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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.

OBJECTIVES Magnitudes of morphological integration may constrain or facilitate craniofacial shape variation. The aim of this study was to analyze how the magnitude of integration in the skull of Macaca fascicularis changes throughout ontogeny in relation to developmental and/or functional modules. MATERIALS AND METHODS Geometric morphometric methods were used to analyze the magnitude of integration in the macaque cranium and mandible in 80 juvenile and 40 adult M. fascicularis specimens. Integration scores in skull modules were calculated using integration coefficient of variation (ICV) of eigenvalues based on a resampling procedure. Resultant ICV scores between the skull as a whole, and developmental and/or functional modules were compared using Mann-Whitney U tests. RESULTS Results showed that most skull modules were more tightly integrated than the skull as a whole, with the exception of the chondrocranium in juveniles without canines, the chondrocranium/face complex and the mandibular corpus in adults, and the mandibular ramus in all juveniles. The chondrocranium/face and face/mandibular corpus complexes were more tightly integrated in juveniles than adults, possibly reflecting the influences of early brain growth/development, and the changing functional demands of infant suckling and later masticatory loading. This is also supported by the much higher integration of the mandibular ramus in adults compared with juveniles. DISCUSSION Magnitudes of integration in skull modules reflect developmental/functional mechanisms in M. fascicularis. However, the relationship between "evolutionary flexibility" and developmental/functional mechanisms was not direct or simple, likely because of the complex morphology, multifunctionality, and various ossification origins of the skull.

Objectives Variation in primate masticatory form and function has been extensively researched through both morphological and experimental studies. As a result, symphyseal fusion in different primate clades has been linked to either the recruitment of vertically directed balancing-side muscle force, the timing and recruitment of transversely directed forces, or both. This study investigates the relationship between jaw muscle activity patterns and morphology in extant primates to make inferences about masticatory function in extinct primates, with implications for understanding the evolution of symphyseal fusion. Materials and methods Three-dimensional mandibular landmark data were collected for 31 extant primates and nine fossil anthropoids and subfossil lemur species. Published electromyography (EMG) data were available for nine of the extant primate species. Partial least squares analysis and phylogenetic partial least squares analysis were used to identify relationships between EMG and jaw shape data and evaluate variation in jaw morphology. Results Primates with partial and complete symphyseal fusion exhibit shape-function patterns associated with the wishboning motor pattern and loading regime, in contrast to shape-function patterns of primates with unfused jaws. All fossil primates examined (except Apidium) exhibit jaw morphologies suggestive of the wishboning motor pattern demonstrated in living anthropoids and indriids. Discussion Partial fusion in Catopithecus, similar to indriids and some subfossil lemurs, may be sufficient to resist, or transfer, some amounts of transversely directed balancing-side muscle force at the symphysis, representing a transition to greater reliance on transverse jaw movement during mastication. Furthermore, possible functional convergences in physiological patterns during chewing (i.e., Archaeolemur) are identified.

OBJECTIVES To increase understanding of the subsistence practices of the first Americans through analysis of the near-complete dentition of a young woman dating to the terminal Pleistocene of the Yucatan Peninsula, Mexico. MATERIALS AND METHODS The skeleton is that of "Naia" a 15 to 17-year-old female from the submerged natural trap of Hoyo Negro found in association with remains of numerous extinct species of megafauna. Superbly preserved remains included the skull with 28 teeth, which are analyzed for evidence of caries, periodontal disease, wear patterns, and malocclusion. RESULTS Naia exhibits a high frequency of dental caries, along with aggressive periodontal disease that threatened all her teeth, particularly her incisors. Dental attrition was extremely light for a hunter-gatherer, reaching to four on the Molnar scale on only one tooth. Lack of wear is associated with severe mandibular retrognathia, and low masticatory forces. DISCUSSION Naia's dental condition is compared with that of other northern Paleoamericans, mostly females, dating before 11,000 cal BP. These exhibit a high degree of variability in both caries and tooth wear. All, however, exhibit rapid anterior wear owing to technological use of the front teeth. Naia exhibits the highest rate of caries, similar to that of the earliest South Americans, and one of the lowest rates of attrition. This demonstrates that she had a nonabrasive diet that was at least seasonally rich in carbohydrates. This does not mean her diet was low in meat, however, because similarly light dental attrition is seen in the Arch Lake female, a Paleoamerican from a big-game hunting society.

1 Objectives Examining the relationships among foraging behavior, food mechanical properties, and masticatory morphology is a bourgeoning research topic among behavioral ecologists and functional morphologists. The purpose of this study was to assess the degree to which leaf toughness influences foraging efficiency with regard to ingestion rate, masticatory investment, and masticatory rate. 2 Methods Diet and feeding data were collected on adults from three groups of Colobus angolensis palliatus in the Diani Forest, Kenya, from July 2014 to December 2015. Ingestion rates were estimated by counting the number of items consumed during feeding bouts and multiplying this value by the mean mass of a particular food item. The number of mastications was also counted during 3–5 minute focal periods. Mechanical toughness of commonly eaten young leaves (n = 27 species) and mature leaves (n = 13 species) was recorded using a toughness tester equipped with a razor blade. 3 Results Ingestion rates (g/min) negatively correlated with leaf toughness (r2 = 0.73; p

Objectives Changes to soft- and hard-tissue components of the masticatory complex during development can impact functional performance by altering muscle excursion potential, maximum muscle forces, and the efficiency of force transfer to specific bitepoints. Within Macaca fascicularis, older individuals exploit larger, more mechanically resistant food items and more frequently utilize wide-gape jaw postures. We therefore predict that key architectural and biomechanical variables will scale during ontogeny to maximize bite force and gape potential within older, larger-bodied individuals. Materials and methods We analyzed 26 specimens of M. fascicularis, representing a full developmental spectrum. The temporalis, superficial masseter, and deep masseter were dissected to determine muscle mass, fiber length, and physiologic cross-sectional area (PCSA). Lever-arm lengths were also measured for each muscle, alongside the height of the temporomandibular joint (TMJ) and basicranial length. These variables were scaled against two biomechanical variables (jaw length and condyle-molar length) to determine relative developmental changes within these parameters. Results During ontogeny, muscle mass, fiber length, and PCSA scaled with positive allometry relative to jaw length and condyle-molar length within all muscles. TMJ height also scaled with positive allometry, while muscle lever arms scaled with isometry relative to jaw length and with positive allometry (temporalis) or isometry (superficial and deep masseter) relative to condyle-molar length. Conclusion Larger individuals demonstrate adaptations during development towards maximizing gape potential and bite force potential at both an anterior and posterior bitepoint. These data provide anatomical evidence to support field observations of dietary and behavioral differences between juvenile and adult M. fascicularis.

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. Am J Phys Anthropol, 2015. © 2015 Wiley Periodicals, Inc.

Objectives Platyrrhines span two orders of magnitude in body size and are characterized by diverse feeding behaviors and diets. While size plays an important role in primate feeding behavior and masticatory apparatus morphology, we know little about size-correlated changes in the force-generating (physiologic cross-sectional area; PCSA) and excursion/stretch (fiber length; Lf) capabilities of the jaw-closing muscles in platyrrhines. Methods We examined scaling relationships of the superficial masseter and temporalis muscles in 21 platyrrhine species. Previous work suggests that larger platyrrhines are at a mechanical disadvantage for generating bite forces compared with smaller platyrrhines. We hypothesize that scaling of jaw-muscle fiber architecture counters this size-correlated decrease in mechanical advantage. Thus, we predicted that jaw-muscle PCSAs and muscle weights scale with positive allometry while Lfs scale with negative allometry, relative to load-arm estimates for incisor/molar biting and chewing. Results Jaw-muscle PCSAs and Lfs appear to scale with negative allometry relative to load-arm estimates and body size. Negative allometry of jaw-muscle weights partially accounts for the size-correlated decreases in PCSA and Lf. Estimates of bite force also scale with negative allometry. Conclusion Large-bodied platyrrhines (e.g., Alouatta) are at a relative disadvantage for generating jaw-muscle and bite force as well as jaw-muscle stretch, compared with smaller species (e.g., Callithrix). The net effect is that larger platyrrhines likely produce relatively smaller maximal bite forces compared with smaller taxa. Relative to small- and intermediate-sized platyrrhines, large-bodied platyrrhines feed on some of the least mechanically challenging foods, consistent with the size-correlated decrease in relative muscle and bite forces across the clade. Am J Phys Anthropol, 2015. © 2015 Wiley Periodicals, Inc. Am J Phys Anthropol 158:242–256, 2015. © 2015 Wiley Periodicals, Inc.

