Experimental stress analysis of topographic diversity in early hominid gnathic morphology

Reconstructing the biomechanics of early hominid mastication is a key element in most models of hominid differentiation. Traditionally, ostelogical features marking muscle attachment surfaces have served as a reference system from which the vector geometry of the masticatory force system and resultant force distributions could be predicted. To augment traditional morphological and computational approaches, we developed a simulation system capable of replicating human and non-human primate chewing motions. The forces of occlusion are recorded as photoelastic fringes in a urethane alveolar process. Simulation experiments evaluating the functional correlates of topographic diversity in zygomatic root position and mandibular ramus height in early hominids indicated that the mandibles and dentitions of robust australopithecines are well adapted to sustain high magnitude, low gradient load distributions.     

Masticatory stress, orbital orientation and the evolution of the primate postorbital bar

A postorbital bar is one of a suite of derived features which distinguishes basal primates from their putative sister taxon, plesiadapiforms. Two hypotheses have been put forward to explain postorbital bar development and variation in circumorbital form: the facial torsion model and visual predation hypothesis. To test the facial torsion model, we employ strain data on circumorbital and mandibular loading patterns in representative primates with a postorbital bar and masticatory apparatus similar to basal primates. To examine the visual predation hypothesis, we employ metric data on orbit orientation in Paleocene and Eocene primates, as well as several clades of visual predators and foragers that vary interspecifically in postorbital bar formation.A comparison of galago circumorbital and mandibular peak strains during powerful mastication demonstrates that circumorbital strains are quite low. This indicates that, as in anthropoids, the strepsirhine circumorbital region is excessively overbuilt for countering routine masticatory loads. The fact that circumorbital peak-strain levels are uniformly low in both primate suborders undermines any model which posits that masticatory stresses are determinants of circumorbital form, function and evolution. This is interpreted to mean that sufficient cortical bone must exist to prevent structural failure due to non-masticatory traumatic forces. Preliminary data also indicate that the difference between circumorbital and mandibular strains is greater in larger taxa.Comparative analyses of several extant analogs suggest that the postorbital bar apparently provides rigidity to the lateral orbital margins to ensure a high level of visual acuity during chewing and biting. The origin of the primate postorbital bar is linked to changes in orbital convergence and frontation at smaller sizes due to nocturnal visual predation and increased encephalization. By incorporating in vivo and fossil data, we reformulate the visual predation hypothesis of primate origins and thus offer new insights into major adaptive transformations in the primate skull.     

Homoplasy and the early hominid masticatory system: inferences from analyses of extant hominoids and papionins

Early hominid masticatory characters are widely considered to be more prone to homoplasy than characters from other regions of the early hominid skull and therefore less reliable for phylogenetic reconstruction. This hypothesis has important implications for current reconstructions of early hominid phylogeny, but it has never been tested. In this paper we evaluate the likely veracity of the hypothesis using craniometric data from extant primate groups for which reliable consensus molecular phylogenies are available.Datasets representing the extant large-bodied hominoid genera and the extant papionin genera were compiled from standard measurements. The data were adjusted to minimise the confounding effects of body size, and then converted into discrete character states using divergence coding. Each dataset was divided into four regional character groups: (1) palate and upper dentition, (2) mandible and lower dentition, (3) face and (4) cranial vault and base. Thereafter, the regional character groups were analysed using cladistic methods and the resulting phylogenetic hypotheses judged against the consensus molecular phylogenies for the hominoids and papionins.The analyses indicated that the regions dominated by masticatory characters-the palate and upper dentition, and the mandible and lower dentition-are no less reliable for phylogenetic reconstruction than the other regions of the skull. The four regions were equally affected by homoplasy and were, therefore, equally unreliable for phylogenetic reconstruction. This finding challenges the recent suggestion that Paranthropus is polyphyletic, which is based on the assumption that masticatory characters are especially prone to homoplasy. Our finding also suggests that, contrary to current practice, there is no a priori reason to de-emphasise the phylogenetic significance of the masticatory similarities between Homo rudolfensis and the australopiths. The corollary of this is that H. rudolfensis is unlikely to be a member of the Homo clade and should therefore be allocated to another genus.     

