Lagothrix Time Allocation Patterns

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International Journal of Primatology [ijop] PP063-295788 May 9, 2001 13:40 Style file version Nov. 19th, 1999
International Journal of Primatology, Vol. 22, No. 3, 2001
Time Allocation Patterns of Lowland Woolly

Monkeys (Lagothrix lagotricha poeppigii)
in a Neotropical Terra Firma Forest
Anthony Di Fiore1,2,3 and Peter S. Rodman2
Received March 24, 2000; revision May 22, 2000; accepted May 31 , 2000
We investigated the time allocation decisions of lowland woolly monkeys

(Lagothrix lagotricha poeppigii) in a terra firma forest in eastern Ecuador
where they occur sympatrically with 9 other primate species. Woolly mon-
keys spent considerable amounts of time searching for and attempting to
procure animal prey—roughly as much time as they spent consuming plant
material: ripe fruits, leaves, and flowers. The amount of time spent foraging
for animal prey is positively related to the habitat-wide availability of ripe
fruits (the predominant component of the woolly monkey diet), and nega-
tively related to both ambient temperature and the abundance of potential
prey items in the habitat. Time spent resting showed exactly the opposite
pattern with respect to these ecological variables. These results suggest that
woolly monkeys follow an energy-maximizing strategy of food acquisition
during times of fruit abundance—focusing on animal foods and perhaps lay-
ing down fat reserves to utilize when ecological conditions worsen—and follow
an energy-minimizing strategy when fruit resources are scarce. Such a strong
and seasonal commitment to animal prey foraging is unique among the ate-
line primates and is not ubiquitous even among lowland woolly monkeys.
We suggest that this foraging strategy, and the greater intragroup cohesion
that characterizes some populations of Lagothrix, are both opportunistic re-
sponses to regional differences in habitat quality. Identifying and accounting
1 Department of Anthropology, University of California, Davis; Department of Anthropology,
New York University; NYCEP (New York Consortium in Evolutionary Primatology).
2 Department of Anthropology, University of California, Davis.
3 To whom correspondence should be addressed at Department of Anthropology, New York
University, Rufus Smith Hall, Room 801, 25 Waverly Place, New York, NY 10003 (e-mail:
anthony.difiore@nyu.edu).
449
0164-0291/01/0600-0449$19.50/0 °
C 2001 Plenum Publishing Corporation
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450 Di Fiore and Rodman
for such intraspecific variation should be a goal of any analysis of comparative

socioecology.
KEY WORDS: Lagothrix lagotricha; woolly monkey; Atelinae; time budgets; ecological
strategies; animal prey foraging; socioecology.
INTRODUCTION
Time is limited for most animals (Dunbar, 1988, 1992). A day’s forag-
ing must provide sufficient resources to avoid starvation, to maintain body
condition, and perhaps to grow or support reproduction. Animals vulner-
able to predation may need to invest time being vigilant or traveling large
distances to sleep in protected areas. Grouping to lower the risk of preda-
tion can necessitate additional foraging time or visits to more food patches
each day (and thus more time traveling) to compensate for the costs of in-
creased feeding competition (Waser, 1977; Isbell, 1991; Wrangham, et al.,
1993; Janson and Goldsmith, 1995). Many social animals, such as primates,
also require time to establish and maintain social relationships that affect
their access to resources or mates (Dunbar, 1992; Janson, 1992).
When these multiple activities cannot be performed simultaneously,
some individuals may allocate time between various behavioral options bet-
ter than others (Dunbar, 1988, 1992; Mangel and Clark, 1986; Krebs and
Kacelnik, 1991). The costs and benefits of these options are likely to change
periodically as the ecological or social environment or the animal’s own phys-
iological state changes. Evolutionary theory predicts that animals should be
sensitive to these changes and adjust time allocation to current conditions
(Schoener, 1971; Mangel and Clark, 1986). Consequently, behavioral ecolo-
gists have examined time budgets as a means of investigating ecological and
social influences on individual behavior.
Time allocation has been studied in a broad array of primate taxa
(callitrichines: Terborgh, 1983; Digby and Barreto, 1996; cebines: Terborgh,
1983; de Ruiter, 1986; Robinson, 1986; Miller, 1996; atelines: Milton, 1980,
1984; Strier, 1987; Symington, 1988; Defler, 1995; colobines: Struhsaker, 1975;
Stanford, 1991; cercopithecines: Post, 1981; van Schaik, et al., 1983; Isbell and
Young, 1993; hominoids: Goodall, 1986; Watts, 1988; Clutton-Brock, 1977b).
In some species, individual activity budgets vary seasonally in response to
changes in the abundance, quality, or distribution of important food re-
sources (Robinson, 1986; Altmann and Muruthi, 1988; Isbell and Young,
1993). Particular aspects of individual time budgets also covary with social
variables such as group size and group composition. For example, in some
primates, time allocated to movement (Clutton-Brock and Harvey, 1977;
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Time Allocation by Woolly Monkeys 451
Waser, 1977; Isbell, 1991; Wrangham, et al., 1993; Janson and Goldsmith,
1995) or to foraging (de Ruiter, 1986; Miller, 1996) increases with group
size. Primate time budgets are also influenced by a variety of other demo-
graphic and environmental factors such as age, sex, social rank, reproductive
condition, and the degree of human disturbance in the region (Whitten, 1983;
Muruthi, et al., 1991). Together, these results suggest an important interplay
between ecological, social, and demographic influences on primate time al-
location patterns.
Below we examine time allocation by lowland woolly monkeys (Lago-
thrix lagotricha poeppigii). We first characterize the basic time budget and ex-
amine how time budgets vary seasonally. We then investigate how changing
environmental variables, e.g., resource abundance, temperature, and rainfall,
influence time allocation decisions, we consider whether these influences are
consistent with predictions derived from basic foraging theory, and we offer
a reassessment of woolly monkey foraging strategies that incorporates and
extends previous work. Finally, we compare time allocation of woolly mon-
keys to both close phylogenetic relatives and more distantly related primate
taxa and discuss the implications of this comparison for understanding the
comparative socioecology of ateline primates.
METHODS
Study Area
We worked between August 1994 and April 1996 in Yasunı́ National

Park in the Amazon region of Ecuador (Figures 1A, 1B). The study site is
south of the Rı́o Napo, 47 km along a petroleum access road recently con-
structed through the park’s western margin (75◦ 280 W, 0◦ 420 S). The roughly
350-ha study area (Fig. 1C) consisted of intact, primary tropical rain forest
spread out over a series of ridges and minor drainages feeding several small
but permanent streams. In contrast to many other primate research sites in
Amazonian South America, the area is inland from the nearest major river
and comprised entirely nonfloodable terra firme forest. Surveys conducted in
the area during the summers of 1992 and 1993, before the road was built, re-
vealed flourishing primate populations of 10 species (Di Fiore and Rodman,
unpublished data). Before petroleum development in the region, the area
was subject to only occasional human impact by nomadic indigenous hunters
and gatherers. Both hunting and swidden agriculture in the region have in-
creased in recent years, but hunting in and around the study site is minimal
because of an accord we established in 1994 with local families.
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Fig. 1. (A) Geographic distribution of four subspecies of lowland woolly monkeys and the
locations where they have been subjects of long-term studies, modified from Fooden (1963)
and Defler and Defler (1996). (B) Map of Yasunı́ National Park showing the study site and
surrounding area. (C) Map of the study area showing the locations of trails and botanical
transects.
A profile of meteorological conditions at the site is in Fig. 2A. During

the primary study period—from April 1995 to March 1996—rainfall totaled
3274 mm with an average of 273 mm per month. Variation in temperature
was less pronounced than variation in rainfall, though the monthly average
of daily temperature maxima measured in the shade ranged from just over
34◦ C to nearly 38◦ C. Thus, while not as seasonal as other neotropical forests
farther from the Equator, the forest at Yasunı́ nonetheless shows a short,
but marked, dry and hot season each year and several cooler months of very
heavy precipitation.
Study Species and Groups
Woolly monkeys are large-bodied, highly frugivorous primates that

live in multimale-multifemale groups containing between 20 and 40 indi-
viduals (Ramirez, 1980, 1988; Nishimura, 1990; Peres, 1994a; Defler and
Defler, 1996; Kinzey, 1997). At least five social groups, comprising close
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Fig. 2. (A) Monthly changes in average minimum and maximum temperature (measured in
the shade) and rainfall. (B) Monthly changes in three indices describing the habitat-wide
availability of ripe fruits.
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Table I. Composition of the principal study groups

