White-faced Capuchins Research Articles

Figs Are More Than Fallback Foods: The Relationship betweenFicusandCebusin a Tropical Dry Forest

In many studies on primate feeding ecology, figs (Ficusspp.) are characterized as fallback foods, utilized only when preferred sources of food are unavailable. However, for white-faced capuchin monkeys (Cebus capucinus) living in northwestern Costa Rica, figs are a consistently important resource and may increase groupwide energy intake. We investigated whether visits to figs affect ranging and behavioural patterns of capuchins. Although daily range length and average travel speed do not differ on days when fig trees are visited, capuchins spend more time in directed travel and more time stationary on “fig days”. Capuchins also increase time spent foraging for fruit and decrease time spent foraging for invertebrates on days when figs trees are visited. Capuchins experience higher energy intake and lower energy output on “fig” days. Thus, the patterns of foraging for figs support an energy-maximization strategy and constitute an important nutritional resource for capuchins.

The effects of provisioning and crop‐raiding on the diet and foraging activities of human‐commensal white‐faced Capuchins (Cebus capucinus)

Non-human primates are coming into increasingly frequent contact with humans and with human-modified environments. The potential for monkeys to survive in such modified landscapes is questionable, and is likely related to a species' behavioral plasticity, particularly as it relates to diet. In this study, I explore the ways in which white-faced capuchins (Cebus capucinus) adjust their diet and foraging behaviors in response to anthropogenic impact. I compare a troop of human-commensal monkeys and a similar troop of wild-feeding monkeys living within the Curú Wildlife Refuge in western Costa Rica for differences in overall diet composition and activity budgets to evaluate the impact of habitat change in this context. The commensal-living white-faced capuchins rely on raided coconut (Cocos nucifera) and oil palm (Elaeis guineensis) crops and provisioned or stolen human foods for over one-half of their total diet. Regardless of this highly anthropogenic diet, the two study troops do not significantly differ in their activity budgets, and the human-commensal troop maintains wild-foraging activities consistent with those of the wild-feeding troop. These data suggest that the white-faced capuchins at this site are responding to anthropogenic disturbance primarily through the exploitation of human food resources, but they do not yet appear to have lost the foraging skills required to survive in this modified landscape on their own. This study adds to our growing body of knowledge on primate survival in matrix habitats, and will hopefully inform primate management plans throughout the Neotropics.

Two girls for every boy: The effects of group size and composition on the reproductive success of male and female white‐faced capuchins

Many factors have been hypothesized to affect the size and adult sex ratios of primate groups and these, in turn, have been argued to influence birth rates. Using park-wide census data collected on a population of capuchins over a 25-year period, we examined whether group size and adult sex ratio affect the per capita reproductive success of male and female white-faced capuchins (Cebus capucinus) in Santa Rosa National Park, Costa Rica. We found that the reproductive success of females (measured as the observed minus the expected ratio of immatures to adult females in the group) decreased with increasing group size, whereas that of males was independent of group size. The proportion of adult males residing in groups had significant, yet contrasting effects on males and females. Male reproductive success was negatively associated with the proportion of males residing in groups whereas female reproductive success increased with the proportion of males. The latter finding supports the intersexual conflict hypothesis, which suggests that a conflict of interest occurs between males and females over adult sex ratios. The effects of group size and composition on the reproductive success of capuchins, a male-dispersed omnivorous species, are similar to those reported for howlers, a bisexually-dispersed folivorous species. One common factor between these taxa is that groups with low ratios of males to females are at greater risk of takeovers and resultant infanticide. Our results suggest that regardless of dietary preference and dispersal pattern, the threat of infanticide can constrain primate group size and composition.

Male dominance and reproductive success in wild white‐faced capuchins (Cebus capucinus) at Lomas Barbudal, Costa Rica

Theory and a growing body of empirical evidence suggest that higher ranking males experience reproductive advantages in group-living mammals. White-faced capuchins (Cebus capucinus) exhibit an interesting social system for investigating the relationship between dominance and reproductive success (RS) because they live in multimale multifemale social groups, in which the alpha males can have extraordinarily long tenures (i.e. they coreside with daughters of reproductive age). Genetic paternity was determined from fecal samples for 120 infants born into three social groups of wild C. capucinus at Lomas Barbudal Biological Reserve, Costa Rica. Alpha males produced far more offspring than expected by chance, and significantly high Nonac's B indices (a measure of deviation from a random distribution of RS among potentially breeding individuals) were a feature of six out of eight male tenures. The likelihood of the alpha male siring a particular offspring was predicted by the kin relationship between the mother and the alpha male, as well as the total number of males and females in the group. The almost complete lack of father-daughter inbreeding [Muniz et al., 2006] constitutes an impediment to alpha male reproductive monopolization in this population, particularly toward the end of long alpha male tenures.

