International Journal of Primatology, VoL 17, No.

6, 1996

Lion-Tailed Macaques (Macaca silenus) in a

Disturbed Forest Fragment: Activity Patterns
and Time Budget
Shaily M e n o n l~ and F r a n k E. Poirier l~
Received January 4, 1995; revised May 22, 1995, and June 13, 1996; acceptedJuly 1, 1996.

We describe the activity patterns and time budget of a feral group of lion-tailed
macaques that were confined to a disturbed forest fragment of 65 ha and
compare the results with those obtained for groups in protected forests. The
degraded nature of the study site was reflected in low tree densities, low specific
diversity, gaps in the girfh distribution of trees, and frequent disturbance by
humans. The study group of 43 subjects was twice as large as lion-tailed
macaque groups in protected habitats. They spent the most time ranging
(34.0%), followed by foraging (23.7%), feeding (17.9%), resting (16.0%), and
other activities such as social interactions (8.4%). Monthly variations are
significant for all activity categories except ranging. Times spent resting and
foraging are negatively correlated (r = -0.5) and show significant seasonal
differences. Foraging time was highest from September to November, when key
food sources such as C u l l e n i a and Artocarpus were absent or marginally
available. The study group spent most time (40.4%) at canopy levels between
21 and 30 m. They spent more time each day ranging than resting or feeding
and more time terrestrially compared with groups in protected forests. Large
group size, poor habitat quality, and seasonal variation in food availability
were the major variables affecting their time budget, and these variables
accounted for differences from the time budgets of groups in protected forests.
KEY WORDS: lion-tailed macaque; Macaca silenus; forest fragment; time budget.

IDepartment of Anthropology, The Ohio State University, 245 Lord Hall, 124 West 17th
Avenue, Columbus, Ohio 43210.
2present address: Department of Biology, University of Massachusetts, Boston, Boston,
Massachusetts 02125.
3I'o whom correspondence should be addressed.

969
0164-0291/96/1200-0969509.50/0C 1996 Plenum PublishingCorporation
970 Menon and Poirier

INTRODUCTION

Lion-tailed macaques are endemic to the Western Ghats, India. Dur-

ing the fourth international symposium on lion-tailed macaques, the
working group on census and distribution estimated that 3660 individuals
(183 groups) survive in the wild (Anonymous, 1995). This estimate is > 10
years old, based on fieldwork conducted in the 1980s. Previous researchers
examined the time budget and activity patterns of lion-tailed macaque
groups inhabiting large protected forests (Kumar, 1987; Kurnp and Kumar,
1993). However, 40-50% of the total feral lion-tailed macaque population
inhabits forest fragments ranging from 0.5 to 20 km 2 (Kumar 1987). Many
of the smaller forest fragments are isolated, degraded, and subjected to
ongoing disturbances. The management of groups confined to forest frag-
ments has implications for the long-term conservation of the species.
Primate species inhabiting increasingly isolated and marginal forests
face several ecological constraints: restricted home ranges, low tree densi-
ties, low food availability, high predation risk, and increased hunting
pressure. These constraints impinge on the diet, mortality rates, reproduc-
tive rates, and ultimately survival How primate groups apportion their time
can provide clues to the species' ability to survive under such constraints
and to the ecological limits of their tolerance. Time budgets and activity
patterns of primates are influenced by group size (van Schaik et al. 1983;
Stacey, 1986; Altmann, 1987, Watts, 1988; Barton et al., 1992; Dunbar,
1992; Isbell and Young, 1993; Waser, 1977), habitat quality (Oates, 1977;
Marsh, 1981a; Iwamoto and Dunbar, 1983; Watts, 1988; Dunbar, 1992; Is-
bell and Young, 1993); and proximity to human settlements and/or
provisioning (Altmann and Murnthi, 1988; Eley et al., 1989, Singh and Vi-
nanthe, 1990).
We describe the activity patterns and time budgets of a lion-tailed ma-
caque group in a forest fragment and compare the results with those
obtained for groups in protected forests.