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.

Evolutionary and functional significance of the human chin has long been explored from various perspectives including masticatory biomechanics, speech, and anterior tooth size. Recent ontogenetic studies have indicated that the spatial position of internally forming anterior teeth partially constrains adult mandibular symphyseal morphology. The present study therefore preliminarily examined the size and placement of developing anterior teeth in immature Neanderthal mandibles of Dederiyeh 1 and 2, compared with similarly-aged modern humans (N = 16) and chimpanzees (N = 7) whose incisors are comparatively small and large among extant hominids, respectively. The Dederiyeh 1 mandible is described as slightly presenting a mental trigone and attendant mental fossa, whereas Dederiyeh 2 completely lacks such chin-associated configurations. Results showed that, despite symphyseal size being within the modern human range, both Dederiyeh mandibles accommodated overall larger anterior dentition and displayed a remarkably wide bicanine space compared to those of modern humans. Dederiyeh 2 had comparatively thicker deciduous incisor roots and more enlarged permanent incisor crypts than Dederiyeh 1, but both Dederiyeh individuals exhibited a total dental size mostly intermediate between modern humans and chimpanzees. These findings potentially imply that the large deciduous/permanent incisors collectively distended the labial alveolar bone, obscuring an incipient mental trigone. It is therefore hypothesized that the appearance of chin-associated features, particularly of the mental trigone and fossa, can be accounted for partly by developmental relationships between the sizes of the available mandibular space and anterior teeth. This hypothesis must be, however, further addressed with more referential samples in future studies. Am J Phys Anthropol 156:482–488, 2015. © 2014 Wiley Periodicals, Inc.

Ecological factors can be important to shape the patterns of morphological variation among human populations. Particularly, diet plays a fundamental role in craniofacial variation due to both the effect of the nutritional status-mostly dependent on the type and amount of nutrients consumed-on skeletal growth and the localized effects of masticatory forces. We examine these two dimensions of diet and evaluate their influence on morphological diversification of human populations from southern South America during the late Holocene. Cranial morphology was measured as 3D coordinates defining the face, base and vault. Size, form, and shape variables were obtained for 474 adult individuals coming from 12 samples. Diet composition was inferred from carious lesions and δ(13) C data, whereas bite forces were estimated using traits of main jaw muscles. The spatial structure of the morphological and ecological variables was measured using correlograms. The influence of diet composition and bite force on morphometric variation was estimated by a spatial regression model. Cranial variation and diet composition display a geographical structure, while no geographical pattern was observed in bite forces. Cranial variation in size and form is significantly associated with diet composition, suggesting a strong effect of systemic factors on cranial growth. Conversely, bite forces do not contribute significantly to the pattern of morphological variation among the samples analyzed. Overall, these results show that an association between diet composition and hardness cannot be assumed, and highlight the complex relationship between morphological diversification and diet in human populations.

Variation in recent human mandibular form is often thought to reflect differences in masticatory behavior associated with variation in food preparation and subsistence strategies. Nevertheless, while mandibular variation in some human comparisons appear to reflect differences in functional loading, other comparisons indicate that this relationship is not universal. This suggests that morphological variation in the mandible is influenced by other factors that may obscure the effects of loading on mandibular form. It is likely that highly strained mandibular regions, including the corpus, are influenced by well-established patterns of lower facial skeletal integration. As such, it is unclear to what degree mandibular form reflects localized stresses incurred during mastication vs. a larger set of correlated features that may influence bone distribution patterns. In this study, we examine the relationship between mandibular symphyseal bone distribution (i.e., second moments of area, cortical bone area) and masticatory force production (i.e., in vivo maximal bite force magnitude and estimated symphyseal bending forces) along with lower facial shape variation in a sample of n = 20 living human male subjects. Our results indicate that while some aspects of symphyseal form (e.g., wishboning resistance) are significantly correlated with estimates of symphyseal bending force magnitude, others (i.e., vertical bending resistance) are more closely tied to variation in lower facial shape. This suggests that while the symphysis reflects variation in some variables related to functional loading, the complex and multifactorial influences on symphyseal form underscores the importance of exercising caution when inferring function from the mandible especially in narrow taxonomic comparisons.

Maximum bite force affects craniofacial morphology and an organism's ability to break down foods with different material properties. Humans are generally believed to produce low bite forces and spend less time chewing compared with other apes because advances in mechanical and thermal food processing techniques alter food material properties in such a way as to reduce overall masticatory effort. However, when hominins began regularly consuming mechanically proc- essed or cooked diets is not known. Here, we apply a model for estimating maximum bite forces and stresses at the second molar in modern human, nonhuman pri- mate, and hominin skulls that incorporates skeletal data along with species-specific estimates of jaw muscle archi- tecture. The model, which reliably estimates bite forces, shows a significant relationship between second molar bite force and second molar area across species but does not confirm our hypothesis of isometry. Specimens in the genus Homo fall below the regression line describing the relationship between bite force and molar area for non- human anthropoids and australopiths. These results suggest that Homo species generate maximum bite forces below those predicted based on scaling among australopiths and nonhuman primates. Because this decline occurred before evidence for cooking, we hypothesize that selection for lower bite force production was likely made possible by an increased reliance on nonthermal food processing. However, given substantial variability among in vivo bite force magnitudes meas- ured in humans, environmental effects, especially varia- tions in food mechanical properties, may also be a factor. The results also suggest that australopiths had ape-like bite force capabilities. Am J Phys Anthropol 151:544- 557, 2013. V C 2013 Wiley Periodicals, Inc.

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.

Jaw-joint height (JJH) above the occlusal plane is thought to be influenced by cranial base angle (CBA) and facial angulation during growth. To better understand how JJH relates to midline craniofacial form, we test the hypothesis that relative increases in JJH are correlated with increasing CBA flexion and facial kyphosis (i.e., ventral bending) across primates. We compared JJH above the occlusal plane to CBA and the angle of facial kyphosis (AFK) across adults from 82 species. JJH scales with positive allometry relative to a skull geometric mean in anthropoids and most likely strepsirrhines. Anthropoid regressions for JJH are elevated above strepsirrhines, whereas catarrhines exhibit a higher slope than platyrrhines. Semipartial correlations between relative JJH and both CBA and AFK show no association across a small strepsirrhine sample, limited associations among catarrhines and anthropoids, but strong correlations in platyrrhines. Contrary to our hypothesis, however, increases in relative JJH are correlated with relatively less flexed basicrania and more airorhynch faces (i.e., reduced ventral bending) in platyrrhines. The mosaic pattern of relationships involving JJH across primate clades points to multiple influences on JJH across primates. In clades showing little association with basicranial and facial angles, such as strepsirrhines, the potential morphological independence of JJH may facilitate a relative freedom for evolutionary changes related to masticatory function. Finally, failure to associate relative JJH and basicranial flexion in most clades suggests that the relatively taller JJH and more flexed basicrania of anthropoids compared to strepsirrhines may have evolved as an isolated event during the origin of anthropoids.

