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Objectives The function of the browridge in primates is a subject of enduring debate. Early studies argued for a role in resisting masticatory stresses, but recent studies have suggested sexual signaling as a biological role. We tested associations between circumorbital form, diet, oral processing, and social behavior in two species of colobus monkey-the king colobus (Colobus polykomos) and western red or bay colobus (Piliocolobus badius). Materials and methods We quantified circumorbital size and dimorphism in a sample of 98 crania. Controlling for age and facial size, we tested whether variation in circumorbital morphology can be explained by variation in diet, oral processing behavior, masticatory muscle size, and mating system. To contextualize our results, we included a broader sample of facial dimorphism for 67 anthropoid species. Results Greater circumorbital thickness is unrelated to the stresses of food processing. King colobus engages in longer bouts of anterior tooth use, chews more per ingestive event, and processes a tougher diet, yet circumorbital thickness of C. polykomos is reduced compared to P. badius. Differences in circumorbital development do not vary with wear or facial size. Greater sexual dimorphism is present in P. badius; comparisons across anthropoids indicated patterns of circumorbital dimorphism were decoupled from overall size dimorphism. Conclusions The expanded circumorbits of male red colobus monkeys evolved in response to intense male-male competition. This hypothesis is consistent with the pattern across anthropoid primates and highlights the underappreciated role of sexual selection in shaping the primate face.

The sooty mangabey (Cercocebus atys) practices year-round durophagy. A large part of the C. atys diet consists of the oily nut of Sacoglottis gabonensis, which is accessed by post-canine crushing of the hard, protective seed coat. During a typical foraging bout, some seeds are discarded after initial crushing attempts using isometric biting, but the reason mangabeys reject some seeds and break into others is unclear. Although C. atys is sexually dimorphic, little is known about whether differences between males and females affect the selectivity of mechanically protected foods. We studied C. atys feeding on S. gabonensis in the Tai National Park, Cote d’Ivoire, in July and August 2016. Nuts discarded after an initial crushing attempt were collected and their hardness measured using a Shore D durometer. Measurements were taken in the region of the nut where monkeys attempted to crush it. Hardness values of nuts rejected by adult male (n = 79) and adult female (n = 104) C. atys were compared to those of a control assemblage of nuts collected randomly on the forest floor (n = 69). Nuts rejected by either sex do not differ statistically from the random sample; however, they do differ from each other, with females rejecting harder nuts. This suggests that males are more effective at broaching harder seed husks, and discard seeds based on other factors.

One way bone adapts to its mechanical environment is by Haversian remodeling, a repair process in which existing bone is resorbed and replaced by new bone. Haversian remodeling forms interconnected, cylindrical structures called secondary osteons. The amount of remodeling that occurs is related to the nature of mechanical loading and accrual of microdamage, but it is uncertain whether habitual loads of high magnitude versus high frequency result in more remodeling. The answer to this question is important if remodeling is to be a tool for inferring loading environments, and thus behavior, in past populations. Here, secondary osteon population density (OPD), osteon cross-sectional area (On.Ar), and percent Haversian bone (%HAV) were compared among mid-diaphysis femora, tibia, fibulae, and mid-level ribs of five adult crab-eating macaques (Macaca fascicularis). Ribs experience relatively low strains but have a high daily loading frequency (~33 times per minute). Limb bones are loaded for fewer cycles per day, but the femur and tibia have high load magnitudes due to gravitational forces. Strain magnitudes in the fibula are a fraction of those in the femur and tibia. Analyses of variance demonstrated significant differences in OPD (P = 0.010) and On.Ar (P 0.001) among the bones. Pairwise t-tests revealed greater OPD but lower On.Ar in the rib than all other bones. The high rib OPD suggests that Haversian remodeling is more responsive to load frequency than strain magnitude. The fact that osteons are smaller in ribs than any other bone may be an effect of remodeling in comparatively narrow cortices.

Limb bone articular and diaphyseal proportions have been shown to relate to locomotor behavior in broad comparisons across catarrhines, but comparisons among phylogenetically and functionally more closely related species may be particularly useful in investigating form-function relationships that can be applied to fossil taxa. Here we compare inter- and intra-limb proportions of diaphyseal strength and articular surface area and breadth of the femur and humerus with frequencies of leaping and vertical climbing behavior in 13 cercopithecid species. Leaping frequency is highly positively correlated with femoral/humeral diaphyseal strength, moderately positively correlated with femoral/humeral articular breadth, and less highly correlated with femoral/humeral articular surface area. These results are consistent with predicted higher bending loads as well as joint reaction forces on the femora of leapers. Surface areas may show a weaker association because they also directly impact joint excursion and are thus more influenced by other aspects of locomotion, including climbing. Climbing frequency is positively correlated with humeral head articular surface area/diaphyseal strength, but weakly negatively correlated with femoral head articular surface area/diaphyseal strength. These combined trends lead to a strong negative association between climbing and femoral/humeral head surface area. Femoral/humeral diaphyseal strength and distal articular breadth are not correlated with climbing frequency. The climbing results are consistent with greater shoulder mobility in more frequent vertical climbers. The lack of such a relationship in the femur among these taxa contrasts with earlier findings for catarrhines more generally, including hominoids, and may be a result of different climbing kinematics in cercopithecoids involving less hip abduction than in hominoids. Different use of the forelimb during climbing in cercopithecoids and hominoids may also explain the lack of association between femoral/humeral diaphyseal strength and climbing in the present study, in contrast to comparisons across catarrhines more generally.

Aye-ayes (Daubentonia madagascariensis) locate and acquire invertebrates from within woody substrates at all levels of the rainforest; yet how their locomotion helps them accommodate this diet has not been explored in detail. We studied the locomotor behavior of an adult female (N = 1,085) and juvenile male (N = 708) aye-aye in the undisturbed forest of Torotorofotsy, Madagascar from May to December 2017. We used bout sampling to record locomotion during foraging and travel of the two radio-collared individuals. We used χ 2 tests to compare overall locomotion, travel, and foraging, as well as strata and support use. We performed a correspondence analysis to examine relationships between individual behaviors, strata, and support types. Leaping accounted for 47.9% and 50.1% of all locomotor activity in the adult female and juvenile male, respectively. Leaping was the most common behavior during travel in both individuals (59.2% and 53.9%, respectively), whereas head-first descent was most frequent during foraging (35.0% and 48.0%, respectively). For all three locomotor categories, the main canopy (40.3%-79.6%) was used most frequently and trunks were the most frequently used support type (50.7%-60.0%). There is a strong association between strata and support use overall and during travel. Quadrupedal walking was significantly associated with the main canopy, as was head-first descent with the low canopy. Our analysis demonstrates that aye-ayes use a variety of locomotor behaviors to forage for invertebrates. Aye-ayes' ability to repurpose their positional repertoire to acquire other resources in degraded forests should not obscure the importance of invertebrates to this species.