Parasitosis intestinal en monos capuchinos cariblancos Cebus capucinus (Primates: Cebidae) de un área protegida en la provincia de Limón, noreste de Costa Rica

Deforestation of tropical forests is threatening monkey biodiversity and their health status, dependent of an ecologically undisturbed area. To asses this relationship, we analyzed parasite occurrence in their intestines. The study was conducted at the Estación Biológica La Suerte (EBLS), Limón, Costa Rica. The group of white-faced capuchin monkeys (Cebus capucinus) was observed between March and December of 2006. A total of 75 feces samples were obtained. Once a sample was collected, the eaten plant type was identified to family and species level, and feces were processed in the laboratory to determine parasite incidence. Results showed that Moraceae was the most represented family in the samples. Among parasites, Strongyloides spp. and Acanthocephala were the most common. Positive prevalence of parasites was found similar and independent of sex and age of capuchin individuals. Microsporids were mainly reported in feces associated with Piperaceae. A low presence of these parasites was found in samples associated with Myrtaceae, with possible anti-parasite active components. The occurrence of parasites was relatively high in EBLS, when compared to other regions in Costa Rica. The higher occurrence of parasites observed in capuchins at EBLS may be due to the fact that this rain forest is surrounded by areas affected by human activities. We suggest the promotion of research in neotropical primates parasitology, for a better comprehension of the parasite-host relationship, and in a long term, being able to understand the ecosystems where they coexist, and consequently, preserve the biodiversity of the whole region.

Does watching a monkey change its behaviour? Quantifying observer effects in habituated wild primates using automated radiotelemetry

In studies of animal behaviour, researchers have long been concerned that their presence may change the conduct of their study subjects. To minimize observer effects, researchers often habituate their study animals. The premise of this method is that, with sufficient neutral exposure, animals will stop reacting to humans. While numerous studies demonstrate that negative responses to humans decrease over time, the fact that an animal does not flee from or behave aggressively towards observers cannot be taken as evidence that it is not altering its behaviour in other, more subtle ways. Because remotely monitoring the behaviour of wild animals is difficult, it has not been possible to answer the critical question: do habituated animals change their behaviour when researchers are present? Here, we use data from an automated radiotelemetry system that remotely monitored the movement and activity of radiocollared animals to test whether observers affected the behaviour of seven habituated white-faced capuchins, Cebus capucinus. We found no evidence that observers influenced the ranging behaviour or activity patterns of their study subjects. Capuchins did not move faster, stop to rest less frequently, or display higher levels of activity when they were being followed compared to when they were alone. It has been suggested that researchers may embolden habituated study subjects, artificially increasing their relative dominance, but we found no relationship between observer presence and proximity to neighbouring social groups. Although it remains possible that observer effects existed but were too subtle to be detected with the remote sensing technology we used, the results of this study nevertheless provide reassuring evidence that humans can observe habituated wild animals without overly influencing the animals’ activity and movement patterns.

Can color vision variation explain sex differences in invertebrate foraging by capuchin monkeys?

Abstract Invertebrates are the main source of protein for many small-to-medium sized monkeys. Prey vary in size, mobility, degree of protective covering, and use of the forest, i.e. canopy height, and whether they are exposed or embed themselves in substrates. Sex-differentiation in foraging patterns is well documented for some monkey species and recent studies find that color vision phenotype can also affect invertebrate foraging. Since vision phenotype is polymorphic and sex-linked in most New World monkeys - males have dichromatic vision and females have either dichromatic or trichromatic vision - this raises the possibility that sex differences are linked to visual ecology. We tested predicted sex differences for invertebrate foraging in white-faced capuchins Cebus capucinus and conducted 12 months of study on four free-ranging groups between January 2007 and September 2008. We found both sex and color vision effects. Sex: Males spent more time foraging for invertebrates on the ground. Females spent more time consuming embedded, colonial invertebrates, ate relatively more “soft” sedentary invertebrates, and devoted more of their activity budget to invertebrate foraging. Color Vision: Dichromatic monkeys had a higher capture efficiency of exposed invertebrates and spent less time visually foraging. Trichromats ate relatively more “hard” sedentary invertebrates. We conclude that some variation in invertebrate foraging reflects differences between the sexes that may be due to disparities in size, strength, reproductive demands or niche preferences. However, other intraspecific variation in invertebrate foraging that might be mistakenly attributed to sex differences actually reflects differences in color vision.