Masticatory loading is one of the main environmental stimuli that generate craniofacial variation among recent humans. Experimental studies on a wide variety of mammals, including those with retrognathic postcanine teeth, predict that responses to masticatory loading will be greater in the occlusal plane, the inferior rostrum, and regions associated with the attachments of the temporalis and masseter muscles. Here we test these experimentally-derived predictions on an extinct human population from the middle and upper Ohio valley that underwent a marked shift from hunting-gathering to extensive farming during the last 3,000 years and for which we have good archaeological evidence about diet and food processing technology. Geometric morphometric methods were used to detect and measure the putative effect of diet changes on cranial shape independent of size. Our results partially confirm only some of the experimental predictions. The effect of softer and/or less tough diets on craniofacial shape seem to be concentrated in the relative reduction of the temporal fossa and in a displacement of the attachment of the temporal muscle. However, there were few differences in craniofacial shape in regions closer to the occlusal plane. These results highlight the utility of exploring specific localized morphological shifts using a hierarchical model of craniofacial integration.

Information about food ingestion and mastication behavior during the lifespan of an individual is encoded in the dental occlusal wear pattern. To decode this information, we describe a new method called occlusal fingerprint analysis (OFA). Structural parameters of wear facets on the occlusal surface of teeth are quantified from digitized casts for the interpretation of occlusal aspects. The OFA provides an individual three-dimensional dental occlusal compass that indicates the major pathways of interaction between antagonists, revealing information about development, spatial position, and enlargement of wear facets. Humans develop a very similar overall pattern of crown contacts, although specific characteristics of wear facets reflect an individual's occlusal relationships and masticatory behavior. We hypothesize that the wear pattern is a unique character and therefore valuable for individual identification. Furthermore we suggest that OFA, when further developed, may be useful for identification of behavioral, biological, and chemical factors affecting crown morphology.

Research on the evolution and adaptive significance of primate craniofacial morphologies has focused on adult, fully developed individuals. Here, we investigate the possible relationship between the local stress environment arising from masticatory loadings and the emergence of the supraorbital torus in the developing face of the crab-eating macaque Macaca fascicularis. By using finite element analysis (FEA), we are able to evaluate the hypothesis that strain energy density (SED) magnitudes are high in subadult individuals with resulting bone growth in the supraorbital torus. We developed three micro-CT-based FEA models of M. fascicularis skulls ranging in dental age from deciduous to permanent dentitions and validated them against published experimental data. Applied masticatory muscle forces were estimated from physiological cross-sectional areas of macaque cadaveric specimens. The models were sequentially constrained at each working side tooth to simulate the variation of the bite point applied during masticatory function. Custom FEA software was used to solve the voxel-based models and SED and principal strains were computed. A physiological superposition SED map throughout the face was created by allocating to each element the maximum SED value from each of the load cases. SED values were found to be low in the supraorbital torus region throughout ontogeny, while they were consistently high in the zygomatic arch and infraorbital region. Thus, if the supraorbital torus arises to resist masticatory loads, it is either already adapted in each of our subadult models so that we do not observe high SED or a lower site-specific bone deposition threshold must apply.

Cercopithecoid monkeys experience relatively high strains along the lingual aspect of the mandibular symphysis because of lateral transverse bending of the mandibular corpora ("wishboning") during mastication. Hylander (Am J Phys Anthropol 64 (1984) 1-46; Am Zool 25 (1985) 315-330) demonstrated that the distribution of strains arising from wishboning loads is comprehensible with reference to the mechanics of curved beams. Theory of curved beams suggests that lingual tensile strains are some multiple of labial compressive strains, yet limitations of experimental methods and uncertainty in estimating parameters needed for theoretical calculations have confounded attempts to characterize the magnitude of this disparity of normal strains. We evaluate the theoretical disparity of normal strains in wishboning in comparison to in vitro strains collected under controlled loads for a sample of mandibles representing two colobine species (N = 6). These data suggest that in colobine monkeys, maximum normal lingual strains should be at least twice maximum labial strains. In addition, we reexamine the distribution of symphyseal stress under an assumption of asymmetric bending, a general approach for calculation of stress appropriate for members that lack a plane of symmetry and are bent along an axis that is not coincident with the member's principal axes. Under asymmetric bending in colobine mandibles, the effect of symphyseal inclination on lingual strain is mitigating at the superior transverse torus and exacerbating at the inferior transverse torus. Relative compliance of colobine mandibular bone further supports the hypothesis that the structural and material properties of the colobine mandibular symphysis do not represent a morphological strategy for minimizing masticatory strain.

Common (Callithrix jacchus) and pygmy (Cebuella pygmaea) marmosets and cotton-top tamarins (Saguinus oedipus) share broadly similar diets of fruits, insects, and tree exudates. Marmosets, however, differ from tamarins in actively gouging trees with their anterior dentition to elicit tree exudates flow. Tree gouging in common marmosets involves the generation of relatively wide jaw gapes, but not necessarily relatively large bite forces. We compared fiber architecture of the masseter and temporalis muscles in C. jacchus (N = 18), C. pygmaea (N = 5), and S. oedipus (N = 13). We tested the hypothesis that tree-gouging marmosets would exhibit relatively longer fibers and other architectural variables that facilitate muscle stretch. As an architectural trade-off between maximizing muscle excursion/contraction velocity and muscle force, we also tested the hypothesis that marmosets would exhibit relatively less pinnate fibers, smaller physiologic cross-sectional areas (PCSA), and lower priority indices (I) for force. As predicted, marmosets display relatively longer-fibered muscles, a higher ratio of fiber length to muscle mass, and a relatively greater potential excursion of the distal tendon attachments, all of which favor muscle stretch. Marmosets further display relatively smaller PCSAs and other features that reflect a reduced capacity for force generation. The longer fibers and attendant higher contraction velocities likely facilitate the production of relatively wide jaw gapes and the capacity to generate more power from their jaw muscles during gouging. The observed functional trade-off between muscle excursion/contraction velocity and muscle force suggests that primate jaw-muscle architecture reflects evolutionary changes related to jaw movements as one of a number of functional demands imposed on the masticatory apparatus.

The Argentine Center-West was the southernmost portion of the Andes where domestication of plants and animals evolved. Populations located in the southern portion of this area displayed a hunter-gatherer subsistence economy up to historical times, and coexisted with farmers located to the north. Archaeological and biological evidence suggests that the transition to food production was associated with the consumption of a softer diet and a more sedentary way of life. This study tests the hypothesis that diet-related factors influenced morphological differentiation, by comparing functional cranial components of farmers and hunter-gatherers. Three-dimensional changes on eight minor functional components (anteroneural, midneural, posteroneural, otic, optic, respiratory, masticatory, and alveolar) were measured on skulls derived from both subareas. Volumetric and morphometric indices were calculated to estimate the absolute and relative size of components, respectively. Results of a paired t-test indicated that farmers have a smaller craniofacial size than hunter-gatherers. The components that varied the most were masticatory and posteroneural, showing smaller absolute and relative sizes in farmers. Discriminant analyses indicated that lengths and widths were the most affected dimensions of these and other components. The pattern of differentiation, which involves specific components, enabled us to exclude differential gene flow and stochastic mechanisms as the main causes. Instead, results support the hypothesis that diet-related factors associated with both subsistence economies influenced craniofacial morphology. A proportion of the observed variation associated with size differences can be explained by two systemic factors: the lesser quality of nutrition due to a low protein content in the diet, and a decrease of growth hormone circulation induced by a lower mobility due to sedentism. However, differentiation is better explained by a localized factor: the reduction in the masticatory and posteroneural components in farmers resulted from a decrease of masticatory stresses and workload on the head and neck, linked to the consumption of a softer diet.

