The functional significance of primate mandibular form.

A stress analysis of the primate mandible suggests that vertically deep jaws in the molar region are usually an adaptation to counter increased sagittal bending stress about the balancing-side mandibular corpus during unilateral mastication. This increased bending stress about the balancing side is caused by an increase in the amount of balancing-side muscle force. Furthermore, this increased muscle force will also cause an increase in dorso-ventral shear stress along the mandibular symphysis. Since increased symphyseal stress can be countered by symphyseal fusion and as increased bending stress can be countered by a deeper jaw, deep jaws and symphyseal fusion are often part of the same functional pattern. In some primates (e.g., Cercocebus albigena), deep jaws are an adaptation to counter bending in the sagittal plane during powerful incisor biting, rather than during unilateral mastication. The stress analysis of the primate mandible also suggests that jaws which are transversely thick in the molar region are an adaptation to counter increased torsion about the long axis of the working-side mandibular corpus during unilateral mastication. Increased torsion of the mandibular corpus can be caused by an increase in masticatory muscle force, an increase in the transverse component of the postcanine bite force and/or an increase in premolar use during mastication. Patterns of masticatory muscle force were estimated for galagos and macaques, demonstrating that the ratio of working-side muscle force to balancing-side muscle force is approximately 1.5:1 in macaques and 3.5:1 in galagos during unilateral isometric molar biting. These data support the hypothesis that mandibular symphyseal fusion is an adaptative response to maximize unilateral molar bite force by utilizing a greater percentage of balancing-side muscle force.     

Allometric relations of teeth and jaws in man

Static adult intraspecific allometry of jaws and teeth was investigated in a sample of 100 Negro crania. The relations between tooth area, postcanine surface, incisor surface, and four viscerocranial measures were examined separately for males and females. Our results indicate a marked lack of morphological integration between P-sets within the orofacial subregion and a similar lack of correspondence between jaw size and tooth size. Allometric analyses indicate that mandibular length scales negatively allometric to maxilloalveolar length and to bigonial width, that canine base area scales positively to upper and lower jaw length, and that all the other teeth scale negatively to jaw length. The postcanine surface area was found to be negatively allometric to the canine base area, which in turn scaled isometrically to incisor surface. Hence, any lengthening of the mandible will tend to be associated with a relative shortening of the maxilla, with relatively larger canines and a relative reduction of the cheek teeth.     

Static intraspecific allometry of jaws and teeth in Cercopithecus aethiops

Adult static intraspecific allometry of jaw size and tooth area was evaluated in a sample of 100 Cercopithecus aethiops crania (50 male, 50 female). Tooth areas were calculated from mesiodistal and buccolingual measurements of all the teeth in both arcades and were scaled to four viscero-cranial measurements: bimaxillary breadth, maxillo-alveolar length, mandibular length and bigonial width. Allometric coefficients calculated for jaw dimensions alone indicate tighter viscerocranial integration in females than in males. A finding of note was that half of the variation in maxillo-alveolar length may be accounted for by variation in mandibular length: females are isometric, males negatively allometric.
A similar degree of allometric mosaicism was found when maxillary incisor size was scaled to maxillary length and width. In females, the relationship was negatively allometric, whilst incisor size in males was found to be unrelated to either. Negative allometry characterized the relationship of canine base area to jaw length in both sexes, with males additionally being positively allometric to mandibular width.
The scaling of postcanine tooth areas to jaw length was characterized by a dichotomous pattern: males showed significant mandibular integration whilst females showed only significant maxillary integration. Compensatory tooth size interaction between maxillary canine base area and the summed incisor and postcanine areas was suggested by the significant negative allometric relation between them.     

