Reconciliation in wild white-faced capuchins (Cebus capucinus)

The likelihood of reconciliation (defined as preferential peaceful contact among former opponents following conflicts) has been predicted to vary positively with relationship value and compatibility, and negatively with relationship security. Long-term data on wild white-faced capuchins (Cebus capucinus) indicate that dyads consisting of an adult female and an alpha male have high value and compatibility, but low security. Two studies of C. capucinus postconflict (PC) behavior were carried out at Lomas Barbudal Biological Reserve, Costa Rica. One study consisted of 30-min PC and matched control (MC) follows. The second study extracted PC and MC periods from long follows, yielding PC/MC periods averaging 105 min. In study 2, but not study 1, significantly more PC/MC pairs were attracted (former opponents affiliated with each other sooner in the PC period than in the MC period) than were dispersed (former opponents affiliated with each other sooner in the MC period than in the PC period). Reconciliation in study 2 could not be explained as a by-product of former opponents' tendency to seek affiliative contact with conspecifics generally, or of the spatial proximity of opponents following conflicts. Attempted reconciliation was less likely to be followed by renewed aggression when reconciliation attempts were delayed following conflicts. The data were insufficient for a formal test of differences in conciliatory tendency (the difference between the number of attracted and dispersed PC/MC pairs, divided by the total number of pairs) among dyad types to be conducted.     

Intergroup encounters in wild white-faced capuchins (Cebus capucinus)

Wrangham (1980) hypothesized that knowledge of the nature of intergroup encounters is crucial to understanding primate social relationships and social organization. I studied a single social group of wild white-faced capuchins over a period of 26 months and observed 44 encounters between social groups during 3703 hr of observation. All intergroup encounters consisted of predominantly hostile social interactions. However, nonaggressive interactions between males of different social groups occurred in a few cases. Adult males were the sole participants in 39 encounters and the primary participants in all 44 encounters. The alpha male was the most frequent participant. High-ranking females participated aggressively in five encounters, and low-ranking females never participated. There was no stable intergroup dominance hierarchy. I hypothesize that the need for male-male cooperation in intergroup aggression is an important factor influencing the quality of intragroup male-male relationships. Behavior during intergroup encounters is consistent with the idea that intergroup behavior is related to male reproductive strategies, but inconsistent with the idea that intergroup aggression is related to female defense of resources. The possibility that males are “hired guns” (Wrangham, 1980) cannot be ruled out.     

Cross-site differences in foraging behavior of white-faced capuchins (Cebus capucinus)

Researchers have identified a variety of cross-site differences in the foraging behavior of free-ranging great apes, most notably among chimpanzees (Pan troglodytes) and more recently orangutans (Pongo pygmaeus), that are not due to obvious genetic or ecological differences. These differences are often referred to as "traditions." What is not known is whether this high level of interpopulation variation in behavior is limited to hominoids. In this study, we use long-term data from three Costa Rican field sites that are geographically close and similar ecologically to identify potential foraging traditions in white-faced capuchins (Cebus capucinus). Foraging traditions are predicted in Cebus because of many behavioral and morphological convergences between this genus and the great apes. The processing techniques used for the same food species were compared across sites, and all differences found were classified as present, habitual, or customary. Proximity data were also analyzed to determine if social learning mechanisms could explain variation in foraging behavior. Of the 61 foods compared, we found that 20 of them are processed differently by capuchins across sites. The differences involve pound, rub, tap, "fulcrum," "leaf-wrap," and "army ant following." For most of the differences with enough data to analyze, the average proximity score of the "matched" dyads (two individuals within a group who shared a "different" processing technique) was statistically higher than the average proximity score of the remaining "unmatched" dyads.     

Carrying and hand-use patterns in Panamanian white-faced capuchins (Cebus capucinus)

The aim of this project was to examine the potential influence of postural regulation on capuchin hand-use patterns by focusing on tasks that involved the carrying of objects. Two months were spent on Barro Colorado Island, Panama, collecting data on ten white-faced capuchins (Cebus capucicus) (four adult males/six adult females). Over 215 contact hours were spent with the monkeys, and a total of 213 carries were recorded. No hand preference bias at the population level was found for the carry task; and no detectable hand-use patterns for carry were correlated with the weight of the object carried (except a right-hand bias for carrying medium-sized objects), the plane of the monkey's movement, its locomotor pattern, or its height above the ground. Therefore, the results of this study do not support the hypothesis that the need for postural regulation while carrying an object influences hand-use patterns in free-ranging capuchins. Our results are especially important because they are contrary to the results of the only other free-ranging capuchin laterality study conducted to date (Panger, 1998).     

