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Benefits and costs of resident males to females in white-faced capuchins Cebus capucinus.

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American Journal of Primatology 32235-248 (1994)
Benefits and Costs of Resident Males to Females in
White- Faced Capuchins, Cebus capucinus
Department of Anthropology, University of Alberta, Edmonton, Alberta, Canada
Some benefits and costs of resident males to females are examined in
white-faced capuchins (Cebus capucinus) at Santa Rosa National Park,
Costa Rica. A total of 380 hours of focal data were collected on adults in
two groups, between January and July 1991. The results of this study
suggest that for females, males provide some greater benefits, and impose
some higher costs than do other females. Males are more vigilant than
females, and are somewhat more successful in detecting predators. To the
extent that predator protection is a major benefit of group living, this
benefit seems to derive more from males than from females. Increased
contest competition is the major cost of group living, and the study suggests that females bear a higher proportion of this cost than males. More
foraging related agonism occurs between males and females than between
females, more aggression occurs between than within sexes, and female
foraging success is negatively associated with agonistic interactions involving males. However, female foraging success is negatively affected by
the proximity of other females, and not by the proximity of males. Differences in the distribution of male benefits and costs according to female
dominance rank are suggested. o 1994 Wiley-Liss, Inc.
Key words: Cebus capucinus, sex differences, vigilance, competition
Female reproductive strategies are considered to be the predominant factor
shaping primate social systems, since male strategies depend largely on the distribution and behavior of females [Emlen & Oring, 1977; Wrangham, 1980; Mitchell et al., 19911. However, the presence of resident males within a social group
provides several benefits for females. These include male participation in group
defense against predators, which decreases the probability of mortality for females
and infants, and the exclusion of non-group conspecifics, which contributes indirectly to female reproductive success by maintaining or increasing a group’s access
to resources [Robinson, 1988; van Schaik, 1989; O’Brien, 19911. In addition, male
Received for publication July 2, 1993; revision accepted October 18, 1993.
Address reprint requests to Lisa M. Rose, Department of Anthropology,WashingtonUniversity, Campus
Box 1114, One Brookings Drive, St. Louis, MO 63130-4899.
0 1994 Wiley-Liss, Inc.
236 I Rose
vigilance may provide protection for females and their infants, allowing them more
time for foraging and social activity. Males also may contribute to infant care by
carrying, grooming, or “babysitting” infants, and providing protection from conspecifics [Altmann, 1980; Izawa, 19801. Concomitantly, the addition of males to a
group imposes costs on females, primarily by increasing feeding competition. Individual success in contest competition is strongly influenced by social factors,
especially dominance rank and aggressive behavior [Janson, 1985; 1990al. In
many primate species, males are dominant over females, and ma.y obtain a disproportionate share of food resources through their greater success in contest competition.
This study examines particular short-term benefits and costs of resident males
to females in two groups of white-faced capuchins, Cebus capucinus. These arboreal neotropical primates live in multi-male, multi-female groups averaging about
15 individuals [Fedigan et al., 1985; Massey, 19871. Most groups have well-defined
home ranges, and while overlaps do occur, a t least some boundaries are defended
[Oppenheimer, 1982; Buckley, 19831. Females generally remain and breed in their
natal groups, whereas most males migrate to other groups prior to maturity [Robinson & Janson, 19871. Males are 25-35% larger than females and are individually
dominant over them, although the alpha female ranks immediately below the
alpha male in some groups [Fedigan, 19931.
I: For females, the benefits made available by males within a group
exceed the benefits made available by existing female group members. In
this section, I compare the extent to which males and females respectively provide
particular benefits for the females within a social group. Predictions associated
with this hypothesis are: 1)Males are more vigilant than females, thereby relieving requirements for female vigilance and increasing the likelihood of predator or
conspecific group detection. 2) Males are more active than females in detecting and
confronting predators, thereby playing a greater role than females in predator
protection. 3) Males are more active and aggressive than females in intergroup
encounters, thereby playing a greater role in intergroup competition.
