the fact that in nature (forest floor) most of the tively. While Group C had peak between 0700 and 1000 hours
seeds were either lost, damaged by unknown predator(s), or washed which constituted 81% of the bouts.
out or inundated by rain water, while in the garden it was controlled. The distribution of songs varied among groups throughout the
However, in both places 100% germination was recorded for Prunus day due mainly to territoriality and distribution of food sources, and
ceylanica (Family Rosaceae), possibly because these seeds, which for SF (solitary female) it may be that she is alone. Except for C
are comparatively hard, have a high chance of germination and less
chance of being damaged by the gibbons’ teeth.
Although germination rate was low (10-20%), it proves that the
seeds (of some species of plants) recovered from gibbons’ feces
may be viable for at least one year. On the other hand, the seeds,
which passed through the guts of gibbons, started germinating ear-
lier and also had higher rates of germination than those seeds, that
were gathered from the trees.
Therefore, fecal seed viability lasts for at least one year. Early
germination of fecal seeds compared to seeds from trees signify the
hoolock gibbons as seed dispersal agents, and hence, play a signif-
icant role in the process of natural forest regeneration.

SONG AND SINGING BEHAVIOR

H oolock’s song/call is a complex one, and it is difficult to know

whether sounds are different or if different spellings are sim-
ply due to different cultures. However, the following distinctive
units and notes of hoolocks were recognized: (1) Female growling:
preparation of pre-opening sequence of 1 to 8 min (mean 4.6 min,
n=10) duration; (2) Hoo: a short and single-syllable note; (3) Great
call: a regular series of sudden, very rapid disyllable notes rising
very quickly to a climax then, after a short while, descending; (4)
Semi-great call: rapid who-as; (5) Repair and aborted call; (6) Or-
ganizing (interlude) sequence—(a) female tuunk: like hoos; (b) hoo-
u: longer than a single (hoo-yu or hoo-oo); (c) hoouk: deeper than
hoos of different combination (hoo-u, hu-uk, hoo-hook, hoo-yu-
hook, etc.); (d) whoo: an exhalation mostly given by female at the
end like oowhoo-u; (e) hoo-koo: a figure of 2/3 notes together, like
hoo-koo-yu, hoo-koo yu whoo, whoo-koo-oo, who-koo-yu, hoo-u-
hook; and (f) tuunk: a high pitch—(f.1) male tuunk: a high- and
deep- pitch note and (f.2) others high-pitch note (tuunk): given by
other than male member; (7) Alarm call: rapid u-hoo; (8) Cry: dis-
tressed wailing note; (9) Whimper: given by juvenile male or Figure 8. Variation in the units of song bout (top-bottom: female small
subadult female when approaching adult of the same sex; (10) tuunk, male tuunk, great call, organizing sequence and alarm call).

294

Group, all other (including SF) groups home ranges overlapped with
neighboring groups. If there was intergroup competition for re-
sources, they called throughout the day as a measure of defense.
Group C did not have such a competition problem; which may be the
reason that they did not call after 1200 h. Further, SF also called in
the early morning to advertise her loneliness during a quieter period.
Starting time of the first call of the day: The median time when
the first call of the day was given varied considerably among
groups, which showed a significant positive correlation with dawn
and sunrise except for Group C. The monthly median start of the
first call of the day varied considerably according to dawn and sun-
rise, which showed a significant positive correlation in both sites
(p<0.001).
Overall the median starts of first call of the day were 0910 h,
0915 h, 0656 h, and 0829 h respectively for L1, L2, SF, and C
Groups. SF started to call first and C was last. None of the groups
(including SF) called from the sleeping sites/trees on the following
morning, but the agile (Gittins 1979) and Kloss (Whitten 1980)
called from sleeping sites of the previous night before starting their
other diurnal activities.
Frequency of singing: Hoolocks did not sing every day. On aver-
age, Group L1 sang two days out of three, L2 three out of five, SF
three times a day, and C sang once every three days. The mean fre-
quency of singing did not vary significantly among groups (L1, L2,
SF, and C) (c2=4.517, df=3, p>0.05).
