More than a simple fixed action pattern: Yawning in drills.

In the last decade, increasing attention has been devoted to exploring some aspects of yawning in non-human animals. With their chin red mark, bony paranasal swellings, male large brains and long canines, drills (Mandrillus leucophaeus) offer a robust model for testing hypotheses on the phenomenon. We identified two yawn variants (covered, YCT and uncovered teeth, YUCT) which differ in terms of recruitment of muscular action units (AUs). We tested the effects of several variables (sex, dominance rank, context) on the duration of the yawn and the probability of YCT or YUCT occurrence. We found that males performed longer and more YUCT than females. These findings support the Brain Cooling Hypothesis suggesting that those species showing large brains tend to display larger and longer yawns. We also tested the State Changing Hypothesis predicting the presence of a temporal association of yawning and ongoing behavioral transitions. A sequential analysis revealed that after 30 s following a yawn, drills were significantly more likely to change their behavioral state. Through the observation of yawning, conspecifics might gain knowledge of impending state changes. Seeing other's yawns increased the probability of a similar response in the observers, thus suggesting the presence of yawn contagion in drills. Although the dataset needs to be expanded, our findings indicate that yawning is variable in drills, it can be associated with subjects' state changes, and the imminent shifts can be perceived/processed by conspecifics.

Dental microstructure records life history events: A histological study of mandrills (Mandrillus sphinx) from Gabon.

Accentuated lines in dental microstructure are hypothesized to correlate with potentially stressful life history events, but our understanding of when, how and why such accentuated lines form in relation to stressful events is limited. We examined accentuated line formation and life history events in the teeth of three naturally deceased mandrills (Mandrillus sphinx, Cercopithecidae), for whom we had detailed life history information. We determined the ages at formation of accentuated lines in histological tooth sections and used dates of birth and death to calibrate dental histology to calendar time and individual age. We found accentuated lines that matched their mother's resumption of sexual cycles in two individuals, and possibly in the third individual. The subjects also formed lines when their mothers were mate-guarded by males or wounded. Accentuated lines matched the birth of the next sibling in one of two cases. Both females formed accentuated lines when they experienced their own sexual swelling cycles, but lines did not match all sexual swelling cycles. Mate-guarding matched an accentuated line in one case, but not in another. Lines matched all three parturitions in the two females. Changes in alpha male and captures did not consistently coincide with accentuated line formation, but repeated captures were associated with lines. Using simulated data, we show that the observed number of matches between lines and events would be very unlikely under a null hypothesis of random line formation. Our results support the hypothesis that some life history events are physiologically stressful enough to cause accentuated line formation in teeth. They contribute to our understanding of how primate life histories are recorded during dental development and enhance our ability to use teeth to reconstruct life history in the absence of direct observation.

Seasonality and interindividual variation in mandrill feeding ecology revealed by stable isotope analyses of hair and blood.

Mandrills are large-bodied terrestrial forest primates living in particularly large social groups of several hundred individuals. Following these groups in the wild to assess differences in diet over time as well as among individuals is demanding. We here use isotope analyses in blood and hair obtained during repeated captures of 43 identified free-ranging mandrills (Mandrillus sphinx) from Southern Gabon, to test how dietary variation relates to the season as well as an individual's age and sex. We measured the stable carbon (δ13 C‰) and nitrogen (δ15 N‰) isotope ratios in 46 blood and 214 hair section samples as well as from a small selection of mandrill foods (n = 24). We found some seasonal isotopic effects, with lower δ13 C values but higher δ15 N values observed during the highly competitive long dry season compared to the fruit-rich long rainy season. Variation in δ13 C was further predicted by individual age, with higher δ13 C values generally found in younger individuals suggesting that they may consume more high canopy fruit than older individuals, or that older individuals consume more low canopy foliage. The best predictor for δ15 N values was the interaction between age and sex, with mature and reproductively active males revealing the highest δ15 N values, despite the observation that males consume substantially less animal food items than females. We interpret high δ15 N values in these mature male mandrill blood and hair sections to be the result of nutritional stress associated with intense male-male competition, particularly during mating season. This is the first study showing isotopic evidence for nutritional stress in a free-ranging primate species and may spark further investigations into male mandrill diet and energy balance.

The multidimensionality of female mandrill sociality-A dynamic multiplex network approach.