Age-sex class Group 1 Group 2
Males Adult 3/2a 4/4a

Subadult, large 3/2 2/2
Subadult, small 3/3 3/3
Juvenile 1/1 4/4
Infant 0/0 0/0
Females Adult, with dependent 5/6 5/5
offspring
Adult, no associated 6/5 4/3
offspring
Juvenile 4/4 0/0
Infant 0/1 0/1
Total Including Juveniles 25/24 23/23
Total Excluding Juveniles 20/18 18/17
Ratio of Nonjuveniles to Juveniles 4.0:1/3.0:1 4.5:1/3.4:1
Sex Ratio Including Juvenilesb 0.67/0.50 1.44/1.44
Sex Ratio Excluding Juvenilesb 0.82/0.64 1.00/1.13
a Numbers before and after slash indicate counts at start and end of study respectively.
b Calculated as number of males per female.
to 110 independent individuals, utilized various portions of the study site,

and the biomass of woolly monkeys in area is the highest reported for any
neotropical site (circa 1 kg/hectare: Di Fiore, 1997). The two principal study
groups (1 and 2) ranged nearest to the petroleum access road and their age-
sex compositions are in Table I. Group 1 changed from 25 adults and juveniles
at the start of the study to 23 by the end, due to the death of a subadult male
and the disappearance of an adult male. Group 2 initially contained 23 adults
and juveniles, but one adult female was killed by a hunter partway through
the research period. One female infant was born in each group during the
study.
Data Collection
From April 1995 through March 1996, we spent 16–20 days each month
conducting full-day follows of the two principal study groups. We also occa-
sionally followed a third group whose home range extensively overlapped
those of groups 1 and 2 and collected some data opportunistically on two
additional groups whose ranges abutted that of group 1. We divided monthly
data collection into four 5-day periods, two of which were spent with each
of the principal study groups. We chose the focal group for the first and
third periods each month at random and switched groups for the second and
fourth periods. Each 5-day period comprised two days of instantaneous scan
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sampling and three days of focal sampling (Altmann, 1974). We limit our
analyses here to data collected during group scans.
Group scans began 5 min after a group was first encountered or at
0630 h if the subjects were followed to their sleeping sites the previous
evening, and we took scans at 10-min intervals thereafter until 1730 h. Scans
lasted for 5 min and were followed by 5 min of inactivity until the next
scan. We always took a group scan if ≥1 individual came into view during
the scan period. During a scan, we recorded the identity, activity, and near-
est neighbors of each woolly monkey that came into view, with particular
attention paid to foraging and social behaviors. We refer to these obser-
vations of each scanned animal as a behavioral record. We tried to collect
behavioral records on as many different subjects as possible during a scan
by changing position under the group frequently. Group members were of-
ten spread out over an area of >1 ha, so only a fraction of the group was
sampled during any given scan (mean = 4.76, SD = 1.79, range = 1 to 15,
N = 4180).
Whenever possible, we recorded the individual identity of a scanned
subject. If individual identification was impossible, we assigned the individual
to one of five age-sex classes: adult males, adult females associated with
dependent offspring, i.e., females either carrying an infant or young juvenile
or interacting with a young juvenile shortly before or after being scanned,
unburdened adult females that were not associated with dependent juveniles,
subadult males, or juveniles of either sex. We watched each subject for 5 sec
after it came into view and recorded its behavior during that interval. In
most cases, a single, predominant activity was clear, but in some instances,
such as during bouts of rapid social interaction, a scanned subject might have
engaged in >1 recognized behavior sequentially. In this case, we recorded
the sequence of behaviors within those 5 sec.
We assigned each behavioral record to one of seven mutually exclusive
activity categories: eat, forage, move, rest alone, rest social, social activity,
and other nonsocial (Table II). Whenever subjects were eating, we recorded
the type of food being consumed: ripe fruit, immature fruit, seeds, leaves,
flowers, or animal prey. In some cases, it was difficult to visually confirm
consumption of animal prey, but we felt confident scoring a food item as
animal prey if an individual chewed immediately following characteristic
prey-searching behavior, e.g., closely inspecting and unrolling dead leaves.
We determined monthly diets by calculating the proportional representation
of each food type in the set of eat records collected that month. We then
averaged these across months to derive the yearly diet.
When recording nearest-neighbor data for a given individual, we noted
the identity or age-sex class of all other woolly monkeys within 2 m of the
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Table II. Behavioral categories and definitions

Behavior Definition
Eat To handle, process, or consume either plant food items or animal prey.
Forage To search for animal prey while stationary or moving, to actively
manipulate a substrate in search of prey (e.g., unrolling leaves, breaking
apart branches, inspecting vine tangles) or to actively attempt to procure
prey items from the substrate.
Move To change positions, either within or between tree crowns, exclusive of
that movement taking place incidentally while searching a substrate.
Rest alone To be inactive in either a sitting, lying, or standing posture. Included in
this category is time spent scanning the environment (e.g., vigilance)
that was not obviously associated with foraging behavior.
Rest social To be inactive as during rest alone while in close proximity (less than
two meters) of another individual who was either also resting or
otherwise stationary and not interacting directly with the scanned animal.
Social To be engaged in any behavior such as grooming or mating that
activity involved one or more other individuals, usually in close proximity.
Other To perform any of a variety of infrequent, self-directed behaviors
nonsocial (e.g., defecation and urination, self-grooming, solo play, scent marking,
and inspection of scent marks) or to watch or display at the observer.
scanned individual. Beyond this distance, we recognized two additional prox-

imity categories: >2–5 m and >5–10 m. If multiple neighbors were present
in the closest proximity category, we recorded all of their identities. Finally,
for each scan we noted the general activity of the group, the context within
which the scan took place, e.g., during an intergroup encounter, while rest-
ing, while foraging, or while traveling, and the location of the group relative
to mapped trail locations and known feeding trees.
We suspended sampling during periods of heavy rain because the mon-
keys generally ceased activity until the rain let up, but we continued to
sample during light showers since the subjects remained engaged in typical
behavior. We adopted this procedure because we were primarily interested
in how woolly monkeys allocate their time among the alternative behaviors
available to them, and we reasoned that several of these behaviors might
be precluded by heavy rain. For instance, the opportunity for them to for-
age for fruits or insects is likely to be reduced during rainstorms when vi-
sual and olfactory cues are potentially obscured and when prey items are
likely to be less active. However, because omitting data on behavioral in-
activity during heavy rains could potentially bias our results towards under-
representation of resting time, we scored all behavior during heavy rains
post hoc as rain sheltering and examined the effects of ecological variables
on time allocation to rest both including and excluding this additional activity
category.
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Determination of Time Budgets
We conducted a total of 4296 scans in 95 days over the 12 mo of the study.