Amount of time spent singing: Overall, SF spent the most time
singing, 23.8 min and C the least, 7.1 min each day (while L1 spent
11.3 min and L2 17.7 min); (ANOVA one factor analysis F=7.59,
df=3/67, p=0.0002). L1 and L2 showed similar patterns of variation
in time spent singing while others varied.
Duration of song bouts: Overall, the mean song duration in L site
(except SF) was 14.4 min (range 1-44 min, n=430) and in C site 21
min (range 5-42, n=21). Monthly mean duration of bouts differed
significantly among groups (ANOVA one factor analysis F=7.081,
df=4/86, p<0.0001).
Singing sites: Singing sites comprised tall or emergent trees on the
hillslope or hillocks, mostly located near boundary or overlapped
home range areas. Altogether, 41 singing sites for L1, 25 for L2, and
14 for C Groups were located. In the overlapped home range area of
L1 and L2, five sites were common for both groups.
Hoolocks never sang from their sleeping sites on the following
morning, although sometimes later in day, they sang from sleeping
trees that they had used a few days prior. Agile (Gittins 1979) and
Kloss (Whitten 1980) gibbons sang from sleeping sites on the fol-
lowing morning. The distribution of singing and sleeping trees was
analyzed within the home ranges and found that singing sites are
mostly concentrated in tall trees near potential food sources areas at
the edge boundary (Ahsan 1994).
Counter singing and circumstances eliciting duets: L Groups
sang in several situations. Eliciting circumstances noted were: (1)
adjacent group singing near edge boundary; (2) territorial conflict
within overlapped home ranges; (3) competition over shared food
sources; (4) neighbor group (whose home ranges did not overlap)
singing; (5) responses to songs or calls of other animal species; and
(6) sudden train whistles when gibbons were close to the railway
line. Animal calls that elicited gibbon songs were: (a) domestic fowl
295
IV. Field Research
(Gallus gallus domesticus); (b) red jungle fowl (G. g. gallus); (c)
crested serpent eagle (Spilornis cheela); (d) jungle crow (Corvus
macrorhynchos); (e) kalij-pheasant (Lophura leucomelana); and (f)
Indian pied hornbill (Anthracoceros malabaricus). Other bird
species called close to gibbons, but the gibbons reacted only by
growling. They also growled when train whistled and trucks hooted
more than 100 m away.
Factors influencing singing: Environmental factors such as rain,
cold, thunderstorms, and food availability influence hoolock singing
behavior, of which three major weather conditions—rain, cloud
(overcast sky), and clear (sunny sky) were most important. Overall,
at L site hoolocks sang on 60.7% of days during the study period
and in C site on only 25% of the days—less at C because of the
death of the mated female and lack of territorial conflict with neigh-
bors. Hoolocks sang on more than 50% of the sunny days and either
did not sing during rainy days (as in Chunati) or on less than 10%
(9.1%) (as in Lawachara). When gibbons sang during bad weather,
it varied considerable from the normal singing but did not vary sig-
nificantly in the duration of song bouts between sunny and cloudy
(includes rainy) days within groups (M-W U-test, p>0.05).
On days with either thunderstorms or heavy rain, hoolocks did
not sing at all (Gittins and Tilson 1984). Rain and cloudy weather
also depressed calling in siamang, lar, and agile gibbons in Malaysia
(Chivers 1974, 1978; Gittins 1979).
Functions of hoolock gibbon songs: The present study suggests
that hoolock songs showed signs of the following functions: (1)
maintenance of pair bond during duets; (2) concentration and rein-
forcement of social ties within group; (3) mate attraction and pair
formation; (4) mate defense; (5) advertisement for a mate; and (6)
territorial advertisement, defense, and fitness.
TERRITORIALITY AND MONOGAMY
T erritoriality and monogamy are the basic features of gibbon so-
ciety. Functions of territoriality are probably multiple and the
most important are: space, pair-mate, food and shelter. While the
functions of monogamy are not very clear, the important factors in-
clude: exclusive breeding access to the pair-mate, mate guard, and
aid in defending territory from conspecifics. Territoriality and
monogamy were only studied in Lawachara because Group C lived
in an isolated forest pocket.