Anthrax is a worldwide zoonotic disease caused by the spore-forming bacterium Bacillus anthracis. Primarily a disease of herbivores, human infections often result from direct contact with contaminated animal products (cutaneous and inhalational anthrax) or through consumption of infected meat (gastrointestinal anthrax). The genetic near neighbor, Bacillus cereus biovar anthracis (Bcbva), causes an anthrax-like illness in the wildlife and livestock of west and central Africa due to the presence and expression of B. anthracis-specific virulence factors in this background. While Bcbva infections have not been reported in humans, a recent seroprevalence study detected Bcbva antibodies in the rural population around Ta&iuml, National Park. This work describes the development of new TaqMan multiplex PCRs for the simultaneous detection of B. anthracis and Bcbva. The assays are designed to amplify Ba-1, capB, and lef markers in B. anthracis and genomic island IV (GI4), capB, and lef in Bcbva. Our assays allow for the rapid discrimination of B. anthracis and Bcbva and will provide insights into the molecular epidemiology of these two important pathogens that share an overlapping geographical range in west and central Africa.

Polyspecific associations occur when species overlap in their environment by chance, converge at common resources, or in response to predation pressure. However, because larger groups may themselves attract the attention of predators, species forming associations must balance the costs and benefits of comingling. Experimental and observational research suggests red colobus (Piliocolobus badius) associate with Diana monkeys (Cercopithecus diana) in Tai National Park, Cote d’Ivoire in response to seasonally shifting predation pressure from chimpanzees (Pan troglodytes verus). Research on this system has focused on immediate behavioral effects of comingling by red colobus and Diana monkeys. We expand on these analyses and explore longer-term changes in Diana monkey behavior. We use scan samples (N = 7025) collected on three Diana monkey groups over 5 years (N = 380 days) to assess differences in diet, activity budget, and strata use in relation to associating with red colobus and chimpanzee hunting seasonality and test for interaction effects between these two variables. We found limited evidence showing Diana monkeys make more than ephemeral behavioral changes in response to comingling with red colobus during chimpanzees’ hunting season (September–November). Synergistic effects of association and hunting season include expanded use of the main canopy, decreased fruit consumption, and increased invertebrate consumption. We find little evidence indicating Diana monkeys minimize behaviors that increase risk of predation from chimpanzees while associated with red colobus during the season when chimpanzees hunt most often. We conclude that when there are few member costs to forming mixed groups, participating species may tolerate associations even without accruing any benefits.

Enamel formation front (EFF) angles represent the leading edge of enamel matrix secretion at particular points in time. These angles are influenced by rates of enamel extension (the rates at which tooth crowns grow in height), rates of enamel matrix secretion and the angles that prisms make with the enamel-dentine junction. Previous research suggests, but has not yet established, that these angles reflect aspects of primate biology related to their pace of growth and development, most notably brain and body size. The present study tested this possibility on histological sections using phylogenetically-controlled and Bonferroni-corrected analyses spanning a broad taxonomic range. Ten species were represented in the analysis of anterior teeth; 17 in the analysis of posterior (postcanine) teeth (with varying sample sizes). Also, tested was the relationship of EFF angles to striae of Retzius periodicity (long period growth rhythms in enamel) and degree of folivory, as both factors are related to primate developmental rates. Finally, several analyses were conducted to investigate whether tooth size (operationalized as EDJ length) might mediate these relationships. Central results are as follows: (1) Relationships between EFF angles and brain weight (anterior teeth) and between EFF angles and body mass (anterior and posterior teeth) are statistically significant and (2) Mid-crown EFF angles are not statistically significantly related to EDJ lengths. These results suggest that tooth size does not mediate relationships between EFF angles and brain weight/body mass and are discussed with respect to underlying enamel growth variables (especially rates of enamel extension and secretion). Anat Rec, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 301:125-139, 2018. © 2017 Wiley Periodicals, Inc.

Reconstructing diet is critical to understanding hominin adaptations. Isotopic and functional morphological analyses of early hominins are compatible with consumption of hard foods, such as mechanically-protected seeds, but dental microwear analyses are not. The protective shells surrounding seeds are thought to induce complex enamel surface textures characterized by heavy pitting, but these are absent on the teeth of most early hominins. Here we report nanowear experiments showing that the hardest woody shells – the hardest tissues made by dicotyledonous plants – cause very minor damage to enamel but are themselves heavily abraded (worn) in the process. Thus, hard plant tissues do not regularly create pits on enamel surfaces despite high forces clearly being associated with their oral processing. We conclude that hard plant tissues barely influence microwear textures and the exploitation of seeds from graminoid plants such as grasses and sedges could have formed a critical element in the dietary ecology of hominins.

Primates are hypothesized to “fall back” on challenging-to-process foods when preferred foods are less available. Such dietary shifts may be accompanied by changes in oral processing behavior argued to be selectively important. Here, we examine the oral processing behavior of Diana monkeys (Cercopithecus diana) in the Taï Forest across their dietary breadth, testing relationships among food intake, fruit availability, preference, and oral processing behaviors including those involved in food ingestion and breakdown. We conducted 1,066 focal follows from April 2016 to September 2017 documenting frequencies of incisor, canine, and cheek tooth mastications (i.e., chews) per ingestive action (n = 11,906 feeding events). We used phenological survey and scan sample data collected between 2004 and 2009 to examine dietary preference and food availability. Our analyses show that Diana monkeys processed foods in significantly different ways (H2 = 360.8, p < 0.001), with invertebrates requiring the least oral processing, fruit requiring intermediate amounts, and leaves requiring the most oral processing. There was no relationship between fruit availability and consumption of preferred or nonpreferred fruits (p > 0.05); however, preferred fruits were processed with significantly fewer mastications (i.e., less chewing) than nonpreferred fruits (U = 6,557, p < 0.001). We thus demonstrate that, when preferred foods are scarce, Diana monkeys do not fall back on difficult-to-process foods. Changes in processing profiles occurred throughout the year and not solely when preferred foods were in short supply. Though preferred fruits required less processing than nonpreferred fruits, we found no relationship between fruit preference and fruit availability. Diana monkeys’ lack of readily identifiable fallback foods may be attributable to the relatively high tree diversity and productivity of the Taï Forest. We conclude that Diana monkeys engage in resource switching, consuming a relatively easy-to-process diet year-round.

Objectives We previously found that differing degrees of forelimb flexion, elevation, and abduction during nonlocomotor foraging activities covaried with scapular morphology among four sympatric cercopithecids. The purpose of this study is to investigate whether features of the proximal humerus are similarly related to forelimb elevation during foraging. Methods Our sample consists of humeri (n = 42) of adult Cercocebus atys, Cercopithecus diana, Colobus polykomos, and Piliocolobus badius collected from Cote d'Ivoire's Tai National Park. Features of the proximal humerus known to correspond with varying degrees of forelimb mobility were examined using a combination of 2-dimensional caliper-based indices and 3-dimensional analyses. Results Some features of the proximal humerus are consistent with the varying frequencies of forelimb elevation during foraging. For instance, the red colobus (P. badius) has a variety of features consistent with more pronounced shoulder mobility and forelimb flexibility including a wider, more distally projecting deltoid plane, greater tuberosity positioned below the humeral head articular surface, and larger intertuberosity angle. Conclusions The colobines, particularly P. badius, display a number of humeral and scapular features indicative of greater arm flexion, elevation, and abduction compared to the two cercopithecines. We conclude that idiosyncrasies in forelimb use during foraging are reflected in functionally relevant features of the cercopithecid proximal humerus, highlighting that postural behavior, in additional to locomotor behavior, can exert selective pressures on primate skeletal form. This result should provide for more informed reconstructions of the full positional repertoires in fossil taxa.