In calculating time budgets, we omitted from analysis all scans collected dur-
ing aggressive intergroup disputes (N = 42), but included scans collected on
aggregations of >1 social group wherein intergroup aggression was absent.
The results presented here are thus derived from a set of 4254 group scans
representing 709 hours of data and comprising 19,693 behavioral records.
Of these scans, 2242 (373.7 hours) are on group 1, and 1602 (267.0 hours)
are on group 2. We collected the remaining scans either on other groups in
the area, on occasional multigroup associations involving group 1 or 2, or on
rare solitary individuals.
To reduce the chance of biasing our estimates of time allocation towards
over-representation of conspicuous, aggregate activities such as feeding in
the crowns of large fruiting trees, we determined time budgets following
Clutton-Brock (1977a). Thus, we first constructed a separate time budget
for each scan, scoring time allocation to each activity as the proportion of
scanned individuals engaged in that activity, and then used each of these
scan budgets as individual data points in subsequent analyses. This method
is particularly appropriate when variable numbers of individuals are seen
per scan or when behavior within the group is synchronized such that the
behavior of a few individuals recorded during a given scan is likely to rep-
resent the whole group at that time (Clutton-Brock, 1977a). To correct for
minor differences in the number of scans collected during different months
of the year and at different times of the day, we first used the scan budgets
to calculate time budgets for each hour of the day and then averaged them
within each month to construct monthly time budgets. The grand mean of
the set of monthly budgets provided the overall yearly time budget.
Ecological Sampling
Plant Resources
To provide an independent estimate of the density, distribution, and

abundance of plant resources available to the woolly monkey population,
we established five 1-hectare belt transects at random locations relative the
trail system (Fig. 1C). Each transect comprised 100 blocks of 10 m × 10 m,
half of which we selected for monthly phenological sampling according to a
stratified random design in which two blocks out of every four were desig-
nated for monitoring. The 2.5 total ha of phenological blocks contained 1492
trees with a diameter at breast height (DBH) of ≥10 cm.
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Each month, from March 1995 to April 1996, an assistant recorded the
abundance of new leaves or leaf flush, flowers, and fruits in the crown of
each of the trees and in any lianas or other associated epiphytic plants. If a
plant was fruiting, we estimated the maturity of the fruit (immature vs. ripe)
and the size of the fruit crop by counting the number of individual fruits in
several subsections of the crown and then extrapolating from these to the
full crown volume. We converted raw estimates of crop size to abundance
categories based on an exponential scale further subdivided into thirds, fol-
lowing Leighton (1993), which reflected the degree of accuracy with which
fruit counts could reasonably be estimated, e.g., 1 to <4 fruits, 4 to <7 fruits,
7 to <10 fruits, 10 to < 40 fruits, 40 to <70 fruits, etc. We then derived
three indices of habitat-wide fruit abundance from these phenological data:
[1] the tree index, calculated as the proportion of phenology trees bearing
ripe fruits each month, [2] the basal area index, calculated as the summed
basal area of trees bearing ripe fruits each month divided by the total basal
area of trees monitored that month, and [3] the fruit crop index, calculated
as summed logarithms of the midpoint values of the abundance categories
recorded for all trees bearing ripe fruits, divided by the total number of
monitored trees.
Potential Prey Resources
We were unable to measure seasonal variation in the abundance of

potential animal prey items of woolly monkeys directly. However, if Yasunı́
is similar to other neotropical sites, then insect abundance should peak during
the transition from the dry to wet season, probably associated with the flush
of new leaves that follow the first rains (Wolda, 1978, 1982; Smythe, 1982;
Robinson, 1986; Janson and Emmons, 1990). Rainfall may thus serve as
one crude index of insect abundance. We derived a second index of insect
abundance as follows. First, we compared climatological data for Yasunı́
(Fig. 2A) to that for Manu National Park, Peru (71◦ 220 W, 11◦ 540 S), the closest
(and only) Amazonian site for which published data on rainfall and crude
estimates of seasonal insect abundance are both available. The three-month
dry season we recorded in Yasunı́ during July to September corresponded
perfectly with that seen in Manu (Gentry, 1990). Therefore, we used data
on seasonal changes in insect abundance at Manu (Janson and Emmons,
1990) to derive rough estimates of monthly insect abundance in Yasunı́.
We extrapolated weights for the insect catch collected on several nights of
biweekly light-trapping during 1981 from their Figure 17.3 and averaged
the weights for each month of the year to derive monthly insect abundance
scores. To derive a score for January, for which no datum is given by Janson
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and Emmons (1990), we averaged the insect catches from December and
February.
Besides insects, the only other animal prey items that the woolly mon-
keys consumed were tree frogs and, very rarely, snakes (Di Fiore, 1997). The
limited data that exist regarding seasonal changes in herpetofaunal density in
lowland neotropical forests suggest that abundance is correlated with periods
of heavy rainfall (Duellman, 1995). Moreover, Rodriguez and Cadle (1990)
noted that breeding activity of neotropical anurans typically increases with
the start of the rainy season. Given their short maturation times, population
growth—and thus prey abundance—for frogs should thus follow the onset
of the rainy season fairly quickly. These observations, coupled with the fact
that vertebrates contributed <5% of the animal diet of woolly monkeys (Di
Fiore, 1997), suggest that the two indices of insect prey abundance we derived
may serve equally well as crude indices of overall animal prey abundance.
RESULTS
Time Allocation Patterns
Table III shows the monthly and yearly time budgets of woolly mon-
keys. Over the course of a year, most time (36.2%) was spent either feeding
or searching for and attempting to capture animal prey. Moving constituted
another 34.5% of the yearly budget, and total rest time—solitary plus so-
cial rest, combined with rain sheltering—another 23.2%. The remainder
of the budget (6.1%) included various self-directed and social behaviors,
including mating, grooming, and play. The proportion of time woolly mon-
keys allocated to various activity categories varied from month to month.
Separate G tests (Sokal and Rohlf, 1981) for each behavioral category re-
vealed significant intermonthly variation in eat, forage, total rest, and other
nonsocial time (Table III). If time spent inactive during heavy rains is ex-
cluded from total resting time, there is still significant intermonthly varia-
tion in time allocated to the combined activities of rest alone + rest social
(G11adj = 49.93, P < 0.01). Only the proportions of move and social activity
time showed no significant variation across the year.
Diets and Time Allocation
Monthly and yearly diets of woolly monkeys are shown in Table IV.
Across the year, the predominant component of the diet was ripe fruit
(76.3%) followed by animal prey (9.7%). If eat time each month is separated
into two components—time spent consuming plant resources and time spent
consuming animal prey—and the latter is added to forage time, we are left
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Table III. Monthly and yearly time budgets as the percentage of time spent in each activity category
Activity category
Rest
International Journal of Primatology [ijop]
Month Eat Forage Move Alone Social Rain sheltering Total restb Social activity Other nonsocial No. of scans
Apr 1995 15.5 18.5 32.8 12.7 3.5 4.5 20.7 8.0 4.4 330
May 1995 21.0 19.0 32.8 11.5 2.0 8.9 22.3 2.6 2.3 327
Jun 1995 15.6 20.0 35.8 12.9 3.7 4.4 21.1 5.9 1.6 315
Jul 1995 19.6 17.0 35.6 17.9 4.2 0.7 22.7 4.4 0.8 301
Aug 1995 20.6 13.2 32.6 23.0 4.5 2.2 29.7 3.4 0.6 323
PP063-295788
Sep 1995 21.3 10.4 34.9 20.7 6.1 1.6 28.3 3.6 1.5 320
Oct 1995 17.4 11.9 35.9 20.5 6.6 3.0 30.1 3.8 0.9 396
Nov 1995 17.5 13.1 39.5 20.2 3.8 1.6 25.6 3.5 0.8 509
Dec 1995 13.8 17.6 35.7 18.7 3.7 3.2 25.6 6.0 1.4 409
Jan 1996 16.2 21.0 40.1 13.3 2.3 0.3 15.9 5.6 1.1 329
Feb 1996 21.7 23.6 30.8 12.4 2.1 2.9 17.4 5.6 0.8 350
Mar 1996 26.9 21.8 27.4 16.7 2.5 0.0 19.2 4.2 0.6 345
May 9, 2001
Yearly mean 19.0 17.2 34.5 23.2 4.7 1.4 354.50