The mandibular corpus of female primates: taxonomic, dietary, and allometric correlates of interspecific variations in size and shape

Measurements were taken on skulls of 253 adult female anthropoid primates from 32 species, in order to determine patterns and possible causes for variation among species in the cross-sectional size and shape of the mandibular corpus under M1. When all 32 species are considered as a group, there is a tendency for corpus shape to become more robust with increasing body size. However, this does not hold for colobines or cercopithecines evaluated separately. When diets are classified into the general categories of folivory or frugivory, neither size-adjusted measurements of mandibular corpus breadth and height, nor estimates of the second moments of inertia or the polar moment of inertia of the mandibular cross section, show any relationship to dietary variation among species. Species reported to include hard nuts in their diets have larger mandibular cross sections than other species, and the size of the corpus is significantly correlated with size of the dentition and molar enamel thickness. A biomechanical model taking into account frictional effects of tooth-to-tooth contact indicates that mandibular corpus robusticity may not be related to a large horizontal component of force during mastication.     

Static intraspecific maxillofacial allometry in the chacma baboon

Adult static intraspecific allometry of jaw size and tooth area was evaluated in a sample of 104 Papio ursinus crania (52 male, 52 female). Tooth areas were calculated from mesiodistal and buccolingual measurements of all the teeth in both arcades and were scaled to four viscerocranial measurements: bimaxillary width, maxillo-alveolar length, mandibular length and bigonial width. Craniodental allometric analyses indicate that larger animals will tend to have proportionately shorter and narrower lower jaws. From the log-transformed interspecific analyses between P. ursinus and C. aethiops we conclude that males and females within each species share a common exponential value for jaw length. Hence increased sexual dimorphism for muzzle length in P. ursinus is attributable to increased divergence between the male and female slopes. Post-canine area was found to be significantly correlated to maxillary length and to canine size only in females, with exponential values similar to those reported for the same bivariate regressions in C. aethiops. A hypothesis of nutritional equivalence is advanced to account for these observations. Canine base area and the area of P3 were the only tooth areas that scaled in a positively allometric fashion to jaw size--but only in males. Hence the existence of a canine complex is confirmed in the male Chacma baboon, the size of which is related to jaw length.     

Mandibular morphology and diet in the genusCebus

The influence of hard-object feeding on the size and shape of the mandibular corpus was investigated through a comparative biomechanical analysis of the jaws of adult femaleCebus apella andCebus capucinus. Computed tomography (CT) was used to discern the amount and distribution of cortical bone at M₂ and symphyseal cross sections. From these data, the biomechanical properties of the mandibular corpus were determined to assess the structural rigidity of the jaw with respect to the bending, torsional, and shear stresses that occur during mastication and incision. The mandibles ofC. apella are demonstrably more robust than those ofC. capucinus in terms of biomechanical rigidity; differences in corporeal size rather than shape largely account for the enhanced robusticity in the sample ofC. apella. The differences that separate the two taxa probably represent a structural response to the mechanical demands of durophagy inC. apella. These observations suggest that specialization on a diet of hard objects may be expected to result in an overall hypertrophy of bony contours throughout the mandibular corpus.     

Interspecies difference in placement of developing teeth and its relationship with cross-sectional geometry of the mandibular symphysis in four primate species including modern humans

The form of the anthropoid mandibular symphysis has recently been addressed in association with spatial requirements for the forming anterior teeth. To evaluate potential relationships between the symphyseal shape and teeth further, the growth patterns of the symphyseal region and the positioning of the tooth crypts were examined using CT data, comparing four primate species (modern humans, chimpanzees, Japanese monkeys, and hamadryas baboons) with varied symphyseal curvature and tooth size. First, results showed that interspecies differences in overall mandibular shape including symphyseal inclination and bicanine width are consistently expressed throughout postnatal ontogeny, although local symphyseal configurations related to the superior transverse torus (STT) tended to change considerably during growth in chimpanzees. Second, the four species were found to exhibit differentiated formation positions of the incisor and canine crypts. In particular, I2 developed between I1 and C in humans with a broad bicanine space and small teeth, whereas it was positioned posterior to I1 and above C in the cercopithecines with an extremely narrow bicanine space. In chimpanzees, despite the large bicanine width, I1 and I2 grew with a large antero-posterior overlap owing to their large size. These results indicate that the dental positioning is determined in concert with the size balance of the available mandibular space and forming teeth. Finally, the positions/contours of I2 crypt were shown to correspond strongly with the STT across the taxa. This suggests that interspecies differences in symphyseal shape should be interpreted partially by the species-specific positional relationships of the developing anterior teeth.     