Strain in the galago facial skull

Little experimental work has been directed at understanding the distribution of stresses along the facial skull during routine masticatory behaviors. Such information is important for understanding the functional significance of the mammalian circumorbital region. In this study, bone strain was recorded along the dorsal interorbit, postorbital bar, and mandibular corpus in Otolemur garnettii and O. crassicaudatus (greater galagos) during molar chewing and biting. We determined principal-strain magnitudes and directions, compared peak shear-strain magnitudes between various regions of the face, and compared galago strain patterns with similar experimental data for anthropoids. This suite of analyses were used to test the facial torsion model (Greaves [1985] J Zool (Lond) 207:125-136; [1991] Zool J Linn Soc 101:121-129; [1995] Functional morphology in vertebrate paleontology. Cambridge: Cambridge University Press, p 99-115). A comparison of galago circumorbital and mandibular peak strains during powerful mastication indicates that circumorbital strains are very low in magnitude. This demonstrates that, as in anthropoids, the strepsirhine circumorbital region is highly overbuilt for countering routine masticatory loads. The fact that circumorbital peak-strain magnitudes are uniformly low in both primate suborders undermines any model that emphasizes the importance of masticatory stresses as a determinant of circumorbital form, function, and evolution. Preliminary data also suggest that the difference between mandibular and circumorbital strains is greater in larger-bodied primates. This pattern is interpreted to mean that sufficient cortical bone must exist in the circumorbital region to prevent structural failure due to nonmasticatory traumatic forces. During unilateral mastication, the direction of epsilon(1) at the galago dorsal interorbit indicates the presence of facial torsion combined with bending in the frontal plane. Postorbital bar principal-strain directions during mastication are oriented, on average, very close to 45 degrees relative to the skull's long axis, much as predicted by the facial torsion model. When chewing shifts from one side of the face to the other, there is a characteristic reversal or flip-flop in principal-strain directions for both the interorbit and postorbital bar. Although anthropoids also exhibit an interorbital reversal pattern, peak-strain directions for this clade are opposite those for galagos. The presence of such variation may be due to suborder differences in relative balancing-side jaw-muscle force recruitment. Most importantly, although the strain-direction data for the galago circumorbital region offer support for the occurrence of facial torsion, the low magnitude of these strains suggests that this loading pattern may not be an important determinant of circumorbital morphology.     

Telemetry System for Assessing Jaw-Muscle Function in Free-ranging Primates

In vivo laboratory-based studies describing jaw-muscle activity and mandibular bone strain during mastication provide the empirical basis for most evolutionary hypotheses linking primate masticatory apparatus form to diet. However, the laboratory data pose a potential problem for testing predictions of these hypotheses because estimates of masticatory function and performance recorded in the laboratory may lack the appropriate ecological context for understanding adaptation and evolution. For example, in laboratory studies researchers elicit rhythmic chewing using foods that may differ significantly from the diets of wild primates. Because the textural and mechanical properties of foods influence jaw-muscle activity and the resulting strains, chewing behaviors studied in the laboratory may not adequately reflect chewing behaviors of primates feeding in their natural habitats. To circumvent this limitation of laboratory-based studies of primate mastication, we developed a system for recording jaw-muscle electromyograms (EMGs) from free-ranging primates so that researchers can conduct studies of primate jaw-muscle function in vivo in the field. We used the system to record jaw-muscle EMGs from mantled howlers (Alouatta palliata) at Hacienda La Pacifica, Costa Rica. These are the first EMGs recorded from a noncaptive primate feeding in its natural habitat. Further refinements of the system will allow long-term EMG data collection so that researchers can correlate jaw-muscle function with food mechanical properties and behavioral observations. In addition to furthering understanding of primate feeding biology, our work will foster improved adaptive hypotheses explaining the evolution of primate jaw form.     

Occlusal forces and mandibular bone strain: Is the primate jaw “overdesigned”?