Object-use in free-ranging white-faced capuchins (Cebus capucinus) in Costa Rica

Chimpanzees and capuchins demonstrate greater varieties and higher rates of tool-use when compared to other non-human primates. Although capuchins have been studied extensively in captivity, data on their tool-using behavior under free-ranging conditions are limited. This is the first long-term field research to systematically study complex object manipulation in capuchins. The aims of this research are 1) to examine the types, rates, and contexts of tool- and object-use in free-ranging capuchins and 2) to determine if free-ranging capuchins' object manipulation behavior is comparable to the behavior exhibited by captive individuals. Data on 3 troops of white-faced capuchins (Cebus capucinus) were collected from February 1995 to January 1996 at Palo Verde, Costa Rica. Data were collected using focal animal and ad libitum sampling techniques. Any observed incident of tool-use and object-use was recorded. No tool-use was observed during the 11-month study. Object-use (pound, rub, and fulcrum-use) occurred at a rate of 0.19/hr and made up less than 1% of the monkeys' time (there were no differences among the age/sex classes). The results indicate that free-ranging capuchins do not exhibit the range of tool-using behavior demonstrated by their captive counterparts. This may be the result of differential motivational responses to objects, arboreal lifestyle, absence of adequate tool material, and/or absence of food resources that require extraction involving tool-use. Am J Phys Anthropol 106:311–321, 1998. © 1998 Wiley-Liss, Inc.     

Predictors of reproductive success in female white-faced capuchins (Cebus capucinus)

Early investigations into variable reproductive success in nonhuman primates tended to focus on the benefits conferred by high dominance rank. However, the effect of high rank on individual reproductive success has been found to vary both intra- and interspecifically, requiring researchers to expand their investigations to include additional factors. Here we examine the age and rank of the mother, sex of the infant, group size, number of close kin, replacement of group males, and resource availability as possible predictors of female reproductive success in white-faced capuchins (Cebus capucinus) in the Santa Rosa sector of the Area de Conservación Guanacaste, Costa Rica. We examine the length of interbirth intervals (IBI) and infant survivorship as measures of individual reproductive success for the 31 adult females that resided in our three study groups between 1986 and 2007. The greatest predictor of IBI length was whether or not the first infant in the interval survived (number of matrilineal kin and resource availability were also significant predictors); while infant survivorship was most significantly predicted by the occurrence of a turnover in group males in the year following the birth of an infant (infant sex was also a significant factor). Based on these findings, we conclude that male and female reproductive strategies are at odds in this species, with male strategies strongly influencing female reproductive success.     

Behavioral indicators of ovarian phase in white-faced capuchins (Cebus capucinus)

In many primate species, conspicuous behavioral and/or morphological changes are indicators of the fertile phase of the female cycle. However, several primate species, such as the white-faced capuchin, lack these cues. This is referred to as "concealed ovulation," and is argued to be a reproductive strategy that confuses paternity and lowers the risk of infanticide. We studied 10 adult female white-faced capuchins in Santa Rosa National Park, Costa Rica, from January to June 2002. We determined their ovarian cycling patterns by analyzing fecal ovarian hormones, and compared simultaneously collected behavioral data to determine which, if any, cues females use to signal their fertile phases. We found that four females cycled during the study period but ceased to cycle without becoming pregnant. We considered several explanations for the lack of conception during our study, including reproductive seasonality. We found that female C. capucinus showed only small increases in rates of affiliative/proceptive behaviors directed toward adult males during their periovulatory phases. The best indicator of cycle phase was a significant increase in male affiliative behaviors (e.g., following and grooming bouts) and sexual behaviors (e.g., copulations and courtship displays) directed toward females during the periovulatory phase compared to the nonovulatory phase. Our finding that females exhibit little proceptive behavior, but that copulations and male courtship are nonetheless concentrated in periovulatory phases suggests that even though females do not provide behavioral and morphological cues to ovulation, males are still able to detect it. Infanticide occurs with some frequency in these monkeys, and there is evidence for postconceptive mating as a female strategy to lower risk of infanticide via paternity confusion. However, despite this occurrence of nonconceptive mating and the absence of female cues to ovulation, truly concealed ovulation does not appear to be characteristic of this study population of white-faced capuchins.     