Mandibular form and function in North American and European Adapidae and Omomyidae

Previous experimental and comparative studies among a wide variety of primate and nonprimate mammals provide a unique source of information for investigating the functional and phylogenetic significance of variation in the masticatory apparatus of Eocene primates. To provide a quantitative study of mandibular form and function in Eocene primates, the scaling of jaw dimensions and the development of symphyseal fusion was considered in a broad sample of North American and European Adapidae and Omomyidae. Statistical analyses indicate a significant size-related pattern of symphyseal fusion across Eocene primates, with larger taxa often having a greater degree of fusion than smaller species; this trend is also evident at the family level. As adapids are mostly larger than omomyids and these taxa show allometry of symphyseal fusion, this may explain why no omomyids evince complete fusion. Controlling for jaw size, species with greater symphyseal fusion tend to have more robust jaws than those with a lesser amount of fusion. Upon further examination, a primary reason why adapids have more robust mandibles than omomyids is associated with the presence of taxa with fused symphyses, and thus more robust jaws, in the adapid sample, whereas no omomyids have fused symphyses. In addition, there is little indication of a dietary effect, as measured by molar shear-crest development, on symphyseal fusion. Moreover, as there is no correlation between molar shear-crest development and skull size, this also points to the absence of a size-related pattern of dietary preference underlying the allometry of symphyseal fusion. Based on the interspecific and ontogenetic allometry of symphyseal ossification in Eocene primates, jaw-scaling patterns are used to further examine the functional determinants of fusion in this group. This study indicates that greater dorsoventral shear during mastication is a more likely factor than lateral transverse bending (“wishboning”) in the evolution of symphyseal fusion among “late-fusing” mammals like adapids and omomyids. Given that wishboning is an important functional determinant of symphyseal form in recent anthropoids, apparently the evolutionary development of marked wishboning occurs only in taxa that shift the timing of fusion to a growth stage preceding the onset of weaning (before adult masticatory patterns are fully developed) and perhaps first ossified the symphysis to counter elevated dorsoventral shear stress. As early anthropoids probably consisted of members varying interspecifically and ontogenetically in the degree of ossification, it is especially informative to analyze the adaptive setting in which anthropoid symphyseal fusion evolved from a similar primitive “prosimian” perspective. © 1996 Wiley-Liss, Inc.     

Tooth size and arcadal length correlates in man

Intra-arcadal mesiodistal and buccolingual tooth size correlations were evaluated in a sample of 125 caucasoids with ideal occlusion. Dental dimensions were corrected for arcade mength (as a measure of jaw size) by a series of regression analyses of each mesiodistal dimension on the sum of the mesiodistal dimensions within each arcade. Regression coefficients of tooth dimension on arcade length were calculated to gain an insight into the dimensional sensitivity of individual teeth to arcade length variation. The data presented here suggest a strong association between arcadal length (jaw size) dependence, and the dimensional stability of individual teeth. When corrected for arcade length, a definite pattern of tooth size correlation emerges: postcanine maxillary and mandibular teeth are negatively correlated to the anterior teeth and are positively correlated to one another. The hypothesis is developed that anterior and postcanine teeth should be viewed as two separate and negatively size-correlated units, beyond the boundaries of the four morphological tooth classes. Recognition of this basic dichotomous size arrangement within each jaw allows for a reassessment of some of the problems associated with hominid dental evolution.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.     

