James D. Pampush

Hard-object feeding in sooty mangabeys (Cercocebus atys) and interpretation of early hominin feeding ecology.

Morphology of the dentofacial complex of early hominins has figured prominently in the inference of their dietary adaptations. Recent theoretical analysis of craniofacial morphology of Australopithecus africanus proposes that skull form in this taxon represents adaptation to feeding on large, hard objects. A modern analog for this specific dietary specialization is provided by the West African sooty mangabey, Cercocebus atys. This species habitually feeds on the large, exceptionally hard nuts of Sacoglottis gabonensis, stereotypically crushing the seed casings using their premolars and molars. This type of behavior has been inferred for A. africanus based on mathematical stress analysis and aspects of dental wear and morphology. While postcanine megadontia, premolar enlargement and thick molar enamel characterize both A. africanus and C. atys, these features are not universally associated with durophagy among living anthropoids. Occlusal microwear analysis reveals complex microwear textures in C. atys unlike those observed in A. africanus, but more closely resembling textures observed in Paranthropus robustus. Since sooty mangabeys process hard objects in a manner similar to that proposed for A. africanus, yet do so without the craniofacial buttressing characteristic of this hominin, it follows that derived features of the australopith skull are sufficient but not necessary for the consumption of large, hard objects. The adaptive significance of australopith craniofacial morphology may instead be related to the toughness, rather than the hardness, of ingested foods.

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.

Introducing molaR: a New R Package for Quantitative Topographic Analysis of Teeth (and Other Topographic Surfaces)

Researchers studying mammalian dentitions from functional and adaptive perspectives increasingly have moved towards using dental topography measures that can be estimated from 3D surface scans, which do not require identification of specific homologous landmarks. Here we present molaR, a new R package designed to assist researchers in calculating four commonly used topographic measures: Dirichlet Normal Energy (DNE), Relief Index (RFI), Orientation Patch Count (OPC), and Orientation Patch Count Rotated (OPCR) from surface scans of teeth, enabling a unified application of these informative new metrics. In addition to providing topographic measuring tools, molaR has complimentary plotting functions enabling highly customizable visualization of results. This article gives a detailed description of the DNE measure, walks researchers through installing, operating, and troubleshooting molaR and its functions, and gives an example of a simple comparison that measured teeth of the primates Alouatta and Pithecia in molaR and other available software packages. molaR is a free and open source software extension, which can be found at the doi:10.13140/RG.2.1.3563.4961 (molaR v. 2.0) as well as on the Internet repository CRAN, which stores R packages.

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

AbstractVervet 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.

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. J. Morphol. 313–327, 2014.

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 chins 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 centurys worth of chin hypotheses and discuss future research avenues that may provide greater insight into this human peculiarity.

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.

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.

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.