The structure and dynamics of primate social groups are shaped by the social relationships of its members. These relationships are based on different types of interactions and vary in relation to the identity of the interactants and over time. Social network analysis tools represent a powerful and comprehensive method to characterise social interactions and recent methodological advances now allow the study of the multidimensionality of sociality via multilayer networks that incorporate multiple types of interactions. Here, we use a multidimensional network approach to investigate the multidimensionality of sociality of females in a captive group of mandrills. We constructed two multiplex networks based on agonistic, proximity and grooming interactions of 6-7 mature females to analyse the multidimensionality of relationships within two independent observation periods; and three multiplex networks (one for each interaction type) to examine how relationships changed between periods. Within each period, different individuals were the most central in each layer and at the multiplex level, and different layers (i.e., interaction types) contributed non-redundant information to the multilayer structure. Across periods, relationships based on the same interaction type also contained non-redundant information. These results indicate that female mandrills engage in multidimensional and dynamic relationships, suggesting that in order to represent the full complexity of relationships, networks need to be constructed from more than a single type of interaction and across time. Our results provide evidence for the potential value of the multilayer network approach to characterise the multidimensionality of primate sociality.

Monitoring the social behavior of a bachelor mandrill (Mandrillus sphinx) dyad participating in touchscreen-mediated cognitive testing.

Cognitive testing of primates in zoos is becoming increasingly common. Cognition experiments are generally thought to be beneficial as they provide participants with an opportunity to engage in species-specific cognitive functioning, perhaps more so than with traditional forms of environmental enrichment. However, testing may increase competition and aggression between conspecifics if it has monopolizable features or creates social tension within groups. The purpose of this study was to monitor the social behavior of a bachelor mandrill (Mandrillus sphinx) dyad participating in a touchscreen-mediated cognition study. The mandrills' behavior was monitored before and after testing sessions for 8 months. Positive changes in the mandrills' affiliative behavior were observed. Rates of play, presentations, and silent bared-teeth face increased posttesting. No change in rates of agonism were observed between pre- and posttesting conditions. The observed positive changes in affiliative behavior suggest cognitive testing was enriching for the mandrills and participating in testing improved their welfare. Zoos beginning cognitive studies should monitor participant behavior to ensure their welfare is not compromised and is, ideally, enhanced.

Seasonal glucocorticoid production correlates with a suite of small-magnitude environmental, demographic, and physiological effects in mandrills.

OBJECTIVES: The activation of the hypothalamic-pituitary-adrenal axis is a neuroendocrine response to external and internal changes that animals face on a predictable or unpredictable basis. Across species, variation in glucocorticoid production has been related to such changes. In this study, we investigated the predictable, seasonal sources of variation in the levels of fecal glucocorticoid metabolites (fGCM) in a large natural population of mandrills (Mandrillus sphinx) in Southern Gabon. MATERIALS AND METHODS: Using five years of regular behavioral monitoring and hormone analyses performed on 1,233 fecal samples collected on 99 individuals of both sexes and all ages and General Linear Mixed Models, we studied the three main seasonal predictors of fGCM concentrations: (i) weather conditions, (ii) number of adult males, and (iii) female reproductive status. These three predictors all vary seasonally in mandrills. RESULTS: We first showed an increase in fGCM concentrations during the short dry season while controlling for other factors. Pregnant females, which include the large majority of adult females at this time of the year, mainly drove this increase, although a combination of other small-magnitude, season-related effects linked to climatic events and demographic changes also partly explained this seasonal trend. Indeed, fGCM concentrations increased with both low temperatures (and low rainfall) and high numbers of adult males present in the group. These seasonal changes, while correlated, held true throughout the studied years and when restricting our analyses to a given season. Finally, we found that older mandrills showed on average higher fGCM concentrations than younger ones and that medium-ranked females exhibited the highest levels of fGCMs. DISCUSSION: The observed patterns suggest that plasticity in mandrills' metabolism in the form of glucocorticoid production allows them to adjust to predictable changes in climatic, demographic and physiological conditions by mobilizing and redirecting energetic resources toward appropriate, calibrated seasonal responses.

Dental microwear textural analysis as an analytical tool to depict individual traits and reconstruct the diet of a primate.