STUDY SITE

Much of our knowledge about feral lion-tailed macaque activity pat-

terns and ecology comes from intensive ecological studies conducted in the
Anaimalai region of the Western Ghats. The Anaimalai Wildlife Sanctuary,
established in 1972, was later renamed the Indira Gandhi Wildlife Sanctu-
ary and National Park. The sanctuary is hilly with an altitudinal gradient
of 350 to 2500 m above sea level. Anakunthi and Varagaliar are two of
several relatively undisturbed forests inside the sanctuary containing lion-
 Lion-Tailed Macaques in a Forest Fragment 971

tailed macaque groups. Anakunthi and Varagaliar forests have areas of 2.34
and 16 km 2, respectively, and occur at an elevation of approximately 650
m a.s.l. Both are contiguous with larger protected forest tracts; they are
the sites of previous ecological studies on lion-tailed macaques (Kumar,
1987; Kurup and Kumar, 1993). Outside the sanctuary is a heterogeneous
matrix of cashcrop cultivation, human habitation, and forest fragments cre-
ated during the previous century when large forest tracts were converted
to tea and coffee monocultural plantations. Several of them retain isolated
patches of rain forest. The main study site--Puthuthotam Cardamom For-
est (PCF)--is a 65-ha disturbed forest patch surrounded on three sides by
tea and coffee plantations and on one side by a heavily used road. PCF is
located at 10~20'N and 76~ approximately 12 and 17 km south-east of
Varagaliar and Anakunthi forests, respectively. PCF is at a higher elevation
of 1085 m a.s.l. It is a private forest underplanted with cardamom and cof-
fee. The forest floor is cleared for planting, but mature trees remain to
provide shade and humidity. PCF also has a history of selective logging.

STUDY GROUP

The lion-tailed macaque study group at PCF included 43 individuals.

It is unusually large compared with groups in the relatively undisturbed
Anakunthi and Varagaliar forests. The age-sex composition of the lion-
tailed macaque groups, however, was similar at PCF and in the protected
forests (Table I). The following demographic changes occurred in the group
during the study period: In December 1990, when consistent counts were
obtained for the total number of animals and the age-sex classes, the group
comprised 41 animals. There were four infants, one of which was newborn
and three >6 months. By June 1991, the 3 older infants could be classed
as juveniles and there were 2 new births, bringing the total to 43. One
infant died just before the study ended in August 1991.

METHODS

We divided the study area into numbered plots by overlaying a grid

on a base map and collected data on vegetation structure and composition
at PCF in a 40 x 40 m plot, 20 10 x 10-m plots, and 3 400 x 5-m transects.
We selected plots randomly from the grid and established transects along
existing trails. We collected data similarly at Varagaliar in a 40 x 40-m plot
and a 400 x 5-m transect, for comparative purposes. We tagged trees and
lianas with a gbh (girth at breast height) ___10cm and recorded their species,
family, gbh, and height.
972 Menon and Poirier

TableI. Compositions of Lion-Tailed Macaque Groups [Data for AnakunthJ and

Varagaliar are from Kurup and Kumar (1993)]
Group Adult Adult
Group Year size males females Subadults Juveniles Infants
PCF 1990 41 2 12 13 10 4
PCF 1991 43 2 12 13 13 3
Anakunthi
(AS) 1974 22 2 6 7 6 1
Anakunthi
(AS) 1981 31 3 12 9 5 2
Varagaliar
(VG) 1978 12 1 5 0 5 1
Varagaliar
(VG) 1980 14 1 5 2 4 2

W e estimated seasonal availability of food at PCF from phenological

data collected for 1 year on 102 trees belonging to 20 species that contrib-
uted to the lion-tailed macaque diet. We selected species for phenological
sampling after preliminary observation of lion-tailed macaque diet at P C F
and from information gleaned from the literature and from local people.
W e selected representative trees or shrubs of each species randomly and
marked them. For 6 of the 20 species we located only one to four indi-
viduals. We noted the presence of flush (new leaves), flowers, unripe fruit,
and ripe fruit for each tree in the phenological sample every month on the
day before observations on the study group began. We calculated the den-
sity (di) of each species at PCF from vegetational data in all transects and
plots and estimated the monthly availability of fruit from each plant species
relative to other species in the phenological sample with the following for-
mula:

in which
Ai = percentage availability of fruit and flower of species/,
= number of trees of species i fruiting in the sample,
di = density of species i at PCF,
ni = number of trees of species i in the phenological sample, and
k = total number of tree species in the phenological sample.
The formula provides a crude estimate of plant food availability be-
cause the sample sizes are small and we collected only presence/absence
 Lion-Tailed Macaques in a Forest Fragment 973