St. dev. 3.63 4.23 3.53 4.69 1.53 1.07 58.22
Ga11adj ** ** NS ** NS **
13:40
aG statistic for each activity category using all scans each month and adjusted with Williams’ correction for a more conservative test (Sokal and
Rohlf 1981).
b Calculated as sum of rest alone, rest social, and rain sheltering.
∗ P < 0.05.
∗∗ P < 0.01.
Di Fiore and Rodman
Style file version Nov. 19th, 1999
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Table IV. Monthly and yearly diets as the percentage of feeding records spent on each food type and rescaling of time allocation to
subsistence on plant versus prey resources
Other plant Time allocation to Time allocation to
Month Fruits Leaves Flowers material Animal prey plant resourcesa animal preyb
Apr 1995 78.2 5.1 0.0 3.9 12.8 13.5 20.5

May 1995 79.5 5.3 0.0 1.7 13.6 18.2 21.8
Time Allocation by Woolly Monkeys
Jun 1995 78.0 7.9 0.0 1.2 12.9 13.6 22.0

Jul 1995 88.7 3.6 2.0 1.0 4.6 18.7 17.9
Aug 1995 67.0 14.7 10.8 3.9 3.6 19.9 13.9
Sep 1995 65.0 14.5 11.1 3.0 6.4 19.9 11.7
PP063-295788
Oct 1995 75.3 9.4 3.8 2.6 8.8 15.8 13.4

Nov 1995 71.6 12.2 3.8 4.5 8.0 16.1 14.5
Dec 1995 63.6 5.7 10.4 7.0 13.3 12.0 19.4
Jan 1996 76.8 4.3 0.7 2.0 16.2 13.6 23.7
Feb 1996 84.3 4.5 0.6 2.1 8.5 19.9 25.4
Mar 1996 87.5 2.3 0.0 2.1 8.1 24.7 23.9
May 9, 2001
Yearly mean 76.3 7.5 3.6 2.9 9.7 17.2 19.0

St. dev. 8.31 4.28 4.54 1.68 3.95 3.71 4.65
13:40
a Calculated as % eat time—(proportion of diet consisting of animal prey × eat time).

b Calculated as % forage time + (proportion of diet consisting of animal prey × eat time).
461
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two new variables that more accurately describe the way subsistence activity
is partitioned between these two classes of resources (Table IV). Overall,
woolly monkeys devote slightly more of their total subsistence activity to
foraging for animal prey than to consuming plant resources (19.0% versus
17.2%).
Ecological Influences on Time Budgets
If part of the monthly variance in time budgets of Lagothrix is due to

adjusting time allocation to changing ecological conditions, time budgets
should covary with particular environmental variables. As a starting point, it
seems likely that woolly monkey time allocation decisions should be closely
related to variation in the distribution and abundance of important food
resources. Assuming that woolly monkeys forage each day to maintain a
certain daily energy intake, it follows that as the overall abundance of food
resources in the habitat drops, their subsistence behavior should change to
compensate. Of several possible changes, three obvious ones would be [1] to
increase the total time allocated to subsistence, assuming that sufficient food
is out there, just harder to find or process, [2] to increase the time allocated
to feeding per se, assuming that the animals must consume a greater volume
of lower quality food to meet the same nutritional needs, or [3] to minimize
energy expenditure and reduce metabolic activity by decreasing time de-
voted to high cost behaviors such as moving or by resting more (Schoener,
1971). Since ripe fruits constitute the majority of the yearly woolly monkey
diet (Table IV), we would predict that as the habitat-wide abundance of ripe
fruit drops, either total subsistence time (eat + forage) or eat time by itself
would increase if hypothesis [1] or [2] were correct, respectively. Alterna-
tively, if hypothesis [3] were correct, we would expect move time to decrease
or rest time to increase as fruit abundance drops.
Figure 2B is a plot of variation in the three indices of monthly habitat-
wide ripe fruit availability. In contrast to the predictions of hypothesis [1]
and [2], neither the total amount of time spent in subsistence activity nor
eat time by itself is significantly negatively correlated with any of the indices
(Table V). This suggests that woolly monkeys do not adjust time spent in
subsistence activity to the overall abundance of ripe fruit in the habitat by
either of the first two mechanisms suggested above. Hypothesis [3] received
support to the extent that there is a significant negative relationship between
two of the indices of ripe fruit availability and the proportion of time spent
resting each month, whether rest alone, rest alone + rest social, or total rest
is considered, though the predicted positive relationship between ripe fruit
availability and time spent moving is lacking (Table V).
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Table V. Spearman rank correlation coefficients (corrected for ties) between

activity categories and three indices of habitat-wide ripe fruit abundancea
Activity category Tree index Fruit crop index Basal area index
Eat −0.070 −0.042 −0.343

Eat + Forage 0.573* 0.559* 0.028
Forage 0.818** 0.790** 0.413
Move −0.287 −0.273 −0.021
Rest alone −0.790** −0.776** −0.399
Rest alone + Rest social −0.755** −0.734** −0.350
Rest total −0.797** −0.769** −0.420
a N = 12 for all cells, all tests are 1-tailed.
∗ P < 0.05.
∗∗ P < 0.01.
The relationship between ripe fruit availability and time devoted to

subsistence activity deserves further attention because the observed corre-
lation is in the opposite direction to that predicted. Specifically, the amount
of time that woolly monkeys devoted to foraging for animal prey is signif-
icantly positively rather than negatively related to two of the three indices
of monthly fruit availability (Table V). In fact, in a simple linear regression
model, >50% of the variance in time spent foraging for animal prey is ex-
plained by either of these two indices of ripe fruit abundance (tree index:
R2adj = 0.521, F1,10 = 12.942, P < 0.01; fruit crop index: R2adj = 0.518, F1,10 =
12.836, P < 0.01), and recalculating these regressions using the total amount
of subsistence time devoted to animal prey (forage time plus time spent eat-
ing prey: Table IV) rather than forage time alone as the dependent variable
yielded slightly higher R2adj values (Fig. 3).
Other ecological variables besides ripe fruit availability also undoubt-
edly underlie the variation in time allocation by woolly monkeys. For ex-
ample, temperature conditions can influence time allocation by imposing
direct energetic costs on animals, since extremes of heat or cold force them
to deal with the issue of thermoregulation. Ambient temperature, in fact,
is one of a handful of important predictors of variation in savanna baboon
time budgets and day ranges (Dunbar, 1992). Additionally, even though in-
sects and other animal prey constitute a relatively small proportion of the
diet (Table IV), woolly monkey time budgets may nonetheless be sensitive
to seasonal changes in these resources as well as ripe fruits, particularly
given that time devoted to animal prey foraging is clearly flexible. We there-
fore employed multiple regression to investigate how woolly monkey time
allocation to animal prey foraging and resting was affected by additional
ecological variables. We used the tree index as a single independent vari-
able describing the habitat-wide abundance of ripe fruit because that index
is commonly used in other studies and because it explained only slightly
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Fig. 3. Relationship between time allocation to subsistence on animal prey