We collected frequency data on oral processing behaviors during feeding in habituated groups of Western red colobus, Piliocolobus badius, and Western black and white, Colobus polykomos, ranging in the Ivory Coast's Tai National Park. During the sampling period, the diet of red colobus consisted of approximately 75% leaves compared to approximately 47% leaves and buds in black and white colobus. Black and white colobus chewed more frequently per ingestive event than did red colobus. Black and white colobus also employed their anterior teeth much more frequently than did red colobus, a difference attributed to the frequent consumption by C. polykomos of Pentaclethra macrophylla seeds and pods. A material analysis of these food items reveals that both the seed coating and seed flesh are quite soft; however, the pod housing the seeds is very tough. We argue that the pod's toughness, geometry, and fiber orientation collectively result in a food that is very difficult to process, resulting in long handling times and frequent, aggressive use of the incisors. We compare these data with those collected on another Tai primate—the sooty mangabey, Cercocebus atys—and demonstrate that during feeding, both colobine species use their incisors less than the mangabey, but that the cercopithecine chews less than either colobine. Combining data on oral processing behaviors with those on the material properties of items being ingested should lead to more informed interpretations of dentognathic morphology.

Objectives Independent lines of evidence suggest that osteonal bone remodeling is a function of both mechanical (i.e., changes in stress) and non-mechanical (i.e., metabolic needs related to calcium liberation) factors. The degree to which secondary bone reflects mechanical factors, however, is incompletely understood despite the common assumption that the stress environment mediates remodeling activity. Here, we investigate whether there are remodeling differences between regions of primate mandibular bone known to have distinct stress environments. Methods Osteon density, osteon fragment density, and relative osteonal area are measured as indicators of remodeling activity from postcanine and symphyseal thin sections of four sympatric monkey species (N = 20 total) from Tai Forest, Cote dʼIvoire: Piliocolobus badius, Colobus polykomos, Cercocebus atys, Cercopithecus diana. Subfamily and regional effects were assessed by two-way ANOVA. Results Symphyseal bone has lower osteonal density, fragment density and relative osteonal area than postcanine bone in all species, indicating relatively low remodeling activity in symphyseal bone, despite the likelihood of relatively high stresses in its lingual cortex. Subfamily differences in postcanine remodeling are significant in that colobines exhibit greater remodeling than cercopithecines. Discussion The data suggest that high strains are not requisite for bone remodeling. How the lingual symphysis negotiates a high strain environment without the reparative benefit of remodeling is yet to be elucidated, but the data prompt reevaluation of the relationship of targeted remodeling to stress histories. Differences in osteonal bone density between taxa might reflect feeding or dietary influences on remodeling activity, but sample composition prevents parsing behavioral from other taxon-specific effects.

Antemortem enamel chipping in living and fossil primates is often interpreted as evidence of hard-object feeding (i.e., ‘durophagy’). Laboratory analyses of tooth fracture have modeled the theoretical diets and loading conditions that may produce such chips. Previous chipping studies of nonhuman primates tend to combine populations into species samples, despite the fact that species can vary significantly in diet across their ranges. Chipping is yet to be analyzed across population-specific species samples for which long-term dietary data are available. Here, we test the association between enamel chipping and diet in a community of cercopithecid primates inhabiting the Tai Forest, Ivory Coast. We examined fourth premolars and first molars (n = 867) from naturally deceased specimens of Cercocebus atys, Colobus polykomos, Piliocolobus badius, Procolobus verus, and three species of Cercopithecus. We found little support for a predictive relationship between enamel chipping and diet across the entire Tai monkey community. Cercocebus atys, a dedicated hard-object feeder, exhibited the highest frequencies of (1) chipped teeth and (2) chips of large size; however, the other monkey with a significant degree of granivory, Co. polykomos, exhibited the lowest chip frequency. In addition, primates with little evidence of mechanically challenging or hard-food diets—such as Cercopithecus spp., Pi. badius, and Pr. verus—evinced higher chipping frequencies than expected. The equivocal and stochastic nature of enamel chipping in the Tai monkeys suggests nondietary factors contribute significantly to chipping. A negative association between canopy preference and chipping suggests a role of exogenous particles in chip formation, whereby taxa foraging closer to the forest floor encounter more errant particulates during feeding than species foraging in higher strata. We conclude that current enamel chipping models may provide insight into the diets of fossil primates, but only in cases of extreme durophagy. Given the role of nondietary factors in chip formation, our ability to reliably reconstruct a range of diets from a gradient of chipping in fossil taxa is likely weak.

Bone remodeling is at least partially mediated by the mechanical environment created by an animal's behavior. Here, we test the hypothesis that bone remodeling is primarily induced by high magnitude loads, likely encountered during leaping/bounding behaviors. Osteon population density (OPD), osteon cross-sectional area (On.Ar), and relative osteonal area (%HAV) were measured from femoral and humeral midshaft thin sections of four cercopithecids (N = 5 per species) from Tai Forest, Cote d'Ivoire: Colobus polykomos, Piliocolobus badius, Cercopithecus diana, and Cercocebus atys. All species are generalized quadrupeds but vary in leaping frequency and overall activity budget. Differences between taxa with high (C. polykomos and P. badius) and low leaping frequency (C. diana and C. atys) were assessed via a phylogenetically informed generalized linear mixed model using Markov Chain Monte Carlo methods. Femoral OPD and %HAV are greater in the high frequency leapers than in low frequency leapers, suggesting that frequent high magnitude loads engender remodeling, however, there is no similar pattern in the humerus, which presumably also experiences high magnitude loads during leaping. Additionally, OPD and %HAV are greater in the humerus than the femur, despite load magnitude being presumably higher in the femur. These results provide conflicting support for hypotheses about load magnitude and load frequency as they relate to bone remodeling activity. Future work is proposed to parse out the respective effects of load magnitude and frequency on bone remodeling. Anat Rec, 302:1116-1126, 2019. © 2018 Wiley Periodicals, Inc.

