Emotional states following grooming in female mandrills.

Grooming is a common cooperative behavior whose exact costs and benefits are still to be fully elucidated. In this study, we evaluated the emotional consequences of giving and receiving grooming in mandrills (Mandrillus sphinx), and how these may change along time after the termination of grooming. We used scratching as a behavioral indicator of anxiety-like emotions. Groomees showed increased scratching immediately after the termination of grooming, while in the subsequent minutes scratching decreased below baseline. The initial increase was larger after longer grooming events, suggesting it represented a case of postinhibitory rebound. The subsequent decline in scratching rates was larger after grooming received by a kin, suggesting interactions with kin are particularly relaxing. Scratching rates shown by groomers were unaffected by grooming interactions. These results highlight that the emotional states following grooming can have a complex time course, and may contribute to explain the inconsistencies found in the previous literature.

Comparative performance of orangutans (Pongo spp.), gorillas (Gorilla gorilla gorilla), and drills (Mandrillus leucophaeus), in an ephemeral foraging task.

A goal of the comparative approach is to test a variety of species on the same task. Here, we examined whether the factors that helped capuchin monkeys improve their performance in a dichotomous choice task would generalize to three other primate species: orangutans, gorillas, and drill monkeys. In this task, subjects have access to two options, each resulting in an identical food, but one (the ephemeral option) is only available if it is chosen first, whereas the other one (the permanent option) is always available. Therefore, the food-maximizing solution is to choose the ephemeral option first, followed by the permanent option for an additional reward. On the original version (plate task), the options were discriminated by the color and pattern of the plates holding the food, while on two subsequent versions we used altered cues that we predicted would improve performance: (1) the color of the foods themselves (color task), which we hypothesized was relevant to primates, who choose foods rather than substrates on which foods are found when foraging, and (2) patterned cups covering the foods (cup task), which we hypothesized would help primates avoid the prepotent response associated with visible food. Like capuchins, all three species initially failed to solve the plate task. However, while orangutans improved their performance from the plate to the color task, they did not for the cup task, and only a few gorillas and no drills succeeded in either task. Unfortunately, our ability to interpret these data was obscured by differences in the subjects' level of experience with cognitive testing and practical constraints that precluded the use of completely identical procedures across species. Nonetheless, we consider what these results can tell us, and discuss the value of conducting studies across multiple sites despite unavoidable differences.

Monitoring of group mates in relation to their behavior in mandrills.

Primates are known to have considerable knowledge about the social relationships that link their group mates and are likely to derive this information from observing the social interactions that occur in their social group. They may, therefore, be hypothesized to pay particular attention to the social interactions involving group mates. In this study, we evaluated how the attention captive mandrills (Mandrillus sphinx) devote to their group mates was modulated by the behavior of the latter. Mandrills looked most frequently at foraging individuals and least frequently at sleeping individuals. Mandrills also looked at grooming individuals more than at individuals that were simply sitting in contact. Grooming dyads were looked at regardless of the social rank and kinship of the individuals involved. These results contribute to our understanding of how primates obtain their social knowledge.

Improving the Welfare of a Zoo-Housed Male Drill (Mandrillus leucophaeus poensis) Aggressive Toward Visitors.

Improving the welfare of nonhuman animals in captivity and maintaining behavioral competence for future conservation purposes is of the highest priority for zoos. The behavior of an aggressive male drill (Mandrillus leucophaeus poensis) was assessed in Barcelona Zoo. The 2-year study presented in this article examined the effects of introducing changes in the exhibit of the drill to improve his welfare by analyzing scan behaviors. First, a partial visual barrier was applied and proved to be insufficient to decrease the long-term stress indicators assessed. Next, a feeding enrichment program was implemented. The results supported the hypothesis that feeding and explorative activities would increase, whereas apathetic and stereotypic behaviors would decrease. However, visitor-directed aggression did not vary, indicating that more profound structural modifications were needed to reduce the negative impact of the agonistic interactions between the drill and the public. The study emphasized the usefulness of environmental enrichment evaluations in assessing captive animal welfare.

Androgens in a female primate: Relationships with reproductive status, age, dominance rank, fetal sex and secondary sexual color.