data. Ai estimates the abundance of the fruit or flowers of species i relative

to the other species fruiting or flowering in the sample.
We defined seasons at PCF based on temperature and precipitation:
winter (December to February), summer (March to May), southwest mon-
soon (June to August), and northeast monsoon (September to November).
We used the instantaneous scan sampling method (Altmann, 1974) to
collect data on time budgets and activity patterns in the study group.
Menon observed the group for 6 days every month from September 1990
to August 1991 (Table II). Heavy rain (>1000 mm/month) delayed or dis-
rupted observations during June, July, and August. Data collection was
possible only for 3, 2, and 4 days in each of these months, respectively.
We recorded data for several activity and diet parameters for all individuals
observable within a period of 5 min and repeated this procedure every 15
min. Five-minute scans alternating with 10-rain scan intervals started at
dawn before the study group left its sleeping trees and continued for 12
hr each day. Each 5-min period of data collection is a scan and data per-
taining to a single individual within a 5-rain scan are a record. Menon
observed the study group for a total of 659 hr. The average number of
scans per day--46.8--generated a total of 14,171 records. W e have data
on the following activity categories: (1) resting is inactive sitting or standing;
(2) ranging is directional travel and includes movements from one canopy
level to another (vertical travel) or over long distances (horizontal travel);
(3) feeding is the actual manipulation or intake of food items; (4)foraging
is active search for and selection of food items, which includes local travel
within the canopy while searching for food items; and (5) other is collec-
tively other activities including social interactions, agonistic behavior, and
predator alarm reactions. We collected additional data on the height at
which the individual was located in the canopy and the height of the canopy
above the individual. If the individual's activity was feeding, then we col-
lected data on the diet type (plant or animal), plant part, and plant species
upon which the individual fed.
We estimated the percentage of daytime spent on an activity by the
study group as the percentage of the number of records for that activity
out of the total number of records per day. The instantaneous scan sam-
pling method allows standardized comparisons between months within a
study and between studies conducted by different researchers in different
study sites (Marsh, 1981b). Accordingly, we selected 6 days of monthly
scanning and 5-min scans to obtain data comparable with previous studies
on lion-tailed macaques in the protected forests of Anakunthi and Vara-
galiar by Kumar (1987) and Kurup and Kumar (1993). However, we
compared data on time budgets only with those of Kurup and Kumar
(1993) because their definitions of activity categories are similar to ours.
 974 Menon and Poirier

Table II. Summary of Instantaneous Scan Sampling Data Collected at PCF

No. of No. of Total Mean scans
Year Month days hours records per day

1990 September 6 65.5 769 44.3

October 6 51.25 573 34.3
November 6 72 1,180 48
December 6 68 1,321 46.3
1991 January 6 72 1,594 48
February 6 71 1,778 47.3
March 6 72 1,913 47
April 6 72 1,786 47
May 6 65 1,887 43.2
June 3 19.25 498 26
July 2 5 165 10.5
August 4 26 707 26.3
Total 63 659 14,171 46.82

RESULTS

The sample plot (0.16 ha) yielded a lower value for tree density per
hectare at PCF (812) compared to the protected forest of VG (1063). Pas-
cal (1988) obtained a value of 1520 for tree density/ha from a comparable
plot (0.2 ha) at Attapadi Reserve Forest. Furthermore, the species com-
position was very different in the disturbed study area compared with
protected forests. For example, a pioneer species, Maesa perotettiana, and
a cultivated species, Coffea arabica, constituted the largest proportion (24.6
and 14.6%, respectively) of the total species in the sample plot at PCF. At
Attapadi Reserve Forest (Pascal, 1988), two climax evergreen species
formed the largest proportion of species in the sample plot (Aglaia anamal-
layana, 21.4%; and Cullenia excelsa, 17.1%). Apart from lower tree densities
and altered species composition, PCF had fewer emergents and tall trees
(>31 m). Several intermediate girth classes were missing, and the middle
and upper canopy layers were discontinuous at PCF because of selective
logging and systematic weeding of the undergrowth (Menon, 1993). The
canopy cover at VG was more continuous than that at PCF but also showed
some gaps due to selective logging in the past.
Food sources belonging to four key genera--Cullenia, Artocarpus,
Ficus, and Litsea--contributed >70% to the diet of the study group. Phe-
nological data indicate that availability of fruit/seed from these four genera
was higher from March to August and lower from September to February
(Fig. 1). Food availability was lowest in September, October, November,
Lion-Tailed Macaques in a Forest Fragment 975

and December, when fruit/seed of the four key genera were either absent
or marginally available.

