Dental sculpting and compensatory shearing crests demonstrated in a Wear series of Presbytis rubicunda (Cercopithecoidea, Colobidae) with dental topography analysis.

OBJECTIVES: Maintaining effective and efficient occlusal morphology presents adaptive challenges for mammals, particularly because mastication produces interactions with foods and other materials that alters the geometry of occlusal surfaces through macrowear and/or catastrophic failure (i.e. "chipping"). Altered occlusal morphologies are often less effective for masticating materials of given diet-but not always-some species exhibit dental sculpting, meaning their dentitions are set up to harness macrowear to hone their occlusal surfaces into more effective morphologies (i.e. secondary morphologies). Here we show that dental sculpting is present in the folivorous Presbytis rubicunda of Borneo. METHODS: Thirty-one undamaged lower second molars of P. rubicunda exhibiting various stages of macroscopic wear were micro-CT scanned and processed into digital surfaces. The surfaces were measured for convex Dirichlet normal energy (vDNE, a measure of surface sharpness), and degree of surface wear. Regression analyses compared surface sharpness with several measures of wear to test for the presence and magnitude of dental sculpting. RESULTS: Positive correlations between the wear proxies and vDNE reveal that P. rubicunda wear in such a way as to become sharper, and therefore more effective chewing surfaces by exposing enamel-dentine junctions on their occlusal surfaces and then honing these junctions into sharpened edges. Compared to another primate folivore in which increasing surface sharpness with macrowear has been demonstrated (i.e., Alouatta palliata), the worn surfaces are similarly sharp, but the dental sculpting process appears to be different. DISCUSSION: The results presented here suggest that not only do some primates exhibit dental sculpting and the attendant secondary morphology, but that there appear to be multiple different morphological configurations that can achieve this result. P. rubicunda has thicker enamel and a more stereotyped wear pattern than A. palliata, although both show positive correlations of occlusal surface sharpness (vDNE) with various wear proxies. These findings shed light on the varied approaches for the maintenance of effective and efficient occlusal surfaces in primates.

Dental topography of the Oligocene anthropoids Aegyptopithecus zeuxis and Apidium phiomense: Paleodietary insights from analysis of wear series.

Fossil primate dietary inference is enhanced when ascertained through multiple, distinct proxies. Dental topography can be used to assess changes in occlusal morphology with macrowear, providing insight on tooth use and function across the lifespans of individuals. We measured convex Dirichlet normal energy-a dental topography metric reflecting occlusal sharpness of features such as cusps and crests-in macrowear series of the second mandibular molars of two African anthropoid taxa from ∼30 Ma (Aegyptopithecus zeuxis and Apidium phiomense). Wear was quantified via three proxies: occlusal dentine exposure, inverse relief index, and inverse occlusal relief. The same measurements were calculated on macrowear series of four extant platyrrhine taxa (Alouatta, Ateles, Plecturocebus, and Sapajus apella) to provide an analogical framework for dietary inference in the fossil taxa. We predicted that Ae. zeuxis and Ap. phiomense would show similar patterns in topographic change with wear to one another and to extant platyrrhine frugivores like Ateles and Plecturocebus. The fossil taxa have similar distributions of convex Dirichlet normal energy to one another, and high amounts of concave Dirichlet normal energy 'noise' in unworn molars-a pattern shared with extant hominids that may distort dietary interpretations. Inverse relief index was the most useful wear proxy for comparison among the taxa in this study which possess disparate enamel thicknesses. Contrary to expectations, Ae. zeuxis and Ap. phiomense both resemble S. apella in exhibiting an initial decline in convex Dirichlet normal energy followed by an increase at the latest stages of wear as measured by inverse relief index, lending support to previous suggestions that hard-object feeding played a role in their dietary ecology. Based on these results and previous analyses of molar shearing quotients, microwear, and enamel microstructure, we suggest that Ae. zeuxis had a pitheciine-like strategy of seed predation, whereas Ap. phiomense potentially consumed berry-like compound fruits with hard seeds.

Using BayesModelS to provide Bayesian- and phylogenetically-informed primate body mass predictions.

