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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.

We review feeding and mandibular anatomy in a community of West African monkeys. We use field observations, food material property data, and skeletal specimens from the Ivory Coast's Taï Forest to explore the factors that shape mandibular architecture in colobines and cercopithecines. Despite excellent geographic control across our sample, the fit between bone form (as conventionally described) and functional activity (as we perceive it) is not spectacular. We present a thought experiment to assess how well we could reconstruct diet in the Taï monkeys if we only had skeletons and teeth to study. This exercise indicated that we would be correct about half the time. Our analyses reinforce the notion that diet is anything but a monolithic variable and that better success at relating mandibular form to food must incorporate information on ingestive and processing behavior, geometric and material properties of foods, and both material and structural data on jaws themselves.

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.

The olive colobus (Procolobus verus) is the smallest extant colobine. Based on the axiom that folivory is associated with larger body mass, the olive colobus is expected to be less folivorous than its sister taxon Piliocolobus badius, but previous studies show that the opposite is true. Here we test the hypothesis that masticatory and ingestive effort in the olive colobus is greater due to allometric factors related to bite force scaling and throughput of ingested foods. We analyzed oral processing data collected on olive colobus in the Taï Forest, Ivory Coast, between May 2016 and May 2018. We compare these with previously published data on P. badius and Colobus polykomos from Taï. In terms of overall feeding effort, olive colobus invest more effort (i.e., chewing cycles) than the larger colobines. When contrasts are restricted to commonly consumed foods, this greater energetic investment is not consistently observed. Ingestion of young leaves is associated with a reduced number of masticatory cycles in all three colobine species. A slightly elevated average effort in the olive colobus during young leaf feeding suggests this food source is more challenging in smaller monkeys, but mature leaf processing effort is generally the same among Taï colobines. Thus, for olive colobus, leaf ductility may be more problematic than leaf toughness in terms of masticatory effort. While there may be an allometric cost to being a small colobine, food selectivity is an important mitigating factor.

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.

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.

Two similarly-sized colobine species living sympatrically in the Ivory Coast's Taï Forest that differ in both diet and oral processing behavior provide an opportunity to explore the strength of associations between feeding behavior and dental wear patterns. Here we test the hypothesis that vigorous processing of tough, hard Pentaclethra macrophylla pods by Colobus polykomos manifests in greater anterior tooth wear relative to that observed in Piliocolobus badius, which does not exploit this resource. We assessed levels of anterior tooth wear in a sample of 160 upper incisors and 131 lower incisors from 18 adult Colobus polykomos and 62 adult Piliocolobus badius naturally deceased individuals from Taï National Park. We operationalized tooth wear by dividing the area of exposed dentin by total occlusal crown area. To assess relative degrees of incisor wear, we regressed incisor wear against molar wear (sample = 105 upper molars, 135 lower molars) for the pooled Colobus polykomos and Piliocolobus badius wear data and compared the number of individuals from each species that fell above and below the pooled regression curve for each model using Chi-square tests of independence and odds ratios. Under our hypothesis, we would expect more Colobus polykomos points above the pooled regression curve than Piliocolobus badius, indicating higher incisor wear relative to molar wear in Colobus polykomos. Nine of sixteen interspecific comparisons demonstrated this predicted pattern; however, none of the Chi-square tests or odds ratios were significant, indicating no difference between Colobus polykomos and Piliocolobusbadius incisor wear relative to molar wear. The absence of significant differences in incisor wear relative to molar wear highlights the challenge of identifying idiosyncratic feeding behavior in fossil taxa and the necessity for continued exploration of the relationship between diet and macrowear.

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 The relationship between radiographic grayscale and elastic modulus was determined using mandibular cortical bone of colobine monkeys. Quantification of this relationship is critical for establishing absolute measures of structural rigidity of skeletal elements. Materials and methods We determined the Vickers microhardness values in mandibular bone from two species of African colobine monkeys and related these values to elastic modulus through an empirically determined correlation. We also determined radiographic grayscale values from microcomputed tomographic scans of the mandible in the same regions in which microhardness was sampled. We then correlated modulus to grayscale with a power law relationship. Results We found that elastic modulus scaled with negative allometry with respect to grayscale with an exponent of 0.77. Discussion Our results suggest a single exponent can effectively capture the relationship of grayscale to elastic modulus and facilitate development of heterogeneous structural models for use in comparative and computational biomechanical studies.

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.

According to the IUCN [1], four of the nine anthropoid species found in Liberia are classified as either Vulnerable or Endangered and this number is likely to rise in coming years due to an increas...

Primate canines function in social displays but are also recruited for biting in agonistic encounters. Although the precise nature of the loads such behavior places on the canine crown is unknown, it is thought that bending is a major component of such loads. To date, modeling of canine bending strength has relied on idealized geometric representations. Accounting for the tapering of the crown as well as shape changes along an apical-basal axis provides a more realistic model for assessment of bending stress in canines. We provide such an accounting and evaluate the hypothesis that the morphology of the cercopithecoid canine represents a structural solution for maintaining constant maximum bending stress under apical or distributed loading in a parasagittal plane. This isostress hypothesis is analogous to a design criterion of minimum mass for a given structural requirement. Examining permanent maxillary canines from males and females representing eight West African cercopithecoid species, we reconstructed crown geometry from apex to base using microcomputed tomography. From reconstructed cross-sections, we determined section moduli about a buccolingual centroidal axis. We then determined what the taper should be for a variety of parasagittal loading distributions if the isostress hypothesis were true and compared these theoretical tapers to actual crown geometry. We found that a variety of loading distributions can be accommodated by the canines, particularly among males. These results suggest that our sample of canines are not optimized for resisting particular biting loads, but effectively limit stress gradients associated with a range of behaviors.

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.

Researchers have increasingly recognized the need to quantify the material properties of primate food items, particularly hardness (H) and stiffness (E), which is measured as elastic modulus. Assessing E in the field is particularly difficult because the typical equipment needed to perform the requisite analyses is expensive and cumbersome. Alternatively, researchers can use hand-held, relatively inexpensive, portable durometers that measure H on Shore scales. Shore-D durometers show a reliable ability to characterize H in harder-stiffer materials, and Shore-D measures in these materials can be reliably converted to E. Shore-A durometers-employed in past field studies of food properties-do not accurately characterize the properties of harder-stiffer materials, which are likely to be those materials exerting the greatest mechanical demands on primate masticatory morphology. We offer recommendations for Shore-D durometer usage in the field.

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.