OBJECTIVES: Dental microwear is a promising tool to reconstruct animals' diet because it reflects the interplay between the enamel surface and the food items recently consumed. This study examines the sources of inter-individual variations in dietary habits in a free-ranging population of mandrills (Mandrillus sphinx) using a combination of feeding monitoring and in vivo dental microwear textural analysis (DMTA). METHODS: We investigated the impact of seasonality and individual traits on four DMTA parameters. In parallel, we further studied the influence of the physical properties of the food items consumed on these four parameters, using three proxies (mechanical properties, estimates of phytolith and external grit contents). RESULTS: We found that seasonality, age, and sex all impact DMTA parameters but those results differ depending on the facet analyzed (crushing vs. shearing facets). Three DMTA parameters (anisotropy, complexity, and heterogeneity of complexity) appear sensitive to seasonal variations and anisotropy also differs between the sexes while textural fill volume tends to vary with age. Moreover, the physical properties of the food items consumed vary seasonally and also differ depending on individual sex and age. CONCLUSION: Considering the interplay between the tested variables and both dental microwear and diet, we reaffirm that food physical properties play a major role in microwear variations. These results suggest that DMTA parameters may provide valuable hints for paleoecological reconstruction using fragmentary fossil dental remains.

Buccal dental-microwear and dietary ecology in a free-ranging population of mandrills (Mandrillus sphinx) from southern Gabon.

Analyses of dental micro- and macro-wear offer valuable information about dietary adaptations. The buccal surface of the teeth does not undergo attrition, indicating that dental microwear may directly inform about food properties. Only a few studies have, however, investigated the environmental and individual factors involved in the formation of such microwear in wild animals. Here, we examine variation of buccal microwear patterns of mandibular molars in a large free-ranging population of mandrills (Mandrillus sphinx). We first explore the influence of seasonality and individual's sex, age and tooth macrowear-expressed as the percent of dentine exposure (PDE)-on six microwear variables. Second, we analyze the interplay between individual's diet and PDE. In a last analysis, we revisit our results on mandrills in the light of other primate's microwear studies. We show that the average buccal scratch length and the frequency of vertical buccal scratches are both higher during the long dry season compared to the long rainy season, while we observe the inverse relationship for disto-mesial scratches. In addition, females present more disto-mesial scratches than males and older individuals present higher scratch density, a greater proportion of horizontal scratches but a lower proportion of vertical scratches than young animals. PDE yields similar results than individual's age confirming earlier results in this population on the relationship between age and tooth macrowear. Because seasonality and individual characteristics are both known to impact mandrills' diet in the study population, our results suggest that buccal microwear patterns may inform about individual feeding strategies. Furthermore, PDE increases with the consumption of potentially abrasive monocotyledonous plants, independently of the individuals' age, although it is not affected by food mechanical properties. Finally, buccal scratch densities by orientation appear as relevant proxies for discriminating between different primate taxa.

Sexual Selection and the differences between the sexes in Mandrills (Mandrillus sphinx).

Sexual selection has become a major focus in evolutionary and behavioral ecology. It is also a popular research topic in primatology. I use studies of mandrills (Mandrillus sphinx), a classic example of extravagant armaments and ornaments in animals, to exemplify how a long-term, multidisciplinary approach that integrates field observations with laboratory methods can contribute to on-going theoretical debates in the field of sexual selection. I begin with a brief summary of the main concepts of sexual selection theory and the differences between the sexes. I then introduce mandrills and the study population and review mandrill life history, the ontogeny of sex differences, and maternal effects. Next, I focus on male-male competition and female choice, followed by the less well-studied questions of female-female competition and male choice. This review shows how different reproductive priorities lead to very different life histories and divergent adaptations in males and females. It demonstrates how broadening traditional perspectives on sexual selection beyond the ostentatious results of intense sexual selection on males leads to an understanding of more subtle and cryptic forms of competition and choice in both sexes and opens many productive avenues in the study of primate reproductive strategies. These include the potential for studies of postcopulatory selection, female intrasexual competition, and male choice. These studies of mandrills provide comparison and, I hope, inspiration for studies of both other polygynandrous species and species with mating systems less traditionally associated with sexual selection.

Sternal Gland Scent-Marking Signals Sex, Age, Rank, and Group Identity in Captive Mandrills.

Mandrills are one of the few Old World primates to show scent-marking. We combined ethological and chemical approaches to improve our understanding of this behavior in 3 zoo-managed groups. We observed the olfactory behavior performed by adults and adolescents (N = 39) for 775h. We investigated the volatile components of sternal scent-marks using gas chromatography-mass spectrometry and compared volatile profiles with traits of the signaler. Males marked more than females and within each sex the frequency of scent-marking was related to age and dominance status, but alpha males scent-marked most frequently and particularly in specific areas at the enclosure boundaries. We identified a total of 77 volatile components of sternal gland secretion, including compounds functioning as male sex pheromones in other mammals, in scent-marks spontaneously released on filter paper by 27 male and 18 female mandrills. We confirmed our previous findings that chemical profiles contain information including sex, male age and rank, and we also found that odor may encode information about group membership in mandrills. Our results support the hypotheses that scent-marking signals the status of the dominant male as well as playing territorial functions but also suggest that it is part of sociosexual communication.