The human face differs from most other primates in that it lacks a rostrum and is oriented primarily in the coronal plane. These differences may influence biomechanical resistance to torsional, shearing, and bending forces generated during mastication that are known to affect the growth of the maxillary and mandibular arches. We tested the effects of facial retraction on facial strain patterns using in vivo strain data collected from rock hyraxes (Procavia capensis) and pigs (Sus scrofa). The postcanine tooth row in hyraxes is retracted beneath the orbits, as in humans, but lies anterior to the orbit under a rostrum in pigs. Average shear strains recorded for hard foods on the anterior dorsal rostrum (lateral to the midline) were 65 μ€ in the pigs, and 25 μ€ in the hyraxes. Average shear strains on the dorsal interorbital surface (lateral to the midline) were 125 μ€ in the pigs, and 320 μ€ in the hydraxes. Although some bending occurred, torsion about an A-P axis was the major mode of deformation during mastication in both species. The lower rostral strains and higher caudal strains in the hyrax relative to the pig support the notion that facial retraction decreases resistance to masticatory forces in the superior portion of the face, but this effect is only moderate in the experiment. In both species, applied stresses are effectively generated in the rostrum and/or lower face, suggesting that a similar pattern may be inferred for the human face.

One important limitation of mechanical analyses with strain gages is the difficulty in directly estimating patterns of stress or loading in skeletal elements from strain measurements. Because of the inherent anisotropy in cortical bone, orientation of principal strains and stresses do not necessarily coincide, and it has been demonstrated theoretically that such differences may be as great as 45° (Cowin and Hart, 1990). Likewise, relative proportions of stress and strain magnitudes may differ. This investigation measured the elastic properties of a region of cortical bone on both the buccal and lingual surfaces of the lower border of the macaque mandible. The elastic property data was then combined with macaque mandibular strain data from published and a new in vivo strain gage experiment to determine directions and magnitudes of maximum and minimum principal stresses. The goal was to compare the stresses and strains and assess the differences in orientation and relative magnitude between them. The main question was whether these differences might lead to different interpretations of mandibular function. Elastic and shear moduli, and Poisson's ratios were measured using an ultrasonic technique from buccal and lingual cortical surfaces in 12 macaque mandibles. Mandibular strain gage data were taken from a published set of experiments (Hylander, 1979), and from a new experiment in which rosette strain gauges were fixed to the buccal and lingual cortices of the mandibular corpus of an adult female Macaca fascicularis, after which bone strain was recorded during mastication. Averaged elastic properties were combined with strain data to calculate an estimate of stresses in the mandibular corpus. The elastic properties were similar to those of the human mandibular cortex. Near its lower border, the macaque mandible was most stiff in a longitudinal direction, less stiff in an inferosuperior direction, and least stiff in a direction normal to the bone's surface. The lingual aspect of the mandible was slightly stiffer than the buccal aspect. Magnitudes of stresses calculated from average strains ranged from a compressive stress of - 16.00 GPa to a tensile stress of 8.84 GPa. The orientation of the principal stresses depended on whether the strain gage site was on the working or balancing side. On the balancing side of the mandibles, maximum principal stresses were oriented nearly perpendicular to the lower border of the mandible. On the working side of the mandibles, the orientation of the maximum principal stresses was more variable than on the balancing side, indicating a larger range of possible mechanisms of loading. Near the lower border of the mandible, differences between the orientation of stresses and strains were 12° or less. Compared to ratios between maximum and minimum strains, ratios between maximum and minimum stresses were more divergent from a ratio of 1.0. Results did not provide any major reinterpretations of mandibular function in macaques, but rather confirmed and extended existing work. The differences between stresses and strains on the balancing side of the mandible generally supported the view that during the power stroke the mandible was bent and slightly twisted both during mastication and transducer biting. The calculated stresses served to de-emphasize the relative importance of torsion. On the working side, the greater range of variability in the stress analysis compared to the strain analysis suggested that a more detailed examination of loadings and stress patterns in each individual experiment would be useful to interpret the results. Torsion was evident on the working side; but in a number of experiments, further information was needed to interpret other superimposed regional loading patterns, which may have included parasagittal bending and reverse parasagittal bending.

Major transformations in the skull and masticatory system characterized the evolution of crown anthropoids. To offer further insight into the phylogenetic and arguably adaptive significance of specific primate mandibular loading and kinematic patterns, allometric analyses of metric parameters linked to masticatory function are performed within and between 47 strepsirhine and 45 recent anthropoid species. When possible, basal anthropoids are considered. These results are subsequently integrated with prior experimental and morphological work on primate skull form. As compared to strepsirhines, crown anthropoids have a vertically longer ascending ramus linked to a glenoid and condyle positioned relatively higher above the occlusal plane. Interestingly, anthropoids and strepsirhines do not exhibit different mean ratios of condylar to glenoid height, which suggests that both clades are similar in their ability to evenly distribute occlusal contacts and perhaps forces along the postcanine teeth. Thus, given the considerable suborder differences in the scaling of both glenoid and condylar height, we argue that much of this variation in jaw-joint height is linked to suborder differences in relative facial height due in turn to increased encephalization, basicranial flexion, and facial kyphosis in anthropoids. Due to a more elongate ascending ramus, anthropoids evince more vertically oriented masseters than like-sized strepsirhines. Having a relatively longer ramus and a more medially displaced lateral pterygoid plate, crown anthropoids exhibit medial pterygoids oriented similar to those of strepsirhines, but with a variably longer lever arm. As anthropoid masseters are less advantageously placed to effect transverse movements/forces, we argue that balancing-side deep-masseter activity underlying a wishboning loading regime serves to increase, or at least maintain, transverse levels of jaw movement and occlusal force at the end of the masticatory power stroke. Crown anthropoids are also more isognathic and isodontic than strepsirhines. A consideration of early anthropoids suggests that the crown anthropoid masticatory pattern, i.e., more vertical masseters due to a high condyle as well as greater isognathy and isodonty, occurred stepwise during stem anthropoid evolution. This appears to correspond to a more transverse, and perhaps progressively larger, power stroke across oligopithecids, parapithecids, and propliopithecids.

In a previous study, we found systematic differences in the biomechanical behavior of modern human molars using finite element stress analyses (FESA), which led us to propose that molars are adapted to differently-directed loads depending on their position within the mouth (Spears and Macho (1998) Am. J. Phys. Anthropol. 106:467-482 ). While the FESA results thus derived have not been verified experimentally, such an interpretation seemed reasonable. To refine the model previously presented, this study assessed the effects of 1) food particle size on the biomechanical behavior of molars, and those of 2) differences in morphology, particularly enamel thickness, on stress distribution. In order to appraise the evolutionary significance of the findings, the FESA results for modern humans were subsequently compared with those obtained for molars of one individual of Pan and Pongo, respectively. Bearing in mind limitations imposed by the FESA models created and analyzed in this study, constant cleavage-type loads and cuspal tip loads at different directions were employed on all teeth: this facilitated comparisons of patterns of stress distribution across molars and species. In Pan and Homo, cleavage-type loads exerted by big food particles tended to be better dissipated anteriorly than posteriorly, although trends in Pongo were less clear-cut. Furthermore, similar to modern humans, the buccal cusps of mandibular molars appeared to be able to dissipate the loads associated with a pestle-type action, while maxillary molars were better designed to dissipate the loads which would result if they acted as mortars against which the food is crushed/ground. While increases in enamel thick- ness lowered the overall stress values in teeth only slightly, changes in outer morphology could have a more profound effect on these stress levels. Overall, Pan appeared to be most generalized, while Homo and Pongo showed a number of unique specializations, which are in accordance with what is currently understood about their respective masticatory apparatus and dietary niche. Am J Phys Anthropol 109:211-227, 1999. r 1999 Wiley-Liss, Inc.