Molecular, morphological, and behavioral data indicate that the primates commonly referred to as mangabeys do not comprise a natural group. The Cercocebus mangabeys are seven species of African monkeys also referred to as “drill-mangabeys.” Cercocebus is the sister taxon of Mandrillus (mandrills and drills). The five species of Lophocebus mangabeys, also known as baboon mangabeys, form a clade with Papio, Rungwecebus, and Theropithecus. The Cercocebus–Mandrillus clade is united by a suite of skeletodental features and behavioral adaptations related to the acquisition and processing of hard-object foods collected from the forest floor. Mangabey diphyly highlights the considerable homoplasy among African papionins. Keywords: conservation; physical/biological anthropology; primatology

Elastic modulus of bone from the anterior mandibular corpus was determined via microindentation in a mixed-sex ontogenetic sample (N = 14) of Macaca fascicularis. This investigation focused on the hypothesis that material heterogeneity in the macaque mandibular symphysis—provided an accounting of age and sex variation—is explicable as a means to homogenize strains in this region. Experimental data and theoretical models of masticatory loading indicate that in the absence of material compensation, large strain gradients exist in the anterior mandibular corpus of macaques, particularly between lingual and labial cortical plates owing to the effects of lateral transverse bending. Microindentation data indicate that juvenile macaques possess less stiff bone than their subadult and adult counterparts; however, sex differences in elastic modulus are not apparent. Anisotropy variation is idiosyncratic; that is, there is not a common pattern of variation in stiffness sampled among orthogonal planes across individuals. Similarly, differences in stiffness between lingual and labial cortical plates, as well as differences among alveolar, midcorpus, and basal regions are inconsistently observed. Consequently, we find little evidence in support of the hypothesis that spatial variation in bone stiffness functions to homogenize strains in the anterior corpus; in fact, in some individuals, this spatial variation serves to exacerbate, rather than to minimize, strain gradients. The mechanical benefit of elastic modulus variation in the macaque mandibular symphysis is unclear; this variation may not confer adaptive benefit in terms of structural integrity despite the fact that such variation has discernible functional consequences. Am J Phys Anthropol 156:649–660, 2015. © 2014 Wiley Periodicals, Inc.

We conducted surveys in Cote d'Ivoire's Bandama–Sassandra interfluvial region to assess the abundance of primates and investigate further a population of black-and-white colobus monkey whose taxonomic position has been debated. Primates were encountered in five of eight forests surveyed. The most common primates were Campbell's guenons and lesser spot nosed monkeys. Roloway monkeys were located in one reserve (Dassioko FR), sooty mangabeys in two (Dassioko and Port Gauthier FR) and olive colobus in three reserves (Dassioko, Port Gauthier, Bakadou) as well as one forest patch was considered a sacred grove (Grebouo I FG). Red colobus and chimpanzees were never observed. The only black-and-white colobus encountered were located in a forest adjacent to the Grebouo I village. The coat of this population is similar to that of Colobus vellerosus; however, vocal data are more complex. In one vocal attribute, the Grebouo I colobus is similar to Colobus polykomos while in another it is similar to C. vellerosus. We hope that by investigating the taxonomy of the Grebouo 1 colobus population, highlighting the conservation value of sacred forests and advocating for more active law enforcement in protected reserves, we can prevent the extirpation of all primates within the Bandama–Sassandra interfluvial zone region. Resume Nous avons mene des inventaires dans la region interfluviale situee entre les fleuves Sassandra et Bandama en Cote d'Ivoire, pour estimer l'abondance des primates et etudier une population de colobe noir et blanc dont la classification taxonomique fait debat. Des primates ont ete observes dans cinq des huit forets visitees. Les plus communs etaient les cercopitheques de Campbell et les cercopitheques petauristes. En outre, les cercopitheques de Roloway ont ete rencontres dans une seule foret (la foret classee de Dassioko) alors que les cercocebes couronnes l'ont ete dans deux forets (les forets classees de Dassioko et de Port- Gauthier). Par ailleurs, le colobe vert a ete observe dans trois forets (Dassioko, Port-Gauthier, Bakadou) ainsi que dans un ilot forestier considere comme une foret sacree (foret sacree de Grebouo 1). Les colobes bais et les chimpanzes n'ont pas ete observes. Seule la foret sacree de Grebouo 1 abrite encore les colobes noir et blancs. Le pelage de cette population est semblable a celui de Colobus vellerosus, mais l'interpretation des donnees sur leur vocalisation reste complexe. Pour un des parametres de vocalisation, le colobe de Grebouo 1 est semblable a Colobus polykomos alors que pour un autre, il est semblable a C. vellerosus. Nous esperons qu'en etudiant la taxonomie de la population du colobe de Grebouo 1, en mettant en evidence la valeur des forets sacrees en vue de leur conservation et en defendant une application plus stricte de la loi dans les aires protegees, nous arriverons a eviter la disparition des primates dans la region interfluviale Bandama-Sassandra.

We present information on food hardness and monthly dietary changes in female sooty mangabeys (Cercocebus atys) in Tai Forest, Ivory Coast to reassess the hypothesis that thick molar enamel is parsimoniously interpreted as a response to consumption of hard foods during fallback periods. We demonstrate that the diet of sooty mangabeys varies seasonally, but that one food—Sacoglottis gabonensis—is the most frequently consumed food every month and year round. This food is the hardest item in the sooty diet. Given that this species has among the thickest enamel within the primate order, a plausible conclusion is that thick enamel in this taxon evolved not in response to seasonally critical function or fallback foods, but rather to the habitual, year round processing of a mechanically protected foodstuff. These data serve as a caution against de rigueur interpretations that reliance on fallback foods during lean periods primarily explains the evolution of thick enamel in primates. Am J Phys Anthropol 154:413–423, 2014. © 2014 Wiley Periodicals, Inc.

Evidence is accumulating that bone material stiffness increases during ontogeny, and the role of elastic modulus in conditioning attributes of strength and toughness is therefore a focus of ongoing investigation. Developmental changes in structural properties of the primate mandible have been documented, but comparatively little is known about changes in material heterogeneity and their impact on biomechanical behavior. We examine a cross-sectional sample of Macaca fascicularis (N = 14) to investigate a series of hypotheses that collectively evaluate whether the patterning of material stiffness (elastic modulus) heterogeneity in the mandible differs among juvenile, subadult and adult individuals. Because differences in age-related activity patterns are known to influence bone stiffness and strength, these data are potentially useful for understanding the relationship between feeding behavior on the one hand and material and structural properties of the mandible on the other. Elastic modulus is shown to be spatially dependent regardless of age, with this dependence being explicable primarily by differences in alveolar versus basal cortical bone. Elastic modulus does not differ consistently between buccal and lingual cortical plates, despite likely differences in the biomechanical milieu of these regions. Since we found only weak support for the hypothesis that the spatial patterning of heterogeneity becomes more predictable with age, accumulated load history may not account for regional differences in bone material properties in mature individuals with respect to the mandibular corpus. Am J Phys Anthropol 153:297–304, 2014. © 2013 Wiley Periodicals, Inc.

Most scholars agree that avoiding predators is a central concern of lemurs, monkeys, and apes. However, given uncertainties about the frequency with which primates actually become prey, the selective importance of predation in primate evolution continues to be debated.[1-9] Some argue that primates are often killed by predators,[5, 6] while others maintain that such events are relatively rare.[2, 7, 9] Some authors have contended that predation's influence on primate sociality has been trivial[10, 11]; others counter that predation need not occur often to be a powerful selective force.[12-14] Given the challenges of documenting events that can be ephemeral and irregular, we are unlikely ever to amass the volume of systematic, comparative data we have on such topics as feeding, social dynamics, or locomotor behavior. Nevertheless, a steady accumulation of field observations, insight gained from natural experiments, and novel taphonomic analyses have enhanced understanding of how primates interact with several predators, especially raptors, the subject of this review.