II: For females, the costs imposed by males within a group exceed the
costs imposed by existing female group members. In this section, I compare
the extent to which males and females respectively impose particular costs on
females within a social group. Predictions associated with this hypothesis are: 1)
Females engage in more agonistic interactions with males than with other females, especially in a foraging context, suggesting greater contest competition
between females and males. A counter-prediction is that agonism reflects patterns
of proximity between and within the sexes. 2) Males tend to direct aggression
toward females, while females direct submissive signals toward males, suggesting
that females are disadvantaged in agonistic interaction with males. 3) Female
foraging is more often interrupted by males than by females. 4) Female foraging
success decreases with increased time spent in proximity to males, suggesting that
the presence of males has a negative impact on female foraging. A counter prediction is that foraging success is similarly reduced by proximity to others in all
sex combinations. 5) Female foraging success decreases with increased agonism
involving males, suggesting that male agonism has a negative impact on female
foraging. A counter-prediction is that foraging success is similarly reduced by
agonism in all sex combinations. The two hypotheses are not mutually exclusive,
and may be independently accepted or rejected.
Benefits and Costs of Males I 237
Study Site and Groups
Santa Rosa National Park is situated near the Pacific coast of Costa Rica,
about 35 km northwest of Liberia, Guanacaste. Santa Rosa covers approximately
10,800 ha of dry tropical forest, semi-evergreen forest, and reclaimed pasture in
varying stages of secondary succession. Potential predators on capuchins at Santa
Rosa include constricting snakes, felids, tayras, and raptors [Freese, 1983; Chapman, 19861. The study focused on two white-faced capuchin groups occupying nonoverlapping home ranges. Group Los Valles (LV) included 3-4 adult males, 4-5
adult females, 2-4 immatures, and 4-6 infants. Group Cerco de Pietra (CP) included 1-3 adult males, 4 adult females, 4 immatures, and 3-4 infants. Both
groups were habituated to human observers. Visibility varied throughout the two
ranges, but observation distances of five to ten metres were common. Individuals
were identified by size, variation in brow color and length, and facial characteristics. Dominance hierarchies were constructed for each sex from the direction of
aggression and submission within dyads.
Data Collection and Analysis
Preliminary observations for this study were made in May-July 1990 and
January 1991. Intensive behavioral sampling of adults was conducted from February to July 1991. I collected 380 hours of focal data in the form of ten-minute
continuous-time sessions [Altmann, 19741, and additional ad libitum data on comparatively rare events such as predator mobbing and intergroup encounters. Data
were collected in alternating ten-day blocks for each group. Given the difficulty of
locating particular group members in this arboreal species, it was not practical to
follow a rigid sampling regime. I attempted t o sample each focal subject within a
group before beginning a new round of observations, and kept a continuous record
of the number of samples obtained for each individual to ensure that sampling was
evenly distributed. If a focal subject was lost from view for more than 90 seconds,
the session was ended and the data discarded. The nearest three neighbors within
ten metres of the focal animal were noted immediately before and after each
session. Agonistic behaviors were scored as “directed” or “received,” and other
participants noted. Most behavioral categories were standard ethogram units [see
Rose, 1992 for full description]. Vigilance behavior (SCAN) occurred when an
animal remained stationary and alert, gazing beyond the immediate vegetation.
ALARM calls [GYRRAHS; Boinski, 19931 were included in the analyses only if a
potential predator was subsequently seen by the observer, or if the call elicited a
response from other group members. MOBBINGS involved all or most group members gathering near and repeatedly threatening a potential predator, typically
with repeated barking vocalizations, branch shaking and breaking, and openmouthed threats [Oppenheimer, 19681. AVOID was scored if an individual clearly
moved away from an approaching group member, and was typically preceded by
slight body tension and rapid glances toward the approaching individual. Since an
AVOID was only recorded if I was able to see the associated approach, this category
may be somewhat under-represented in the data.