An accurate prediction of the body mass of an extinct species can greatly inform the reconstruction of that species' ecology. Therefore, paleontologists frequently predict the body mass of extinct taxa from fossilized materials, particularly dental dimensions. Body mass prediction has traditionally been performed in a frequentist statistical framework, and accounting for phylogenetic relationships while calibrating prediction models has only recently become more commonplace. In this article, we apply BayesModelS-a phylogenetically informed Bayesian prediction method-to predict body mass in a sample of 49 euarchontan species (24 strepsirrhines, 20 platyrrhines, 3 tarsiids, 1 dermopteran, and 1 scandentian) and compare this approach's body mass prediction accuracy with other commonly used techniques, namely ordinary least squares, phylogenetic generalized least squares, and phylogenetic independent contrasts (PICs). When predicting the body masses of extant euarchontans from dental and postcranial variables, BayesModelS and PICs have substantially higher predictive accuracy than ordinary least squares and phylogenetic generalized least squares. The improved performances of BayesModelS and PIC are most evident for dentally derived body mass proxies or when body mass proxies have high degrees of phylogenetic covariance. Predicted values generated by BayesModelS and PIC methods also show less variance across body mass proxies when applied to the Eocene adapiform Notharctus tenebrosus. These more explicitly phylogenetically based methods should prove useful for predicting body mass in a paleontological context, and we provide executive scripts for both BayesModelS and PIC to increase ease of application.

Technical note: Comparing dental topography software using platyrrhine molars.

OBJECTIVES: There remain many idiosyncrasies among the values calculated for varying dental topography metrics arising from differences in software preferences among research groups. The aim of this work is to compare and provide potential conversion formulae for dental topography metrics calculated using differing software platforms. METHODS: Three software packages: ArcGIS, Surfer Manipulator, and molaR were used to calculate orientation patch count rotated (OPCR), Dirichlet normal energy (DNE), occlusal relief (OR), slope (m), and angularity (a) on platyrrhine second upper molars. Values derived from the various software packages were compared for distributional consistency and correlation. Where appropriate, formulae for conversion between like measures calculated on different software platforms were developed. RESULTS: When compared with the same measurement across software, OPCR, OR, and slope were all highly correlated. However, only OR demonstrated distributional consistency (i.e., nearly consistent mean, median, max, and min). Slope and OPCR were both higher when calculated by molaR as compared to Surfer Manipulator and ArcGIS calculations, conversion formulae are provided for these measures. DNE is only weakly correlated with angularity; but is correlated with orientation patch count across taxa. DISCUSSION: We explore why there is variation in the dental topography values calculated among the various software packages. The conversion formulae provided in this work will make possible direct comparisons among studies conducted across multiple research groups.

The Influence of Leaping Frequency on Secondary Bone in Cercopithecid Primates.

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 Taï Forest, Côte 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.

Oblique human symphyseal angle is associated with an evolutionary rate-shift early in the hominin clade.

The rate of change in primate mandibular symphyseal angles was modeled with particular aim of locating a rate-shift within the hominin clade. Prior work noted that the human symphyseal angle must have experienced a rapid rate of change in order to assume the modern human form, suggestive of the non-random work of natural selection. This study indicates that the rate of symphyseal evolution rose dramatically between Australopithecus anamensis and Australopithecus afarensis and continued throughout the diversification of the hominin clade. Noting the timing of this event, we speculate as to what ecological factors could have been at play in driving this rearrangement of the anterior mandible, contributing to the eventual appearance of the human chin.

Do Mid-Crown Enamel Formation Front Angles Reflect Factors Linked to the Pace of Primate Growth and Development?

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.

Functional morphology of the douc langur (Pygathrix spp.) scapula.