Two groups (L2 and L3) and two solitary females (one exclud-
ed from L1) were involved in territorial disputes with L1. With L2,
territorial disputes occurred with three groups (L1, L3, and L4) and
the same two solitary females. The home range of one SF did not
overlap with the home range of L2, but did with L1.
Territorial behavior: Usually disputes start when two groups range
close to the boundary of their territories and see each other. Disputes
start less frequently when one group hears its neighbors singing near
a boundary. There were 11 records of one group singing within the
overlapped boundary of the neighbors. The neighboors then ceased
activities and moved very fast to reach to its own boundary.
Although the adult males were the main protagonists in terri-
torial disputes (80% cases), occasionally other members joined in.
Interestingly, when L1 and L2 adult males were engaged in territo-
rial disputes in the upper and/or middle canopies, the juvenile male
of L1 and the juvenile female-I and II of L2 were found to play to-
gether in the lower canopy during most of the dispute observations.
Overall, 2% of their diurnal times, L1 and L2 were spent in territo-
rial dispute.
The Apes: Challenges for the 21st Century
Upon detecting each other, the two males approach each other
and sit in the high canopy silently about 5–25 m apart (closest one
recorded was about 2 m in the same tree without severe physical con-
tact) in the same and/or different trees depending on the location,
branching systems, and canopy area of the tree(s) from which they
display. Usually females and immature rest about 10–30 m behind
the respective males and do not take part in the proceedings, except
in a few cases. Often they drift away after 10–30 min and continue
foraging, while young play with siblings or even sometimes with
young of other groups. Males display in different ways: (1) locomote
on branches and return to the same point repeatedly; (2) climb up and
down the trees(s); (3) try to be higher (in height) than the opponent;
(4) fluff body hair; (5) hang with outstretched limbs like a “star;” (6)
present a large surface area, as much as possible, to the opponent; (7)
stand on a branch supported by arms and/or hanging; and (8) display
scrotum and/or back to the opponent. All these behaviors seem to be
rather casual, but close observations reveal their movement to be
rigid and lacking their natural rhythms.
After 10–30 min of display, chases occur back and forth across
the boundary. Often these are proceeded by mock charges, but not
deep into each other’s territory. These chases happened so fast that
it was difficult to observe details through the foliage and branches.
They appeared to be displays of strength without physical contact.
Besides females, only the juvenile-I and II of L2 occasionally
took part in the territorial disputes standing behind their fathers.
Both females of L1 occasionally participated in territorial disputes
by displaying their backs to opposing groups.
Once after 20 min of disputes between males of L1 and L2, the
light colored female of L1 initiated an intense charging/chasing
bout. The event happened so quickly that it could not be followed
properly as trees and foliage prevented precise observations. At that
time, fighting, screams, squeals, and whistles of several animals
were heard. After that event, juvenile-II of L2 produced distress
shrill-hoos for the next 10 min. Several animals dropped partially or
completely to the ground during the first bout of chasing/fighting.
No serious injury to any individuals was observed.
Song: Morning loud songs of hoolocks are the best means of ad-
vertising occupancy and defending territory. Besides the usual
singing encounters, in 75% of encounters singing occurred imme-
diately pre-or post-dispute time. Overall, 41% of the dispute cases
ended through counter singing.
Intergroup encounters: Of the 45 territorial disputes recorded dur-
ing scan samples, 82% occurred between groups L1 and L2.
Frequency of territorial disputes: Overall, territorial disputes be-
tween L1 and L2 occurred two out of five days and about 91% of
disputes happened before 1200 h of the day. There were marked dif-
ferences in the monthly distribution of the number of disputes be-
tween L1 and L2. No dispute occurred in November 1989 possibly
due to the abundant supply of teak skeletonizer caterpillar (Hyblaea
puera) which the hoolocks relish. Disputes increased markedly dur-
ing March 1990 because many figs (Ficus gibbosa and F. variega-
ta), the two preferred food species for L Groups and jora gota
(Prunus ceylanica), the fifth preferred fruit, had ripe fruits within
the overlapped home range of L1 and L2.