Finite element modelling of the function of the periodontium and surrounding alveolar bone suggests these tissues are subjected to unusually large strains in comparison with the bone of the basal mandibular corpus. These studies, in addition to certain experimental investigations, have led to the suggestion that the strains experienced in the basal mandibular corpus are not functionally important. Under this view, size and shape of the basal corpus are not functionally linked to masticatory forces. Since previous comparative investigations have been premised on the assumption that masticatory strains in the basal corpus are functionally important, the assertion that masticatory stresses are concentrated primarily in the alveolar process undermines the credibility of this body of work. The hypothesis that the biomechanical effects of masticatory forces are localized in the alveolar process can be evaluated by reference to a number of bone strain investigations, as well as through consideration of current understanding of bone biology and behavior. Experimental studies indicate that the effects of occlusal forces during mastication are quite apparent in alveolar bone, although relatively large strains are also observed in regions well-removed from a loaded alveolus. It is also apparent that both alveolar and basal mandibular bone are subject to bending and twisting strains associated not only with occlusal forces, but also with muscular and condylar reaction forces. The result is that strain levels in alveolarvs.basal bone may be roughly similar, in contradiction to some published theoretical models. Based on empirical evidence and theoretical considerations, it is premature to conclude that mandibular corpus size and shape are not functionally linked to the biomechanics of chewing and biting.     

A finite element analysis of masticatory stress hypotheses

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 simple 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 simple global loading regimes predict strains that approximate the craniofacial deformation pattern observed during mastication. Loading regimes, including those simulating 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 loading 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 regime satisfactorily accounts for the strain pattern found in the validated FEM. Instead, we propose that FE models replace simple cranial models when interpreting bone strain data and formulating hypotheses about craniofacial biomechanics.     

The influence of experimental manipulations on chewing speed during in vivo laboratory research in tufted capuchins (Cebus apella)

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.     

Mechanical and spatial determinants of Paranthropus facial form

It is well documented in the anthropological literature that the distinctive morphology of the “robust” hominid facial skeleton reflects its dietary specialization. Rak (1983) has provided the most comprehensive evaluation of Paranthropus facial morphology and this important study concluded that bone strain generated during mastication was responsible for the scaling of measures of facial height and breadth. The present study evaluated Rak's analysis by examining the relationship between bizygomatic breadth and facial height in an ontogenetic series of Pan and Gorilla crania. Results of this analysis indicate that facial height and breadth dimensions were not mechanically scaled in the “robust” australopithecines. Structural analysis of African ape facial maturation was also used to examine alternative spatial methods of malar elongation in Paranthropus. It is concluded that the increased height of the malar region in these specimens is not related to either vertical expansion of the posterior facial skeleton or to expansion of the temporal fossa. Malar elongation is, however, consistent with a derived pattern of facial growth in crania possessing a thickened hard palate. © 1994 Wiley-Liss, Inc.     

Complexity of ruminant masticatory evolution

The evolution of robust jaws, hypsodont teeth, and large chewing muscles among grazing ruminants is a quintessential example of putative morphological adaptation. However, the degree of correlated evolution (i.e., to what extent the grazer feeding apparatus represents an evolutionary module), especially of soft and hard tissues, remains poorly understood. Recent generation of large datasets and phylogenetic information has made testing hypotheses of correlated evolution possible. We, therefore, test for correlated evolution among various traits of the ruminant masticatory apparatus including tooth crown height, jaw robustness, chewing muscle size, and characters of the molar occlusal surfaces, using phylogenetic and nonphylogenetic comparative methods as well as phylogenetic evolutionary model selection. We find that the large masseter muscles of grazing ruminants evolved with the inclusion of grass in the diet, an increase in the proportion of occlusal enamel bands oriented parallel to the chewing stroke, and possibly hypsodonty. We suggest that the masseter evolved under two evolutionary regimes: i) selection for higher masticatory forces during chewing and ii) flattening of the tooth profile, which resulted in reduced tooth guidance and, thus, a requirement for more chewing muscle activity during each chewing stroke, in agreement with previous research. The linear jaw metrics (depth of the mandibular angle, mandibular angle width, and length of the superficial masseteric scar) all show correlated evolution with hypsodonty and the proportion of enamel bands oriented parallel to the chewing stroke. We suggest that changes in the shape of the mandible represent the combined effects of selection for a reorientation of the chewing stroke, so as to emphasize horizontal translation of the teeth, and accommodation of high‐crowned teeth. Our analyses show that the ruminant feeding apparatus is an evolutionary mosaic with its various components showing both correlated and independent evolution. J. Morphol. 275:1093–1102, 2014. © 2014 Wiley Periodicals, Inc.     