Vigilance behavior was expressed as a proportion of observation time for each
focal subject. The proportions were transformed using the arcsine method, and
tested for sex and group differences using a two-way analysis of variance. Significance was set at the P < 0.05 level. Hourly rates of agonism involving males and
females respectively, were calculated for each individual, and divided by the num-
238 I Rose
ber of potential partners available. Since some males were not present for the
entire study period, the number of potential partners in male-female and malemale interactions reflected the proportion of focal samples for which each male was
available. Hourly rates of directed aggression and submission were similarly calculated for all sex combinations. A Wilcoxon-Mann-Whitney test [Siegel & Castellan, 19881 was used to examine the adjusted agonism rates for intergroup differences. Wilcoxon matched-pairs signed-rank tests [Siegel & Castellan, 19881
were used to compare respective interaction rates with males and females for each
sex (female interactions with males versus females; male interactions with males
versus females). Wilcoxon-Mann-Whitney tests were used to compare male-male
and female-female interaction rates, and male-female versus female-male rates
of aggression and submission. Individual mean hourly rates of foraging interruptions were calculated from frequencies of supplantation and avoidance during feeding, and subjected to Wilcoxon tests (interrupts by males and females respectively).
Foraging success was expressed as the percentage of foraging time spent feeding
[Janson, 1990bl. Rates of male and female proximity were calculated from scans
made at the start and end of each focal session. Female proximity was calculated
as the total score for females observed within ten metres of the focal animal,
divided by the number of samples obtained for the focal animal and by the number
of possible female interactants. Male proximity was similarly calculated, allowing
for the factor of male availability. A Spearman’s rank order correlation test [Siegel
& Castellan, 19881was used to test the relationships between proximity, agonism,
and foraging success.
I: Male Benefits to Females
1. Vigilance behavior. As predicted, males were more vigilant than females.
On average, males devoted 8.0% (SE 0.80) of their time to vigilance behavior
(SCAN), significantly more than the 3.2%(SE 0.17) spent by females (F = 42.47,
P < =0.001). No significant intergroup difference was detected (F = 1.87, P =
0.199). The alpha male was the most vigilant individual in each group.
2. Predator detection and encounters. Males tended to give more confirmed predator alarms than females per sample period (Table I), but there were
insufficient data for statistical analysis. I was able to discern the initial caller in
two mobbing situations; in both cases, the CP alpha male gave the initial alarm.
I observed a total of six confirmed snake mobbings (three involving boa constrictors) and several mobbings in which I was unable to confirm the target’s identity.
Typically, the entire group participated in mobbings. Adult males typically approached the predator most closely, and were more boisterous and more persistent
in threatening and following the predator than females.
3. Intergroup encounters. Males were involved in slightly more agonistic
intergroup encounters than females. However, I observed only six cases, all involving the LV group and a small neighboring group in areas of range overlap.
Three episodes involved only males, one primarily females, and two included both
sexes. Contrary to what was predicted, females involved in intergroup encounters
appeared to be more aggressive than males. I observed several females fighting,
whereas male encounters mainly consisted of prolonged threats and chases. However, males were more persistent than females, and tended to patrol the area,
scanning intently, for up to 30 min following an actual encounter. Aggression
during intergroup encounters occurred almost exclusively within sex classes.
Benefits and Costs of Males I 239
TABLE I. Confirmed Predator Alarms by Males and Females in Each Group, Including
Focal and ad lib Observations
LV males
LV females
(n = 4)
CP males
(n = 3)
CP females
(n = 4)
Total males
(n = 7)
Total females
(n = 8)
"Total alarms by each sex divided by number of individuals of that sex. For males, reflects percentage of total
observation time that each male was present in group.
11: Male Costs to Females
1. Agonism between and within sexes. As predicted, females were involved
in more agonistic interactions with males than with other females, both in total
(Wilcoxon T + = 34, n = 8, P = 0.012) and during foraging (T+ = 36, n = 8, P
= 0.004). Males tended t o have more agonistic interactions with females than with
other males, but the difference was not significant for either total agonism (T =
21, n = 7, P = 0.148) or foraging-related agonism (T+ = 17.5, n = 7, P = 0.344).