Dominance among female white-faced capuchin monkeys (Cebus capucinus): hierarchical linearity, nepotism, strength and stability

Research on Old World primates provided the foundation for understanding competitive strategies resulting from social and ecological pressures. The neotropical primate, Cebus capucinus shares many social patterns with Old World cercopithecines (e.g., female philopatry, male dispersal), which may contribute to similar expression of competitive strategies. To clarify the nature of dominance patterns among female white-faced capuchins we examined hierarchical linearity, rank acquisition, matrilineal rank inheritance, hierarchical strength and stability. We collected focal data on 22 adult females (2008) and long-term dominance data (1986–2008) on 33 adult females in Sector Santa Rosa, Costa Rica. Females displayed linear hierarchies based on the direction of dyadic submission. At sexual maturity females quickly acquired rank positions beneath their mother and older sisters. Hierarchies were considered strong based on high proportions of food-related agonism, short latency to detection of hierarchies (21 h/female) and low directional inconsistency scores (<5%). Hierarchies were considered stable based on lack of tied submissive interactions (indicative of uncontested rank positions), low rates of rank change (0.510 changes/year), and long-term stability in matrilineal rank order. These findings enhance our understanding of capuchin social systems and how the competitive strategies of white-faced capuchins compare to those of Old World primates.

Sleep tree use by white-faced capuchins (&lt;i&gt;Cebus capucinus&lt;/i&gt;): implications for differences in seedling composition

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Fig Foraging by Dichromatic and Trichromatic Cebus capucinus in a Tropical Dry Forest

Figs are important resources for frugivores, and Ficus is an ideal taxon for evaluating patterns of primate foraging related to food color. Ficus spp. can be classified as conspicuous (color change from greenish to reddish during ripening) or cryptic (green throughout ripening). To investigate the effect on foraging of color vision phenotype variation for these 2 types of figs, we conducted a 20-mo study on 4 groups of white-faced capuchins (Cebus capucinus) in the Santa Rosa Sector of the ACG, Costa Rica between May 2004 and September 2008. We genotyped all individuals and collected behavioral data on feeding rates, acceptance indices, and foraging sequences. We found a significant effect of fig type; feeding rates and acceptance indices were higher for conspicuous figs than for cryptic figs, and subjects sniffed cryptic figs more often than conspicuous figs. We also found that dichromats sniffed more figs and had longer foraging sequences than trichromats, especially for cryptic figs. Among 6 subtypes of dichromats and trichromats, monkeys possessing the trichromat phenotype with the most spectrally separated L-M opsin alleles showed the highest acceptance index for conspicuous figs, though there were no differences in feeding rates among phenotypes. We conclude: 1) conspicuous figs are visually salient not only for trichromats but also for dichromats, 2) olfaction is important for evaluating edibility of cryptic figs, and 3) the reliance on olfaction for selecting fruit is greater in dichromats. These results indicate divergent foraging strategies among color vision phenotypes for assessing food items.

Collective decision-making in white-faced capuchin monkeys.

In group-living animals, collective movements are a widespread phenomenon and occur through consensus decision. When one animal proposes a direction for group movement, the others decide to follow or not and hence take part in the decision-making process. This paper examines the temporal spread of individual responses after the departure of a first individual (the initiator) in a semi-free ranging group of white-faced capuchins (Cebus capucinus). We analysed 294 start attempts, 111 succeeding and 183 failing. Using a modelling approach, we have demonstrated that consensus decision-making for group movements is based on two complementary phenomena in this species: firstly, the joining together of group members thanks to a mimetic process; and secondly, a modulation of this phenomenon through the propensity of the initiator to give up (i.e. cancellation rate). This cancellation rate seems to be directly dependent upon the number of followers: the greater this number is, the lower the cancellation rate is seen to be. The coupling between joining and cancellation rates leads to a quorum: when three individuals join the initiator, the group collectively moves. If the initiator abandons the movement, this influences the joining behaviour of the other group members, which in return influences the initiator's behaviour. This study demonstrates the synergy between the initiator's behaviour and the self-organized mechanisms underlying group movements.