Duration of territorial dispute: The mean duration of territorial
disputes involving L1 and L2 and their neighbors was 56 min (me-
dian 50 min, range 6-138 min, n=37), which did not vary signifi-
cantly (M-W test, U=189, z=1.203, p=0.2289). Furthermore, there
296
was no significant difference in the duration of territorial disputes at
different times of the day (K-W 1-way ANOVA, H=6.133, df=9,
n=45, p=0.7165).
Role of food sources: Occupancy of food sources is the main cause
of territorial disputes. It was found that hoolocks did not visit some
places of their home ranges until they was a potential and favored
food item available. When one of these places had ripe figs (espe-
cially F. gibbosa), almost every day one or both groups used that
source separately. For example, 42% of the total disputes (n=48) oc-
curred for the competition of small figs (F. gibbosa) and the rest
(58%) for other food sources.
Role of sleeping trees: Sometimes hoolock groups slept in the
perimeters of their home ranges and the following morning they
traveled long distances either to patrol and/or to forage for foods at
the other end of their home ranges. Furthermore, members of a
group (when more than two) dispersed in different sleeping trees
covering a large area for the same night. This may be a hoolock
strategy to maintain sole ownership of the area while minimizing
energy used in territorial disputes.
Monogamy: Like other gibbons, hoolocks maintained monogamy
through exclusion of potential reproductive females. This occured
in Group L1, and later the L2 subadult female left. Of the five hy-
potheses given by Wittenberger and Tilson (1980), the last two (H4:
monogamy should evolve though polygyny threshold is exceeded if
aggression by mated females prevent males from acquiring addi-
tional mates; and H5: monogamy should evolve when males are less
successful with two mates than with one) fit in this study. First,
while the group RLB male was copulating with a solitary female,
the growling warning of the RLB female brought him back to her.
When the L1 group had two females, neither female gave birth (i.e.,
they were not reproductively successful) which fits the H4 hypoth-
esis. After exclusion of one female from L1, the resident female
gave birth, which fits the H5 hypothesis. The instance of the expul-
sion of the L2 subadult female also fits the definition of monogamy
given by Emlen and Oring (1977); monogamy occurs when “neither
sex has the opportunity to monopolize additional members of the
opposite sex.”
Ecological factors: Ecological factors such as habitat disturbance,
population size, forest fragment, etc. influence hoolocks’ social sys-
tems to be monogamous or polygamous. For example, the adult fe-
male of Group C (living in a man-made fragmented forest patch) did
not die and the juvenile female (who formed a group with her fa-
ther) had to stay with the natal group, which would possibly direct
the adult male to polygyny.
Availability of opposite sex: An excess of females may lead to
males being involved in extra-pair copulation (e.g., L1, RLB, and
V1) and a shortage of males may initiate polygyny, either through
an incestuous conception or brother-sister pair formation (e.g., L1
group). A shortage of females may allow father-daughter pair for-
mation (e.g., C Group). When a female is sexually unreceptive (ei-
ther in advance stage of parturition or with an unweaned infant), the
biological needs of a male may lead to cuckoldry or incestuous sex-
relationship either with floating females or adolescent daughter(s)
(e.g., RLB and L2). Finally, possible small scale human predation
may influence the social system of hoolocks and may often lead to
extra-pair copulation (e.g., excess females in Lawachara may be one
of the probable reasons).

HUMAN IMPACT
T he reasons for rapid deforestation taking place in Bangladesh
are complex. The physical alterations to the landscape of
Bangladesh are largely due to human impact. The rate of forest con-
version has reached levels at which environmental catastrophe is
likely. The principal causes of deforestation in Bangladesh are: (1)
small land area and limited resources; (2) over population; (3) clear-
felling and subsequent planting of short-rotation exotic tree species;
(4) illegal timber extraction, poaching and over-exploitation of fire-
wood for commercial purpose; and (5) poor management.
Bangladesh is a small developing country, having an area of
144,054 km2 including about 7% forest coverage. It is densely pop-
ulated having 764 people/km2 totaling 110 million in 1991 census
(BBS 1992); 85% of the people live in the rural communities, farm
or keep livestock (21.5 million cattle, 0.6 million buffalo, 13.7 mil-
lion goats, and 0.7 million sheep [BBS 1983-84]). The demands of
people and livestock on the land have marked effects on the decline
of the country’s forest cover. Annual losses of closed forest in
Bangladesh averaged 0.4% from 1981 to 1985, yielding a total loss
of 80 km2 (IIEDWRI 1986).