A comprehensive understanding of the role of androgens in reproduction, behavior and morphology requires the examination of female, as well as male, hormone profiles. However, we know far less about the biological significance of androgens in females than in males. We investigated the relationships between fecal androgen (immunoreactive testosterone) levels and reproductive status, age, dominance rank, fetal sex and a secondary sexual trait (facial color) in semi-free-ranging female mandrills (Mandrillus sphinx), using samples collected from 19 reproductively mature females over 13months. Fecal androgens varied with reproductive status, being highest during gestation. Fecal androgens began to increase at 3months of gestation, and peaked at 5months. This pattern is more similar to that found in a platyrrhine than in other cercopithecine species, suggesting that such patterns are not necessarily phylogenetically constrained. Fecal androgens did not vary systematically with rank, in contrast to the relationship we have reported for male mandrills, and in line with sex differences in how rank is acquired and maintained. Offspring sex was unrelated to fecal androgens, either prior to conception or during gestation, contrasting with studies of other primate species. Mean facial color was positively related to mean fecal androgens across females, reflecting the same relationship in male mandrills. However, the relationship between color and androgens was negative within females. Future studies of the relationship between female androgens and social behavior, reproduction and secondary sexual traits will help to elucidate the factors underlying the similarities and differences found between the sexes and among studies.

Self-protective function of post-conflict bystander affiliation in mandrills.

BACKGROUND: Affiliative interactions exchanged between victims of aggression and individuals not involved in the original aggression (bystanders) have been observed in various species. Three hypothetical functions have been proposed for these interactions: consolation, self-protection and substitute reconciliation, but data to test them are scanty. METHODOLOGY/PRINCIPAL FINDINGS: We conducted post-conflict and matched control observations on a captive group of mandrills (Mandrillus sphinx). We found that victims often redirected aggression to bystanders, that they received most affiliation from those bystanders that were frequently the target of redirection, and that bystander affiliation reduced the likelihood of redirection. Bystander affiliation did not reduce the victim's distress (as measured by its scratching rates) and was not received primarily from kin/friends. Finally, bystander affiliation did not reduce the likelihood of renewed aggression from the original aggressor. CONCLUSIONS/SIGNIFICANCE: These results provide support for the self-protection hypothesis but not for the consolation and substitute reconciliation hypotheses.

Observation of tool use and modification for apparent hygiene purposes in a mandrill.

Tool making or modification to produce a tool of apparent improved functionality has rarely been reported in monkeys, especially when tools are used outside the context of food acquisition. We report on an observation of selection, modification and use of splinters for hygiene purposes in a male mandrill. The zoo-housed animal was video-recorded breaking splinters in sequence to use them underneath his toenails. This record brings forward new evidence that the ability to use and modify tools is not limited to apes and some New World monkeys but is also apparent in Old Word monkeys.

Status of diurnal primate populations at the former settlement of a displaced village in Cameroon.

To understand whether or not primate populations recover in areas of abandoned human settlements, data are required that allow for site-specific comparison over time. Here, we present baseline information on the presence/absence and relative abundance of primate species at the abandoned settlement of the Ekundukundu village, relocated out of the Korup National Park in 2000. Between July 2007 and March 2008, 62 km of transects was surveyed for sightings and calls of primates. All eight species of diurnal primates reported in the KNP were confirmed: Cercopithecus nictitans ludio, C. mona, C. erythrotis camerunensis, C. pogonias, Procolobus pennantii preussi, Cercocebus torquatus, Mandrillus leucophaeus leucophaeus, and Pan troglodytes vellerosus. At old Ekundukundu, C. nictitans accounted for 65% of all primate group sightings. Overall, sighting frequency of primates (0.55 groups/km) was not significantly different from other park sectors surveyed by a previous observer (J. Linder) in 2004-2005. The data reported here will be useful in the long-term monitoring of primate populations in regenerating forest habitats of earlier human settlements.

Informative breath: olfactory cues sought during social foraging among Old World monkeys (Mandrillus sphinx, M. Leucophaeus, and Papio anubis).