Studies of dental macrowear can be useful for understanding masticatory and ingestive behavior, life history, and for inferring dietary information from the skeletal material of extinct and extant primates. Such studies to date have tended to focus on one or two teeth, potentially missing information that can be garnered through examination of wear patterns across the tooth row. Our study measured macrowear in the postcanine teeth of three sympatric cercopithecid species from the Tai Forest, Cote d'Ivoire (Cercocebus atys, Procolobus badius, and Colobus polykomos), whose diets have been well-described. Inter-specific analyses suggest that different diets and ingestive behaviors are characterized by different patterns of wear across the molar row, with Cercocebus atys emphasizing tooth use near P4 -M1 , P. badius emphasizing a large amount of tooth use near M2 -M3 , and Colobus polykomos exhibiting wear more evenly across the postcanine teeth. Information regarding differential tooth use across the molar row may be more informative than macrowear analysis of isolated teeth for making inferences about primate feeding behavior.

Although predation avoidance is the most commonly invoked explanation for vertebrate social evolution, there is little evidence that individuals in larger groups experience lower predation rates than those in small groups. We compare the morphological and behavioural traits of mammal prey species in the Taï forest, Ivory Coast, with the diet preferences of three of their non-human predators: leopards, chimpanzees and African crowned eagles. Individual predators show marked differences in their predation rates on prey species of different body sizes, but clear patterns with prey behaviour were apparent only when differences in prey habitat use were incorporated into the analyses. Leopard predation rates are highest for terrestrial species living in smaller groups, whereas eagle predation rates are negatively correlated with group size only among arboreal prey. When prey predation rates are summed over all three predators, terrestrial species incur higher predation rates than arboreal species and, within both categories, predation rates decline with increasing prey group size and decreasing density of groups in the habitat. These results reveal that it is necessary to consider anti-predator strategies in the context of a dynamic behavioural interaction between predators and prey.

Given that something as fundamental as food acquisition is subject to selection pressure, it follows that morphological and behavioral diversity among primates is reflective of a range of adaptations to diet, feeding, and foraging. The recognition of these adaptations, however, is operationally difficult because it is the interaction between morphological and ecological variables that serves to define the particular adaptation. Researchers have addressed this problem of recognition of adaptation by integrating functional and biomechanical measures of morphological performance with observations of foraging and feeding behavior of primates in natural habitats. These disparate approaches traditionally resided in separate laboratory and field domains, but technological and analytical advances have blurred the distinction between them. The success with which this integration of approaches has elucidated the nature of primate foraging adaptations is reviewed with respect to (a) ingestive strategies, (b) locomotor diversity, (c) hard-object feeding in papionin primates, and (d) the influence of “fallback foods” on behavior and morphology.

Alveolar bone is functionally involved with tooth support and must also be competent in accommodating masticatory forcess. Data from a variety of perspectives – including observations of anatomy, metabolic activity and determination of mechanical properties – suggest that alveolar bone is more compliant than the more inferiorly situated basal bone of the mandibular corpus. The nature of stiffness variation within the alveolar process, however, remains largely unexplored. The technique of microindentation is used to estimate bone stiffness variation in the cortical alveolar bone of a sample of eight adult mandibles from two species of West African colobus monkeys. The null hypothesis under investigation is that bone stiffness variation in the alveolar process is random with respect to species, sex, and location. Alternative hypotheses are evaluated with respect to species, sex and locality effects, including those associated with cortical thickness variation and masticatory loading. Microindentation data do not indicate species or sex differences. Spatial variation in alveolar bone, however, is nonrandom. The stiffest bone is localized in the superior alveolar process, and significant differences between buccal and lingual cortical plates are found in five of eight specimens. Evidence for a superoinferior stiffness gradient is found in half the specimens but is confined to either the buccal or lingual cortical plate. Covariation between bone stiffness and bone thickness is weak. Relative compliance of the alveolar process represents a biomechanical solution for enhancing toughness of cortical bone. The relationship of the spatial patterning of bone stiffness to the masticatory loading environment is unclear.

Members of the Cercocebus-Mandrillus clade are united by several morphological features, including expanded premolars which are argued to be associated with a preponderance of hard objects in the diet. We test the association between premolar expansion and hard object feeding by examining how different dental regions are used during food processing. We examined the diet and oral processing activities of sooty mangabeys (Cercocebus atys) in the Ivory Coast's Tai forest from August 2008 to September 2009. In addition to compiling diet profiles, we recorded the frequency that individuals performed four activities: 1) incising, 2) canine puncturing/scraping, 3) postcanine crushing (i.e., isometric biting), and 4) routine mastication (chewing cycles). Sooty mangabeys have a relatively narrow diet that consists largely of nuts/seeds, fruits, and invertebrates. While there are age and sex differences in diet, the most frequently consumed foods are similar across age and sex classes. The most frequently consumed foods are seeds of Sacoglottis gabonensis which are the hardest items in the sooty mangabey diet. Patterns of ingestive behavior vary with food type, but adults and nonadults (excluding infants dependent on mothers) of both sexes process similar foods. Premolar expansion in Cercocebus atys is associated with powerful crushing of hard objects of specific size and durophagy is a constant feature of sooty mangabey feeding ecology throughout ontogeny.

Recent studies suggest a large raptor such as the crowned eagle (Stephanaoetus coronatus) was responsible for collecting at least a portion of the primate fauna from the South African fossil site of Taung, including its lone hominin specimen. This taphonomic signature at Taung is currently regarded as a unique and, most likely, isolated case in primate and human evolution. However, the activities of large, carnivorous birds should also be detectable at other primate fossil localities in Africa if raptors have been a strong selective force throughout primate evolution. Over the last 60 years, a collection of extinct cercopithecids has been assembled from several cave breccias on the Humpata Plateau in southern Angola. The material, dated near the Plio-Pleistocene boundary, includes an assortment of craniodental and postcranial remains variably assigned to Papio (Dinopithecus) cf. quadratirostris, Parapapio, Cercopithecoides, and Theropithecus. We compare the Angolan and Taung material to remains of extant primates killed by crowned eagles in the Ivory Coast's Tai National Park. Our analysis indicates that the size distribution and composition of fauna from the localities is quite similar and that there are striking consistencies in damage to the crania from each site. The absence of large bodied (>20 kg) primates and other mammalian taxa at the Taung hominin locality and Tai, and their rarity in Angola, combined with the strong likelihood that raptor nests were positioned near fissure openings at both fossil localities, provides additional support for eagle involvement. On the basis of this evidence, we conclude that at least some of the Angolan cercopithecids were most likely raptor prey and hypothesize that raptor predation has been a strong and perhaps underappreciated selective force during the course of primate evolution.