From frugivore to folivore: Altitudinal variations in the diet and feeding ecology of the Bioko Island drill (Mandrillus leucophaeus poensis).

Variation in the quality and availability of food resources can greatly influence the ecology, behavior, and conservation of wild primates. We studied the influence of altitudinal differences in resource availability on diet in wild drill monkeys (Mandrillus leucophaeus poensis) on Bioko Island, Equatorial Guinea. We compared fecal samples (n = 234) collected across three consecutive dry seasons for drills living in lowland (0-300 m asl) forest with nearby (18 km distance) drills living in montane forest (500-1000 m asl) in the Gran Caldera Southern Highlands Scientific Reserve. Lowland forest drills had a frugivorous diet very similar to that reported from studies on nearby mainland drills (M. l. leucophaeus) and mandrills (M. sphinx), with fruits comprising 90% of their dried fecal samples. However drills living in montane forest had a more folivorous diet, with herbaceous pith, leaves and fungi comprising 74% of their dried fecal samples and fruit becoming a minor component (24%). Furthermore, a dietary preference index indicated that the differences in the proportion of fruit and fibrous vegetation in the diets of lowland compared to montane drills was not simply a result of relative availability. Montane drills were actively consuming a higher mass of the available fruits and fibrous vegetation, a condition reflected in the greater mass of their fresh feces. Our results demonstrate the unexpected flexibility and complexity of dietary choices of this endangered species in two adjacent habitat types, a comparison of considerable importance for many other limited-range species faced with habitat loss and climate change.

Social organization and space use of a wild mandrill (Mandrillus sphinx) group.

Mandrills (Mandrillus sphinx) are enigmatic Old World primates whose social organization and ecology remain poorly known. Previous studies indicated, for example, that groups are composed of only adult females and their young or that several units composed of one adult male and several females make up larger permanent social units. Here, we present the first data on group composition and male ranging patterns from the only habituated wild mandrill group and examine how home range size and daily path length varied with environmental and demographic factors over a 15-month period. Our study site is located in southern Gabon where we followed the group on a daily basis, collecting data on presence, ranging, behavior, and parasite load of its individual members. Throughout the study, the group was made up of about 120 individuals, including several non-natal and natal adult and sub-adult males. One-male units were never observed. The mandrills traveled an estimated 0.44-6.50 km/day in a home range area of 866.7 ha. Exploratory analyses revealed that precipitation, the number of adult males present, and the richness of protozoan parasites were all positively correlated with daily path length. These results clarify the social system of mandrills and provide first insights into the factors that shape their ranging patterns.

New evidence from observations of progressions of mandrills (Mandrillus sphinx): a multilevel or non-nested society?

African papionins are well known for the diversity of their social systems, ranging from multilevel societies based on one-male-multifemale units (OMUs) to non-nested societies. However, the characteristics of Mandrillus societies are still unclear due to difficult observational conditions in the dense forests of central Africa. To elucidate the characteristics of mandrill societies and their social systems, I analysed the age-sex compositions, behaviours, and progression patterns of their horde/subgroups using videos of them crossing open places. The progressions were very cohesive, and the very large aggregations (169-442 individuals) had only 3-6 adult males (1.4-1.8 % of all individuals) and 11-32 subadult males (6.5-7.2 %). No herding behaviours were observed in the males, and most of the small clusters within the progressions were not analogous to the OMUs of a multilevel society but instead consisted of only adult females and immatures. The progressions of alert mandrills showed patterns similar to those observed in a non-nested social system: females with dependent infants were concentrated toward the rear and adult and subadult males toward the front. These results suggest that cohesive aggregations and a female-biased sex ratio are common characteristics of mandrill species. Mandrills may form female-bonded and non-nested societies, although their fission-fusion dynamics may be different from those typical of savannah baboons.

Social structure of a semi-free ranging group of mandrills (Mandrillus sphinx): a social network analysis.