More agonism occurred in group CP than in group LV (Wilcoxon-Mann-Whitney
Wx = 40, m = 7, n = 8, P = 0.0361, but the pattern of greater agonism between
sexes is consistent in both groups (Fig. la,b).
Patterns of agonism did not reflect patterns of proximity in either group (Fig.
2). In both groups, females had other females in proximity more often than males
(Wilcoxon T + = 31, n = 8, P = 0.039).However, males had females in proximity
more often than other males (T+ = 26, n = 7, P = 0.0231, and I therefore
examined the relationship between individual proximity and agonism rates. Correlations were non-significant for both males (proximity of females and female
agonism: rho = -0.500; proximity of males and male agonism, rho = 0.696) and
females (proximity of females and female agonism: rho = -0.258; proximity of
males and male agonism: rho = 0.09).
2. Direction of aggressive and submissive signals. There was a tendency
for males to direct more aggression toward females than females directed toward
males in group LV (Fig. 3a), but male to female aggression did not significantly
exceed female to male aggression in either group (LV: Wilcoxon-Mann-Whitney
Wx = 15, m = 4, n = 4, P = 0.243; CP: Wx = 11, m = 3, n = 4, P = 0.314). The
greater proportion of female to male aggression in group CP was primarily due to
frequent aggression by one female. Aggression between males was more common
than aggression between females in group LV (Wx = 11, m = 4, n = 4, P = 0.0291,
but not in group CP (Wx = 9, m = 3, n = 4, P = 0.200). The greater proportion
of female-female aggression in group CP was again largely due to one female, and
the absence of any aggressive interactions between two LV females. In both
240 I Rose
Group LV
Group CP
Group LV
Group CP
Fig. 1. Distribution of (a) agonism and (b)foraging-related agonism between and within sex classes, expressed
as mean hourly rate (and SE)per dyad. Rates adjusted to reflect partner availability. LV: n = 4 females, 4
males; CP: n = 4 females, 3 males.
groups, more aggression occurred between sexes than within sexes (Fig. 3a). Females directed more aggression to males than to females (Wilcoxon T + = 27, n =
8, P = 0.016),and males directed more aggression to females than to males (T+
= 33, n = 7 P = 0.02). Males rarely injured females, but a male in group CP killed
a two-day-old infant shortly after rising from beta to alpha position in the male
dominance hierarchy.
In both groups, females directed more submissive signals to males than males
directed to females (Fig. 3b; Wilcoxon-Mann-Whitney Wx = 28, m = 7, n = 8, P
= 0.002). The few submissive signals directed by males to females (n = 5) were
from low ranking males toward an alpha female. All females directed more sub-
Benefits and Costs of Males / 241
Group LV
Group CP
Fig. 2. Proximity between and within sex classes, expressed a s mean rate (and SE)per sample per dyad. Rates
adjusted to reflect partner availability. LV n = 4 females, 4 males; CP: n = 4 females, 3 males.
missive signals toward males than toward other females (Wilcoxon T + = 36, n =
8, P = 0.004). Submissive signals within sexes occurred too rarely to permit statistical analysis, but the number of submissive signals between females (n = 17)
exceeded those between males (n = 3). All observed cases were directed by low
ranking individuals to high ranking individuals.
3. Foraging interruptions. As predicted, female foraging bouts were more
often interrupted by males than by other females (T+ = 36, n = 8, P = 0.004).
Interruptions by males were broadly distributed, with the alpha female in each
group having the fewest interruptions. Only low ranking females had their foraging interrupted by other females. Interruptions to male foraging bouts were too few
(n = 7) to permit statistical analysis. In three cases, low ranking males were
interrupted by a higher ranking male, in three cases, low ranking males were
interrupted by an alpha female, and in one case, a male was supplanted from a
small fruit tree by two mid-ranking females.