and the habitat-wide availability of ripe fruits as measured by the tree index
(A) and the fruit crop index (B).
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less of the variance in time allocation patterns than the fruit crop index did
(<1% difference). Nonetheless, the fundamental results we report here are
essentially unchanged if the latter index is used instead.
Multiple linear regressions using the tree index, monthly average max-
imum temperature in the shade, rainfall, and animal prey abundance as
independent variables are highly significant and explain nearly 70% of the
variance in forage time (R2adj = 0.692, F4,7 = 7.174, P < 0.05) and nearly 77%
of the variance in forage + eat animal time (R2adj = 0.769, F4,7 = 10.098,
P < 0.01). Similarly, >82% of the variance in rest alone time (R2adj = 0.822,
F4,7 = 13.706, P < 0.01), nearly 90% of the variance in rest alone + rest
social time (R2adj = 0.897, F4,7 = 25.008, P < 0.001), and >60% of the vari-
ance in total rest time (R2adj = 0.606, F4,7 = 5.229, P < 0.05) are explained
by these four variables. Moreover, in all but one of these regressions—that
for total rest—the tree index displayed a significant individual P value (less
than 0.05), and the individual P value for our measure of prey abundance is
also significant in the rest alone and rest alone + rest social regressions.
We then proceeded to use stepwise regression to try to tease out which
ecological variables were the best predictors of time allocation to subsistence
activity and resting. We first used a forward selection procedure wherein in-
dependent variables were added to the regression model in order of their
ability to explain variance in the dependent variable until additional vari-
ables no longer explained a significant proportion of that variance. Then,
to examine the robustness of the results of the forward procedure, we per-
formed backward deletion regression, in which all possible variables were
initially included in the model and then those that explained the least amount
of variance were removed one by one until only significant predictors re-
mained.
The results of the stepwise analyses are in Table VI.A. Three ecological
variables remained significant predictors of forage + eat animal, rest alone,
and rest alone + rest social time following the forward stepwise procedure:
the habitat-wide abundance of ripe fruit, the average maximum tempera-
ture in the shade, and our crude estimate of animal prey abundance. In the
backward stepwise models, rainfall again was the only variable not included
when either rest alone or rest alone + rest social time were the dependent
variables, but temperature and prey abundance rather than rainfall dropped
from the regression involving forage + eat animal time. For forage time
alone, the tree index and rainfall are included in both the forward and back-
ward stepwise model. Similar results are obtained if the average minimum
temperature in the shade is included in the stepwise analysis as an addi-
tional variable (Table VI.B.). Taken together, these results implicate ripe
fruit abundance and, to a lesser extent, prey abundance as important factors
affecting the time allocation decisions of Lagothrix, and they further suggest
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466
Table VI. A. Results of stepwise regression analysis of time allocation to animal prey subsistence and to resting on four
ecological variables
Standardized regression coefficients (ß values)

for ecological variables retained in model Overall model
Tree index Prey abundance Max temp Rainfall F p R2adj
Forage
Forward 0.634 Not entered Not entered 0.411 11.55 <0.01 0.657
PP063-295788
Backward 0.634 Removed Removed 0.411 11.55 <0.01 0.657

Forage + Eat animal
Forward 0.500 −0.313 −0.406 Not entered 12.87 <0.01 0.764
Backward 0.640 Removed Removed 0.448 15.37 <0.01 0.723
Rest alone
Forward −0.375 0.372 0.521 Not entered 19.33 <0.001 0.833
Backward −0.375 0.372 0.521 Removed 19.33 <0.001 0.833
May 9, 2001
Rest alone + Rest social

Forward −0.369 0.390 0.546 Not entered 36.58 <0.001 0.907
Backward −0.369 0.390 0.546 Removed 36.58 <0.001 0.907
Total rest
13:40
Forward −0.718 0.378 Not entered Not entered 9.81 <0.01 0.616
Backward −0.718 0.378 Removed Removed 9.81 <0.01 0.616
Di Fiore and Rodman
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Table VI. B. Results of stepwise regression analysis of time allocation to animal prey subsistence and to resting on five ecological variables
Standardized regression coefficients (ß values) for ecological variables
retained in model Overall model
Tree index Prey abundance Max temp Min temp Rainfall F p R2adj
Time Allocation by Woolly Monkeys
Forage
Forward 0.634 Not entered Not entered Not entered 0.411 11.55 <0.01 0.657
Backward 0.828 −0.295 Removed −0.344 0.335 13.30 <0.01 0.817
Forage + Eat animal
PP063-295788
Forward 0.500 −0.313 −0.406 Not entered Not entered 12.90 <0.01 0.764
Backward 0.809 −0.264 Removed −0.299 0.379 15.90 <0.01 0.845
Rest alone
Forward −0.375 0.372 0.521 Not entered Not entered 19.33 <0.001 0.833
Backward −0.375 0.372 0.521 Removed Removed 19.33 <0.001 0.833
Rest alone + Rest social
Forward −0.369 0.390 0.546 Not entered Not entered 36.57 <0.001 0.907
May 9, 2001
Backward −0.369 0.390 0.546 Removed Removed 36.57 <0.001 0.907

Total rest
Forward −0.932 0.380 Not entered 0.393 Not entered 10.31 <0.01 0.717
Backward −0.932 0.380 Removed 0.393 Removed 10.31 <0.01 0.717
13:40
467
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that either ambient temperature or rainfall (or some other climatic variable
related to these two) also plays an important role. The stepwise regression
analysis for total rest time, which incorporates inactivity during heavy rains,
again includes both the abundance of ripe fruit and the abundance of prey in
the model. Maximum ambient temperature, however, is no longer included
as a third significant predictor. This is not surprising given that maximum
temperatures on days of heavy rain tend to be lower than at other times.
This result underscores the fact that rain sheltering is unlikely to be func-
tionally equivalent to other forms of resting, which was our initial rationale
for suspending activity sampling during periods of heavy rain.
The standardized regression coefficients (Table VI) show that woolly
monkeys devoted more time to subsistence on animal prey and less time to
rest when ripe fruit was more abundant (as shown in the initial bivariate
regressions discussed above), during cooler or wetter months, and when po-
tential prey items were less abundant. They devoted more time to resting and
less time to animal prey foraging during hotter portions of the year, when
potential prey were more common, and when ripe fruit was less abundant.
Ripe fruit abundance was the most important predictor of time allocation
to foraging—and to rest when rain sheltering was included in that activity
category—while maximum ambient temperature is the most important pre-
dictor of time spent resting otherwise. The fact that time spent foraging for
animal prey is negatively associated with presumed prey availability in one
of the foraging regressions might suggest that woolly monkeys are forced
to spend more time looking for animal food when prey are scarce, partic-
ularly if they need to maintain a minimal daily intake of animal nutrients.
However, woolly monkeys in Yasunı́ also consumed more animal prey in
months when they spent more time looking for it, despite prey scarcity: the
partial correlation between monthly time allocation to foraging and abso-
lute prey consumption—measured as the number of prey feeding events
seen per hour of observation each month—while holding prey abundance
constant is r = 0.772 (N = 12, P < 0.01). This indicates that they were not
simply spending more time foraging to acquire the same amount of prey.
DISCUSSION
Time Allocation Patterns and Foraging Strategy
The analysis of woolly monkeys’ time budgets revealed significant sea-

sonal variation in the proportion of time devoted to various activities and
demonstrated that it was related to ecological variables characterizing food
availability and climatic conditions. Rather than adjusting their basic feeding
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time or their overall amount of time spent in subsistence activity in response