Conformism in the food processing techniques of white-faced capuchin monkeys (Cebus capucinus)

Researchers of “culture” have long been interested in the role of social learning in establishing patterns of behavioral variation in wild animals, but very few studies examine this issue using a developmental approach. This 7-year study examines the acquisition of techniques used to process Luehea candida fruits in a wild population of white-faced capuchin monkeys, Cebus capucinus, residing in and near Lomas Barbudal Biological Reserve, Costa Rica. The two techniques for extracting seeds (pounding or scrubbing) were approximately equal in efficiency, and subjects experimented with both techniques before settling on one technique—typically the one they most frequently observed. In a sample of 106 subjects that had already settled on a preferred technique, the females adopted the maternal technique significantly more often than expected by chance, but the males did not. Using a longitudinal approach, I examined the acquisition of Luehea processing techniques during the first 5 years of life. Regression analysis revealed that the technique most frequently observed (measured as proportion of Luehea processing bouts observed that used pounding as opposed to scrubbing) significantly predicted the technique adopted by female observers, particularly in the second year of life; the amount of impact of the observed technique on the practiced technique was somewhat less significant for male observers. These results held true for (a) observations of maternal technique only, (b) observations of technique used by all individuals other than the mother, and (c) observations of maternal and non-maternal techniques combined.Electronic supplementary materialThe online version of this article (doi:10.1007/s10071-009-0230-3) contains supplementary material, which is available to authorized users.

The effects of infant births on male–female relationships in Cebus capucinus

Most primates are characterized by cohesive male-female bonds that are maintained year round. While recent studies have addressed the selective pressures influencing the evolution of male-female relationships in primates, we know relatively little about the proximate mechanisms affecting them. It has been demonstrated that newborn white-faced capuchins (Cebus capucinus) attract the attention of other group members and this may be an important mechanism influencing male-female relationships. We studied two groups of C. capucinus in the Area de Conservación Guanacaste, Costa Rica, between February and July 2007. A total of 348 hr of focal data were collected on all adult males (n=6) residing in each of the study groups. During our study, 13 of the 14 group females were either pregnant or lactating, and 9 infants were born. We calculated an average daily affiliation rate between all group males combined and each adult female four weeks before and four weeks after the birth of her infant. Repeated measures ANOVAs revealed no significant changes in affiliation following infant births (F=2.262, df=1, P<0.176). Results remained nonsignificant for rank (F=1.550, df=1, P<0.260) and group membership (F=0.729, df=1, P<0.429). Infant sex was the only variable with a significant effect on affiliation rates between males and females (F=10.020, df=1, P<0.019). Adult males increased their affiliation with all adult females that gave birth to male infants (n=4), while their rates decreased with all but one of the adult females with female infants (n=4). While preliminary, these results indicate that the adult males may cultivate relationships with other males at a young age.

Field Anesthesia and Health Assessment of Free-ranging Cebus capucinus in Panama

Health and disease are critical factors for understanding primate evolution and for developing effective conservation and management strategies. However, comprehensive health assessments of wild primate populations are rare, in part because of the difficulty and risk of chemically immobilizing subjects to obtain the necessary biological samples. We report here the results of a health assessment, as well as the methods and drug dosages used to capture white-faced capuchins (Cebus capucinus) on Barro Colorado Island, Panama. We captured and measured 24 capuchins as part of an ongoing radiotelemetry study, and collected biological samples from 9 of them for a comprehensive health survey. Telazol® was very effective for immobilizing and capturing capuchins, although the doses we used were higher than those recommended for captive capuchins. High Telazol doses immobilized individuals quickly, ensuring that we were able to find and recover the sedated monkeys. The capuchins were generally in good condition, and had few ecto-, hemo-, or intestinal parasites. However, all but 1 of the adults had substantial dental abnormalities. In addition, 1 juvenile had indeterminate genitalia that we believe to be the result of hypospadias. Seven of the capuchins tested positive for exposure to Herpesvirus tamarinus and all individuals tested positive for exposure to Cebus cytomegalovirus. Hematology, serum chemistry, and plasma mineral levels from the wild individuals were, for the most part, comparable to those recorded for captive Cebus capucinus, and contribute to establishing baseline health values for the species.