Population growth, economic opportunities, and political prob-
lems stimulate the conversion of forest to other uses. Due to short-
age of land, reforestation schemes cannot expand properly (other
than roadsides). Impoverished, often landless, rural households have
moved into forested regions in search of productive land. For in-
stance, during the late 1970s and early 1980s the government of
Bangladesh established new settlements of the landless peoples, es-
pecially those displaced by riverbank erosion areas into the natural
forests of the greater Chittagong Hill-Tracts Districts to control po-
litical issues. Government policies frequently ignore the efficient
use of resources and overemphasize timber harvest at the expense of
other political benefits.
The other important human impacts on the two study sites are
discussed below.
Traditional use: Both Lawachara and Chunati forests receive in-
tensive use by human and livestock. Some people, living in those
areas, are mostly pastoralists, owning herds of cattle, buffalo, goats,
and sheep. Lawachara forests for instance, are used daily to supply
supplementary foods and to provide medicinal plants. Other tradi-
tional uses of the forest include growing betel leaf and betel nut in
the forest, cultivation of agricultural crops through clearing the for-
est edge and patches, and heavy fuel wood extraction. In Chunati,
forest edges and patches have been cleared to established betel leaf,
betel nut gardens, and rice fields. Regenerated trees and bamboos
are used for fencing and poles, and roofing as sunshade for the gar-
dens. Fuel wood extraction is also common.
Shifting cultivation: When an area is commercially logged, vege-
tation is cut and burned. Sometimes rice is cultivated, and vegeta-
bles grown between the remaining stumps. Simultaneously, the
forest department plants forest tree species, but the policy of plant-
ing short-rotation tree species makes the situation worse than shift-
ing cultivation.
Introducing exotic tree species: Natural and secondary forests,
and wildlife habitats are declining due to the introduction of exotic
tree species through clearing existing forests and practicing mono-
culture (e.g., teak [Tectona grandis]). Malakana (Paraserianthes
falcatoria), eucalyptus (Eucalyptus spp.), ipil ipil (Leucaena leuco-
cephala), minjiri (Cassia simea), pine (Pinus spp.), akashmoni
297
IV. Field Research
(Acacia monaliformes), and mangium (Acacia mangium) are the
most important exotic species planted in Lawachara since 1974.
Malakana is being harvested in 10-year rotations and being supplied
to the plywood factory at Sylhet. This is a continuous process that
may induce harmful effects on the habitat. Harvesting these short-
rotation species through clear-felling and then slash-and-burn sys-
tems changes the landscape of West Bhanugach Reserve Forest. It
increases the amount of silt entering rivers, especially during heavy
rains. As a result, wildlife is concentrated into a limited area which
increases inter- and intraspecies interactions and may result in the
loss of certain species from that area.
Fuel wood extraction: In Bangladesh, about 24.2 million people
rely on fuel wood ([Ahsan 1995] complied with the BBS 1983-84
and 1991 census [BBS 1992]). Fuel wood is gathered daily from
Lawachara forest by every family living in the neighboring villages,
at least at the rate of 5 kg/family/day (Ahsan 1995). Removing dead
and fallen trees for timber and fuel wood does not contribute direct-
ly to the killing of trees but has a harmful effect on the soil, depriv-
ing it of nutrition and ground cover. Cutting naturally grown trees
for fuel wood has a tremendous effect on the forest, primates, and
other wildlife. My field experience in Bangladesh since 1979 sug-
gests that fuel wood used by the forest villagers for their own cook-
ing has not affected the forest, but the illegal commercial
exploitation of fuel wood by cutting standing trees has had a
tremendous effect. The forest canopy becomes open, resulting in
more bushy areas (Ahsan 1995).
About 170 people (both sexes) collected fuel wood daily from
the Lawachara Beat area (3 Beats form Moulvibazar Range) and sell
it on the local market. On average > 50 people gathered fuel wood
every day from my study block (approx. 120 ha) with an estimate of
1,250 kg/day and annually about 456 tons to 1,330 tons from the
whole Lawachara Beat area.