to changes in food availability, woolly monkeys specifically adjust time al-
location to foraging for and feeding on animal prey. They appear to limit
animal prey foraging during hot or dry periods of the year, which may be a
strategy to reduce thermoregulatory stress or, alternatively, may result from
a generally lower abundance of arthropods in neotropical forests during
drier months (Smythe, 1982; Wolda, 1978, 1982). More importantly, how-
ever, woolly monkeys seem to concentrate on animal prey foraging during
months of high fruit availability, when nutritional requirements are other-
wise easily met.
Dunbar (1992) suggested that ecologically stressed groups of baboons
may reduce rest and social time for increased foraging time under poor
ecological circumstances. Woolly monkeys seem to make a slightly different
tradeoff, opting specifically for greater prey foraging time under ecologically-
permissive conditions and resting more under conditions of low fruit abun-
dance and high ambient temperatures. This suggests that woolly monkeys
are following a mixed strategy of [1] minimizing energy use during peri-
ods of resource scarcity, and [2] taking advantage of periods of ripe fruit
abundance—when they may be able to meet daily nutritional requirements
more efficiently—to forage for greater quantities of animal prey. Strier
(1992) proposed that muriquis (Brachyteles arachnoides) might follow a sim-
ilar mixed strategy, maximizing energy intake when ripe fruits and flowers
are abundant and minimizing energy expenditure when these high quality
resources are scarce. We suggest that for woolly monkeys, nutrients oppor-
tunistically consumed in excess of those required for survival are perhaps
laid down as fat stores. This idea is consistent with published observations
of marked seasonal variation in body condition in some populations of
Lagothrix (Yost and Kelly, 1983; Peres 1991, 1994b) and with the reports
of indigenous hunters in Ecuador, who told us that they preferentially hunt
woolly monkeys when they are fat.
This strategy of trading-off time allocation to animal prey foraging
versus rest can be contrasted with several alternative responses to vary-
ing resource abundance. First, under conditions of resource scarcity woolly
monkeys could try to increase the rate at which new patches of resources are
encountered by traveling faster or farther each day. However, if the distance
traveled each day were increased without a substantial increase in travel rate,
time moving should increase as food abundance declines, which it did not in
this study (Table V). Secondly, the rate of encountering new patches could
be increased, without concomitant changes in time allocation, by increasing
group spread and thereby widening the search radius of a group. Moreover,
increasing group spread (or splitting into subgroups for foraging) should
reduce intragroup competition over limited resources without necessarily
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forcing individuals to alter time allocation patterns (Symington, 1987, 1988;

Janson, 1992; Peres, 1996).We did not systematically collect data on group
spread, but the groups did seem to spread out over larger areas during pe-
riods of low food abundance, as reported by Peres (1996) for a population
of woolly monkeys in western Brazil, which suggests that this strategy may
complement the time allocation tradeoff. Thirdly, woolly monkeys could try
to deal with resource scarcity by broadening their diet to include different
or less-preferred items when more-preferred items become scarce. For ex-
ample, they might increasingly feed on leaves or other nonfruit items as the
habitat-wide abundance of ripe fruits decreases (Schoener, 1971; Charnov,
1976; Stephens and Krebs, 1986). A negative, but nonsignificant, relationship
between fruit availability and the proportion of the diet comprising nonfruit
items suggests that such a dietary shift may occur in our study population. Fi-
nally, during periods of scarcity, woolly monkeys could try to increase energy
intake over the short term without altering feeding time by feeding faster
when in a food patch, i.e., by consuming more food items per unit time dur-
ing periods when they are actually feeding. We did not collect quantitative
data on individual feeding rates, so we are unable to test the extent to which
woolly monkeys utilize this strategy.
The foraging strategy of woolly monkeys in Yasunı́ is quite different
from that of other populations of Lagothrix and from other frugivorous
platyrrhines. Ripe fruit pulp is still by far the major component of the diet of
Lagothrix lagotricha poeppigii in Yasunı́. Nonetheless, they spend roughly
as much time searching for animal prey as they spend feeding, and animal
prey constitutes almost 10% of the overall diet (Table IV). In this respect,
their foraging resembles that of capuchins (Cebus olivaceous: Robinson,
1986; Miller, 1992) and other generalist primates. Isbell (1991) has pointed
out that broad dietary classifications such as frugivore or folivore, based
simply on the relative proportions of different items in the diet, may not
adequately reflect which critical food resources shape foraging decisions.
Based on dietary proportions alone, woolly monkeys rank among the most
highly frugivorous primates. If, however, we classify diets by the proportion
of time that individuals allocate to different food items (Isbell, 1991), then
animal prey clearly takes precedence over ripe fruits as the single most
important dietary item (Table IV).
This is not the first study to conclude that foraging for animal prey
is a critical component of the foraging ecology of woolly monkeys. Milton
and Nessimian (1984) recovered portions of least 133 individual insects of
≥12 families and 6 orders from the stomach of a single wild-caught adult
female Lagothrix lagotricha lagotricha in northwestern Brazil. Moreover,
they noted that at least some of the types of insects were probably difficult
ones to capture, e.g., some coleopterans and homopterans, leading them to
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conclude that woolly monkeys may actively seek out prey during the late
wet to early dry season transition when the woolly monkey specimen was
collected. Milton and Nessimian (1984) also suggested that because of the
buffering effect of large body size, woolly monkeys may be able to afford the
extra energy needed to seek out insect prey, even if the abundance of that
prey type is low. Perhaps significant is the observation that woolly monkeys
in Yasunı́ travel an average of 2.5 km each day, a distance over four times the
diameter of the home range (Di Fiore, 1997), which may constantly bring
them into new areas where the local abundance of insect prey has not been
depleted.
Intraspecific Comparisons
Several long-term studies of other subspecies of woolly monkeys have

recently been published; thus, it is possible to compare populations of
Lagothrix across their geographic range (Figure 1A), and to place them in a
broader comparative perspective within the Atelinae and within the subor-
der Platyrrhini (Table VII). Of particular interest is the strong commitment
to animal prey foraging in woolly monkeys from the western end of their
geographic range. In addition to the Yasunı́ population, woolly monkeys in
the Tinigua National Park in southeastern Colombia (Stevenson, et al., 1994)
show a large commitment to subsistence on animal prey. In that population,
23% of the diet consisted of arthropods. In contrast, invertebrates consti-
tuted only 4.9% of the diet of Lagothrix lagotricha lagotricha from the central
Amazon at Caparú (Defler and Defler, 1996), though published data on ad-
ditional time spent foraging for animal prey is not available. Finally, animal
prey comprised <0.1% of the diet of a group of Lagothrix lagotricha cana
living close to the Rı́o Urucu in western Brazil, where feeding data compa-
rable to the Yasunı́ data were collected (Peres, 1994a). Part of the difference
in the use of animal resources may be attributable to differences in body
size among the four subspecies of lowland woolly monkeys—Yasunı́ and
Tinigua host the two smaller subspecies (Lagothrix lagotricha poeppigii and
L. l. lugens), while the two larger subspecies (L. l. lagotricha and L. l. cana)
occur at Caparú and Urucu, respectively (Peres, 1994b). The limited data on
body weights of wild caught woolly monkeys indicate that body size differ-
ences may be considerable: adult male Lagothrix lagotricha cana can weigh
>30% more than male L. l. poeppigii (mean 9.5 versus 7.1 kg), while adult
female L. l. cana can weigh >65% more than female L. l. poeppigii (7.65 ver-
sus 4.53 kg) (Peres, 1994b; Lu, 1999). Individuals of larger subspecies may be
less able to rely on insects as a source of protein, or they may be less efficient
at finding or capturing insects, than members of smaller subspecies are.
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Table VII. Time budget data for woolly monkeys and other New World primates
Activity categorya
Total Total Total
Genus Species Subspecies E F Subsistence M RA RS Rain Rest SA ON Other Referenceb
Lagothrix lagotricha poeppigii 19.0 17.2 36.2 34.5 16.7 3.8 2.7 23.2 4.7 1.4 6.1 1
Lagothrix lagotricha lagotricha 25.8 38.8 29.9 4.9 0.7 5.5 2
Lagothrix lagotricha lugens 36.0 24.0 36.0 4.0 3
Ateles belzebuth 22.2 9.7 61.0 7.1 4
Ateles chamek 29.0 26.0 45.0 0.0 5
Ateles geoffroyi 10.8 27.6 54.1 7.5 6
Alouatta fusca 17.0 11.0 72.0 0.0 7
Alouatta palliata 16.2 10.2 66.2 7.3 8
PP063-295788
Alouatta seniculus 12.7 5.6 78.5 3.2 9