4. Proximity of others and foraging success. Contrary to what was predicted, there was no significant relationship between female foraging success and
male proximity (rho = -0.167). However, female foraging success was negatively
correlated with female proximity (rho = -0.762, n = 8, P < 0.05, two-tailed).
There was also a tendency for male foraging success to decline with increased
proximity of females, but this did not reach statistical significance (rho = -0.678).
There was no significant relationship between male foraging success and proximity of other males (rho = 0.143).
5. Agonism involving males and female foraging success. As predicted,
female foraging success decreased with increased foraging-related agonism involving males (Fig. 4; rho = -0.714, n = 8, P < 0.10, two-tailed). Female foraging
success was not significantly correlated with foraging-related agonism involving
other females (rho = -0.077). Male foraging success was not significantly correlated with agonism involving either females (rho = 0.473) or other males (rho =
-0.571). The females with the highest foraging success were the highest and
lowest ranked female in each group (Fig. 4).
242 I Rose
E .El
-2 . M
2 .f
Group LV
Group CP
Fern to Fern
E .a
-h m
8 .$
J = M
Group LV
Group CP
Fig. 3. Distribution of (a) aggressive and (b)submissive signals directed between and within sex classes,
expressed as mean hourly rate (and SE) per dyad. Rates adjusted to reflect partner availability. LV: n = 4
females, 4 males. CP: n = 4 females, 3 males.
Male Benefits to Females
This study suggests that males afford greater benefits to females than do other
females in the form of vigilant behavior, with qualitative trends toward greater
predator detection and defense. Since predation risk may be the primary pressure
selecting for group living in primates [van Schaik, 1983; Terborgh & Janson, 1986;
Mitchell et al., 19911, these results suggest that for C. capucinus females, a major
benefit of group living derives largely from resident males. Comparatively greater
vigilance by males has been reported for capuchins and other primate species
[Fedigan, 1993; Fragaszy, 1986,1990; de Ruiter, 1986; Robinson, 1988; van Schaik
& van Noordwijk, 1989; Gautier-Hion, 1980; Cheney & Seyfarth, 1981; Boinski,
1988aI. It cannot be assumed that male vigilance is directed solely (or even pri-
Benefits and Costs of Males I 243
m 2
0 2
Female Foraging success
(% of foraging time spent feeding)
Fig. 4. Relationship between foraging-related agonism (hourly rate per female-male dyad, adjusted for partner
availability) and female foraging success. n = 8 females; numbers indicate female dominance rank. Rho =
-0.714, P < 0.05, two-tailed test.
marily) toward predator detection, since vigilance can also serve a social function
[Caine & Marra, 19881 or be directed toward conspecific groups [Baldellou &
Henzi, 19921. The high level of vigilance by alpha males, and the tendency for
males to focus on extragroup males during intergroup encounters suggest that
male-male competition is a significant factor influencing male vigilance in the
study groups. The object of male vigilance in Santa Rosa capuchins is currently
under investigation [Rose & Fedigan, in press]. However, as Baldellou and Henzi
[1992] have noted, the intent of male vigilance is irrelevant from a female’s perspective, provided that it results in her receiving warning of potential dangers.
Since the samples of predator and intergroup encounters are small and data
are primarily qualitative, the findings in the following sections are tentative.
However, the tendency for more vigilant individuals to be more successful in detecting predators is consistent with previous findings for other capuchin species
[van Schaik & van Noordwijk, 19891. Males, particularly the CP alpha male, performed better than females in detecting snakes. Since males spend considerably
more time than females on and near the ground [Gebo, 1992; Rose, 19941, their
success in snake detection may reflect differential substrate use as well as higher
levels of vigilance. Constricting snakes are a confirmed predator on young capuchins at the study site [Chapman, 19861, and the contribution of males in detecting
them is thus likely to be significant. The persistence of males in pursuing and
harassing large snakes may provide additional warning and protection for females
and their infants. Male white-faced capuchins have been reported to kill a venomous snake by throwing branches at it [Boinski, 1988bl.