Do homoiologies impede phylogenetic analyses of the fossil hominids? An assessment based on extant papionin craniodental morphology

Homoiologies are phylogenetically misleading resemblances among taxa that can be attributed to phenotypic plasticity. Recently, it has been claimed that homoiologies are widespread in the hominid skull, especially in those regions affected by mastication-related strain, and that their prevalence is a major reason why researchers have so far been unable to obtain a reliable estimate of hominid phylogeny. To evaluate this "homoiology hypothesis", we carried out analyses of a group of extant primates for which a robust molecular phylogeny is available-the papionins. We compiled a craniometric dataset from measurements that differ in their susceptibility to mastication-related strain according to developmental considerations and experimental evidence. We used the coefficient of variation and analysis of variance with post hoc least significant difference comparisons in order to evaluate the variability of the measurements. The prediction from the homoiology hypothesis was that dental measurements, which do not remodel in response to strain, should be less variable than low-to-moderate-strain measurements, and that the latter should be less variable than high-strain measurements. We then performed phylogenetic analyses using characters derived from the measurements and compared the resulting phylogenetic hypotheses to the group's consensus molecular phylogeny. The prediction was that, if the homoiology hypothesis is correct, the agreement between the craniometric and molecular phylogenies would be best in the analyses of dental characters, intermediate in the analyses of low-to-moderate-strain characters, and least in the analyses of high-strain characters. The results of this study support the suggestion that mastication-related mechanical loading can result in variation in hominid cranial characters. However, they do not support the hypothesis that homoiology is a major reason why phylogenetic analyses of hominid crania have so far yielded conflicting and weakly supported hypotheses of relationship. These findings are consistent with a recent test of the homoiology hypothesis using craniodental data from extant hominoids, and cast doubt on the validity of the homoiology hypothesis, as originally formulated.     

Time analysis of hard and soft bolus processing

Objective: Clinical observations and mathematical models show that dental implants are influenced by the magnitude of loading. Therefore, the knowledge of mandible movement during mastication is important to assess occlusal and masticatory force vectors. The purpose of this study was to detect the path of movement of the lower jaw and to distinguish stages of mastication, duration of bolus processing and peak amplitude of mastication. Method: Motion analysis was used to record three-dimensional mandible movements. Individualized sensors were rigidly attached to the mandible of 51 study participants. At the beginning of the measurement, all subjects were asked to move the mandible in extreme positions (maximal opening and maximal lateral movements). Then, each subject masticated a bite of hard and soft food. Duration of bolus mastication and peak amplitude of mastication movement in mesio-distal, cranio-caudal and vestibulo-oral axes related to peak amplitude of marginal movements were evaluated for each subject. The chewing record of each subject was divided into three phases (chopping, grinding and swallowing), and the duration of mastication and number of closing movements were evaluated. Results: The findings of this pilot study suggest that masticatory movements vary in individuals. Bolus character influences the process duration, but not the frequency of closing movements. Neither gender nor age had any influence on either the time or frequency of bolus processing. Conclusion: Relationships to directions and magnitudes of acting chewing force should be more precisely examined since transversally acted forces during grinding are important factors in tooth/implant overloading.     

Palatal thickening and facial form in Paranthropus: Examination of alternative developmental models

Paranthropus is distinctive among hominoids in its possession of a greatly thickened hard palate. Although traditionally considered a structural adaptation to counter high-magnitude masticatory stress, alternative developmental models are equally viable. Three models of palatal thickening were evaluated in this study. A mechanical model interprets palatal thickening as a compensatory response to increased instability of the midpalatal suture effected by an anterior placement of the masseteric muscle mass. This model predicts that palatal thickness is correlated with the length of the palate posterior to the masseteric tubercle. Two non-mechanical models consider the thickness of the hard palate to be structurally related to and therefore correlated with either 1) the degree to which the premaxilla overlaps the hard palate in the subnasal region or 2) the height of the posterior facial skeleton. The correlation of craniofacial variables was assessed intraspecifically in ontogenetic series of great ape and human crania. Tests of correlation were performed for each comparison using both residuals calculated from reduced major axis regression of the variable of interest against a measure of cranial size and shape ratios. A significant correlation of palatal thickness with posterior facial height in Pan suggests that the unusually thick hard palate of Paranthropus is directly related to the increased posterior facial height characteristic of this taxon. Further evaluation suggests that extreme palatal thickening in these specimens occurred by virtue of their possession of a nasal septum morphology in which the vomer extends onto the superior nasal surface of the premaxilla. Such a morphology would have constrained the palatal nasal lamina to maintain the approximate level of the premaxillary nasal lamina throughout the growth process thereby promoting palatal thickening. Am J Phys Anthropol 103:375–392, 1997. © 1997 Wiley-Liss, Inc.     