Most colobine monkeys primarily move through their arboreal environment quadrupedally. Douc langurs (Pygathrix spp.), however, are regularly observed to use suspensory behaviors at the Endangered Primate Rescue Center (EPRC) in Northern Vietnam. Previous work has linked variation in scapular morphology to different modes of primate arboreal locomotion. Here we investigate whether the shape of the Pygathrix scapula resembles obligate brachiators (gibbons) or obligate arboreal quadrupeds (other cercopithecoids). Using a MicroScribe G2X 3D digitizer, the positions of 17 landmarks were recorded on 15 different species of nonhuman primates (n = 100) from three categories of locomotor behavior: brachiator, arboreal quadruped, and unknown (Pygathrix). All analyses were conducted in the R package geomorph. A Procrustes analysis uniformly scaled the shape data and placed specimens into the same morphospace. A Principal Component Analysis was used to examine scapular shape and a Procrustes ANOVA was conducted to test for shape difference in the scapulae. A pairwise analysis was used to compare the means of the locomotor categories and identify any statistically significant differences. A phylogenetically controlled Procrustes ANOVA was also conducted using a phylogeny from 10kTrees. Results show Pygathrix scapular morphology is significantly different from both arboreal colobine quadrupeds (p < 0.01) and hylobatid brachiators (p < 0.01). It does, however, share some features with each including a long vertebral border, like other cercopithecoids, and a more laterally projecting acromion process, like the hylobatids. The principal difference segregating Pygathrix from both the arboreal quadrupeds and the brachiators is the more medially placed superior angle. These nuanced morphological characteristics associated with suspensory behaviors may be useful for inferring suspensory locomotion in the primate fossil record.

Smooth operator: The effects of different 3D mesh retriangulation protocols on the computation of Dirichlet normal energy.

OBJECTIVES: Dirichlet normal energy (DNE) is a metric of surface topography that has been used to evaluate the relationship between the surface complexity of primate cheek teeth and dietary categories. This study examines the effects of different 3D mesh retriangulation protocols on DNE. We examine how different protocols influence the DNE of a simple geometric shape-a hemisphere-to gain a more thorough understanding than can be achieved by investigating a complex biological surface such as a tooth crown. MATERIALS AND METHODS: We calculate DNE on 3D surface meshes of hemispheres and on primate molars subjected to various retriangulation protocols, including smoothing algorithms, smoothing amounts, target face counts, and criteria for boundary face exclusion. Software used includes R, MorphoTester, Avizo, and MeshLab. DNE was calculated using the R package "molaR." RESULTS: In all cases, smoothing as performed in Avizo sharply decreases DNE initially, after which DNE becomes stable. Using a broader boundary exclusion criterion or performing additional smoothing (using "mesh fairing" methods) further decreases DNE. Increasing the mesh face count also results in increased DNE on tooth surfaces. CONCLUSIONS: Different retriangulation protocols yield different DNE values for the same surfaces, and should not be combined in meta-analyses. Increasing face count will capture surface microfeatures, but at the expense of computational speed. More aggressive smoothing is more likely to alter the essential geometry of the surface. A protocol is proposed that limits potential artifacts created during surface production while preserving pertinent features on the occlusal surface.

Wear and its effects on dental topography measures in howling monkeys (Alouatta palliata).

OBJECTIVES: Three dental topography measurements: Dirichlet Normal Energy (DNE), Relief Index (RFI), and Orientation Patch Count Rotated (OPCR) are examined for their interaction with measures of wear, within and between upper and lower molars in Alouatta palliata. Potential inferences of the "dental sculpting" phenomenon are explored. MATERIALS AND METHODS: Fifteen occluding pairs of howling monkey first molars (15 upper, 15 lower) opportunistically collected from La Pacifica, Costa Rica, were selected to sample wear stages ranging from unworn to heavily worn as measured by the Dentine Exposure Ratio (DER). DNE, RFI, and OPCR were measured from three-dimensional surface reconstructions (PLY files) derived from high-resolution CT scans. Relationships among the variables were tested with regression analyses. RESULTS: Upper molars have more cutting edges, exhibiting significantly higher DNE, but have significantly lower RFI values. However, the relationships among the measures are concordant across both sets of molars. DER and EDJL are curvilinearly related. DER is positively correlated with DNE, negatively correlated with RFI, and uncorrelated with OPCR. EDJL is not correlated with DNE, or RFI, but is positively correlated with OPCR among lower molars only. DISCUSSION: The relationships among these metrics suggest that howling monkey teeth adaptively engage macrowear. DNE increases with wear in this sample presumably improving food breakdown. RFI is initially high but declines with wear, suggesting that the initially high RFI safeguards against dental senescence. OPCR values in howling monkey teeth do not show a clear relationship with wear changes.