Temporal Variation in Activities

The lion-tailed macaque group at PCF showed significant variation in

time spent in different activities (Kruskal-Wallis test, p < 0.001). They
spent the largest proportion of time each day ranging (34%). The second
most common activity was foraging (23.7%), followed by feeding (17.9%)
and resting (16%). The group spent 8.4% of its time in other activities.
In Table III we compare the time budgets of lion-tailed macaques at
PCF with those obtained by Kurup and Kumar (1993) in the protected
forests of Varagaliar and Anakunthi. Ranging, the most common activity
(34%) at PCF, was one of the least common activities (15%) in the pro-
tected forests of VG/AS. Foraging was the next most frequent activity at
PCF, followed by feeding and resting. The groups at VG/AS spent almost
an equal amount of time resting, foraging, and feeding.
Time spent resting by the study group is significantly lower in the early
morning (0600-1000) than during the remainder of the day (Kruskal-Wallis
test, p < 0.01). Differences in time spent ranging, foraging, feeding, and
in other activities are not statistically significant at different times of the

60

[ ] Cullenia
50
9 Artocarpus

40 [ ] Ficus

9 Litsea
30

~_ 20

0 p
J F M A M J J A S 0 N D
Month
Fig. 1. Monthly variation in availability of four key food resources at PCF. Percentage
availability of each species is an estimate of the abundance of the fruit or flowers of that
species relative to the other species fruiting or flowering in the sample.
 976 Menon and Polder

Table IlL Tune Budgets of Lion-Tailed Macaque

Groups in Different Forests
Activity PCF VG/ASa

Resting 16.0 27.8

Foraging 23.7 26.7
Feeding 17.9 27.8
Ranging 34.0 15.0
Other 8.4 3.5
aCombined data for lion-tailed macaques at Varagaliar and
Anakunthi Shola from Kurup and Kumar (1993).

day. These data are similar to those observed for lion-tailed macaques in
protected forests.
The monthly variation (Fig. 2) is significant for time spent resting (p
< 0.01) and time spent foraging and feeding (p < 0.001) by the study
group. However, differences between months in time spent ranging are not
statistically significant.
Plant foods contributed to 96.5% of the diet of the study group. The
number of plant species in the annual diet at PCF is 42, which is consid-
erably less than the 81 species in the annual diet at VG (Kumar, 1987).
During the months of low food availability, the study group fed on
the fruit of Coffea, Macaranga, and Diospyros and on Cullenia flowers. The
amount of time spent foraging was higher from September to December
than during the rest of the year. Time spent feeding was highest in Sep-
tember, during which 80% of the feeding time was spent on flowers of
Cullenia and figs. Both of these plant parts are labor- and time-intensive
food sources compared to the pulp or seeds obtained from larger fruit such
as Artocarpus and Cullenia.
Time spent resting was lowest during months in which there were
peaks in feeding and foraging. In protected forests, lion-tailed macaque
resting time decreased during months of relatively low food availability and
no significant monthly variation occurred in time spent feeding or ranging
(Kurup and Kumar, 1993).
Seasonal variations (Table IV) in time spent ranging, feeding, and in
other activities by the study group are not significant. Only resting and
foraging show significant seasonal variation. Time spent foraging was high-
est from September to February, the period of low food availability. Time
spent resting was lowest in these months. Time spent foraging and resting
over the year is negatively correlated (r = -0.5, p < 0.0001).
Lion-Tailed Macaques in a Forest Fragment 977

a
9~ Resting
35

30

25
G)
E
20
t~

c
o) 15
o

13.
10

J F M A M J J A S O N
Month

= Ranging
45
40
3S
~ 3O
~z5
8u zo
a_ 15
10

F M A M J J A S O N D
Month

Fig. 2. Percentage of daytime spent by the study group in (a) resting, feeding, and foraging,
and (b) ranging and other activities.
 978 Menon and Poirler

Table IV. Seasonal Variation in Percentage of Daytime Spent in Different Activities by

the Study Group at PCF
Winter Summer SW monsoon NE monsoon
(Dec.-Feb.) (Mar.-May) (June-Aug.) (Sep.-Nov.)