It is well established that some observed patterns of force production in the primate masticatory system match those predicted by a simplified lever model. This model is also commonly invoked in adaptive explanations of craniodental diversity. However, systematic studies of the predictive power of this model are missing, leaving open the possibility that factors not traditionally included in the model alter the function and evolution of the masticatory system. One such factor was proposed for mammals generally by Greaves ([1978] J. Zool. (Lond.) 184:271–285), who argued that the temporomandibular joint (TMJ) was poorly suited to being pulled apart. In this constrained lever model, the avoidance of joint distraction leads to limitations on masticatory system form and function. The goal of the present study was to quantify masticatory system diversity in anthropoid primates for comparison with these predictions. Results indicate that all sampled taxa exhibit a form that is consistent with selection against regular distraction of the TMJ. Also apparent from observed patterns of scaling is a regular interaction among a limited set of cranial and dental dimensions, in accordance with the constrained model. However, the data indicate that specific positional relationships among the muscles, joints, and teeth differ from those predicted by Greaves (1978). The pattern of deviation suggests that selection has favored a conservative masticatory system configuration that safeguards the TMJ from distraction during the dynamic processing of irregular foods. The resulting buffered model leads to alternative hypotheses regarding the response of the masticatory system to dietary selection pressures. It may, therefore, improve our understanding of the adaptive significance of primate craniofacial form. Am J Phys Anthropol 108:483–506, 1999. © 1999 Wiley-Liss, Inc.

Comparative investigations of mandibular function among primates have relied upon elementary structural models to estimate states of masticatory stress and strain. In these studies, mandibular corpus morphology is idealized as a homogeneous, isotropic symmetrical body of invariant geometry, and this morphological abstraction is used to infer relative levels of stress and strain in the jaw. In reality, none of the limiting conditions assumed by these models is satisfied; consequently, it is prudent to ask whether this "textbook" engineering approach is valid for the inference of biomechanical behavior. In this study, the predictions of various geometric representations of the mandibular corpus are evaluated against strains recorded in a sample of human jaws loaded in torsion. Symmetrical geometrical models (including various "robusticity" shape indices), although convenient, are probably not consistently reliable for predicting the distribution of strains in the corpus. The experimental data suggest that variations in cortical thickness within sections play a significant role in determining the profile of relative strains. For comparative applications, characterization of the corpus as an asymmetrical hollow ellipse (i.e., with differing thickness of medial and lateral cortical plates) may provide a reasonable portrayal of relative strains.

Ten male Saimiri sciureus boliviensis (Cebidae), born at the Centro Argentino de Primates (CAPRIM), were grown in captivity. At weaning (6 months old), five individuals were fed ad libitum on a 20% protein diet (controls). The other five animals were fed ad libitum on a 5% protein diet (malnourished). Animals were radiographed monthly. The length, width, and height of the anterior, middle, and posterior components of the neurocranium, and those of the masticatory, respiratory, and optic components of the face were measured. A pattern of high growth rate was observed in both the three facial and the middle neural components. The anterior and posterior neural components showed a pattern of low rate of growth. The growth behavior of each variable was also different. Lengths grew more than widths and heights in the facial components, whereas widths grew more than heights and lengths in the neurocranium. Malnutrition delayed growth in size and altered the normal shape changes. High-patterned variables, such as masticatory and respiratory lengths, and the anterior and middle neural widths were particularly affected. The masticatory and the middle neural components underwent the greatest growth arrest. The optic and the respiratory components suffered a mild effect. The anterior and the posterior neural components were affected to a lesser degree.

It has been proposed that the mammalian facial skeleton is optimized for countering or dissipating masticatory stress. As optimized load-bearing structures by definition exhibit maximum strength with a minimum amount of material, this hypothesis predicts that during chewing and biting there should be relatively high and near uniform amounts of strain throughout the facial skeleton. If levels of strain in certain areas of the facial skeleton are relatively low during these behaviors, this indicates that the amount of bone mass in these areas could be significantly reduced without resulting in the danger of structural failure due to repeated masticatory loads. Furthermore, and by definition, this indicates that these areas are not optimized for countering masticatory stress, and instead their overall morphology and concentration of bone mass has most likely been selected or influenced mainly by factors unrelated to the dissipation or countering of chewing and biting forces. An analysis of in vivo bone strain along the lateral aspect of the zygomatic arch of macaques indicates the clear absence of a high and near uniform strain environment throughout its extent. Instead, there is a steep strain gradient along the zygomatic arch, with the highest strains along its anterior portion, intermediate strains along its middle portion, and the lowest strains along its posterior portion. These data, in combination with earlier published data (Hylander et al., 1991), indicate that levels of functional strains during chewing and biting are highly variable from one region of the face to the next, and therefore it is unlikely that all facial bones are especially designed so as to minimize bone tissue and maximize strength for countering masticatory loads. Thus, the functional significance of the morphology of certain facial bones need not necessarily bear any important or special relationship to routine and habitual cyclical mechanical loads associated with chewing or biting. Furthermore, the presence of these steep strain gradients within the facial skeleton suggests that the amount of bone mass in the low-strain areas may be largely determined by factors unrelated to processes frequently referred to as "functional adaptation," or conversely, that the "optimal strain environment" of bone varies enormously throughout the facial skeleton (cf., Rubin et al., 1994). Based solely on anatomical considerations, it is likely that the zygomatic arch is bent in both the parasagittal and transverse planes and twisted about its long axis. Due to constraints on rosette position, the strain data are incapable of determining if one or more of these loading conditions predominate. Instead, the strain data simply provide limited support for the possible presence of all of these loading regimes. Finally, as the masseter muscle is concentrated along the anterior portion of the zygomatic arch and as the arch has fixed ends, the largest shearing forces and the largest bending and twisting moments are located along its anterior portion. This in turn explains why the largest strains are found along the anterior portion of the zygomatic arch.

Cross-sectional geometric properties of the postcanine mandibular corpus are determined for the only known specimen of Otavipithecus namibiensis, a middle Miocene hominoid from southern Africa. It is shown that Otavipithecus is unique in that several important mechanical properties of its mandible, including maximum and minimum moments of inertia and distribution of cortical bone, differ from patterns seen in both extant hominoids and the early hominids Australopithecus africanus and Australopithecus (Paranthropus) robustus. This is particularly apparent in the mechanical design of the posterior portion of the mandibular corpus for resisting increased torsional and transverse bending moments. Cortical index values at the level of M2 also reveal that both Otavipithecus and A. africanus are similarly designed to resist increased masticatory loads with relatively less cortical bone area, a highly efficient mechanical design.

Subvertical grooves, located on the interproximal facets of most Neandertal posterior teeth, are less frequently noted on the teeth of other hominids, including modern humans. These grooves, 0.1–0.5 mm in width, are strictly localized within the facet area. Scanning electron microscopic (SEM) examination of grooves present on Neandertal teeth from Caverna delle Fate (Liguria, Italy) and Genay (Cote d'Or, France) demonstrated that they were produced during the life of these individuals. Characteristics of the groove surface suggest an erosion-abrasion mechanism of formation. These grooves, which developed in parts of the dentition exposed to marked stress, originated in areas characterized by changes in the orientation of enamel prism bundles (i.e., Hunter-Schreger bands). Observations carried out on modern human molars showed a subvertical disposition of these bands near interproximal ridges facilitating subvertical microfractures. Possible correlations between enamel structure, masticatory stress, and interproximal groove formation in Neandertals are discussed. © 1995 Wiley-Liss, Inc.

Behavioral observations of great apes have consistently identified differences in feeding behavior among species, and these differences have been linked to variation in masticatory form. As the point at which the mandible and cranium articulate, the temporomandibular joint (TMJ) is an important component of the masticatory apparatus. Forces are transmitted between the mandible and cranium via the TMJ, and this joint helps govern mandibular range of motion. This study examined the extent to which TMJ form covaries with feeding behavior in the great apes by testing a series of biomechanical hypotheses relating to specific components of joint shape using linear measurements extracted from three-dimensional coordinate data. Results of these analyses found that taxa differ significantly in TMJ shape, particularly in the mandibular fossa. Chimpanzees have relatively more anteroposteriorly elongated joint surfaces, whereas gorillas tend to have relatively anteroposteriorly compressed joints. Orangutans were most commonly intermediate in form between Pan and Gorilla, perhaps reflecting a trade-off between jaw gape and load resistance capabilities. Importantly, much of the observed variation among taxa reflects differences in morphologies that facilitate gape over force production. These data therefore continue to emphasize the unclear relationship between mandibular loading and bony morphology, but highlight the need for further data regarding food material properties, jaw gape, and ingestive/food processing behaviors.