The anterior mandibular corpus of anthropoid primates is routinely subjected to masticatory loads that result in relatively high local levels of stress and strain. While structural morphological responses to these loads have been extensively explored, relatively little is known about material property variation in mandibular bone of nonhuman primates. Consequently, the role of regional and local variation in bone stiffness in conditioning stress and strain gradients is poorly understood. We sampled elastic modulus variation in the bone of the anterior mandibular corpus in two species (N = 3 each) of sympatric colobine monkeys, Procolobus badius and Colobus polykomos. These monkeys were chosen for comparison owing to their distinctive dietary regimens, as P. badius rarely includes hard objects in its diet while C. polykomos habitually processes obdurate items during feeding. Elastic modulus is determined through bone hardness data obtained via microindentation, which enables the description of stiffness variation on sub-millimeter scales. Labial bone stiffness exceeds that of lingual bone in the sample overall. Female mandibular bone is generally stiffer than that found in males, and overall Procolobus mandibular bone is stiffer than that in Colobus. These results, interpreted collectively, suggest that the material response to elevated masticatory stress is increased compliance of the affected bone.

The Ivory Coast's Tai Forest is home to a diverse primate fauna, including eight monkey species that interact daily with the environment, each other, and a web of predators. These interactions have led to an array of adaptations, some of which conform to theoretical expectations, and others are at odds with current behavioral ecology models. In this paper, we draw on fifteen years of observations to examine two central issues. First, how do broad trends in diet and resource distribution influence group size and levels of sociality? Second, what mechanisms are used to counter pressures exerted by the predator community? The general picture that emerges is that while the pressure exerted by predators at Tai has led to a host of behavioral and cognitive adaptations, it is food preferences and foraging habits, not predation pressure, that are the main determinants, not only of group size, but also its direct consequences - mating systems and sociality levels.

Most investigations of primate scapular morphology use differences in locomotion to explain variation; less is known about how scapular geometry covaries with nonlocomotor behavior. We examined forelimb use during foraging in 4 cercopithecids ranging throughout the Ivory Coast's Tai Forest. During 5-min feeding bouts, we recorded the frequency individuals of Piliocolobus badius, Colobus polykomos, Cercocebus atys and Cercopithecus diana performed 5 forelimb behaviors involved in the acquisition and introduction of food to the oral cavity. Scapulae from these populations were examined to determine whether differences in forelimb use were reflected in features known to correspond with varying degrees of arm flexion, abduction and elevation. Our results reveal that the species differ markedly in forelimb use and that these differences are interpretable via their scapular morphology. For example, P. badius engages in more frequent flexion, abduction and elevation of the arm above the head relative to C. polykomos, and red colobus scapulae are longer craniocaudally and have larger, more cranially directed supraspinous fossae than those of closely related black-and-white colobus. Our attempt to explore how nonlocomotor behavior covaries with skeletal morphology should provide for more informed interpretations of the primate fossil record.

Objective Procolobus badius , Colobus polykomos , and Cercocebus atys are three West African primate species known to have distinctive feeding behaviors. Our objectives were (1) to test whether intra-taxon variation in bone mineral content exists between anterior and posterior regions of the mandible and (2) to determine if interspecific differences are interpretable via feeding and dietary idiosyncracies among the three taxa. Design Twenty-one mandibles from naturally deceased C. polykomos ( n = 7), P. badius ( n = 9), and C. atys ( n = 5) were scanned using cone beam computerized tomography. Alveolar bone (AB) and basal cortical bone (CB) of incisor (I 1 ) and second molar (M 2 ) regions were digitally isolated. Degree of bone mineralization (DBM) distribution was obtained using histograms of CT attenuation values. Mean, standard deviation (SD), the 5th (Low 5 ) and the 95th (High 5 ) percentiles of the DBM histogram were compared between the jaw regions in species. Results The mean and Low 5 of DBM were significantly lower for AB than CB of all species ( p 1 and M 2 of all species ( p > 0.056) except the mean of C. polykomos ( p 5 of CB DBM at M 2 of C. atys was significantly higher than those of C. polykomos and P. badius ( p Conclusions The durophagous C. atys had higher CB DBM value at the M 2 region than C. polykomos and P. badius, which supports the hypothesis that materially stiffer mandibular bone in C. atys can develop in response to the generation of high bite forces during hard object feeding.

Understanding the initial processes of deposition can help with interpretations of fossil assemblages. Here we discuss the taphonomy of primate remains collected under 16 nests of African crowned eagles (Stephanoaetus coronatus) in the Tai Forest, Ivory Coast. From 1,200 bones collected, including 669 primate bones, we calculated minimum number of individuals (MNI), survivability profiles, and damage profiles using methods identical to those employed by Sanders et al. (2003 J. Hum. Evol. 44:87-105) in their analysis of bones from eagle nests in Uganda. Crowned eagles leave a consistent taphonomic signature on their prey remains; hence, results from our analysis of the Tai assemblage are similar to those from the Ugandan sample. Hindlimb and cranial bones are relatively abundant in the sample, while ribs, vertebrae, carpals, and tarsals do not survive well. Primate crania typically display puncture marks around the eye, long bones remain largely intact, and scapulae exhibit raked breakage. These data have implications for understanding the dynamic between extant primates and one of their principle predators, as well as the taphonomy of hominid-bearing caves in South Africa. We concur with Berger and Clarke (1995 J. Hum. Evol. 29:275-299) that a large raptor could have been responsible for the death of the Taung child, Australopithecus africanus.

Miss Waldron’s red colobus (Procolobus badius waldroni) has a restricted distribution in eastern Ivory Coast and western Ghana. There have been no confirmed sightings of them since 1978 and surveys carried out from 1993 to the present have yet to reveal any living individual. Since the announcement of the monkey’s probable extinction (Oates et al., 2000), new evidence from forest in the extreme southeast of Ivory Coast suggests that a handful of individuals have remained undetected to this point. I discuss the evidence—a tail, a skin and a photograph—and results of accompanying surveys. Additional surveys of the Ehy Forest are required to confirm the presence of Procolobus badius waldroni and an Action Plan for the conservation of all red colobus across the continent is needed to prevent the disappearance of other taxa.

Context Duikers play a vital role in maintaining forest ecosystems, as both seed dispersers and prey. In West Africa, duikers are also an important source of food and income; however, it is likely that some species are now being hunted at unsustainable levels. So as to contribute to duiker conservation, we used line-transect data and additional information on dung and track counts to estimate relative abundance and establish distribution patterns of forest duikers in Côte d'Ivoire's Dassioko Sud Forest Reserve (DSFR), a poorly known, remnant coastal rainforest. We also collected basic information on hunting pressure. Methods Forest duiker counts were made twice per month from July 2013 to March 2014 on six line transects of varying lengths (5.6–6.7km) and in different regions of the DSFR, representing different habitat types. The number of sightings per kilometre walked, defined as ‘encounter rate', was used to compute an index of relative abundance. We recorded all evidence of poachers and used these data to generate a poaching index (# poaching signs per kilometre) for different habitat types. Key results The DSFR has lost three of seven total forest duiker species and populations of the four remaining species are declining. Encounter rates were greatest for Philantomba maxwellii (0.46 indices km–1) and accounted for 68% of total encounters. Cephalophus niger, C. dorsalis and C. silvicultor were encountered at similar (0.07 indices km–1) but much lower rates. For all species, mean encounter rates were highest in degraded forest (0.38–0.73 indices km–1), followed by secondary (0.2–0.44 indices km–1) and primary (0.19–0.44 indices km–1) forest. Conclusions Of the four forest duiker species still present in the DSFR, the black, bay and yellow-backed duikers are the most affected by anthropogenic pressures. Duiker abundance in the DSFR varies with habitat type. This variation is apparently influenced by differences in forest structure, poaching pressure and proximity to villages. Implications The study provides baseline data for future managing of ungulate populations in the DSFR. We recommend that a biomonitoring study of duiker populations be initiated immediately to help determine population trends, investigate habitat preferences, and help curb illegal hunting activities in this important forest reserve.