Temperature a Low High Moderate Moderate

Rainfallb Low Low High Moderate
Food availabilit3/ Low High High LOw
Resting 14.9 21.3 19.5 11.9 p < 0.001
Ranging 32.8 37.9 34.3 30.0 n.s.
Foraging 23.3 19.7 19.8 28.6 p < 0.0001
Feeding 17.8 15.1 17_5 21.3 n.s.
Other 11.2 6.0 8.9 8.2 n.s.
aTemperature: low (15-20"C); moderate (21-25"C); high (26-30~
bRainfall/month: low (0-50 ram); moderate (51-300 ram); high (301-1000 ram).
CFood availability: trees were attuned to fruiting before and during the southwest monsoons
and the months from March to August have the higher food availability.

Spatial Variation in Activities

The lion-tailed macaque group at PCF spent the largest proportion of

its time (40.4%) in canopy levels between 21 and 30 m, 26.5% in levels
between 11 and 20 m, 16.7% between 1 and 10 m, 11.5% at levels >31
m, and 4.9% on the ground. The difference in time spent in different can-
opy levels is statistically significant (Z2 test, p < 0.0001). Lion-tailed
macaques in protected forests spent 66% of their time in canopy levels
between 5 and 20 m and only 0.35% of their time on the ground (Kurup
and Kumar, 1993).
The study group did not exhibit significant variation in the percentage
of daytime spent in different canopy levels during different times of the
day except for heights in the canopy between 21 and 30 m. The group
spent the most time between 21 and 31 m during early morning hours and
the least time during midday (Z2 test, p < 0.05).
There are significant differences in times spent in various activities at
different heights (Table V; Z2 test, p < 0.001). Allocations of time spent
resting, foraging, feeding and in other (a table category) activities are simi-
lar in each of the lower and middle canopy levels between 1 and 30 m
(Table V, Fig. 3). However, time spent ranging demonstrates a marked
deviation from this pattern. More than half the time spent ranging was
concentrated in the canopy between 21 and 30 m.
There was also considerable monthly variation in performance of ac-
tivities within each canopy level. Ranging was the most common activity
 Lion-Tailed Macaques in a Forest Fragment 979

50

45

40

35 I--o-- Resting
E 30 /--D-t Foraging
"~" /__e_ Feeding
r 25 /_B_Ranging
o 20 |'~>-- Other
11-15

10

5 ~

0 ,
Ground 1-10m 11-?Om 21-30m >31 m
Canopy height (meters)
Fig. 3. Spatial variation in percentage of daytime spent in different activities by the study
group at PCF.

(X = 32.3%) at the highest canopy level (>31 m) for most of the year
except July when this canopy level was not used and in September and
January when feeding was the most common activity at this level (50%).
Ranging was the most common activity (X = 45%) at heights between 21
and 30 m throughout the year. Ranging was common at levels between 11
and 20 m except for May and October when there were peaks in foraging
at this level (34 and 38.4%, respectively.) In the 2 months with foraging
peaks at this level there were corresponding lows in resting (18 and 8%,
respectively). In contrast, during August, there was a peak in resting (38%)
at this level with a corresponding low in foraging (17%.)

Table V. Spatial Variation in Percentage of Daytime Spent in Different Activities by the

Study Group at PCF
Resting Foraging Feeding Ranging Other

Ground 0.4 5.8 3.7 7.0 6.1

1-10 m 20.5 21.8 20.3 6.5 29.7
11-20 m 28.4 31.8 25.6 21.6 30.3
21-30 m 37.2 32.3 34.9 53.3 27.0
>31 m 13.5 8.3 15.5 11.6 6.9
 980 Menon and Poirier

The monkeys performed all activities to similar extents year-round at

canopy levels between 1 and 10 m except in October when there was a
foraging high (46%) and a low in other activities (0%). In contrast to
groups in protected forests that never used the ground for ranging, the
study group used the ground most frequently for ranging (X = 35%) and
foraging (33%). Resting was the least common terrestrial activity (2.1%).
Other activities performed on the ground at PCF include feeding, play-
ing, and drinking water from streams. The lion-tailed macaques crossed
clearings in the forest by coming to the ground. They also crossed the road
to raid fruit from Artocarpus trees growing in the coffee plantation on the
other side of the road. Human presence in the coffee plantation was some-
times an indirect and often a direct deterrent, in the form of gesturing and
stone-throwing, which prevented the monkeys from spending much time in
the coffee plantation.
More than 90% of the time spent on the ground was in areas without
tree cover, such as the road and large clearings within the forest. Thus,
although the study group spent more time on the ground than groups in
protected forests did, they clearly preferred to stay off the ground if trees
or shrubs were available. Individuals jumped across large gaps in the canopy
even when this seemed to require significant effort. Lion-tailed macaques
fell from the canopy three times at PCF. The first two individuals were
juveniles that fell while clearing gaps in the canopy. Both of them survived.
In the third instance, an infant died after hitting the ground.