The enduring puzzle of the human chin.

Although modern humans are considered to be morphologically distinct from other living primates because of our large brains, dexterous hands, and bipedal gait, all of these features are found among extinct hominins. The chin, however, appears to be a uniquely modern human trait. Probably because of the chin's exclusivity, many evolutionary scenarios have been proposed to explain its origins. To date, researchers have developed adaptive hypotheses relating chins to speech, mastication, and sexual selection; still others see it as a structural artifact tangentially related to complex processes involving evolutionary retraction of the midfacial skeleton. Consensus has remained elusive, partly because hypotheses purporting to explain how this feature developed uniquely in modern humans are all fraught with theoretical and/or empirical shortcomings. Here we review a century's worth of chin hypotheses and discuss future research avenues that may provide greater insight into this human peculiarity.

Symphyseal surface strain during in vitro human mandibular wishboning.

OBJECTIVES: This research theoretically models and empirically records symphyseal surface strain during in vitro human mandibular wishboning (lateral transverse bending) in order to test one aspect of the hypothesis that the chin is an adaptive response to masticatory stresses. From a perspective of optimality, three questions were tested: 1) Do human mandibles function as curved beams during wishboning? 2) Is the presence of a chin associated with lower than predicted curved beam effects? 3) Are there relatively low strain gradients on the lingual and labial symphyseal surfaces respectively? MATERIALS AND METHODS: Based on morphometric criteria, theoretical wishboning strains were calculated for five dentate adult human mandibles. The same mandibles were fitted with strain gauges and subjected to simulated wishboning loads. From the empirically-recorded strains, relative strains were calculated by dividing all strains by the absolute lowest strain in a given specimen. The theoretical and empirical results were compared in order to address the three related questions guiding this research. RESULTS: Human mandibles behave as curved beams during wishboning (question 1). Empirical strain measures showed greater disparity both between and within the labial and lingual symphyseal surfaces than the theoretical models predictions (questions 2 and 3). DISCUSSION: Human symphyseal form, with its distinctive chin, is unlikely to be adapted for countering wishboning loads. Chins are associated with larger than expected strain gradients within and between symphyseal surfaces, which runs counter to the optimality criterion typically invoked in assessing trait performance for signs of adaptation. The implications are twofold: 1) wishboning may not, in fact, be a regular feature of human mastication or 2) wishboning may not pose the same structural risks in human jaws as this load does in other anthropoid primates.

Variation in vervet (Chlorocebus aethiops) hair cortisol concentrations reflects ecological disturbance by humans.

Vervet monkeys (Chlorocebus aethiops) often live in close proximity to humans. Vervets are known to raid crops, homes and gardens in suburban areas leading to human-vervet conflict. In general, primate groups with access to human foods experience increased population densities and intra-group aggression. This suggests high stress loads for vervets living in environments with high levels of human habitat disturbance and close proximity to humans. We tested the hypothesis that populations characterized by high levels of human impact are more physiologically stressed than low human impact populations, and that this increased stress would be reflected in higher concentrations of hair cortisol. We predicted that because females would be less likely to engage in high risk foraging activities, and hence keep more distance from humans than males, their hair cortisol levels should be lower than those in males. We quantified cortisol in the hair of wild caught individuals from populations that experienced different degrees of human habitat disturbance and differences in access to human food. We found that males in high human impact groups had significantly higher hair cortisol concentrations than those in low human impact groups, although this difference was not observed in female vervets. Human impacts on vervet behavioral ecology appear to be a significant source of stress for male animals in particular.

Selection played a role in the evolution of the human chin.

