Red-capped mangabeys (Cercocebus torquatus) adapt their interspecific gestural communication to the recipient's behaviour.

Sensitivity to recipient's attention and responsiveness are critical markers of intentional communication. Although previous research showed that ape gestures can be intentional, few studies have yet addressed this question concerning monkeys. Here, we characterise the effect of a recipient's presence, attentional state and responsiveness on the interspecific gestural communication of captive red-capped mangabeys (Cercocebus torquatus). Previous reports showed that they produced learnt begging gestures towards a human recipient preferentially when the latter was facing them. We used here a novel setup that allows subjects to move around an experimenter and to use different modalities (visual and acoustic) to communicate. We found that when the recipient was not facing them, mangabeys moved to a position in the visual field of their recipient rather than using attention-getters. Interestingly, unlike apes, they did not elaborate their communication visually or acoustically when the experimenter did not respond favourably to their begging. However, our results may suggest that begging gestures were goal-directed, since mangabeys inhibited them when the experimenter was not available to answer immediately (i.e. give a reward). Overall, red-capped mangabeys' interspecific visual communication presented intentionality features, but their use of begging gestures was less flexible than that of great apes in similar situations.

Maternal Care in Free-Ranging Arboreal Grey-Cheeked Mangabeys (Lophocebus albigena johnstoni) in Kibale National Park, Uganda.

Maternal styles have been intensively studied in a variety of terrestrial species of primates, but far less in arboreal species. However, to have a balanced view of the evolution of maternal care, it is necessary to investigate this behaviour in the context of habitat. Here, we investigate whether the mother's parity, age and dominance rank, as well as the infant's age and sex, influence maternal care and mother-infant proximity in arboreal grey-cheeked mangabeys (Lophocebus albigena). We observed 13 mother-infant dyads in four free-ranging groups for 6 months. Our main finding is that maternal care is a dynamic process affected by a mix of mothers' and infants' characteristics. We found that first-time mothers spent more time watching their infants than multiparous mothers, who carried, groomed and protected their infants more often. We also found that low-ranking mothers prevented their infants from leaving them more often than did high-ranking mothers. Moreover, mothers adjusted their care as infants became older. They groomed and protected female infants more than male infants, behaviours common in female-bonded species. Our study shows the ever-changing dynamics of maternal care related to infant age and highlights the role of the mother's parity and rank in this process.

Changes in physiological stress and behaviour in semi-free-ranging red-capped mangabeys (Cercocebus torquatus) following antiparasitic treatment.

Parasites are ubiquitous in wildlife populations, but physiological and behavioural responses of hosts to infection are difficult to measure. We experimentally treated semi-free-ranging red-capped mangabeys (Cercocebus torquatus) in Nigeria with antiparasitic drugs and examined subsequent changes in glucocorticoid production and individual behaviour. Because both parasites and stress impact energy balance and health, we measured (i) behavioural time re-allocation via activity budgets, (ii) social relationships (e.g. social connectivity and dominance hierarchy stability) and (iii) body condition. We collected triplicate faecal samples (n = 441) from 49 individuals prior to and following treatment. Cortisol levels fluctuated in parallel with parasite abundance. Elevations in cortisol, but not parasitism, were related to reduced body condition. Behaviour also shifted according to infection status, with uninfected individuals spending more time foraging and less time resting and vigilant compared with when they were infected. Time spent feeding, travelling or socializing did not differ between pre- and post-treatment time periods. Group cohesion, but not dominance stability, changed following treatment, suggesting parasite-induced social avoidance. Together, these findings show a coordinated response to infection that promotes host tolerance through stress and energy conservation, reduces transmission risk and increases protection when infected hosts are vulnerable.

Emergent patterns of social organization in captive Cercocebus torquatus: testing the GrooFiWorld agent-based model.

We empirically applied the GrooFiWorld agent-based model (Puga-Gonzalez et al. 2009) in a group of captive redcapped mangabeys (Cercocebus torquatus). We analysed several measurements related to aggression and affiliative patterns. The group adopted a combination of despotic and egalitarian behaviours resulting from the behavioural flexibility observed in the Cercopithecinae subfamily. Our study also demonstrates that the GrooFiWorld agent-based model can be extended to other members of the Cercopithecinae subfamily generating parsimonious hypotheses related to the social organization.

Acoustic variability and individual distinctiveness in the vocal repertoire of red-capped mangabeys (Cercocebus torquatus).

Acoustic variability and individual distinctiveness of vocal signals are expected to vary with both their communicative function and the need for individual recognition during social interactions. So far, few attempts have been made to comparatively study these features across the different call types within a species' vocal repertoire. We collected recordings of the six most common call types from 14 red-capped mangabeys (Cercocebus torquatus) to assess intra- and interindividual acoustic variability, using a range of temporal and frequency parameters. Acoustic variability was highest in contact and threat calls, intermediate in food calls, and lowest in loud and alarm calls. Individual distinctiveness was high in contact, threat, loud and alarm calls, and low in food calls. In sum, calls mediating intragroup social interactions were structurally most variable and individually most distinctive, highlighting the key role that social factors must have played in the evolution of the vocal repertoire in this species. We discuss these findings in light of existing hypotheses of acoustic variability in primate vocal behavior.

The use of fruiting synchrony by foraging mangabey monkeys: a 'simple tool' to find fruit.