The difficulty involved in following mandrills in the wild means that very little is known about social structure in this species. Most studies initially considered mandrill groups to be an aggregation of one-male/multifemale units, with males occupying central positions in a structure similar to those observed in the majority of baboon species. However, a recent study hypothesized that mandrills form stable groups with only two or three permanent males, and that females occupy more central positions than males within these groups. We used social network analysis methods to examine how a semi-free ranging group of 19 mandrills is structured. We recorded all dyads of individuals that were in contact as a measure of association. The betweenness and the eigenvector centrality for each individual were calculated and correlated to kinship, age and dominance. Finally, we performed a resilience analysis by simulating the removal of individuals displaying the highest betweenness and eigenvector centrality values. We found that related dyads were more frequently associated than unrelated dyads. Moreover, our results showed that the cumulative distribution of individual betweenness and eigenvector centrality followed a power function, which is characteristic of scale-free networks. This property showed that some group members, mostly females, occupied a highly central position. Finally, the resilience analysis showed that the removal of the two most central females split the network into small subgroups and increased the network diameter. Critically, this study confirms that females appear to occupy more central positions than males in mandrill groups. Consequently, these females appear to be crucial for group cohesion and probably play a pivotal role in this species.

Nasopalatine ducts and flehmen behavior in the mandrill: reevaluating olfactory communication in Old World primates.

Compared to other modes of communication, chemical signaling between conspecifics generally has been overlooked in Old World primates, despite the presence in this group of secretory glands and scent-marking behavior, as well as the confirmed production and perception of olfactory signals. In other mammalian species, flehmen is a behavior thought to transport primarily nonvolatile, aqueous-soluble odorants via specialized ducts to the vomeronasal organ (VNO). By contrast, Old World primates are traditionally thought to lack a functional VNO, relying instead on the main olfactory system to process volatile odorants from their environment. Here, in the mandrill (Mandrillus sphinx), we document unusual morphological and behavioral traits that typically are associated with the uptake of conspecific chemical cues for processing by an accessory olfactory system. Notably, we confirmed that both sexes possess open nasopalatine ducts and, in response to the presentation of conspecific odorants, we found that both sexes showed stereotyped behavior consistent with the flehmen response. If, as in other species, flehmen in the mandrill serves to mediate social or reproductive information, we expected its occurrence to vary with characteristics of either the signaler or receiver. Flehmen, particularly in a given male, occurred most often in response to odorants derived from male, as opposed to female, conspecifics. Moreover, odorants derived during the breeding season elicited more flehmen responses than did odorants collected during the birthing season. Lastly, odorants from reproductively cycling females also elicited more responses than did odorants from contracepted females. Although confirming a link between the nasopalatine ducts, flehmen behavior, and olfactory processing in mandrills would require further study, our observations provide new information to suggest anatomical variability within Old World primates, calling further attention to the underappreciated role of chemical communication in this lineage.

Natural simian immunodeficiency virus transmission in mandrills: a family affair?

Understanding how pathogens spread and persist in the ecosystem is critical for deciphering the epidemiology of diseases of significance for global health and the fundamental mechanisms involved in the evolution of virulence and host resistance. Combining long-term behavioural and epidemiological data collected in a naturally infected mandrill population and a Bayesian framework, the present study investigated unknown aspects of the eco-epidemiology of simian immunodeficiency virus (SIV), the recent ancestor of HIV. Results show that, in contrast to what is expected from aggressive and sexual transmission (i.e. the two commonly accepted transmission modes for SIV), cases of SIVmnd-1 subtype were significantly correlated among related individuals (greater than 30% of the observed cases). Challenging the traditional view of SIV, this finding suggests the inheritance of genetic determinants of susceptibility to SIV and/or a role for behavioural interactions among maternal kin affecting the transmission of the virus, which would highlight the underappreciated role of sociality in the spread of infectious diseases. Outcomes of this study also provide novel insights into the role of host social structure in the evolution of pathogens.

Craniodental features in male Mandrillus may signal size and fitness: an allometric approach.

According to a hypothesis in the broader mammalian literature, secondary sexual characteristics that have evolved to signal fitness and size to other conspecifics should exhibit positive allometry across adult males within a species. Here this hypothesis is tested in the genus Mandrillus. The overbuilding of bony features in larger individuals necessitates a functional explanation as bone is metabolically expensive to produce and maintain. Canine size and size of the maxillary ridge are scaled against a body size surrogate in intraspecific samples of male Mandrillus sphinx (mandrills) and Mandrillus leucophaeus (drills). Areal dimensions are weighted more heavily as they represent the size of a feature as it is viewed by individuals. Measures of the maxillary ridge and canine tooth are significantly correlated with the size surrogate and scale with positive allometry in both samples supporting the hypothesis that these features function to advertise a male's body size and fitness to other males competing for mates and potential discerning females. This is the first study in primates to test for intraspecific positive allometric scaling of bony facial features in adult males based on a theory of fitness signaling and sexual selection.