Effects of gut passage, feces, and seed handling on latency and rate of germination in seeds consumed by capuchins (Cebus capucinus)

One of the key measures of the effectiveness of primary seed dispersal by animals is the quality of seed dispersal (Schupp: Plant Ecol 107/108 [1993] 15-29). We present data on quality of seed dispersal by two groups of white-faced capuchins (Cebus capucinus) in Costa Rica to test the hypothesis that capuchin seed handling results in effective primary dispersal for some fruit species they consume. We examined seed handling for 27 plant species, and germination rates of 18 species consumed by capuchins. For five of the most commonly swallowed seed species, we determined germination rates and average time to germination (latency) for seeds ingested and defecated by capuchins and compared these to seeds removed directly from fruit and planted. For the same five species, we compared germination rates and latency for passed seeds planted in capuchin feces to those cleaned of feces and planted in soil. For three of five species, differences in proportion of germinated seeds were significantly higher for gut passed seeds than for controls. For four of five species, germination latency was significantly faster for gut passed seeds than for controls. Feces had either no effect on seed germination rate or precluded germination. Data presented here support the hypothesis that white-faced capuchins are effective primary dispersers.

Female dispersal in a female-philopatric species, Cebus capucinus

[White-faced capuchins ( Cebus capucinus ) reside in multimale–multifemale groups characterized by female philopatry and frequent male dispersal. However, over the years we have observed five females immigrate into our study groups and 23 disappear/emigrate. We examined long-term demographic and behavioural data on three groups of C. capucinus residing in Santa Rosa, Costa Rica, between 1986 and 2007. During this time, 56 females resided in our study groups and as of June 2007, 23 were still present, ten were confirmed/presumed dead and 23 were missing. Here we review the circumstances surrounding the five immigrations and 23 missing females and evaluate three main hypotheses to explain female dispersal in a female philopatric species: inbreeding avoidance, reduction of intragroup feeding competition and infanticide avoidance. The two main predictions of the inbreeding avoidance hypothesis were not supported by our study; male tenure did not exceed female age at first birth and the majority of dispersers were parous females. The reduction in intragroup feeding competition hypothesis received moderate support; dispersing/disappearing females tend to leave during the dry season and they have fewer matrilineal kin than females remaining in their natal group. Our data were most consistent with the infanticide avoidance hypothesis in that females are more likely to disperse/disappear during years with male replacements, a time when infant deaths are also more common. These data provide further evidence of the large impact that the movement and actions of adult male white-faced capuchins have on the lives of females in this species., White-faced capuchins ( Cebus capucinus ) reside in multimale–multifemale groups characterized by female philopatry and frequent male dispersal. However, over the years we have observed five females immigrate into our study groups and 23 disappear/emigrate. We examined long-term demographic and behavioural data on three groups of C. capucinus residing in Santa Rosa, Costa Rica, between 1986 and 2007. During this time, 56 females resided in our study groups and as of June 2007, 23 were still present, ten were confirmed/presumed dead and 23 were missing. Here we review the circumstances surrounding the five immigrations and 23 missing females and evaluate three main hypotheses to explain female dispersal in a female philopatric species: inbreeding avoidance, reduction of intragroup feeding competition and infanticide avoidance. The two main predictions of the inbreeding avoidance hypothesis were not supported by our study; male tenure did not exceed female age at first birth and the majority of dispersers were parous females. The reduction in intragroup feeding competition hypothesis received moderate support; dispersing/disappearing females tend to leave during the dry season and they have fewer matrilineal kin than females remaining in their natal group. Our data were most consistent with the infanticide avoidance hypothesis in that females are more likely to disperse/disappear during years with male replacements, a time when infant deaths are also more common. These data provide further evidence of the large impact that the movement and actions of adult male white-faced capuchins have on the lives of females in this species.]

Predators at Nests of the Western Slaty Antshrike (Thamnophilus atrinucha)

Predation is the primary cause of nest loss in most passerine species. While nest predation is important, the predator community and behaviors of parents and predators during predation events are poorly documented. I witnessed one and videotaped four predation events at nests of Western Slaty Antshrikes (Thamnophilus atrinucha) in central Panama. Predators included a snake (Pseustes poecilonotus), a monkey (white-faced capuchin [Cebus capucinus]), and three species of birds. Predators spent little time at the nest, yet some returned repeatedly to the same nest. Parents also returned several times to nests after the predation event and parental behaviors varied depending on the predator. Nest disturbance was not an accurate indicator of predator type.