Brachyteles arachnoides 18.8 29.4 49.3 1.6 0.9 2.5 10
Brachyteles arachnoides 27.8 9.7 61.0 1.6 11
Cebus capucinus ∼54 ∼7 ∼15 ∼11 ∼13 ∼24 12
Cebus olivaceus ∼18 ∼28 ∼46 ∼22 ∼15 ∼11 ∼5 ∼16 13
Cebus apella 24.3 64.5 88.8 6.8 4.3 0.2 14
May 9, 2001
Cebus apella 16.0 50.0 66.0 21.0 12.0 1.0 15

Cebus albifrons 22.0 39.0 61.0 21.0 18.0 0.0 15
Saimiri sciureus 11.0 50.0 61.0 27.0 11.0 1.0 15
Saguinus imperator 17.0 34.0 51.0 21.0 25.0 3.0 15
13:40
Saguinus fuscicollis 16.0 16.0 32.0 20.0 44.0 4.0 15

Callicebus moloch 23.0 3.0 26.0 19.0 54.0 1.0 15
Cacajao calvus calvus 36.0 35.0 29.0 0.0 16
aE= eat, F = forage, M = move, RA = rest alone, RS = rest social, Rain = rain sheltering, SA = social activity, ON = other nonsocial.
b 1.
this study; 2. Defler, 1995; 3. Stevenson, et al., 1994; 4. Klein and Klein, 1977; 5. Symington, 1988; 6. Richard, 1970; 7. Mendes, 1985; 8. Milton, 1980; 9.
Gaulin and Gaulin, 1982, 10. Strier, 1987; 11. Milton, 1984; 12. Fedigan, 1993; 13. Robinson, 1986; 14. Janson, 1985; 15. Terborgh, 1983; 16. Ayres, 1986.
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Other significant differences might also differentiate western Ama-

zonian from central Amazonian woolly monkeys. First, although a large
proportion of time is devoted to moving in all well-studied populations
(Table VII), home range sizes are 4–10 times as large for central Amazonian
Lagothrix as they are for the two western subspecies (Stevenson, et al., 1994;
Defler, 1996; Peres, 1996). In fact, the Yasunı́ groups utilized the smallest
home ranges reported for any population of the species (108 to 124 ha),
while also traveling the greatest distances each day (Di Fiore, 1997). Ad-
ditionally, compared to populations of Lagothrix elsewhere, woolly mon-
keys in Yasunı́ and Tinigua seem to live in somewhat smaller and much
more spatially cohesive social groups. The mean monthly group spread
of 431 m for Lagothrix lagotricha cana in western Brazil (Peres, 1996) is
considerably greater than the typical axis of group spread in Yasunı́: 100–
200 m. Moreover, while the Caparú group regularly split into two indepen-
dent subgroups for foraging (Defler, 1996), we observed this only once at
Yasunı́.
The reasons for this striking variation in the natural history of lowland
woolly monkeys across their geographic range are unclear, but subtle dif-
ferences in habitat quality or in the density of potential animal and plant
food sources may provide a partial explanation. Defler and Defler (1996)
noted that the quality of the soil at Caparú is quite poor relative to that
at Tinigua where Lagothrix lagotricha lugens was studied (Stevenson, et al.,
1994), and they suggested that differences in soil quality may be generally im-
portant to explain variation in the foraging ecology of woolly monkey popu-
lations throughout Amazonia. More comparative data are needed, however,
to test the generality of this explanation.
Interspecific Comparisons
Despite this intraspecific variablity, woolly monkeys are uniformly very

active compared to other platyrrhines: between 60% and 75% of their over-
all time budget is spent in subsistence activity or movement, while only
21% to 36% is spent resting. Only capuchins (Cebus), squirrel monkeys
(Saimiri), and some species of tamarins (Saguinus)—three other genera that
forage heavily for insects—allocate less time to rest than woolly monkeys do
(Table VII). With respect to the other atelines, the large amount of time
that all subspecies of lowland Lagothrix devote to movement and the small
amount they devote to resting is perhaps not surprising. Muriquis (Milton,
1984; Strier, 1986, 1987) and howlers (Milton, 1980; Gaulin and Gaulin, 1982;
Mendes, 1985) utilize leaves to a much greater extent than woolly monkeys
do, they seldom forage for insect prey, and they spend a greater proportion
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of the day resting. These observations are consistent with the inference that
Alouatta and Brachyteles follow a more energetically conservative foraging
strategy than that of Lagothrix, as is expected for folivores (Milton, 1980;
Strier, 1992). What is intriguing, however, is how greatly the time budgets of
Lagothrix differ from those of Ateles, which also feed predominantly on ripe
fruits. Typically, ripe fruits constitute ≥70% of the diet of spider monkeys
(Ateles belzebuth: Klein and Klein, 1975; A. geoffroyi: Cant, 1977; Chapman,
1987; A. paniscus: van Roosmalen, 1985, van Roosmalen and Klein, 1988;
A. chamek: Symington, 1987). Similarly, ripe fruit pulp constitutes the single
largest class of items in woolly monkey diets: 60% for Lagothrix lagotricha
lugens at Tinigua (Stevenson, et al., 1994), 67% for L. l. cana at Urucu (Peres,
1994a), 83% for L. l. lagotricha at Caparú (Defler, 1995), and 77% for L. l.
poeppigii both in Yasunı́ (this study) and along the Rı́o Pacaya (Soini, 1990).
In marked contrast to woolly monkeys, spider monkeys seldom (if ever)
specifically forage for animal prey. While Symington (1987) noted Ateles
chamek consuming caterpillars and termites on occasion and Milton (1993)
reported that A. geoffroyi also occasionally ate insects, prey foraging and
consumption by Ateles is largely opportunistic whereas in Lagothrix such
behavior is clearly deliberate and directed. Moreover, while Ateles gener-
ally devote as much time to moving as Lagothrix, the fact that they do not
forage for prey makes the total amount of time they spend on subsistence
activity considerably less than that of Lagothrix and the proportion of time
they spend resting correspondingly much greater.
Spider monkeys and woolly monkeys also differ from each other in
social organization. Contrary to some early reports of subgrouping behavior
in woolly monkeys (Kavanaugh and Dresdale, 1975; Ramirez, 1980, 1988), it
does not appear that they adopt the typical spider monkey pattern of fission-
fusion sociality as a means of coping with changes in resource abundance. For
spider monkeys, foraging party size is very flexible, correlated with both the
overall abundance of resources in the environment (Symington, 1987, 1988)
and with the size and density of resource patches being used (Chapman, 1990;
Chapman, et al., 1995). In contrast, woolly monkey groups remain socially
cohesive while foraging, though an individual’s behavior while foraging is
generally not coordinated with the activities of other individuals except when
multiple individuals coalesce in fruiting trees to feed on ripe fruits.
Stevenson, et al. (1994) suggested that the high proportion of arthropods
in the diet may help to explain the cohesive nature of woolly monkey so-
cial organization relative to that of fission-fusion societies. In particular, they
postulate that insect feeding might mitigate the costs of being displaced from
patches of fruit by conspecifics (Stevenson, et al., 1994). This is a compelling
suggestion, especially given that the two woolly monkey populations with
the highest proportion of animal prey in the diet—Tinigua and Yasunı́—also
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show greater intragroup cohesion and have smaller home ranges than either
the Urucu or Caparú study groups. If this hypothesis were true, however, we
would predict that insect foraging would be greatest during periods of fruit
scarcity. Instead, we found that time allocation to prey foraging and actual
prey consumption in Yasunı́ were, in fact, greatest during periods of fruit
abundance, and a similar pattern is evident for the Tinigua population from
the phenological and diet data presented by Stevenson, et al. (1994).
CONCLUSIONS
Woolly monkey monkeys in Yasunı́ National Park, Ecuador expended