Masticatory-stress hypotheses and the supraorbital region of primates

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.     

Loading patterns and jaw movements during mastication in Macaca fascicularis: a bone-strain, electromyographic, and cineradiographic analysis

Rosette strain gage, electromyography (EMG), and cineradiographic techniques were used to analyze loading patterns and jaw movements during mastication in Macaca fascicularis. The cineradiographic data indicate that macaques generally swallow frequently throughout a chewing sequence, and these swallows are intercalated into a chewing cycle towards the end of a power stroke. The bone strain and jaw movement data indicate that during vigorous mastication the transition between fast close and the power stroke is correlated with a sharp increase in masticatory force, and they also show that in most instances the jaws of macaques are maximally loaded prior to maximum intercuspation, i.e. during phase I (buccal phase) occlusal movements. Moreover, these data indicate that loads during phase II (lingual phase) occlusal movements are ordinarily relatively small. The bone strain data also suggest that the duration of unloading of the jaw during the power stroke of mastication is largely a function of the relaxation time of the jaw adductors. This interpretation is based on the finding that the duration from 100% peak strain to 50% peak strain during unloading closely approximates the half-relaxation time of whole adductor jaw muscles of macaques. The EMG data of the masseter and medial pterygoid muscles have important implications for understanding both the biomechanics of the power stroke and the external forces responsible for the "wishboning" effect that takes place along the mandibular symphysis and corpus during the power stroke of mastication. Although both medial pterygoid muscles reach maximum EMG activity during the power stroke, the activity of the working-side medial pterygoid peaks after the balancing-side medial pterygoid. Associated with the simultaneous increase of force of the working-side medial pterygoid and the decrease of force of the balancing-side medial pterygoid is the persistently high level of EMG activity of the balancing-side deep masseter (posterior portion). This pattern is of considerable significance because the direction of force of both the working-side medial pterygoid and the balancing-side deep masseter are well aligned to aid in driving the working-side lower molars across the upper molars in the medial direction during unilateral mastication.(ABSTRACT TRUNCATED AT 400 WORDS)     

In vivo and in vitro bone strain in the owl monkey circumorbital region and the function of the postorbital septum

Anthropoids and tarsiers are the only vertebrates possessing a postorbital septum. This septum, formed by the frontal, alisphenoid, and zygomatic bones, separates the orbital contents from the temporal muscles. Three hypotheses suggest that the postorbital septum evolved to resist stresses acting on the skull during mastication or incision. The facial-torsion hypothesis posits that the septum resists twisting of the face about a rostrocaudal axis during unilateral mastication; the transverse-bending hypothesis argues that the septum resists caudally directed forces acting at the lateral orbital margin during mastication or incision; and the tension hypothesis suggests that the septum resists ventrally directed components of masseter muscle force during mastication and incision. This study evaluates these hypotheses using in vitro and in vivo bone strain data recorded from the circumorbital region of owl monkeys. Incisor loading of an owl monkey skull in vitro bends the face upward in the sagittal plane, compressing the interorbital region rostrocaudally and “buckling” the lateral orbital walls. Unilateral loading of the toothrow in vitro also bends the face in the sagittal plane, compressing the interorbital region rostrocaudally and buckling the working side lateral orbital wall. When the lateral orbital wall is partially cut, so as to reduce the width of its attachment to the braincase, the following changes in circumorbital bone strain patterns occur. During loading of the incisors, lower bone strain magnitudes are recorded in the interorbital region and lateral orbital walls. In contrast, during unilateral loading of the P³, higher bone strain magnitudes are observed in the interorbital region, and generally lower bone strain magnitudes are observed in the lateral orbital walls. During unilateral loading of the M², higher bone strain magnitudes are observed in both the interorbital region and in the lateral orbital wall ipsilateral to the loaded molar. Comparisons of the in vitro results with data gathered in vivo suggest that, during incision and unilateral mastication, the face is subjected to upward bending in the sagittal plane resulting in rostrocaudal compression of the interorbital region. Modeling the lateral orbital walls as curved plates suggests that during mastication the working side wall is buckled due to the dorsally directed component of the maxillary force which causes upward bending of the face in the sagittal plane. The balancing side lateral orbital wall may also be buckled due to upward bending of the face in the sagittal plane as well as being twisted by the caudoventrally directed components of the superficial masseter muscle force. The in vivo data do not exclude the possibility that the postorbital septum functions to improve the structural integrity of the postorbital bar during mastication. However, there is no reason to believe that a more robust postorbital bar could not also perform this function. Hypotheses stating that the postorbital septum originally evolved to reinforce the skull against routine masticatory loads must explain why, rather than evolving a postorbital septum, the stem anthropoids did not simply enlarge their postorbital bars. © 1996 Wiley-Liss, Inc.     