Resource patch size and flexible foraging in white-faced capuchins (Cebus capucinus)

White-faced capuchins (Cebus capucinus)on Barro Colorado Island, Panama, have a flexible foraging strategy. Typically, foraging party size is small and individuals feed dispersed from one another. When seasonal fruiting of large volume trees occurs, the majority of the group forages simultaneously. As C. capucinusdo not display a rigorous dominance structure and there are few indications that individuals or coalitions monopolize food patches,individuals are expected to display scramble strategies instead of high frequencies of contest competition. I recorded foraging party size (simultaneous foragers), the total number of animals to feed successively, and the diameter at breast height (DBH) of fruit trees used in two habituated troops. Individuals in each group spent a substantial amount of time — 65 and 48% of foraging time for each group — foraging in party sizes of one. Monkeys predominantly foraged alone in small trees (0- to 20- cm DBH), successively in medium trees (21- to 60- cm DBH), and simultaneously in large trees (>61- cm DBH). They used small trees more frequently than all other tree classes. In medium-sized trees, although fruit was plentiful, space was limited. In these trees Cebusforaged successively. In large-volume trees, space and fruit were abundant and several individuals fed together. As the DBH of fruiting trees increased, the average foraging party size increased exponentially. Cebus capucinusat Barro Colorado Island modify their foraging party size to adapt to the seasonal patterns of fruit production.     

Social factors influencing natal dispersal in male white-faced capuchins (Cebus capucinus)

White-faced capuchin males disperse from their natal group at around 4.5 years of age, but there is much variation in dispersal timing: our youngest confirmed disperser was 19 months and the oldest 11 years old. In this study, we investigate possible factors influencing dispersal decisions in this species. Between 1983 and 2010, 64 males were born into three study groups in Santa Rosa National Park, Area de Conservación Guanacaste, and Costa Rica. As of August 2010, 21 died or were presumed dead (<14 months), 13 remained natal residents, and 30 were presumed dispersers. We used backward logistic regression to identify proximate factors that predict the occurrence of male natal dispersal. The occurrence of a takeover (significant positive association) and group size (nonsignificant negative association) were included in the model. Male age, number of maternal brothers, and number of adult males were not significant predictors of natal dispersal. The resultant model correctly classified 97% of dispersed and 89% of resident natal males, for an overall success rate of 95%. The occurrence of a group takeover was the strongest predictor of male dispersal, with natal males being 18.7 times more likely to disperse in the context of a group takeover than during peaceful times. A linear regression model showed that the tenure length of a male's probable father influences the age of natal dispersal, explaining 15% of the observed variation in age. However, when our oldest disperser was removed (an outlier) this effect disappeared. Collectively, these results indicate that group instability, as evidenced by the occurrence of a takeover, shorter tenure length of a natal male's father, and smaller group size, triggers natal dispersal in this species while the converse leads to a delay. These data add to our growing evidence of the enormous impact that takeovers have on the behavioral ecology of this species.     

Explaining variation in affiliative relationships among male white-faced capuchins (Cebus capucinus)

Relationships among coresident male white-faced capuchins are highly variable, ranging from affiliative to aggressive. In this paper I examine the affiliative relationships of all adult and subadult males residing in four social groups in Santa Rosa National Park, Costa Rica. Relationships among males in two study groups were neutral and tolerant, while in the remaining two groups males were highly affiliative. Male-male dyadic affiliative interactions were examined to determine which variables (group size, sex ratio, age relationship, relationship duration, and rank distance) influence the quality of male relationships within the study groups. Group size explained much of the variation, with males in small groups being more affiliative. However, the duration and history of the relationship among coresident males appears to be the most important variable in understanding male-male relationships within social groups.     