Sociality provides a unique opportunity for animals to acquire information and learn from others. Especially during foraging, where trial-and-error food selection might be fatal, conspecifics could act as valuable sources of information. During a six-year study across captive, semifree ranging, and wild Old World monkeys, I investigated whether individuals garnered olfactory-based information from their group mates that could guide their feeding decisions. Each of three study species [mandrills (Mandrillus sphinx), drills (M. leucophaeus), and olive baboons (Papio anubis)] performed a prominent muzzle-muzzle behavior, potentially enabling individuals to smell others' mouths and determine via olfaction what foods their conspecifics had chosen. This muzzle-muzzle behavior (1) was preferentially directed by naïve, younger individuals toward more experienced, older individuals, (2) occurred specifically while recipients were chewing and hence emitting the most potent chemical cues, (3) was typically followed by the actor consuming the very same food type the recipient had been eating, (4) was elicited most often in response to experiments involving novel foods, and (5) occurred less frequently as initially novel foods became more familiar. In contrast to this evidence for information acquisition, there was little support for previous proposals suggesting that muzzle-muzzle functions as a social display. Instead, the omnivorous diets and intensely social lifestyles of mandrills, drills, and baboons, may have each favored a convergent form of information acquisition: seeking out the breath of knowledgeable conspecifics to help decide what foods are safe to eat.

Do captive mandrills invent new gestures?

Studies of intraspecific behavioral variability have documented cases where behaviors are present in some populations or groups but are absent in others. In some cases these differences cannot be explained by recourse to environmental or genetic variation, and may instead represent "traditions". Despite many examples of animal traditions in acoustic communication, relatively few examples exist of gestural traditions. Here I report on a study of communication across eight captive groups of mandrills (Mandrillus sphinx) in which a prominent gesture (Hand extension) was unique to two groups. Habitat variability, genetic differences, and sampling bias were not sufficient to account for the gesture's limited distribution across the study groups. Within the two groups where the gesture did occur only the juveniles in the group performed it, consistently directing it toward adults. Quantitative analysis of the contexts and responses associated with the gesture suggested that juveniles utilized it to provoke adults. Moreover, the gesture appeared to minimize the risk juveniles incurred while inciting adults, suggesting that repeated social interactions shaped the gesture's form. Interestingly, both the groups where the gesture emerged contained few juveniles. With limited play partners, these juveniles may have resorted to harassing adults as an alternative social play outlet. The creation of this novel gesture may thus be due to the combined influence of a shortage of play partners and of the increased free time for playful social exploration afforded by captivity. Although juveniles frequently "eavesdropped" on dyadic interactions involving the gesture and would subsequently initiate an interaction with the recipient of the gesture, there was no definitive evidence for social transmission; the gesture could instead have been independently invented by each juvenile.

Life history in male mandrills (Mandrillus sphinx): physical development, dominance rank, and group association.

We assess life history from birth to death in male mandrills (Mandrillus sphinx) living in a semifree-ranging colony in Gabon, using data collected for 82 males that attained at least the age of puberty, including 33 that reached adulthood and 25 that died, yielding data for their entire lifespan. We describe patterns of mortality and injuries, dominance rank, group association, growth and stature, and secondary sexual character expression across the male lifespan. We examine relationships among these variables and investigate potential influences on male life history, including differences in the social environment (maternal rank and group demography) and early development, with the aim of identifying characteristics of successful males. Sons of higher-ranking females were more likely to survive to adulthood than sons of low-ranking females. Adolescent males varied consistently in the rate at which they developed, and this variation was related to a male's own dominance rank. Males with fewer peers and sons of higher-ranking and heavier mothers also matured faster. However, maternal variables were not significantly related to dominance rank during adolescence, the age at which males attained adult dominance rank, or whether a male became alpha male. Among adult males, behavior and morphological development were related to a male's own dominance rank, and sons of high-ranking females were larger than sons of low-ranking females. Alpha males were always the most social, and the most brightly colored males, but were not necessarily the largest males present. Finally, alpha male tenure was related to group demography, with larger numbers of rival adult males and maturing adolescent males reducing the time a male spent as alpha male. Tenure did not appear to be related to characteristics of the alpha male himself.

Violent coalitionary attack by female mandrills against an injured alpha male.

Female contact aggression against males is relatively rare in species in which the adult males are larger than the females, but it has the potential to influence group structure, male group membership, tenure, and dominance rank. We report an incident in which female mandrills living in a semi-free-ranging group in Franceville, Gabon, attacked a male that was apparently incapacitated after a fight with another male and was unable to escape. The attack involved the alpha male and did not occur in a sexual or infanticidal context. Other adult and adolescent males observed the attack, but when one adult male attempted to participate he was chased away by the females. This observation adds to reports of female coalitions excluding unwanted males from primate groups, or even killing them. The fact that this can also occur in mandrills suggests that females have a degree of control over male group membership, despite the large degree of sexual dimorphism in this species, and highlights the importance of coalitions in primate social organization.