On average, in the Chunati study block, two people (30 kg each)
collected fuel wood daily and supplied it commercially to the tea-stall
nearby, with an annual total of 22 tons. Some illegal wood goes to the
brickfield as well, but estimating that amount is difficult. The study
block also fulfills the demand for fuel wood of the local farmers.
Extracting tree bark: In Lawachara, a small group of people col-
lects bark from some tree species and sells it to local agents. Mos-
quito coils are prepared from this bark by private, small cottage
industries for commercial purposes. The tree species used are main-
ly Litsea sebifera and Dillenia pentagyna, but small amounts are
used from Garcinia cowa, Garcinia sp., and Microcos paniculata.
Thisextraction is illegal and damaging as people cut the bark from
the entire bole depending on the stem. A few months later most of
these trees slowly die. Then the fuel wood collectors cut down the
dead trees for commercial purposes.
Extracting timber: The traditional houses in the Sylhet area are
supported by a framework of wood with a roof of corrugated tin. In
the frame, trees of 15-30 cm dbh (diameter at breast height) are used
as poles instead of pillars and beams. Sylhet villagers and some peo-
ple from nearby small towns collect their building materials illegal-
ly from the surrounding forest. Alhough people use commercially
less important tree species for building frameworks, they are im-
portant ecological components of the natural forest. Doors, win-
dows, and household furniture are also made from timber. Several
factors that encourage illegal timber extraction from the forests of
Bangladesh, including illegal timber trade, local unemployment,
The Apes: Challenges for the 21st Century
mutual monetary benefits between poachers and foresters, and inef-
ficient implementation of forest law.
Livestock herbivory: The actual effect of livestock grazing in and
around two of the study sites has not been quantified, but some of
their feeding activities can be discussed. Around my field station in
Lawachara, three cows destroyed most of the seedlings of the silvi-
culture nursery and some new plantations. Hundreds of cattle tem-
porarily live inside the forests of Kalachara (about 2 km north of the
study area). In Chunati, livestock mostly graze on browse or grass
in or around the study site. Feeding behavior of livestock may be in-
fluenced to some extent by competition for food with primates be-
cause some plant species are common to both. When herbivores are
highly generalized in their food choice, they remove certain critical
plant species that negatively affect the food chain.
Betel leaf plantation: Betel leaf plantations inside or at the edge of
forests also reduces wildlife habitats. In Lawachara, the Khasia tribe
have Khasia-variety betel leaf plantations as their livelihood. The
groundcover is cleared, and the vine is grown creeping up the tree
trunk. The main trunk of all trees inside the plantation is used and
large branches are lopped off. Since these trees are mostly bare, they
do not produce fruit and primates avoid these areas. There are sev-
eral plantations spreading over 10 ha in Lawachara. These planta-
tions last for several years until an epidemic disease infects them.
New plantation areas are then acquired (and sometimes new resi-
dences are also established) and the old ones abandoned.
A different variety of betel leaf vine is grown in Chunati in a
different way. An area is cleared either in or by the forest, then
fenced and roofed with bamboo cane or sticks. All fencing, roofing
and supporting materials are gathered from the forests using bam-
boos and regenerated trees. These gardens vary in size depending on
the demand and usually last for 2 to 5 years. There were 14 such
gardens (about 1 ha) in and around 26 ha block of my study area,
and 215 gardens (782 ha) within the whole Satgarh Beat. As there
are no trees inside such gardens, the habitat of arboreal animals is
reduced by these activities.
Hunting, capturing, and direct harassment: According to the
Bangladesh Wildlife (Preservation) (Amendment) Act, 1974, cap-
turing, selling, and hunting, or any type of harassment of wildlife is
prohibited in the country. In both study sites, primates are not hunt-
ed for meat due to religious beliefs. Occasionally farmers kill pri-
mates and hang them on a tree to discourage remaining group
members from crop raiding, as I observed in Chunati. During my
Lawachara study, I met hunters carrying guns six times (2 to 4 peo-
ple with one gun) who were searching for barking deer (Muntiacus
muntjac). In Chunati on eight occasions I witnessed dozens of peo-
ple with 2 to 3 guns searching for a barking deer. Other wildlife
hunted in both sites are red jungle fowl (Gallus gallus gallus) and
kalij-pheasant (Lophura leucomelana). Occasionally, Khasia tribe
members in Lawachara hunt primates and birds for meat.