The calvaria of an adult Australopithecus boisei from Area 104, Koobi Fora, Lake Turkana, is described. The specimen, KNM-ER 23000, comes from sediments dated to about 1.9 Ma. It consists of the frontal, both parietals, both temporals, most of the occipital as well as two small pieces of sphenoid, and a mandibular tooth root. The specimen is presumed to be an adult male, based on its size and the great development of features associated with the masticatory apparatus. KNM-ER 23000 is close in general size and shape to KNM-ER 406, KNM-ER 13750, and Olduvai Hominid 5 and it has a mixture of features seen in these three roughly contemporaneous crania. The frontal, especially the tori, resembles that of OH 5; the parietals are most like those of KNM-ER 13750; the occipital is like those of the two other Turkana specimens, and the temporals have a mixture of features from all of these, This specimen adds to our knowledge of variability in A. boisei. © 1993 Wiley-Liss, Inc.

The extent to which the mandibular corpus exhibits developmental plasticity has important implications for interpreting variation in adult and juvenile mandibular morphology in the archaeological and paleontological record. Here, we examine ontogenetic changes in mandibular corpus breadth, rigidity, and strength in two population samples with contrasting diets: late prehistoric Tigara from Point Hope, Alaska, characterized by a very demanding masticatory regime, and proto-historic Arikara from the Sully Site in South Dakota, with a less demanding regime. A total of 52 juvenile and 11 adult Tigara, and 32 juvenile and 10 adult Arikara were included in the study. Juveniles ranged in age from 1 to 17 years, with good representation of younger (1-6-year-old) juveniles (20 Arikara, 18 Tigara). Superoinferior and buccolingual external and cortical bone breadths of mandibles were measured at the Pm(4) -M(1) and M(1) -M(2) junctions using calipers and biplanar radiographs, respectively. An asymmetrical hollow beam model was employed to reconstruct cross sections and calculate bending rigidities and strengths in the sagittal and transverse planes. Among adults, Tigara have greater transverse corpus width, bending rigidity, and strength, and ratios of transverse to sagittal dimensions than Arikara. This shape difference develops gradually during growth, with only weak trends among young juveniles, increasing to near-adult contrasts among adolescents. These results support a role for functional mechanical loading of the mandible during growth in producing adult differences in mandibular corpus morphology.

Even though in vivo studies of mastication in living primates are often used to test functional and adaptive hypotheses explaining primate masticatory behavior, we currently have little data addressing how experimental procedures performed in the laboratory influence mastication. The obvious logistical issue in assessing how animal manipulation impacts feeding physiology reflects the difficulty in quantifying mechanical parameters without handling the animal. In this study, we measured chewing cycle duration as a mechanical variable that can be collected remotely to: 1) assess how experimental manipulations affect chewing speed in Cebus apella, 2) compare captive chewing cycle durations to that of wild conspecifics, and 3) document sources of variation (beyond experimental manipulation) impacting captive chewing cycle durations. We find that experimental manipulations do increase chewing cycle durations in C. apella by as much as 152 milliseconds (ms) on average. These slower chewing speeds are mainly an effect of anesthesia (and/or restraint), rather than electrode implantation or more invasive surgical procedures. Comparison of captive and wild C. apella suggest there is no novel effect of captivity on chewing speed, although this cannot unequivocally demonstrate that masticatory mechanics are similar in captive and wild individuals. Furthermore, we document significant differences in cycle durations due to inter-individual variation and food type, although duration did not always significantly correlate with mechanical properties of foods. We advocate that the significant reduction in chewing speed be considered as an appropriate qualification when applying the results of laboratory-based feeding studies to adaptive explanations of primate feeding behaviors. Am J Phys Anthropol, 2011. © 2011 Wiley-Liss, Inc.

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

The modern human mandibular symphysis differs from those of all other primates in being vertically orientated and possessing a chin, but the functional significance of this unique morphology is not well understood. Some hypotheses propose that it is an adaptation to specific loads occurring during masticatory function. This study uses finite element analysis to examine these symphyseal loads in a model of a modern human mandible. By modifying the symphyseal cross-sectional form, the mechanical significance of the presence of the chin and symphyseal orientation is tested, and modern human and Neanderthal symphyseal cross-sections are compared with regard to their ability to withstand different loads. The results show that changes in symphyseal form have profound effects on the strains. The presence of a chin leads to lower symphyseal strains overall, whereas a vertical orientation of the symphysis results in higher strains under wishboning, but not under vertical bending in the coronal plane and dorsoventral shear. Compared to Neanderthals, the modern human symphysis shows higher strains during dorsoventral shear and wishboning, but is as effective as the Neanderthal symphysis in resisting vertical bending in the coronal plane and the loads resulting from simulated incision and unilateral molar biting. In general, the results of this study corroborate prior hypotheses about the mechanical effects of the human chin and vertical symphyseal orientation and support the idea that the relative importance of wishboning and vertical bending in the coronal plane might have played a role in the evolution of modern human symphyseal morphology.

Understanding how the skull transmits and dissipates forces during feeding provides insights into the selective pressures that may have driven the evolution of primate skull morphology. Traditionally, researchers have interpreted masticatory biomechanics in terms of simple global loading regimes applied to sim- ple shapes (i.e., bending in sagittal and frontal planes, dorsoventral shear, and torsion of beams and cylinders). This study uses finite element analysis to examine the extent to which these geometric models provide accurate strain predictions in the face and evaluate whether sim- ple global loading regimes predict strains that approxi- mate the craniofacial deformation pattern observed dur- ing mastication. Loading regimes, including those simu- lating peak loads during molar chewing and those approximating the global loading regimes, were applied to a previously validated finite element model (FEM) of a macaque (Macaca fascicularis) skull, and the resulting strain patterns were compared. When simple global load- ing regimes are applied to the FEM, the resulting strains do not match those predicted by simple geometric models, suggesting that these models fail to generate accurate predictions of facial strain. Of the four loading regimes tested, bending in the frontal plane most closely approximates strain patterns in the circumorbital region and lateral face, apparently due to masseter muscle forces acting on the zygomatic arches. However, these results indicate that no single simple global loading re- gime satisfactorily accounts for the strain pattern found in the validated FEM. Instead, we propose that FE mod- els replace simple cranial models when interpreting bone strain data and formulating hypotheses about craniofa- cial biomechanics. Am J Phys Anthropol 145:1-10, 2011. V

The main purpose of this study is to test the hypothesis that as subjects chew with increasing levels of force, the ratio of the working- to balancing-side jaw-muscle force (W/B) decreases and begins to approach 1.0. We did this by analyzing relative masseter force in Macaca fascicularis using both strain gage and surface electromyographic (EMG) techniques. In addition, we also analyzed: 1) the relationship between jaw position using cineradiographic techniques and relative masseter force, 2) the timing differences between relative masseter force from the working and balancing sides, and 3) the loading and unloading characteristics of the masseter muscle. Our findings indicate that when macaques increase the amount of overall masticatory force during chewing, the W/B ratio for masseter force frequently (but not always) decreases and begins to approach 1.0. Therefore, our working hypothesis is not completely supported because the W/B ratio does not decrease with increasing levels of force in all subjects. The data also demonstrate timing differences in masseter force. During apple-skin mastication, the average peak masseter force on the working side occurs immediately at or slightly after the initial occurrence of maximum intercuspation, whereas the average peak masseter force on the balancing side occurs well before maximum intercuspation. On average, we found that peak force from the balancing-side masseter precedes the working-side masseter by about 26 msec. The greater the asynchrony between working- and balancing-side masseter force, the greater the difference in the relative magnitude of these forces. For example, in the subject with the greatest asynchrony, the balancing-side masseter had already fallen to about one-half of peak force when the working-side masseter reached peak force. Our data also indicate that the loading and unloading characteristics of the masseter differ between the working and balancing sides. Loading (from 50 to 100% of peak force) and unloading (from 100 to 50% of peak force) for the balancing-side masseter tends to be rather symmetrical. In contrast, the working-side masseter takes much longer to load from 50 to 100% of peak force than it does to unload from 100 to 50% of peak force. Finally, it takes on average about 35 msec for the working-side zygoma and 42 msec for the balancing-side zygoma to unload from 100 to 50% of peak force during apple-skin mastication, indicating that the unloading characteristics of the macaque masseter during mastication closely approximates its relaxation characteristics (as determined by muscle stimulation).