We examined the behavior of two arboreal monkeys—Piliocolobus badius (western red colobus) and Cercopithecus diana (Diana monkey)—in the presence and absence of a third, predominately terrestrial monkey, Cercocebus atys (sooty mangabey) in the Ivory Coast's Tai Forest. Via experimental data, we show that sooty mangabeys are effective sentinels for ground predators. Then we compared strata use of red colobus and Diana monkeys in the presence and absence of mangabeys. Our data indicate that red colobus and Diana monkeys descend to low forest levels and to the forest floor significantly more often when mangabeys are spatially intermingled with them, i.e. in polyspecific associations. The niche extension may provide some foraging advantage, especially for red colobus. We identified the specific causal agent—relaxed ground predator pressure—of a temporary shift in habitat use. In this instance, the presence of one species (Cercocebus atys) leads to the temporary expansion of a niche normally avoided by others Piliocolobus badius and Cercopithecus diana.

The functional link between mandibular morphology and masticatory stress has been documented by both experimental and comparative investigation. Somewhat more tenuous is the purported connection between dietary variation and the form of the jaws in primates. Several factors complicate the inference of such a connection, including anecdotal or incomplete dietary data from field studies and allometric effects on skeletal form that may have little to do with diet per se. We compared the jaws of sympatric colobines from West Africa to test the effect of diet on mandibular form. Procolobus badius and Colobus polykomos occupy the same habitat yet differ in diet primarily due to the exploitation of hard seeds by C. polykomos. The fact that the two taxa are comparable in body size also obviates the need for allometric qualifications. Colobus polykomos is expected to possess more robust mandibular corpora than Procolobus badius. In fact, the jaws of Colobus polykomos do not differ consistently from those of Procolobus badius in terms of biomechanical function. This apparent failure of mandibular morphology to reflect differences in diet and feeding behavior may be due to a variety of factors. We suspect that functional demands related to canine tooth support are contributing to obliteration of the expected biomechanical signal. Successful prediction of dietary effects on mandibular form requires consideration of competing structural and functional demands. The influence of diet on mandibular corporeal morphology is not equivalent across different primate species.

We provide the first documented case of the extinction in the twentieth century of a widely recognized primate taxon. During surveys in Ghana and Cote d'Ivoire in 1993–1999, we were unable to find any surviving populations of Miss Waldron's red colobus monkey ( Procolobus badius waldroni), a primate taxon endemic to the forests of this part of West Africa. We conclude that this monkey, which at least one authority considers worthy of species status, is probably extinct. Hunting by humans appears to be the ultimate cause of the extinction. Until our surveys began, little attention had been paid to the plight of this red colobus monkey, despite its listing as endangered by the World Conservation Union. The extinction of other large animals in the Upper Guinea rainforest region is likely to follow soon unless more attention is paid to the full range of endangered forms and more resources are devoted to their rigorous protection.

I studied the positional behavior and habitat use of Cercopithecus petaurista, the lesser spot-nosed monkey, in the Ivory Coast's Tai Forest for 15 months. I compare these data with similar information collected on sympatric groups of Cercopithecus diana and C. campbelli in order to examine further the relationships between locomotion, posture, support use, stratal use, body size, diet, activity patterns and foraging behavior. Spot-nosed monkeys are predominantly quadrupedal primates that frequent the top layer of the understory during all maintenance activities. Locomotion is characteristically slow and cautious; travel takes place on branches and boughs while foraging occurs on twigs and branches. Postural behavior of Cercopithecus petaurista reflects their reliance on more ubiquitously distributed, less mobile food items. The relationship between body size, climbing, leaping and support use among Tai guenons is weak; interspecific differences are more likely functions of strata use and overall behavioral characteristics, e.g. crypticity. I also compare the locomotion and support use of Cercopithecus petaurista with that of C. ascanius from Uganda's Kibale Forest (Gebo and Chapman, 1995a) in order to assess the behavioral similarity of members of the same superspecies. Although overall support use is quite similar, the monkeys differ significantly in frequencies of quadrupedism, leaping and climbing. I present possible reasons for and implications of these differences.

Numerous biomolecular studies from the past 20 years have indicated that the large African monkeys Papio , Theropithecus , and Mandrillus have a diphyletic relationship with different species groups of mangabeys. According to the results of these studies, mandrills and drills ( Mandrillus ) are most closely related to the torquatus – galeritus group of mangabeys placed in the genus Cercocebus , whereas baboons ( Papio ) and geladas ( Theropithecus ) are most closely related to the albigena – aterrimus mangabeys, now commonly placed in the genus Lophocebus . However, there has been very little morphological evidence linking mandrills on the one hand and baboons and geladas on the other with different groups of mangabeys. In a study of mangabey locomotion and skeletal anatomy, we have identified features of the postcranial skeleton and the dentition that support the molecular phylogeny and clearly link mandrills with Cercocebus and Papio with Lophocebus . Moreover, the features linking Cercocebus and Mandrillus accord with ecological studies of these species indicating that these two genera are a cryptic clade characterized by unique adaptations for gleaning insects, hard nuts, and seeds from the forest floor.

The relationships between locomotion, body size, and habitat use in six sympatric Old World monkeys are examined to test whether the associations found are consistent with those demonstrated in previous studies (Fleagle and Mittermeier [1980] Am. J. Phys. Anthropol. 52:301–314; Gebo and Chapman [1995] Am. J. Phys. Anthropol. 97:49–76). Colobus polykomos, C. badius, C. verus, Cercopithecus diana, C. campbelli, and Cercocebus atys were studied for 14 months in the Ivory Coast's Tai Forest. Analyses reveal that (1) larger monkeys tend to frequent those strata containing the greatest densities of large supports while smaller monkeys are more diverse in their canopy use; (2) high frequencies of leaping are not necessarily confined to the understory, and understory specialists are not necessarily frequent leapers; (3) body size does not consistently predict leaping or climbing frequencies; (4) in general, climbing is more frequent during foraging and leaping is more common during travel; (5) larger supports are used during travel while smaller supports are used during foraging; and (6) larger monkeys do not always use larger supports than do smaller monkeys. Some of the factors contributing to the manner that locomotion, body size, and habitat use are related in cercopithecid monkeys are discussed. Am J Phys Anthropol 105:493–510, 1998. © 1998 Wiley-Liss, Inc.