DISCUSSION

Group size and habitat quality are two of the most important influ-
ences on primate time budgets. Our results are consistent with observations
in several other studies that larger primate groups spend more time ranging
or travelling than smaller groups do (van Schaik et aL, 1983; Stacey, 1986;
Altmann, 1987; Barton et aL, 1992; Isbell and Young, 1993). However, our
interpretation is confounded by poor habitat quality and disturbed condi-
tions at PCF: greater time spent ranging may have been determined more
by environmental conditions and human disturbances than by size of the
study group per se.
Large group size confers the advantage of reduced predation risk. This
advantage is offset by stresses imposed on time budgets by increasing group
sizes. Most free-ranging lion-tailed macaque groups fission when group size
approaches 30 individuals (Kumar, 1987). The study group remained intact
at 43 animals, the largest known size for a wild lion-tailed macaque group.
Other disturbed fragments in the area accommodate similarly intact, large
Lion-Tailed Macaques in a Forest Fragment 981

groups. Why are group sizes large in marginal habitats? The average annual
home range for groups in the protected forests of Varagaliar is 1 km 2 (Ku-
mar, 1987) and even higher in other areas (Green and Minkowski, 1977).
In the disturbed PCF area a split group would be forced to remain in the
isolated fragments. Perhaps these forest fragments cannot support the an-
nual home ranges of two groups without large overlaps, which might lead
to high levels of agonistic interactions between groups. Large group sizes
and the absence of fissioning in isolated forest fragments may be a mecha-
nism to avoid intergroup conflict and to maintain group cohesion.
The relationship between habitat quality and time budgets as reported
in the literature is confounded by differences between habitats in group
size, temperature, and proximity to humans. For example, better habitat
quality is associated with decreased feeding, increased resting, and no sig-
nificant difference in time spent moving among baboon groups (Iwamoto
and Dunbar, 1983), decreased feeding and moving in gorilla groups (Watts,
1988), decreased feeding and increased resting among provisioned baboon
groups (Altmann and Muruthi, 1988; Eley et al., 1989), and decreased time
spent foraging in urban and rural bonnet macaque populations (Singh and
Vinanthe, 1990). Contrarily, better habitat quality has also been associated
with increased moving and no difference in time spent feeding among
vervet groups (Isbell and Young, 1993) and increased feeding and de-
creased resting in colobus monkeys (Oates, 1977; Marsh, 1981a, b). The
energy requirement for thermoregulation is another important factor that
may influence time budgets (Oates, 1977; Marsh, 1981a; Iwamoto and Dun-
bar, 1983). The climate at PCF was considerably cooler due to its higher
elevation, especially during winter months, than at Varagaliar and Anakun-
thi Shola, and energy requirements for thermoregulation would be a
plausible explanation for increased ranging instead of resting at PCF. How-
ever, this can be ruled out as a significant factor because the average
amount of daytime spent ranging at PCF did not vary significantly between
months or seasons.
Food availability emerges as a key influence on time budget in both
protected and disturbed forests. At PCF the foraging peak coincided with
months in which the key resources were not available. Key resources such
as Artocarpus, Cullenia, and Litsea were abundant during other months,
thus eliminating the need for extensive foraging. The time budget was com-
pressed during months when these key resources were not fruiting because
additional time devoted to foraging was at the expense of resting. This find-
ing is consistent with Dunbar's (1992) observation that in stressed baboon
groups, resting time might act as a reserve that could be exploited whenever
necessary and converted into additional feeding time.
 982 Menon and Poirier