Few intergroup encounters were observed during the study period, and the
tendency for males to be more active than females is only weakly and qualitatively
supported. The rate of aggressive intergroup encounters was unusually low for this
particular site, perhaps in part because the alpha (and only) male in the nearest
group overlapping the LV range had previously been an LV group member. In
244 I Rose
previous years, such encounters have been recorded far more often, and primarily
involved the males of each group forming opposing lines and repeatedly “bluff
charging” toward each other, until one group retreated and was routed (L.M.
Fedigan, pers. comm.). Long-term data suggest that females may benefit from
exclusion of unfamiliar males by resident males. A number of females and infants
have disappeared following incursions by immigrant males, and one female and
three infant deaths were confirmed during a male takeover of CP group in 1993
(L.M. Fedigan & K. MacKinnon, pers. comm.).
Male Costs to Females
The results of this study suggest that resident males impose a greater cost on
females than do other females in the form of agonistic interactions. Rates of agonism in both study groups were low (334 behaviors in 377 h), but over 70% of adult
agonistic interactions were between males and females. Aggressive signals were
more common than submissive signals because aggressive signals frequently elicited an aggressive response, especially between high ranking or mid-ranking individuals. Most agonistic exchanges consisted of open-mouthed threats. More aggression occurred between sexes than within sexes, but aggression was not
unidirectional, suggesting that females are able to successfully counter male aggression in many cases, particularly if a female has a high rank. However, submission was predominantly directed by females toward males, suggesting that
females are disadvantaged in agonistic interactions.
Almost 75% of adult agonism occurred in a foraging context. Agonism between
males and females occurred three times more often during foraging than at other
times, whereas agonism among females was equally as likely to occur during
foraging as at other times. This suggests that agonism between the sexes was
primarily associated with feeding competition. Agonistic interactions with males
adversely affected female foraging success. One finding was that females often
avoided males during foraging, such that an agonistic signal but no aggression was
recorded. Supplantations were less common than avoidance, especially between
males and low ranking females, suggesting that these females may reduce the risk
of aggression from a male by removing themselves from his vicinity. As avoidance
and supplantation result in similar loss of feeding time and access to feeding sites,
it is more appropriate to consider foraging success in terms of total agonistic
interaction, rather than simply aggression received. Janson 11985, 1990al identified aggression received from the alpha male as the most significant factor reducing female foraging success in Cebus apella. However, in these C. capucinus
groups, the alpha male was not notably more aggressive toward females than other
males. Individual differences in behavior did affect patterns of aggression within
the two groups. For example, the aggressive behavior of the beta female inflated
both female-male and female-female aggression rates in group CP. However, the
higher overall rate of agonism in group CP may have been more broadly associated
with a smaller range and reduced food abundance compared with group LV [Rose,
19941, resulting in increased feeding competition.
Feeding competition is recognized as the primary cost of group living [Janson
& van Schaik, 19881. This study suggests that females bear a higher proportion of
this cost than males, since female foraging success is negatively affected by agonistic interaction with males, while male foraging success is not. Agonistic interactions with females did not significantly affect female or male foraging success.
However, female foraging success (and, to some extent, male foraging success) was
negatively affected by the proximity of other females. This suggests that nonagonstic factors also affect foraging success in C. capucinus social groups, and that
Benefits and Costs of Males / 245
these factors are predominantly associated with females. Subtle effects of female
dominance, greater vigilance by males in proximity to females, and the effect of
infants on adult foraging behavior are among the possible factors that warrant
further investigation.
Males in both study groups were typically tolerant and protective of infants,
and considerable infant care by C . capucinus males has been observed at other
study sites ( S . Perry, pers. comm.). However, the killing of an infant by a male
during this study confirms that infanticide occurs in this species, and thus represents a potential cost associated with male group members. The infanticide occurred shortly after a change in male dominance ranks, but the relationship of the
male to the infant (born to the lowest-ranked female) was unknown. A similar
incident has been reported in Cebus olivaceus [Valderrama et al., 1990, case #2].