Sex differences and intersexual relations in adult white-faced capuchins (Cebus capucinus)

I aim to explicate the pattern of differences and relations between the sexes in two groups ofC. capucinus, in terms of phylogenetic, social and ecological predictors. I use three lines of evidence to develop predictions as to how male and femaleC. capucinus interact and how the sexes differ in behavior: (1) phylogenetic similarities to other species ofCebus; (2) a general model of sex differences in female-bonded social systems; and (3) ecological analogy of Old World monkeys. First, I conclude that phylogenetic affinity is a good predictor in thatC. capucinus are similar to other species ofCebus in many patterns of sex-differentiated behavior. An exception is that, unlikeC. apella andC. olivaceus, in which a single breeding male is reported to be highly conspicuous in each group,C. capucinus live in a decidedly multimale system. Secondly, the general model of sex differences in female philopatric, male-dispersal societies, which was originally developed for Old World species, also accurately predicts several aspects of social behavior inC. capucinus. Thirdly, a proposed ecological analogy betweenCercopithecus ascanius andspecies of Cebus is not substantiated in this study of C. capucinus, though the analogy is apparently well suited to the social dynamics of other species in the genusCebus.     

Evaluating social influences on food-processing behavior in white-faced capuchins (Cebus capucinus)

Interpopulation variability in patterns of food processing, similar to what is described as "traditional" or "cultural" variation in chimpanzees (Pan troglodytes) and orangutans (Pongo pygmaeus), was identified in white-faced capuchins (Cebus capucinus). However, recent comparisons of food processing in capuchins were conducted only at the population level, with relatively little attention given to variability among groups, age/sex classes, or individuals. This paper examines variability in the processing of specific food types within the context of various social networks (i.e., patterns of association, rank, and kinship) among free-ranging Cebus capucinus at Santa Rosa National Park in Costa Rica. We collected data on two groups of white-faced capuchins in 2001, identifying rates of "food interest" for each individual, as well as forms of processing for specific food types. Juveniles exhibited the most interest in the food-processing behavior of other group members, and food interest was directed most frequently toward adult females. We identified distinctive processing techniques for several food items (Luehea candida pods, Sloanea terniflora fruits, and caterpillars) that facilitated comparisons among individuals within groups. Food-processing techniques for Sloanea fruit and caterpillars appeared to vary independently of the social networks examined in this study. However, we found evidence that variation in Luehea candida processing is to some degree linked to both patterns of association and social rank. The potential influence of these variables on observed food processing patterns warrants further scrutiny.     

The stone tools of capuchins (Cebus apella)

We examined the production of stone took by capuchins (Cebus apella). Eleven subjects used five reduction techniques to produce 346 stone tools (48 cores and 298 flakes). They produced a sharp edge on 83% of the cores and largest flakes. Three monkeys later used a sample of these objects as cutting tools. These results demonstrate that monkeys produce lithic tools analogous to those produced by Oldowan hominids.     

The behavior of white-faced capuchins (Cebus capucinus) at a dry-season waterhole

During a study of white-faced capuchins (Cebus capucinus) in Santa Rosa National Park, northwest Costa Rica, the study troop drank daily from a single terrestrial waterhole for approximately three months during the dry season. The drinking order of troop members at the waterhole was not random: adult males tended to be in the first quartile of the drinking progression, adult females generally occurred with equal frequency throughout the progression, and juveniles drank most commonly in the third quartile. There were also differences in the usual drinking ranks among individuals of the same age-sex class. It is hypothesized that those adult males which are usually first in the progression are increasing their personal fitness by reducing the risk of predation on their offspring and on potential mates. The tendency of adult females to precede juveniles may similarly reflect protection of offspring. The daily visits to the waterhole by the study troop did not appear to greatly restrict the use of their range. It is suggested, however, that the availability of standing sources of water may be an important factor affecting the local distribution and density of capuchins in seasonally and chronically dry areas.     

Sex differences in diet and foraging behavior in white-faced capuchins (Cebus capucinus)

I examined sex differences in diet and foraging behavior in two groups of white-faced capuchin monkeys (Cebus capucinus)in a tropical dry forest at Santa Rosa National Park, Costa Rica. I tested three hypotheses: sex differences in diet and foraging behavior are best explained by (1) sexual dimorphism, (2) the energy demands of pregnancy and lactation for females;and (3) avoidance of competition between the sexes. Sexual dimorphism offered the best explanation of sex differences in the diet and foraging behavior of C. capucinus,accurately predicting that males do more strenuous foraging activity, make less use of small foraging supports, and spend more time on or near the ground. Females spent more time foraging than males did but probably obtained a lower protein yield per unit foraging time. Females exploited more small and embedded invertebrates, while males ate more large invertebrates and vertebrate prey. Pregnant and lactating females spent more time resting and less time foraging than other females did, increasing their foraging return by focusing on foods requiring little handling. There was little evidence of competition avoidance between the sexes.     