In constrast to lion-tailed macaques in protected forests, the study

group consistently spent more time ranging than resting b e c a u s e of high
levels of human presence, extent of habitat degradation, and large group
size coupled with the demands of a largely frugivorous diet. PCF had a
very high human presence and the monkeys were often disturbed by the
close proximity of twig pickers and people cutting wood from fallen trees
and walking along the major forest trails. The disturbed conditions together
with the unusually large size of the lion-tailed macaque group at PCF may
have compelled them to move constantly to ensure optimal foraging and
to fulfill their collective dietary requirements. These characteristics--high
human presence, extent of habitat degradation, and group size--were con-
stant throughout the year. Their explanatory power is reinforced by the
fact that among all activities, ranging is the only one that does not vary
significantly with month.
Like lion-tailed macaques in protected forests, the study group spent
most of its time in the middle canopy levels because most of the demands
of their frugivorous diet were met there. Although they spent more time
on the ground than has been previously recorded for the species, the mon-
keys had a clear preference for staying in trees. However, staying in trees
is not without its dangers as evidenced by the accidental infant death at
PCF. Kumar (1987) did not record any instances of falls resulting in deaths
in 6 years of observations on lion-tailed macaques in the protected forest
of Varagaliar. Johns (1981, 1983) noted that infant mortality was high dur-
ing logging and immediately following logging, though he did not list the
exact cause of the deaths.
Their propensity for arboreality makes neighboring forest fragments
inaccessible to groups confined to isolated fragments. Lion-tailed macaques
migrate between groups within interconnected forests (Kumar, 1987), how-
ever, they have not been observed crossing large gaps between forests
separated by cultivation or human habitation. In > 1700 hr of observation,
Green and Minkowski (1977) observed no lion-tailed macaque crossing ad-
joining tea or coffee plantations to the nearest forest patch even if the
plantation was the shortest path between them.
A considerable proportion of the total lion-tailed macaque population
is distributed in genetically isolated, inbreeding groups. The potential for
adaptation to environmental changes declines in inbred populations, which
tend to have decreased fetal, neonatal, and juvenile survivorship, lower
competitive ability, and lower fertility (Harris et al., 1984). The study group
had a birth rate--number of births/number of females--of 0.25, during the
study year, which is close to the mean birth rate/year of 0.29 for eight
groups obtained by Kumar (1987). Southwick et al. (1980) suggested that
a higher birth rate is necessary for long-term survivability of rhesus ma-
 Lion-Tailed Macaques in a Forest Fragment 983

caque populations. However, lion-tailed macaques tend to have higher in-

terbirth intervals than those of other macaque species both in the wild and
in captivity (Kumar, 1987).
Ongoing disturbance and habitat alteration are the most imminent
threats to groups confined in forest fragments. For example, during our
study several wind-fallen trees were extracted from the fragment. A few
months after the study ended, >900 trees were logged (Ahimaaz, 1992).
According to Kumar (personal communication), the PCF group size has
since dropped to 34 individuals, with one instance of predation by a dog.
Viability in each forest fragment containing lion-tailed macaques must
be determined based on ecological, economic, and political considerations.
Groups in forest fragments that are deemed viable should be protected
and monitored and those in nonviable fragments must be translocated to
protected forests wherever possible. These steps are critical for the survival
of isolated groups and the species as a whole.

CONCLUSIONS

(1) Lion-tailed macaques in a marginal forest fragment spent more

time ranging than resting or feeding, unlike groups in protected
forests.
(2) Monthly differences in time spent ranging are not significant.
Thus, time spent ranging was not affected by seasonal variation
in food availability, temperature, and precipitation. The factors
contributing to high amounts of time spent ranging throughout
the year include the large group size, extent of habitat degrada-
tion, and high human presence.
(3) Differences between months in time spent resting, feeding, for-
aging, and in other activities are significant. The peak in time
spent foraging was at the expense of time resting and in other
activities and coincided with months during which the group fed
on nonkey resources.
(4) Lion-tailed macaque groups living in small disturbed forest
patches may not have adequate reproduction for long-term sur-
vivability. They are extremely vulnerable to predation threats as
evidenced by a 20% decline in our study group in recent years.

ACKNOWLEDGEMENTS

This research was supported by grants from the L. S. B. Leakey Foun-

dation, National Science Foundation, World Wildlife Fund--U.S., and The
984 Menon and Polrier

Ohio State University Graduate School Alumni Research Award. We grate-

fully acknowledge the support of the Tamil Nadu Forest Department,
Wildlife Institute of India, and Botanical Survey of India during the re-
search and of the Department of Statistics of The Ohio State University,
during data analysis. We are indebted to Ajith Kumar for introducing S.
Menon to the study area and for his guidance through various stages of
this work. We thank Lori Sheeran, Ajith Kumar, and two anonymous re-
viewers for constructive comments on the manuscript.

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