O’Brien [1991] suggests that Cebus olivaceus females preferentially associate and
mate with the alpha male as a strategy to reduce the risk of infanticide, and that
high ranking females receive greater benefits than low ranking females by “monopolizing” the alpha male.
Distribution of Benefits and Costs Among Females
There was some evidence that the benefits and costs considered in this study
were not equally distributed among the females within each group. High-ranking
females were less likely to avoid or to be supplanted by other individuals than
low-ranking females. As the alpha female in each group ranked immediately below
the alpha male and above other males, she was probably less disadvantaged than
other females in agonistic interactions with males. On the few occasions that a
male avoided a female, the response was directed toward an alpha female. The
alpha female in each group had a strong alliance with the alpha male, and was able
to enlist his support during agonistic interactions and group movement [Rose,
19921. Female success in leading group movement was positively correlated with
dominance rank [Boinksi, 19931. Within each group, the females with highest
foraging success were the highest and lowest ranked. High ranking females tended
to have high proximity to both females and males, while low ranking females were
more peripheral [Rose, 19921. The highest and lowest-ranked females shared the
lowest rate of agonistic interactions with males, suggesting that there may be two
strategies through which a female can improve her foraging success: i) attain high
dominance rank and form strong male bonds, or ii) remain low-ranking and avoid
agonistic encounters with males. However, it is possible that by foraging in less
central areas in the group, low ranking females may forfeit much of the benefit of
male vigilance. High ranking females who forage nearer to males, especially the
highly vigilant alpha male, may benefit more from male vigilance and predator
Balance of Male Benefits and Costs
Primate social systems are often portrayed as a balance between conflict and
cooperation [e.g., Walters & Seyfarth, 1987; O’Brien, 19911. But is the balance an
equal one, or are the options for females more constrained than the options for
males? For example, if the costs of resident males outweigh the benefits that they
provide, could females choose to exclude males from social groups? In this study,
females occasionally formed successful coalitions against a single male, but repeated confrontations would be energetically costly and physically risky as males
are larger and stronger than females, and have larger canines. Continual vigilance
against invading males would also be costly in terms of energy expenditure and
lost foraging or rest time. Thus, excluding males may not be a viable option for C.
246 I Rose
capucinus females, and a mixed-sex social system may persist even if there are
some disadvantages for females. Females may compromise by following strategies
that provide the best balance between the costs of male success in feeding competition and the benefits of male predator protection, in accordance with their rank
in the female dominance hierarchy.
1. Male white-faced capuchins are more vigilant than females, and somewhat
more successful in detecting potential predators.
2. Both males and females are active in intergroup encounters with aggression occurring almost exclusively within sexes.
3. More agonism occurs between than within sexes, especially while foraging,
and female foraging is interrupted more often by males than by females.
4. Female foraging success is negatively correlated with proximity to other
females, but not with proximity to males.
5. Female foraging success is negatively correlated with agonism involving
males, but not with agonism involving other females.
6. Males afford some greater benefits for females than do other females, but
also impose some greater costs.
I thank Linda Marie Fedigan for her support and advice throughout this
project, and the Area de Conservacion Guanacaste for permission to work a t Santa
Rosa. I am grateful to Sue Boinski for sharing her knowledge and observations of
vocalizations, spatial position, and group movement. Marc Koehn gave generously
of his time and expertise in writing the computer programme PRIMA 1.1 for data
entry, and Chuck Humphries assisted with statistical advice. Sue Boinski, Linda
Fedigan, Nancy Lovell, Rick Martin, Jan Murie, and three anonymous reviewers
provided helpful comments on the manuscript. My research was funded by a postmaduate scholarshir, from the Natural Sciences and Engineering Research Council of Canada (NSERCC).
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