Tourism: Tourists bring their unique impacts on forest flora and
fauna. At Lawachara, most come to visit for the day, to spend some
time in a quiet place close to nature, or for recreation, education,
picnics and outings. Tourists of various professions visited
Lawachara mostly for recreational purposes. The highest humber of
tourists visited on a weekend (Friday) during the drier months (No-
vember to March). Usually 2 to 5 groups of visitors came by private
buses and spent some time watching the surroundings. Each group
comprised 15 to 100 people between winter months, November
298
1988 to December 1990. A small number stayed in the Shamoli rest
house and only their representatives wrote comments in the visitors’
book. Other visitors, including forest high officials who visited the
area and spent the night in the rest house, wrote comments in the
visitors’ book. From their comments (between 1952 and 1984),
Chowdhury (1986) determined that the number of visitors in-
creased, while the sighting of wildlife has decreased through the
years. By summarizing visitors’ comments, he suggested that visi-
tors to the rest house are steadily increasing, big game has definite-
ly declined in numbers, and the enjoyment of tree and plant lovers
is gradually decreasing. The last two points indicate that some
deterioration of the natural environment has occurred. Major envi-
ronmental impacts of tourists in Lawachara include leaving nonbio-
degradable wastes, harassment, and polluting the stream source of
drinking water.
Food source: People also consume some of the fruits in the forest
that are eaten by wild animals. A few of them are also occasionally
sold in the local markets. The collection of fruits (e.g., Syzygium
cumini and Caryratia trifolia) occasionally creates direct competi-
tion between man and monkeys. If both species are simultaneously
in a tree, the monkeys are either chased by shouting humans or
move off quickly.
Other factors: Fuel wood collectors use machetes for cutting down
trees, cutting branches, or trimming wood for bundles. The sound
from their cutting activities certainly influence the normal behavior
of gibbons and other wildlife. Gibbons are disturbed by this noise.
For example, when a gibbon family was feeding or approaching a
feeding source, if such noises were made repeatedly, either beneath
or close to the feeding source, the group immediately left the area.
Gibbons were also disturbed by the noise made by public buses, lor-
ries, and trains.
Usually once a year during summer months, some portion (5 to
10%) of the study block in Chunati is affected by man-made fire.
Occasionally fire also causes a considerable damage to the silvicul-
ture research plots in Lawachara. This is done intentionally for the
benefit of the fuelwood collectors because after fire, dry woods and
bamboos can be gathered. Fire may also cause the loss of certain
species of plants (ground and understory) from this area.
CONSERVATION
Threats: Major factors of habitat loss for wildlife in the study sites
are:
1. Pressure on land and demand of forest products squeezing for-
est habitat;
2. Plantation of short-rotation exotic tree species through clearing
primary and/or secondary forests;
3. Destruction of forest through timber poaching;
4. Overexploitation of fire wood for commercial purpose;
5. High human population pressure and limited resources; and
6. Inadequate protection and poor management system.
Recommendations: The following recommendations have been
suggested to conserve biodiversity in the two study sites. These are
also applicable for other areas of Bangladesh:
1. Plantation of short-rotation exotic tree species should be stopped
or at least be selective;

2. Timber poaching must be stopped;
3. Commercial exploitation of fire wood must be stopped;
4. Ensure protection in reality;
5. Public awareness about forests and wildlife program should be
extended to a grassroots level;
6. Wildlife circle must be revived with sufficiently trained man-
power as separate entity from the forest department.
Acknowledgements: I thank The Association of Commonwealth
Universities, International Primatological Society, World Wildlife
Fund-US, Conservation International, Worts Travelling Scholars’
Fund, Chittagong University, University Grants Commission
(Dhaka), and The British Council (Dhaka) for their generous help
during various stages of this work. I gratefully acknowledge The
Chicago Zoological Society and Brookfield Zoo for inviting me to
The Apes: Challenges for the 21st Century conference and also for
their generous logistic support.
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