A preliminary analysis of correlations between chewing motor patterns and mandibular morphology across mammals

The establishment of a publicly-accessible repository of physiological data on feeding in mammals, the Feeding Experiments End-user Database (FEED), along with improvements in reconstruction of mammalian phylogeny, significantly improves our ability to address long-standing questions about the evolution of mammalian feeding. In this study, we use comparative phylogenetic methods to examine correlations between jaw robusticity and both the relative recruitment and the relative time of peak activity for the superficial masseter, deep masseter, and temporalis muscles across 19 mammalian species from six orders. We find little evidence for a relationship between jaw robusticity and electromyographic (EMG) activity for either the superficial masseter or temporalis muscles across mammals. We hypothesize that future analyses may identify significant associations between these physiological and morphological variables within subgroups of mammals that share similar diets, feeding behaviors, and/or phylogenetic histories. Alternatively, the relative peak recruitment and timing of the balancing-side (i.e., non-chewing-side) deep masseter muscle (BDM) is significantly negatively correlated with the relative area of the mandibular symphysis across our mammalian sample. This relationship exists despite BDM activity being associated with different loading regimes in the symphyses of primates compared to ungulates, suggesting a basic association between magnitude of symphyseal loads and symphyseal area among these mammals. Because our sample primarily represents mammals that use significant transverse movements during chewing, future research should address whether the correlations between BDM activity and symphyseal morphology characterize all mammals or should be restricted to this "transverse chewing" group. Finally, the significant correlations observed in this study suggest that physiological parameters are an integrated and evolving component of feeding across mammals.     

Compact bone distribution and biomechanics of early hominid mandibles.

This investigation explores the effects of compact bone distribution on the biomechanical properties of the postcanine mandibular corpus of the fossil hominid taxa Australopithecus africanus and Paranthropus robustus. The mandibles of extant great apes, modern humans, and the fossil hominids are examined by computed tomography (CT), and compact bone contours are used to calculate cross-sectional biomechanical properties (cortical area, second moments of area, and Bredt's formula for torsional strength). The relative amount of compact bone is comparable in the modern and fossil mandibles, but the mechanical properties of A. africanus and P. robustus jaws are distinct in terms of the ratio of minimum to maximum second moments of area. This difference most likely represents a structural response to elevated torsional moments in the fossil hominids. Although the relative amount of compact bone in cross-section does not differ significantly between taxa by statistical criteria, A. africanus utilizes less cortical bone than P. robustus in the same manner in which Pongo is separated from the condition in other extant large-bodied hominoids. It has been suggested that the phenomenon of mandibular "robusticity" (expressed as an index of corpus breadth/corpus height) may be an effect of postcanine megadontia and/or reduced canine size in the australopithecines. Results presented here, however, indicate that it is unlikely that either factor adequately accounts for mandibular size and shape variation in early hominids.     

Relationship between canine dimorphism and mandibular morphology in the hamadryas baboon and the Japanese monkey

To examine morphological interrelationships between canine size and mandibular corpus shape, inter-sex comparisons were made in the hamadryas baboon and the Japanese monkey, known to display extreme and moderate canine dimorphism, respectively. Results of adult comparisons showed that all mandibular dimensions were significantly larger in the males than in the females in both species. In the hamadryas baboon, the males also exhibited a higher ratio of anterior to posterior corpus height than the females. This sex difference in corpus shape was not significant in the Japanese monkey, indicating lack of involvement of canine dimorphism. Analysis of mandibular growth patterns in the hamadryas baboon demonstrated that significant sexual size difference did not occur before incisor eruption, and that the anteriorly high corpus of the adult male mandible was associated with a rapidly increasing symphyseal height after incisor eruption. It was also shown that the female canine started to erupt shortly after incisor eruption, while the forming male canine continued to stay near the mandibular base and developed further in size until eruption. The relative positions of the incisors kept shifting upwards even after eruption in the males, while they hardly changed in the females. It is therefore suggested that the prolonged development and size increase of the male canine is accompanied by further enlargement of the symphysis, resulting in the higher anterior corpus of the adult males compared to the adult females. The results thus indicate the importance of understanding the spatial relationships of the developing teeth in interpreting mandibular morphology.     