Previous research has shown that a considerable number of primates can remember the location and fruiting state of individual trees in their home range. This enables them to relocate fruit or predict whether previously encountered fruit has ripened. Recent studies, however, suggest that the ability of primates to cognitively map fruit-bearing trees is limited. In this study, we investigated an alternative and arguably simpler, more efficient strategy, the use of synchrony, a botanical characteristic of a large number of fruit species. Synchronous fruiting would allow the prediction of the fruiting state of a large number of trees without having to first check the trees. We studied whether rainforest primates, grey-cheeked mangabeys in the Kibale National Park, Uganda, used synchrony in fruit emergence to find fruit. We analysed the movements of adult males towards Uvariopsis congensis food trees, a strongly synchronous fruiting species with different local patterns of synchrony. Monkeys approached within crown distance, entered and inspected significantly more Uvariopsis trees when the percentage of trees with ripe fruit was high compared to when it was low. Since the effect was also found for empty trees, the monkeys likely followed a synchrony-based inspection strategy. We found no indication that the monkeys generalised this strategy to all Uvariopsis trees within their home range. Instead, they attended to fruiting peaks in local areas within the home range and adjusted their inspective behaviour accordingly revealing that non-human primates use botanical knowledge in a flexible way.

Task complexity, posture, age, sex: which is the main factor influencing manual laterality in captive Cercocebus torquatus torquatus?

Behavioural asymmetries reflect brain asymmetry in nonhuman primates (NHP) as in humans. By investigating manual laterality, researchers can study the evolution of brain hemisphere specialisation. Three dominant theories aim to establish an evolutionary scenario. The most recent theory relates different levels of manual laterality to task complexity. Our investigation aimed to evaluate the importance of two extrinsic factors (posture and the need for manual coordination) and two intrinsic factors (age and sex) on the expression of manual laterality by red-capped mangabeys. We observed 19 captive-born mangabeys, in spontaneous situations and under experimental conditions (seven experimental tasks varying in complexity). No directionality was observed in hand preference at the group level whatever the task. But our data revealed an effect of task complexity: more subjects were lateralised than not lateralised for the bipedal task and for the three most complex tasks. Finally, we evidenced an age and a sex effect. We compare our results with data for several other primate species and discuss them in the light of different manual laterality theories.

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.

Postural effect on manual laterality in different tasks in captive grey-cheeked mangabey (Lophocebus albigena).

The authors examined the effects of task complexity and posture on laterality and compared lateralization during different tasks in 9 captive grey-cheeked mangabeys (Lophocebus albigena) during spontaneous food processing and 3 experimental tasks. Comparisons with data of red-capped mangabeys (Cercocebus torquatus torquatus; semiterrestrial species) were used. Less than half the monkeys were lateralized for simple everyday activities, but 6 were lateralized for complex daily activities. Moreover, all the monkeys were lateralized when performing experimental tasks. Laterality at the group level was found for the bipedal task: Mangabeys were right-handed. Complexity of tasks increases laterality at the individual level. Significant differences between the 2 species of mangabeys were found, allowing us to confirm that postural constraints are a major factor in the emergence of group handedness.

Mangabeys (Cercocebus torquatus lunulatus) solve the reverse contingency task without a modified procedure.

Problem solving often relies on generating new responses while inhibiting others, particularly prepotent ones. A paradigm to study inhibitory abilities is the reverse contingency task (Boysen and Berntson in J Exp Psychol Anim Behav Process 21:82-86, 1995), in which two different quantities of food are offered to an individual who receives the array he did not choose. Therefore, mastery of the task demands selecting the smaller quantity to obtain the larger one. Several non-human primates have been tested in the reverse contingency task. To date, only great apes and rhesus monkeys (Macaca mulatta) have succeeded in the original task, with no need of procedural modifications as the large-or-none contingency, correction trials or symbolic stimuli substituting for actual food quantities. Here, four mangabeys were presented with two stimulus arrays of one and four raisins in the context of the reverse contingency task. Three of them learned to perform the task well above chance without a modified procedure. They also reached above-chance performance when presented with two stimulus arrays of zero and four raisins, despite the initial difficulty of choosing a null quantity. After a period of 7-10 months, in which the animals were not tested on any task, all three subjects continued to perform well, even when presented with novel quantity pairs.

Seasonal variation in the feeding ecology of the grey-cheeked mangabey (Lophocebus albigena) in Cameroon.

Seasonal fluctuations in resource abundance often cause primates to change their feeding behavior and ecology. The objective of this study was to examine the response of a largely frugivorous monkey, the grey-cheeked mangabey (Lophocebus albigena), to seasonal variations in fruit abundance. We used 15-min scan sampling to quantify feeding, activity, and habitat use by monkeys between February and December 1998 in the Dja Reserve, Cameroon. L. albigena were found to have omnivorous feeding habits, consuming the fruits, seeds, leaves, and flowers of 132 plant species. Although monkeys fed from many plant species, only five plant species accounted for 45% of all feeding records. The number of feeding observations on a plant species was significantly correlated with its fruit production. L. albigena responded to fruit-lean periods by shifting from a diet dominated by fruit to one dominated by seeds, flowers, and young leaves. This diet shift coincided with greater use of swamp habitat and higher dietary diversity. L. albigena spent the greatest percentage of scan samples feeding and traveling, but activities varied significantly over the day. Individuals spent a significantly higher percentage of scan samples feeding during the fruit-rich season than in the fruit-lean season. Comparing our results to those of studies in Gabon and Uganda, we found that L. albigena differ across regions in the number of plant species they consume and time spent feeding. These differences may be a result of variations in tree diversity or the strength of seasonal fluctuations in resource abundance among sites.