Interspecific perspective on mechanical and nonmechanical models of primate circumorbital morphology.

Linear dimensions and angular orientations of the browridge, postorbital bar, and postorbital septum were obtained from a representative series of primates and compared with variables associated with several nonmechanical and biomechanical/mechanical models put forward to explain the form and function of the circumorbital region. Analyses of the results indicate that face size is the primary determinant of variation in primate circumorbital morphology. Anteroposterior browridge thickness is correlated with neural-orbital disjunction among anthropoid primates, but not among prosimians. This difference appears related to differences in the construction of the upper face and anterior cranial fossa between prosimians and anthropoids. Little support is demonstrated for the anterior dental loading model of browridge development. Mediolateral postorbital bar width and (to a lesser degree) browridge height are correlated with neurofacial torsion during mastication and variation in masticatory muscle size. These analyses further suggest that since circumorbital structures (especially the browridges) are located the farthest away from the chewing apparatus, they are least affected by masticatory stresses.     

Feeding behavior,mastication, and tooth wear in the western tarsier (Tarsius bancanus)

We assessed feeding and masticatory function in western tarsiers, Tarsius bancanus,from field study, from videotaped recordings of the feeding and chewing behavior of wild-caught animals in temporary captivity, from dissections of the muscles of mastication, and from scanning electron microscopic (SEM) examination of wear features of the teeth. Ingestion of large items of animal prey is made possible by the animal’s extremely wide gape. Anterior translation of the knob-like mandibular condyle in the anteroposteriorly elongated mandibular fossa makes possible a gape angle of 60–70‡. We observed two means of ingestion of grasshopper prey: ingestion by mastication, in which the postcanine teeth sever and reduce bites of the food as it is thrust into the mouth cavity, and repeated gape-shove sequences, during which the tarsier pushed grasshoppers of large diameter into the anterior part of its mouth and attempted to sever a bite with its anterior teeth. Morsels were successfully severed after three to five such sequences, and reduced quickly,with relatively few powerful, crushing chews. The insect cuticle was not evenly comminuted during mastication. We observed a marked side-to-side grinding component in the normal chewing cycle of T. bancanuson videotape and confirmed it by SEM. The main jaw adductors are bulky, long-fibered muscles that can accommodate wide grapes and still generate, at wide degrees of gape,the high occlusal pressures necessary to fracture thick chitinous exoskeletons of the scarabid beetles that form a substantial element of the western tarsier’s diet.     

The enigma of the Sangiran 4 palate revisited

The Sangiran 4 palate has been controversial since its discovery in the 1930s because it retains a number of more primitive morphologies such as projecting canines and precanine diastemata. These characters have led some workers to question the hominid status of the palate, suggesting that it is both too large and too primitive to belong to the same individual as the Sangiran 4 cranial fragments. The palate has instead been diagnosed as a new species of Pongo. The present study re-evaluates this controversy through the analysis of new metric data and comparisons with more recently published fossil discoveries. This analysis shows that the Sangiran 4 palate is not unique, and shares several of these putative pongid traits with other Javan hominid fossils as well as recently described hominid specimens from Dmanisi, Georgia. These results suggest that the evolution of the earliest Asians was more complex than has previously been appreciated.