Meaningful gesture in monkeys? Investigating whether mandrills create social culture.

BACKGROUND: Human societies exhibit a rich array of gestures with cultural origins. Often these gestures are found exclusively in local populations, where their meaning has been crafted by a community into a shared convention. In nonhuman primates like African monkeys, little evidence exists for such culturally-conventionalized gestures. METHODOLOGY/PRINCIPAL FINDINGS: Here I report a striking gesture unique to a single community of mandrills (Mandrillus sphinx) among nineteen studied across North America, Africa, and Europe. The gesture was found within a community of 23 mandrills where individuals old and young, female and male covered their eyes with their hands for periods which could exceed 30 min, often while simultaneously raising their elbow prominently into the air. This 'Eye covering' gesture has been performed within the community for a decade, enduring deaths, removals, and births, and it persists into the present. Differential responses to Eye covering versus controls suggested that the gesture might have a locally-respected meaning, potentially functioning over a distance to inhibit interruptions as a 'do not disturb' sign operates. CONCLUSIONS/SIGNIFICANCE: The creation of this gesture by monkeys suggests that the ability to cultivate shared meanings using novel manual acts may be distributed more broadly beyond the human species. Although logistically difficult with primates, the translocation of gesturers between communities remains critical to experimentally establishing the possible cultural origin and transmission of nonhuman gestures.

Do non-human primates synchronise their menstrual cycles? A test in mandrills.

The concept of female cycle (or estrous) synchrony has enduring popular appeal. However, critical reviews of estrous synchrony studies in both humans and non-humans have found that synchrony has not been demonstrated convincingly, due to methodological artifacts and statistical problems. Studies of this phenomenon in animals living under naturalistic conditions are rare. We used long-term records of the timing of the female menstrual cycle in a semi-free-ranging population of mandrills, together with a randomisation procedure, to test hypotheses relating to cycle synchrony in a naturally reproducing primate. We found evidence of significant synchrony of the peri-ovulatory period in only one of 10 group-years - the year in which the largest number of cycles was recorded, both overall and per female. However, this result was no longer significant when we corrected for multiple tests of the same hypothesis. This suggests that mandrills in our study population do not synchronise their cycles, possibly because they usually conceive so quickly that they do not have the opportunity to synchronise. We also tested whether females in the same matriline, which associate with one another more than other females, cycle significantly more closely together in time than unrelated females, finding that they did so in 2 of 10 group-years, but that they were significantly less likely to match their cycles in another group-year. Across 32 matriline-years, patterns of synchrony within individual matrilines (female lineages) never fell outside the distribution based on chance. Thus we found little support for the pheromonal hypothesis for cycle synchrony, which predicts that females that associate with one another should be more likely to cycle together. Overall, our findings are in line with other studies that suggest that cycle synchrony does not occur in non-human primates.

Associations between the drill (Mandrillus leucophaeus) and sympatric monkeys in Korup National Park, Cameroon.

Studies of polyspecific associations among African forest primates have primarily focused on arboreal Cercopithecus and Procolobus/Colobus species. We examined the association frequency of the terrestrial drill (Mandrillus leucophaeus) with six sympatric monkey species in Korup National Park, Cameroon, testing reports that Mandrillus associations are infrequent and transient. We conducted 3,284 km of trail walks for 12 months (February-June 2006; July 2007 to January 2008), recording species composition in 612 primate clusters. Using a Markov chain Monte Carlo test, we compared the observed frequency of dyadic associations against null models of "no association." A novel conservative statistical approach which addresses possible dependence of observations close in time was also used, further strengthening confidence in our findings. Drills associated with all monkeys throughout the study period, and were with at least one other species (range 1-5) in half of the encounters. The association frequency of drills with red-capped mangabeys (Cercocebus torquatus) was greater than expected by chance, which is interesting given the morphological adaptation of the Mandrillus-Cercocebus clade for the exploitation of the same dietary niche, hard seeds. The difference we observed in the use of forest strata by drills and mangabeys may reflect a strategy to reduce food competition while in association. The nature and duration of observed drill associations varied. Although some associations seemed to be chance encounters, others lasted for hours with the involved species foraging together.