The relationship between mandibular form and biomechanical function is a topic of significant interest to morphologists and paleontologists alike. Several previous studies have examined the morphology of the mandible in gouging and nongouging primates as a means of understanding the anatomical correlates of this feeding behavior. The goal of the current study was to quantify the trabecular bone structure of the mandibular condyle of gouging and nongouging primates to assess the functional morphology of the jaw in these animals. High-resolution computed tomography scan data were collected from the mandibles of five adult common marmosets (Callithrix jacchus), saddle-back tamarins (Saguinus fuscicollis), and squirrel monkeys (Saimiri sciureus), respectively, and various three-dimensional morphometric parameters were measured from the condylar trabecular bone. No significant differences were found among the taxa for most trabecular bone structural features. Importantly, no mechanically significant parameters, such as bone volume fraction and degree of anisotropy, were found to vary significantly between gouging and nongouging primates. The lack of significant differences in mechanically relevant structural parameters among these three platyrrhine taxa may suggest that gouging as a habitual dietary behavior does not involve significantly higher loads on the mandibular condyle than other masticatory behaviors. Alternatively, the similarities in trabecular architecture across these three taxa may indicate that trabecular bone is relatively unimportant mechanically in the condyle of these primates and therefore is functionally uninformative. Am J Phys Anthropol, 2010. © 2009 Wiley-Liss, Inc.

The purpose of this study is to test various masticatory-stress hypotheses about the evolution and function of well-developed browridges of higher primates. This was done by measuring and analyzing patterns of in vivo bone strain recorded from three-element rosette strain gages bonded to the supraorbital region and to other portions of the bony face of Macaca fascicularis and Papio anubis during mastication and incision. The magnitude and direction of the principal strains recorded support Endo's hypothesis that the supraorbital region during mastication and incision is bent in the frontal plane (Endo, 1966). Our data do not, however, support his hypothesis that the supraorbital region is bent more during incision than during mastication. The data also demonstrate that overall levels of supraorbital strain are not larger in more prognathic subjects. Most importantly, the data indicate that the supraorbital region of nonhuman catarrhines is strained very little during mastication and incision. This indicates that there is much more supraorbital bone than is necessary both to counter masticatory loads and to provide an adequate safety factor to failure for these loads. This in turn suggests that the macaque and baboon browridges can be considerably reduced in size and still maintain these required structural characteristics. Thus, our experiments provide no support whatsoever for those hypotheses that directly link browridge morphology to masticatory stress (cf. Endo, 1966; Russell, 1983, 1985). A recent review of Endo's original work indicates that this latter statement is also true for humans (Picq and Hylander, 1989). We conclude, therefore, that there is no good reason to believe that enlarged browridges in living and/or fossil primates are structural adaptations to counter intense masticatory forces. The evolution of browridge morphology in primates is best explained on the basis of factors related to the position of the brain relative to the orbits (Moss and Young, 1960). When these structures are widely separated, as in gorillas, the large intervening space must be bridged with bone. In addition, enough bone must be present within the supraorbital and bridged regions to prevent structural failure due to non-masticatory external forces associated with highly active primates (e.g., accidental traumatic forces applied to the orbits and neurocranium). This requirement results in both pronounced browridges and in much more supraorbital bone than is necessary to counter routine cyclical stress during mastication and incision. This in turn explains why bone strains recorded from the supraorbital region are extremely small relative to other portions of the primate face during mastication and incision.

The extensive overlap of the temporal squama on the parietal in KNM-WT 17000 is restricted to a narrow segment of the arc of the squamosal suture. It appears as a long, narrow, posterosuperior extension onto the calvarial wall. The long axis of this extended strip is aligned with what appears to be the most hypertrophied section of the temporalis muscle fan. The substantial differences between the squamosal suture and associated masticatory elements of KNM-WT 17000 and those of Australopithecus boisei provide an opportunity to evaluate the various forces molding the anatomy of the squamosal suture of A. boisei. It is suggested that the extreme flare of the zygomatic arches in the latter accounts for these differences.

Cortical bone distribution of the anthropoid mandibular symphysis has been addressed in relation to mechanical stress generated by mastication. To examine whether or not bone mass and distribution patterns of the human mandibular symphysis could be interpreted as an example of functional adaptation, we compared the skeletal growth series of two populations, prehistoric Jomon, considered to represent a "robust" mandibular morphology associated with a presumed heavier masticatory load, and modern Japanese. Results showed that the adult Jomon symphysis possessed significantly greater bone mass and thicker cortical bone compared to the modern Japanese condition. However, the second moments of area did not differ significantly between the two, indicating comparable rigidity against bending. Furthermore, the Jomon mandibles of the infant to juvenile stages exhibited most of the adult characteristics, in both bone mass/distribution of the symphysis and in mandibular corpus/ramus morphologies. The present study also demonstrated the presence of a growth pattern of symphyseal cortical thickness, common to both the Jomon and the modern Japanese series. In both populations, subsequent to deciduous molar occlusion, cortical bone tends to be thickest at the inferolingual symphysis, at the location where the highest tensile stresses presumably occur during mastication. These findings suggest that the "robust" characteristics of the Jomon mandible are initially manifested early in development, and that the effect of mechanical stimulus to bone mass formation in the human symphysis is largely confined to a regulatory role during growth modeling.

Longitudinal studies of aboriginal children over a 20-year period have drawn attention to the wide variation in morphological features of the dentition and the way in which occlusal relationships develop. This paper summarizes some important determinants of optimal occlusal development, namely, tooth size relationships within and between dentitions, the patterns of alveolar growth, and tooth migrations during the transition from primary to permanent teeth and the nature of growth changes in the dental arches. Dental occlusion constantly changes throughout life in response to changing functional requirements. Observations limited to cross-sectional material provide an incomplete, and sometimes misleading, concept of dental occlusion and masticatory function.

Twenty weanling 6-month-old male squirrel monkeys were allotted to the following treatments: 1) first control animals were killed at weaning; 2) second control animals were killed when 24 months old; and 3) malnourished animals were fed on a low-protein diet and killed at age 24 months. Lateral and vertical teleradiographies were taken. Growth of the neurocranial and splanchnocranial components were measured by volumetric (size estimators) and morphometric (shape estimators) indices. All facial components grew. The neurocranial components showed a heterogeneous behavior: The anteroneural component remained stable, and the increase of the midneural component was compensated by a decrease in the posteroneural component. Malnutrition affected the growths of 1) the craniofacial complex, 2) the splanchnocranium, and 3) the respiratory and midneural components. Growth influenced skull shape through 1) increases of the splanchnocranium and the midneural component relative to the neurocranium; 2) decreases of the masticatory and optic components relative to the splanchnocranium, and 3) decreases of the anteroneural and posteroneural components relative to the neurocranium. Malnutrition influenced skull shape through the relationship between the anteroneural component and the neurocranium. These results confirmed the existence of functional interrelationships among the cranial components. A new approach to craniological studies is suggested.