considerable effort foraging for animal prey, and this effort was greatest
when ripe fruit—the predominant component in the diet—was most abun-
dant in the habitat. This strategy also seems to characterize the Tinigua pop-
ulation of Lagothrix in Colombia, but contrasts with the ecological strategies
of woolly monkeys in the central Amazon where foraging for and consuming
animal prey is uncommon. The strong commitment to animal prey foraging
in some populations of Lagothrix seems to coincide with greater intragroup
cohesion, smaller home range size, and smaller group sizes. We suggest that
woolly monkeys may strategically forage intensively for insects in areas or
seasons of high fruit abundance. Perhaps by thus concentrating on animal
protein, woolly monkeys in western Amazonia are able to sustain higher
reproductive rates and thereby reach higher population densities than those
at more resource-impoverished central Amazonian sites.
However, it is still unclear what ecological conditions either allow or
require some populations to devote so much subsistence activity to insect re-
sources, or what conditions allow Lagothrix in general to associate in larger,
more permanent groups in the same forests where Ateles shows the classic
pattern of fission-fusion sociality. Gross differences in habitat quality be-
tween sites in terms of the availability of ripe fruits are likely to explain some
of the geographic variation in time allocation patterns and intragroup cohe-
sion of Lagothrix. Obviously, more data on soil quality and on the abundance,
distribution, and seasonal variation of potential food plants at different Ama-
zonian sites are needed, as are demographic data on interbirth intervals and
population densities of Lagothrix at presumed resource-rich versus resource-
poor sites. Peres (1991) reports that wild Lagothrix lagotricha cana probably
have interbirth intervals of around three years. While comparable data are
not yet available for Tinigua or Yasunı́, we predict shorter intervals. Simi-
larly, we predict that in western Amazonia the average number of fruiting
trees per hectare fed upon by Lagothrix will be greater or will show less
seasonal variance than in the central Amazon.
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ACKNOWLEDGMENTS
Special thanks are due to the government of Ecuador and to the officials
of INEFAN and the Ministerio de Medio Ambiente for permission to con-
duct this research, and for their continuing interest in our studies of primates
in the Ecuadorean Oriente. Dr. Luis Albuja of the Escuela Politécnica Na-
cional of Quito, who served as our Ecuadorean counterpart in this research,
deserves thanks for the much-appreciated help and friendship he provided
throughout the study. Drs. Laura Arcos Terán and Alberto Padilla of the Pon-
tifı́cia Universidad Catiólica de Ecuador, the administrators of the Estación
Cientı́fica Yasunı́ where we were based for the majority of this work, pro-
vided fundamental logistical support, and Maxus Ecuador, Inc. generously
contributed additional logistical assistance. Special thanks are due to Dr.
Kristin Phillips and Brian Smith who provided invaluable help with many
aspects of the fieldwork. An earlier version of this manuscript benefited
from the suggestions of Drs. Alexander Harcourt, Katherine Milton, Kristin
Phillips, Clifford Jolly, and two anonymous reviewers. Funding for this study
was provided by grants from the University of California, Davis, the National
Science Foundation, the LSB Leakey Foundation, and the Wenner-Gren
Foundation for Anthropological Research.
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International Journal of Primatology, Vol. 22, No. 3, 2001

Time Allocation Patterns of Lowland Woolly

We investigated the time allocation decisions of lowland woolly monkeys

450 Di Fiore and Rodman

for such intraspecific variation should be a goal of any analysis of comparative

Time Allocation by Woolly Monkeys 451

We worked between August 1994 and April 1996 in Yasunı́ National

452 Di Fiore and Rodman

A profile of meteorological conditions at the site is in Fig. 2A. During

Study Species and Groups

Woolly monkeys are large-bodied, highly frugivorous primates that

Time Allocation by Woolly Monkeys 453

454 Di Fiore and Rodman

Table I. Composition of the principal study groups

Males Adult 3/2a 4/4a

to 110 independent individuals, utilized various portions of the study site,

Time Allocation by Woolly Monkeys 455

456 Di Fiore and Rodman

Table II. Behavioral categories and definitions

scanned individual. Beyond this distance, we recognized two additional prox-

Time Allocation by Woolly Monkeys 457

Determination of Time Budgets

We conducted a total of 4296 scans in 95 days over the 12 mo of the study.

To provide an independent estimate of the density, distribution, and

458 Di Fiore and Rodman

Potential Prey Resources

We were unable to measure seasonal variation in the abundance of

Time Allocation by Woolly Monkeys 459

Time Allocation Patterns

Diets and Time Allocation

Yearly mean 19.0 17.2 34.5 23.2 4.7 1.4 354.50

Apr 1995 78.2 5.1 0.0 3.9 12.8 13.5 20.5

Jun 1995 78.0 7.9 0.0 1.2 12.9 13.6 22.0

Oct 1995 75.3 9.4 3.8 2.6 8.8 15.8 13.4

Yearly mean 76.3 7.5 3.6 2.9 9.7 17.2 19.0

a Calculated as % eat time—(proportion of diet consisting of animal prey × eat time).

462 Di Fiore and Rodman

Ecological Influences on Time Budgets

If part of the monthly variance in time budgets of Lagothrix is due to

Time Allocation by Woolly Monkeys 463

Table V. Spearman rank correlation coefficients (corrected for ties) between

Eat −0.070 −0.042 −0.343

The relationship between ripe fruit availability and time devoted to

464 Di Fiore and Rodman

Fig. 3. Relationship between time allocation to subsistence on animal prey

Time Allocation by Woolly Monkeys 465

Standardized regression coefficients (ß values)

Tree index Prey abundance Max temp Rainfall F p R2adj

Backward 0.634 Removed Removed 0.411 11.55 <0.01 0.657

Rest alone + Rest social

Backward −0.369 0.390 0.546 Removed Removed 36.57 <0.001 0.907

468 Di Fiore and Rodman

Time Allocation Patterns and Foraging Strategy

The analysis of woolly monkeys’ time budgets revealed significant sea-

Time Allocation by Woolly Monkeys 469

time or their overall amount of time spent in subsistence activity in response

470 Di Fiore and Rodman

forcing individuals to alter time allocation patterns (Symington, 1987, 1988;

Time Allocation by Woolly Monkeys 471

Several long-term studies of other subspecies of woolly monkeys have

Alouatta seniculus 12.7 5.6 78.5 3.2 9

Cebus apella 16.0 50.0 66.0 21.0 12.0 1.0 15

Saguinus fuscicollis 16.0 16.0 32.0 20.0 44.0 4.0 15