Population variation in tooth, jaw, and root size: a radiographic study of two populations in a high-attrition environment

Radiographs were taken of the jaws of skeletal remains of two populations of different-phenotype Prehistoric Australians from Roonka and Early New Zealanders (Maoris). On these radiographs crown, root, and corpus size were measured. Corpus height was subdivided into alveolar bone height, defined as the bone superior to the mandibular canal, and basal bone height, defined as that inferior to the mandibular canal. Both between and within the two populations there was a significant and negative correlation between crown size and corpus height. The differences between the two populations in corpus height were associated with differences in alveolar bone height rather than basal bone height and support hypotheses associating continued eruption of adult teeth with growth of the alveolar bone. The findings also support previous studies that have shown only a low correlation between crown size, root size, and corpus height.     

The functional morphology of the anterior masticatory apparatus in tree‐gouging marmosets (cebidae,primates)

Although all genera of Callitrichinae feed on tree exudates, marmosets (Callithrix and Cebuella) use specialized anterior teeth to gouge holes in trees and actively stimulate exudate flow. Behavioral studies demonstrate that marmosets use large jaw gapes but do not appear to generate large bite forces (relative to maximal ability) during gouging. Nonetheless, the anterior teeth of marmosets likely experience different loads during gouging compared to nongouging platyrrhines. We use histological data from sectioned teeth, μCTs of jaws and teeth, and in vitro tests of symphyseal strength to compare the anterior masticatory apparatus in Callithrix to nongouging tamarins (Saguinus) and other cebids. We test the hypotheses that (1) marmoset anterior teeth are adapted to accommodate relatively high stresses linked to dissipating gouging forces and (2) the mandibular symphysis does not provide increased load resistance ability compared with closely related nongouging platyrrhines. Differences in decussation between Callithrix and Saguinus are greatest in the anterior teeth, suggesting an increased load resistance ability specifically in incisor and canine enamel of Callithrix. Callithrix lower incisor crowns are labiolingually thicker suggesting increased bending resistance in this plane and improved wedging ability compared with Saguinus. Anterior tooth roots are larger relative to symphyseal bone volume in Callithrix. Anterior tooth root surface areas also are larger in marmosets for their symphyseal volume, but it remains unclear whether this relative increase is an adaptation for dissipating dental stresses versus a growth‐related byproduct of relatively elongated incisors. Finally, simulated jaw loading suggests a reduced ability to withstand external forces in the Callithrix symphysis. The contrast between increased load resistance ability in the anterior dentition versus relatively reduced symphyseal strength (1) suggests a complex loading environment during gouging, (2) highlights the possibility of distinct loading patterns in the anterior teeth versus the symphysis, and (3) points to a potential mosaic pattern of dentofacial adaptations to tree gouging. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.     

Arch form,tooth size,and occlusomandibular kinesis in the Ceboidea

Correlations between dental morphology, arch configuration, and jaw movement patterns were quantitatively investigated in 23 ceboid species to elucidate integrative aspects of occlusal functional anatomy in an adaptive and evolutionary context. Differential maxillary-mandibular arch widths are primary in guiding lateral jaw movements. These movements are characterized according to their associated condylar shifts as either predominantly translatory or rotational. Predominantly translatory movements result from peripheral contact relationships between maxillary arches which are considerably wider posteriorly than their opposing mandibular arches. The greatest degree of mandibular movement is in the molar region in functional association with wide “primitive” maxillary molars, narrow mandibular molars, constricted maxillary intercanine widths, and narrow maxillary incisors. In contrast, predominantly rotational masticatory jaw movements result from differential arch widths which are greatest in the maxillary canine and incisor regions. Here most jaw movement is in the anterior segment and this is reflected in small maxillary-mandibular molar width differences, a high degree of premolarization, wide-set maxillary canine teeth, and wide maxillary incisors. Possible selectional factors in the putative evolution of rotational predominance in mastication from the more primitive translatory pattern are discussed.     