Behavioral adaptations to heat stress and water scarcity in white‐faced capuchins (Cebus capucinus) in Santa Rosa National Park, Costa Rica

We examined thermoregulatory behaviors in a wild population of white-faced capuchins (Cebus capucinus) inhabiting a highly seasonal dry forest in Santa Rosa National Park (SRNP), Costa Rica. The dry season in SRNP lasts approximately 5 months and is characterized by high ambient temperatures regularly exceeding 37 degrees C, low relative humidity, and the near absence of precipitation. This study demonstrates that capuchins rest more and travel shorter distances during the hottest and driest hours of the day, and suggests that they extend their tongues to lower body temperature via evaporative cooling. Seasonal weather patterns and group movement data reported here are based on 940 h of observations on three social groups of capuchins (wet season: 370 h, dry season: 570 h). In the dry season, the proportion of time spent resting increased at higher temperatures whereas the proportion of time spent traveling decreased. Distance traveled between location points taken at half-hour intervals decreased significantly as temperature increased, although the correlation was not strong. Capuchins exposed their tongues during hot, dry, windy conditions, and this behavior was much more frequent in the dry season. Temperature was significantly higher and humidity significantly lower for "tongue-out" events than expected for a random event in the dry season. Finally, as surface water became scarce, home-range areas of heavy use became increasingly centered on the remaining permanent water sources. These results suggest that heat stress and water scarcity are significant influences on the behavior of capuchins in hot, dry conditions.

Vocal Repertoire of Cebus capucinus: Acoustic Structure, Context, and Usage

Researchers studying nonhuman primate vocal repertoires suggest that convergent environmental, social, and motivational factors account for intra- and interspecific vocal variation. We provide a detailed overview of the vocal repertoire of white-faced capuchins, including acoustic analyses and contextual information of vocal production and vocal usage by different age-sex classes in social interactions. The repertoire is a mixture of graded and discrete vocalizations. In addition, there is general support for structural variation in vocalizations with changes in arousal level. We also identified several combined vocalizations, which might represent variable underlying motivations. Lastly, by including data on the social contexts and production of vocalizations by different age-sex classes, we provide preliminary information about the function of vocalizations in social interactions for individuals of different rank, age, and sex. Future studies are necessary to explore the function of combined vocalizations and how the social function of vocalizations relate to their acoustic structure, because social use of vocalizations may play an important role in shaping vocal evolution.

Kin-biased social behaviour in wild adult female white-faced capuchins, Cebus capucinus

*Department of Anthropology and Center for Behavior, Evolution and Culture,University of California, Los AngelesyMax Planck Institute for Evolutionary AnthropologyzDepartment of Psychology, Indiana University(Received 16 October 2007; initial acceptance 30 October 2007;final acceptance 4 January 2008; published online 27 May 2008; MS. number: A10889R)Studies of kin bias in the distribution of social behaviour in group-living matrifocal species generallyunderline the importance of bonds among female kin. However, few studies examine either how kinbias may be affected by variation in the availability of kin or the relevance of paternal kin. In this study,we used genetic and behavioural data to analyse correlates of coalition formation, proximity, groomingand dominance relations among female white-faced capuchins over a 10-year period during which thenumber of adult females in the group varied from 6 to 10. Females sided with the most closely relatedof two opponents when joining coalitions. Both dominance rank and kinship influenced proximity andgrooming patterns. In particular, when group size was small, mean relatedness high and interdyadic var-iation in relatedness low, rank distance was a better predictor of proximity and grooming than was kinshipdistance. However, when group size was large, mean relatedness lower and interdyadic variation in relat-edness higher, females significantly biased their grooming and spatial proximity towards kin. Dominancerank was not so tightly associated with relatedness as in provisioned female-bonded cercopithecines;females did not follow the ‘youngest sister ascendancy rule’. Full sisters, maternal half sisters andmotheredaughter dyads associated at statistically indistinguishable rates, and all associated significantlymore often than paternal half sisters. Paternal half sisters did not associate more often than distantlyrelated femaleefemale dyads (r < 0.125). These and similar results call into question the general impor-tance of paternal kin ties in wild primates.