A common objective in field studies of positional behavior is to establish functional links between locomotion, body size, habitat use, foraging strategies, and maintenance activities. In contrast, there has been relatively little effort to examine posture in a similar, comparative context. Although various studies have shown that particular postures are employed in specific contexts, the theory which could provide the basis for understanding posture on a more general level has not been explicitly stated. This is particularly true for primates lacking specializations such as prehensile tails, claws for clinging, or adaptations for forelimb suspension. Consequently, there are few a priori reasons for predicting postural differences among generalized arboreal quadrupeds. Six sympatric cercopithecid monkeys were studied for 14 months in the Ivory Coast's Tai Forest to determine if more general relationships do exist between posture and other aspects of behavior. The results demonstrate that the postural diversity with these primates can, to varying degrees, be understood within the context of differences in the spatial distribution of preferred food items, activity patterns, support use, and foraging strategies.

Habitat heterogeneity is widely theorized to be important in the evolution of Old World monkeys, yet the impact of architecturally distinct forest types on the positional repertoires of African monkeys is largely unknown. As part of a wider project, this study investigates how the locomotor behavior of five sympatric cercopithecids varies in structurally distinct forest areas. Fieldwork took place in the Tai Forest, Ivory Coast, from June 1993 to August 1994. Data were collected on Colobus badius, C. polykomos, C. verus, Cercopithecus diana, and C. campbelli ranging throughout a common forest mosaic. The behavior of adult females was sampled using an instantaneous time point scheme at 3 min time intervals. Vegetative differences within each 100 x 100 m grid cell used by all species were quantified. Further, the relative numbers of different sized substrates at 10 m intervals within the canopy were recorded in each cell sampled. Data indicate that consistency of locomotor behavior across forest types is maintained because most monkeys select the same set of preferred supports in both forest types. The conservative nature of Tai monkey locomotor behavior is discussed within the context of additional proximate and ultimate influences and constraints.

Context Duikers play a vital role in maintaining forest ecosystems, as both seed dispersers and prey. In West Africa, duikers are also an important source of food and income; however, it is likely that some species are now being hunted at unsustainable levels. So as to contribute to duiker conservation, we used line-transect data and additional information on dung and track counts to estimate relative abundance and establish distribution patterns of forest duikers in Côte d’Ivoire’s Dassioko Sud Forest Reserve (DSFR), a poorly known, remnant coastal rainforest. We also collected basic information on hunting pressure. Methods Forest duiker counts were made twice per month from July 2013 to March 2014 on six line transects of varying lengths (5.6–6.7km) and in different regions of the DSFR, representing different habitat types. The number of sightings per kilometre walked, defined as ‘encounter rate’, was used to compute an index of relative abundance. We recorded all evidence of poachers and used these data to generate a poaching index (# poaching signs per kilometre) for different habitat types. Key results The DSFR has lost three of seven total forest duiker species and populations of the four remaining species are declining. Encounter rates were greatest for Philantomba maxwellii (0.46 indices km–1) and accounted for 68% of total encounters. Cephalophus niger, C. dorsalis and C. silvicultor were encountered at similar (0.07 indices km–1) but much lower rates. For all species, mean encounter rates were highest in degraded forest (0.38–0.73 indices km–1), followed by secondary (0.2–0.44 indices km–1) and primary (0.19–0.44 indices km–1) forest. Conclusions Of the four forest duiker species still present in the DSFR, the black, bay and yellow-backed duikers are the most affected by anthropogenic pressures. Duiker abundance in the DSFR varies with habitat type. This variation is apparently influenced by differences in forest structure, poaching pressure and proximity to villages. Implications The study provides baseline data for future managing of ungulate populations in the DSFR. We recommend that a biomonitoring study of duiker populations be initiated immediately to help determine population trends, investigate habitat preferences, and help curb illegal hunting activities in this important forest reserve.

Traditionally, thick enamel has often been used to infer durophagy (i.e., hard nut and seed consumption) in extinct hominins. These inferences are based on the hypothesis that thick enamel is primarily an adaptation to prevent tooth fracture or chipping resulting from high-stress loads produced during the mastication of large hard foods. An alternative view argues that thick enamel may aid in maintaining tooth function in the face of gradual dental wear from grit, phytoliths and acid, which may be found in foods of widely varying hardness. We use estimates of primate dietary abrasiveness and recorded lifespan to test the hypothesis that enamel thickness is selectively responsive to lifetime dental wear resistance. We use data from the literature to relate enamel thickness to measures of dietary abrasiveness, diet profiles, and longevity for 17 primate species and performed linear regression using several combinations of these variables. We found a positive association between lifetime dietary wear and enamel thickness, suggesting that thick molar enamel in primates may have evolved as a means to resist wear apart from selection to resist tooth fracture. Assuming our estimates of lifetime dietary wear are accurate, we caution against ascribing thick enamel solely to the presence of hard-object feeding in paleoanthropological contexts without also considering primate lifespan and other aspects of feeding ecology.

The documentation of enamel thickness variation across primates is important because enamel thickness has both taxonomic and functional relevance. The Old World monkeys commonly referred to as mangabeys have figured prominently in investigations of feeding ecology and enamel thickness. In this article, we report enamel thickness values for four mangabey taxa (Cercocebus atys, Cercocebus torquatus, Lophocebus aterrimus, and Lophocebus albigena), offer revised interpretation of the significance of thick enamel in papionin evolution, and place our new data in a broader comparative framework. Our data indicate that all mangabeys have thick enamel and that the values obtained for Cercocebus and Lophocebus equal or exceed those published for most extant non-human primates. In addition, new field data combined with a current reading of the dietary literature indicate that hard foods make up a portion of the diet of every mangabey species sampled to date. Clarification on the relationship between diet and enamel thickness among mangabeys is important not only because of recognition that mangabeys are not a natural group but also because of recent arguments that explain thick enamel as an evolved response to the seasonal consumption of hard foods.

Alveolar bone has several mechanical functions, including tooth support and accommodation of occlusal and other masticatory forces. Its unique functional-mechanical environment is reflected by its structural characteristics, but whether alveolar bone is materially distinct from bone elsewhere in the primate facial skeleton is uncertain. This uncertainty is attributable not only to a limited amount of data but also to conflicting findings among these data. We evaluated elastic modulus variation in the mandibular corpus of eight adult specimens of the monkeys Procolobus badius and Colobus polykomos via microindentation to evaluate whether alveolar bone is more compliant than basal bone and to quantify patterns of variation between sexes and species. We sampled Vickers hardness from six serial transverse sections and one coronal section from both the alveolar process and the basal corpus. Hardness values were converted to elastic modulus via regressions specific for bone tissue. Analysis of variance reveals that a plurality of variation is found on a regional scale; i.e., alveolar bone is more compliant than adjacent basal bone. Species affiliation and sex are not significant sources of variation. These findings support a hypothesis that compliance of alveolar bone represents a material solution for avoiding large stress concentrations arising from occlusal loads. Other comparative data suggest important differences between colobine and cercopithecine mandibles in terms of bone stiffness, both overall and in terms of relative stiffness of alveolar and basal cortical bone.