Chins, which are unique to humans, have generated considerable debate concerning their evolutionary origins, yet a consensus has remained elusive. Many have argued that chins are adaptations for chewing stress, speech, or sexual ornamentation. Alternatively, some have suggested that chins are spandrels-byproducts of selection operating elsewhere in the mandible or face. Lastly, chins could be the product of genetic drift. The questions addressed by this study are: [1] whether chins represent an exceptionally derived morphological condition, and [2] if this can be interpreted as the product of natural selection. These questions are important since the chin is one of the features used to define Homo sapiens in the fossil record. Quantitative measures that capture the degree of chin expression were gathered from a sample of 123 primate taxa, and evolutionary rates associated with these measures were reconstructed in the primate phylogeny. The evolutionary rate associated with these measures was reconstructed to be far higher along the Homo tip (∼77 times greater than the primate background rate of evolution) than elsewhere in the primate phylogeny. These results suggest that human symphyseal morphology is exceptionally derived relative to other primates, and selection has been operational in producing the human chin.

The coevolution of circumperineal color and terrestriality.

Old World monkeys (Cercopithecoidea) are unusual among primates for the high percentage of species exhibiting circumperineal coloration, as well as the large percentage of highly terrestrial species. Kingdon [1974, 1980] suggested that circumperineal skin coloration is functionally related to terrestriality, but this hypothesis has not been tested. From the literature, we collected data on habitat use (terrestrial/arboreal) and circumperineal coloration (present/absent) for 78 species. Indeed, among the 78 species surveyed here, 75% of them fall into either the category of colored circumperineals with terrestrial lifestyle, or of uncolored circumperineals with arboreal lifestyle (Χ(2)(1) = 19.550, P < 0.001). However, conventional statistical procedures assume all taxa are equally related-which is not usually the case in multispecies analyses-leading to higher rates of both type I and II statistical errors. We performed Bayesian trait coevolution analyses that show that models of dependent trait evolution are not significantly better than models assuming independent evolution of the two traits (log-likelihood ratio test P = 0.396, Bayes Factor = 1). Bayesian nodal reconstructions of the cercopithecoid phylogeny indicate that relatively few trait transitions are needed to account for the distributions of the two traits. Further, chi-squared distributional tests show that sub-family affiliation (i.e., Cercopithecinae, Colobinae) is an accurate predictor of trait status. The discordance of the analyses may represent the results of a few different evolutionary scenarios, but ultimately circumperineal coloration seems weakly linked to terrestrial ecology.

Functional and evolutionary aspects of axial stability in euarchontans and other mammals.

The presence of a stable thoracolumbar region, found in many arboreal mammals, is considered advantageous for bridging and cantilevering between discontinuous branches. However, no study has directly explored the link between osteological features cited as enhancing axial stability and the frequency of cantilevering and bridging behaviors in a terminal branch environment. To fill this gap, we collected metric data on costal and vertebral morphology of primate and nonprimate mammals known to cantilever and bridge frequently and those that do not. We also quantified the frequency and duration of cantilevering and bridging behaviors using experimental setups for species that have been reported to show differences in use of small branches and back anatomy (Caluromys philander, Loris tardigradus, Monodelphis domestica, and Cheirogaleus medius). Phylogenetically corrected principal component analysis reveals that taxa employing frequent bridging and cantilevering (C. philander and lorises) also exhibit reduced intervertebral and intercostal spaces, which can serve to increase thoracolumbar stability, when compared to closely related species (M. domestica and C. medius). We observed C. philander cantilevering and bridging significantly more often than M. domestica, which never cantilevered or crossed any arboreal gaps. Although no difference in the frequency of cantilevering was observed between L. tardigradus and C. medius, the duration of cantilevering bouts was significantly greater in L. tardigradus. These data suggest that osteological features promoting axial rigidity may be part of a morpho-behavioral complex that increases stability in mammals moving and foraging in a terminal branch environment.

Dental wear among cercopithecid monkeys of the Taï forest, Côte d'Ivoire.

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 Taï Forest, Côte 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.

Homoplasy and thick enamel in primates.

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.

Brief communication: Enamel thickness and durophagy in mangabeys revisited.

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.

Technical note: converting durometer data into elastic modulus in biological materials.

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.