Odontochronologies in male and female mandrills (Mandrillus sphinx) and the development of dental sexual dimorphism.

OBJECTIVES: We examine how dental sexual dimorphism develops in mandrills, an extremely sexually dimorphic primate. We aimed to (a) establish the chronology of dental development (odontochronology) in male and female mandrills, (b) understand interindividual and intersex variation in odontochronologies, and (c) determine how dental sexual dimorphism is achieved. MATERIALS AND METHODS: We prepared histological ground sections from the permanent teeth of four female and four male mandrills from the semi-free ranging colony at the Centre International de Recherches Médicales, Franceville, Gabon. We used the microscopic growth increments in the sections to create odontochronologies. We compared ages at crown initiation, crown formation times (CFT) and crown extension rates (CER) between individuals and sexes to assess interindividual and intersex variation. RESULTS: All mandrill teeth are sexually dimorphic in size. Dental sexual dimorphism in mandrills is achieved via sex differences in the duration of growth (bimaturism) and in growth rates. We also found interindividual and intersex variation in the ages at initiation and completion of crown formation. DISCUSSION: Our results show that the rate of ameloblast differentiation varies between individuals and that selection for both the age at tooth initiation and CER has occurred independently in males and females to ensure that the teeth develop at appropriate times relative to the growth of the sexually dimorphic jaws. They also show that canine dimorphism is achieved through differences in both CER and CFT, unlike extant great apes or Cantius. Given at least three mechanisms for achieving canine dimorphism, we need more information to trace the evolution of this trait in primates.

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.

Age-related tooth wear differs between forest and savanna primates.

Tooth wear in primates is caused by aging and ecological factors. However, comparative data that would allow us to delineate the contribution of each of these factors are lacking. Here, we contrast age-dependent molar tooth wear by scoring percent of dentine exposure (PDE) in two wild African primate populations from Gabonese forest and Kenyan savanna habitats. We found that forest-dwelling mandrills exhibited significantly higher PDE with age than savanna yellow baboons. Mandrills mainly feed on large tough food items, such as hard-shell fruits, and inhabit an ecosystem with a high presence of mineral quartz. By contrast, baboons consume large amounts of exogenous grit that adheres to underground storage organs but the proportion of quartz in the soils where baboons live is low. Our results support the hypothesis that not only age but also physical food properties and soil composition, particularly quartz richness, are factors that significantly impact tooth wear. We further propose that the accelerated dental wear in mandrills resulting in flatter molars with old age may represent an adaptation to process hard food items present in their environment.

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.

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.

Canine tooth size and fitness in male mandrills (Mandrillus sphinx).

Sexual selection theory explains the evolution of exaggerated male morphologies and weaponry, but the fitness consequences of developmental and age-related changes in these features remain poorly understood. This long-term study of mandrill monkeys (Mandrillus sphinx) demonstrates how age-related changes in canine tooth weaponry and adult canine size correlate closely with male lifetime reproductive success. Combining long-term demographic and morphometric data reveals that male fitness covaries simply and directly with canine ontogeny, adult maximum size, and wear. However, fitness is largely independent of other somatometrics. Male mandrills sire offspring almost exclusively when their canines exceed approximately 30 mm, or two-thirds of average adult value (45 mm). Moreover, sires have larger canines than nonsires. The tooth diminishes through wear as animals age, corresponding with, and perhaps influencing, reproductive senescence. These factors combine to constrain male reproductive opportunities to a brief timespan, defined by the period of maximum canine length. Sexually-selected weaponry, especially when it is nonrenewable like the primate canine tooth, is intimately tied to the male life course. Our analyses of this extremely dimorphic species indicate that sexual selection is closely intertwined with growth, development, and aging, pointing to new directions for sexual selection theory. Moreover, the primate canine tooth has potential as a simple mammalian system for testing genetically-based models of aging. Finally, the tooth may record details of life histories in fossil primates, especially when sexual selection has played a role in the evolution of dimorphism.

Muzzle size, paranasal swelling size and body mass in Mandrillus leucophaeus.

The drill (Mandrillus leucophaeus), a forest-living Old World monkey, is highly sexually dimorphic, with males exhibiting extreme secondary sexual characteristics, including growth of paranasal swellings on the muzzle. In this study, the size of the secondary bone that forms the paranasal swellings on the muzzles of drills was assessed in relation to body mass proxies. The relationship between the overall size of the muzzle and surrogate measures of body mass was also examined. In female drills, muzzle breadth was positively correlated with two proxies of overall body mass, greatest skull length and upper M1 area. However, there was no such correlation in males. Paranasal swellings in males also appeared to have no significant relationship to body mass proxies. This suggests that secondary bone growth on the muzzles of male drills is independent of overall body size. Furthermore, this secondary bone appears to be vermiculate, probably developing rapidly and in an irregular manner, with no correlation in the sizes of paranasal swelling height and breadth. However, various paranasal swelling dimensions were related to the size of the muzzle. It is suggested that the growth of the paranasal swellings and possibly the muzzle could be influenced by androgen production and reflect testes size and sperm motility. The size and appearance of the paranasal swellings may thus be an indicator of reproductive quality both to potential mates and male competitors. Further work is required to investigate the importance of the paranasal swellings as secondary sexual characteristics in Mandrillus and the relationship between body size and secondary sexual characteristics. Attention should also be paid to the mechanisms and trajectories of facial growth in Mandrillus.