Craniofacial variation and dietary adaptations of African colobines

African colobine monkeys show considerable craniofacial variation among species, although the evolutionary causes of this diversity are unclear. In light of growing evidence that diet varies considerably among colobine species, we investigated whether colobine craniofacial morphology varies as a function of their diet. We compared craniofacial morphology among five African species: Colobus angolensis, C. guereza, C. polykomos, Piliocolobus badius, and P. verus. Matrix correlation analysis indicated a significant correlation between species-specific morphological distance and dietary distance matrices. The mechanical advantage of the masseter muscle was higher in seed-eaters (C. angolensis and C. polykomos) and lower in those that eat mainly young leaves (C. guereza, P. badius, and P. verus). Canonical correspondence analysis revealed that the durophagous colobines possess relatively wider bigonial breadths, anteroposteriorly shorter faces, shorter postcanine tooth rows, more medially positioned dental batteries, wider bizygomatic arches, and anteroposteriorly longer zygomatic arches. Under the constrained lever model, these morphological features suggest that durophagous colobines have the capacity to generate relatively greater maximum bite forces. However, no consistent relationship was observed between diet and variation in the mandibular corpus and symphysis, implying that robust mandibles are not necessarily adaptations for stress resistance. Factors that may influence mandibular robusticity include allometry of symphyseal curvature and canine tooth support. Finally, linear measures of mandibular robusticity may suffer from error.     

Size and scaling in the mandible of living and extinct apes

The purpose of this study is to fill a gap in our knowledge of dietary and allometric determinants of masticatory function and mandibular morphology in major catarrhine clades. To extend the implications of previous work on variation in mandibular form and function in other primates, a scaling analysis was performed on 20 extinct and 7 living non-cercopithecoid catarrhines or 'dental apes'. Results of allometric comparisons indicate that for a given jaw length, larger apes exhibit significantly more robust corpora and symphyses than smaller forms. This appears linked to size-related increases in dietary toughness and/or hardness, which in turn causes elevated mandibular loads and/or greater repetitive loading during unilateral mastication. Larger-bodied dental apes also display more curved symphyses, which also explains the positive allometry of symphysis width and height. In apes, proconsulids often evince more robust jaws while all hylobatids, Pan and Dryopithecus laietanus possess more gracile cross sections. In propliopithecids, Aegyptopithecus is always more robust than Propliopithecus. In proconsulids, Rangwapithecus and Micropithecus commonly exhibit more robust jaws whereas Dendropithecus and especially Simiolus are more gracile. Most of the larger taxa are folivorous and/or hard-object frugivorous pongids with relatively larger dentaries. Though apes have relatively wider corpora than cercopithecines due to greater axial twisting of the corpora during chewing, they are otherwise alike in robusticity levels. Smaller apes are similar to cercopithecines in evincing a relatively high degree of symphyseal curvature, while larger taxa are like colobines in having less curvature. Larger pongids resemble or even exceed colobine jaw proportions and thus appear to converge on colobines in terms of the mechanical properties of their diets.     

Protective buttressing of the hominin face

When humans fight hand‐to‐hand the face is usually the primary target and the bones that suffer the highest rates of fracture are the parts of the skull that exhibit the greatest increase in robusticity during the evolution of basal hominins. These bones are also the most sexually dimorphic parts of the skull in both australopiths and humans. In this review, we suggest that many of the facial features that characterize early hominins evolved to protect the face from injury during fighting with fists. Specifically, the trend towards a more orthognathic face; the bunodont form and expansion of the postcanine teeth; the increased robusticity of the orbit; the increased robusticity of the masticatory system, including the mandibular corpus and condyle, zygoma, and anterior pillars of the maxilla; and the enlarged jaw adductor musculature are traits that may represent protective buttressing of the face. If the protective buttressing hypothesis is correct, the primary differences in the face of robust versus gracile australopiths may be more a function of differences in mating system than differences in diet as is generally assumed. In this scenario, the evolution of reduced facial robusticity in Homo is associated with the evolution of reduced strength of the upper body and, therefore, with reduced striking power. The protective buttressing hypothesis provides a functional explanation for the puzzling observation that although humans do not fight by biting our species exhibits pronounced sexual dimorphism in the strength and power of the jaw and neck musculature. The protective buttressing hypothesis is also consistent with observations that modern humans can accurately assess a male's strength and fighting ability from facial shape and voice quality.