Hand preferences on unimanual and bimanual tasks in white-faced capuchins (Cebus capucinus)

This study examined hand preference in white-faced capuchins on a unimanual task and on a coordinated bimanual task. For the unimanual task, handedness was assessed by observing simple reaching for small grains. For the bimanual task, tubes lined with chocolate paste inside were presented to the capuchins. The hand and the finger(s) used to remove chocolate paste were recorded. Seven individuals out of eight in the reaching task and 12 out of 13 in the tube task exhibited a hand preference. Moreover, test-retest correlations showed stability in hand use across time for the coordinated bimanual task. We found no significant differences in strength of hand preference between sexes. Finally, as noted in other primate species, the capuchins were more lateralized in the bimanual task compared to the unimanual task.     

Predictors of Parasitism in Wild White-Faced Capuchins (Cebus capucinus)

Parasite infections in wildlife are influenced by many factors including host demography, behavior and physiology, climate, habitat characteristics, and parasite biology and ecology. White-faced capuchins (Cebus capucinus) host a suite of gastrointestinal and pulmonary parasites, yet the mechanisms affecting host susceptibility and parasite transmissibility have not been examined in this host species. We used the information-theoretic approach (Akaike’s information criterion) and traditional null-hypothesis testing to determine which host characteristics, behaviors, or environmental factors affected the presence of two prevalent capuchin parasites (Filariopsis barretoi and Strongyloides sp.) as well as parasite species richness in four groups of wild capuchins from September 2007 to January 2008 and January to August 2009. Older capuchins were more likely to be infected with Filariopsis barretoi and had higher parasite species richness. Age-biased nematode infections may stem from age differences in capuchin behavior and physiology while high species richness likely results from long- term exposure to numerous parasite species. Infections with Strongyloides sp. were more likely to occur in the dry season when capuchins often descend to the forest floor to drink from terrestrial water sources, potentially increasing their risk of infection from soil-borne larvae. Capuchin behaviors were poor predictors of parasitism, as were female rank, host sex, home range size, and habitat quality. Many of our results were atypical for primate parasitology, suggesting that host–parasite interactions, and subsequently infection risk, may differ in highly seasonal habitats such as tropical dry forests where these monkeys occur.     

Experimental field study of problem-solving using tools in free-ranging capuchins (Sapajus nigritus, formerly Cebus nigritus)

Some populations of capuchins are reported to use tools to solve foraging problems in the wild. In most cases, this involves the act of pounding and digging. The use of probing tools by wild capuchins is considerably less common. Here we report on the results of an experimental field study conducted in southern Brazil designed to examine the ability of wild black-horned capuchins (Sapajus nigritus) to use a wooden dowel as a lever or a probe to obtain an embedded food reward. A group of eight capuchins was presented with two experimental platforms, each housing a clear Plexiglas box containing two bananas on a shelf and four inserted dowels. Depending on the conditions of the experiment, the capuchins were required either to pull (Condition I) or push (Conditions II and III) the dowels, in order to dislodge the food reward from the shelf so that it could be manually retrieved. In Condition I, four individuals spontaneously solved the foraging problem by pulling the dowels in 25% (72/291) of visits. In Conditions II and III, however, no capuchin successfully pushed the dowels forward to obtain the food reward. During these latter two experimental conditions, the capuchins continued to pull the dowels (41/151 or 27% of visits), even though this behavior did not result in foraging success. The results of these field experiments are consistent with an identical study conducted on wild Cebus capucinus in Costa Rica, and suggest that when using an external object as a probe to solve a foraging problem, individual capuchins were able to rapidly learn an association between the tool and the food reward, but failed to understand exactly how the tool functioned in accomplishing the task. The results also suggest that once a capuchin learned to solve this tool-mediated foraging problem, the individual persisted in using the same solution even in the face of repeated failure (slow rate of learning extinction).