A post-Pleistocene reduction trend in the dimensions of the masticatory complex followed the transition to agricultural lifestyle in several world regions. A major limitation of previous studies is large temporal gaps between the analyzed skeletal populations, which do not allow the detection and analysis of a diachronic morphological transition. In this work, we analyze a large number of specimens from the southern Levant, where agriculture first emerged in situ and for which there is a good diachronic sequence of the shift from a hunting-gathering way of life to a food producing, farming economy (12,000–7,000 uncalibrated bp). Changes in the masticatory complex are examined in the context of three prevailing dental reduction models: the Probable Mutation Effect (Brace,1963; Brace and Mahler,1971), Increasing Population Density Effect (Macchiarelli and Bondioli,1986) and Selective Compromise Effect (SCE) (Calcagno,1989). A series of linear regressions of dimension vs. time and coefficients of variation for each dimension are analyzed. Our results indicate significant reduction in the buccolingual but not mesiodistal dental dimensions and in the ramus breadth and anterior height dimensions of the mandible but not in its overall size. These findings, taken together with low coefficients of variation for the buccolingual dimensions, suggest selective pressure resulting in reduction of specific dimensions. The observed trend is in partial accordance with the SCE but differs from the trends observed in other regions, and is therefore best explained as a region-specific variant of the SCE. Am J Phys Anthropol, 2008. © 2007 Wiley-Liss, Inc.

In comparative studies of masticatory function and mandibular biomechanics, the mediolateral dimension of the postcanine corpus (corpus breadth) is commonly utilized as a measure of torsional stiffness from which relative torsional strength is inferred. The use of this dimension entails certain assumptions about corpus shape and cortical bone distribution that are invalid. When corpus breadth is related to an appropriate, empirically supported measure of torsional strength, it is revealed that this dimension has limited utility for inference of biomechanical competence under torsion. The use of linear dimensions to infer structural adaptations to specific loading regimes is problematic given that bone tissue is not optimally deployed to minimize strain levels arising from isolated loads. For the inference of the masticatory biomechanical environment, the more reasonable approach is to consider overall size of the corpus (i.e., cross-sectional area) for inference of intra- and inter-specific differences in masticatory forces. Am J Phys Anthropol, 2007 © 2006 Wiley-Liss, Inc.

It was proposed that the power stroke in primates has two distinct periods of occlusal contact, each with a characteristic motion of the mandibular molars relative to the maxillary molars. The two movements are called phase I and phase II, and they occur sequentially in that order (Kay and Hiiemae [1974] Am J. Phys. Anthropol. 40:227-256, Kay and Hiiemae [1974] Prosimian Biology, Pittsburgh: University of Pittsburgh Press, p. 501-530). Phase I movement is said to be associated with shearing along a series of crests, producing planar phase I facets and crushing on surfaces on the basins of the molars. Phase I terminates in centric occlusion. Phase II movement is said to be associated with grinding along the same surfaces that were used for crushing at the termination of phase I. Hylander et al. ([1987] Am J. Phys. Anthropol. 72:287-312; see also Hiiemae [1984] Food Acquisition and Processing, London: Academic Press, p. 257-281; Hylander and Crompton [1980] Am J. Phys. Anthropol. 52:239-251, [1986] Arch. Oral. Biol. 31:841-848) analyzed data on macaques and suggested that phase II movement may not be nearly as significant for food breakdown as phase I movement simply because, based on the magnitude of mandibular bone strain patterns, adductor muscle and occlusal forces are likely negligible during movement out of centric occlusion. Our goal is to better understand the functional significance of phase II movement within the broader context of masticatory kinematics during the power stroke. We analyze vertical and transverse mandibular motion and relative activity of the masseter and temporalis muscles during phase I and II movements in Papio anubis. We test whether significant muscle activity and, by inference, occlusal force occurs during phase II movement. We find that during phase II movement, there is negligible force developed in the superficial and deep masseter and the anterior and posterior temporalis muscles. Furthermore, mandibular movements are small during phase II compared to phase I. These results suggest that grinding during phase II movement is of minimal importance for food breakdown, and that most food breakdown on phase II facets occurs primarily at the end of phase I movement (i.e., crushing during phase I movement). We note, however, that depending on the orientation of phase I facets, significant grinding also occurs along phase I facets during phase I.

Tooth to denture base discrepancy (the discrepancy) is the difference between the dental arch length and the sum of crown diameters of teeth in the jaw, a concept which was originally developed in orthodontics. Since the cause-effect relationship between a soft diet and the discrepancy has been demonstrated, the size of the discrepancy should indicate the amount of load on the masticatory system from chewing foods in jaws from archaeological periods. The dietary condition of 71 citizens compared to that of 186 slaves from the Yin-Shang period of China was reconstructed through a study of the discrepancy. The prevalence of the discrepancy in the Yin-Shang period was around 15%, almost the same as it was during the later Jomon to Yayoi (3000–2000 BP) periods, when rice agriculture was introduced into Japan, and also the same as for present-day pastoralists around Lake Turkana, Kenya. Although the frequency of the discrepancy was slightly higher in male citizens, there were no significant differences in the frequencies between male citizens and female citizens or slaves. The differences in diet may not have been fundamental since the Yin-Shang period would be at the very beginning of the age in which differences of diet according social class began to appear, with implications for the load on the masticatory system. At that time agriculture may not been sufficiently intensified in variety or quantity to have produced a differentiation of the diet between social classes. Am J Phys Anthropol 103:497–505, 1997. © 1997 Wiley-Liss, Inc.

An analysis of in vivo bone strain indicates that the mandibular symphysis of macaques experiences lateral transverse bending or "wishboning" during the power stroke of mastication, and this loading regime results in relatively intense concentrations of stress along the lingual aspect of the symphysis (Hylander 1984, 1985). It has been hypothesized that peak wishboning of the macaque mandible, which probably occurs at the very end of the power stroke, that is, after the initial occurrence of maximum intercuspation, is associated with the late peak activity of the balancing-side deep masseter muscle coupled with the rapid decline in the activity of the balancing-side medial pterygoid and superficial masseter muscles (Hylander et al., 1987). The main purpose of this study is to do a detailed analysis of the electromyographic (EMG) activity of the deep and superficial masseter and medial pterygoid muscles so as to provide a better understanding of the external forces associated with wishboning. This was done by recording and analyzing EMG activity from the superficial masseter, deep masseter, and medial pterygoid muscles in macaques and baboons. EMG activity was recorded from bipolar fine-wire electrodes, and the data were quantified and analyzed using digital techniques. The EMG data clearly support our original observation that the balancing-side posterior deep masseter exhibits peak EMG activity relatively late in the power stroke at a time when activity is rapidly decreasing in the balancing-side medial pterygoid and superficial masseter muscles. Moreover, peak activity of the balancing-side deep masseter occurs at a time when the activity of the working-side deep and superficial masseter and medial pterygoid muscles are also decreasing. Thus, these data are consistent with the hypothesis that wishboning of the mandible is influenced significantly by the late occurrence of force from the balancing-side deep masseter. The EMG data also indicate that residual force from the relaxing superficial masseter may contribute significantly to wishboning. Finally, patterns of wishboning are not fully accounted for by our EMG analysis of the deep and superficial masseter and medial pterygoid muscles. This is probably because the lateral ptergyoids, which can either counter increased wishboning or actually cause reverse wishboning, were not included in our EMG analysis. The EMG data demonstrate that jaw-closing muscle recruitment patterns for macaques and baboons differ from those of humans. Nevertheless, in spite of these differences, it appears, based on previously published EMG data, that the human symphysis may also experience wishboning.(ABSTRACT TRUNCATED AT 400 WORDS)