Balancing function of the masticatory muscles during incisal biting in two murid rodents,Apodemus speciosus and Clethrionomys rufocanus

The functional significance of masticatory muscle direction was estimated using a mechanical model in two murid rodents: the Japanese field mouse (Apodemus speciosus) and the gray red-backed vole (Clethrionomys rufocanus). Theoretical analyses of the data suggest that a balancing mechanism among the muscle forces occurs during incisal power stroke. The activation of the large deep masseter in both murids results in marked tensile separation of two hemimandibles at the flexible mandibular symphysis. Activation of the internal pterygoid decreases this large tensile force at the symphysis more efficiently than other muscles. The lines of action of the deep masseter and internal pterygoid are aligned to produce such a balancing function in both species studied here. The resultant force generated by the deep masseter on both sides is opposite in direction to the reaction force at the lower incisor tip. Therefore, the large deep masseter forms an effective mandibular support mechanism when the reaction forces during biting push the mandible downward. Because of the area of insertion and the line of action, the posterior temporalis appears to have an important role in stabilizing the position of the mandibular condyle in the glenoid fossa during incisal biting. J. Morphol. 236:49–56, 1998. © 1998 Wiley-Liss, Inc.     

Mandibular movement and muscle activity during mastication in the guinea pig (Cavia porcellus)

Optoelectronic analysis of mandibular movement and electromyography (EMG) of masticatory muscles in Cavia porcellus indicate bilateral, unilateral, and gnawing cycles. During bilateral and unilateral cycles, the mandibular tip moves forward, lateral, and down during the lingual phase of the power stroke to bring the teeth into occlusion. EMG activity is generally asymmetric, with the exception of activity of the temporalis muscle during bilateral cycles. During gnawing cycles, the mandible moves in an anteroposterior direction that is opposite that during bilateral and unilateral chew cycles. Bilateral and unilateral cycles of pellets were significantly longer than carrot. With the exception of the width of bilateral cycles, the magnitude of cycle width, length, and height during the mastication of carrots was greater than that during the mastication of pellets. Significant differences exist between EMG durations during mastication of pellets and carrots. The lateral pterygoid displays continuous activity during gnawing cycles. Significant differences also exist in the durations of EMG activity between the working and balancing side during all three cycle types. High level activity of balancing side temporalis and anterior belly of digastric (ABD) during bilateral cycles occurs during rotation and depression of the mandible during the power stroke. The temporalis apparently provides a ?braking”? or compensatory role during closing and power strokes. Differences between Cavia masticatory patterns and those shown by Rattus and Mesocricetus are apparently due to differences in dental morphology, occlusal relationships, and, possibly, the poorly developed temporalis in Cavia. The large number and wide diversity of rodent groups afford students of mammalian mastication an opportunity to investigate and compare different masticatory specializations.     

Structure and direction of jaw adductor muscles as herbivorous adaptations in <Emphasis Type="Italic">Neotoma mexicana</Emphasis> (Muridae,Rodentia)

The herbivorous adaptations of the jaw adductor muscles in Neotoma mexicana were clarified by a comparative study with an unspecialized relative, Peromyscus maniculatus. In P. maniculatus, the anterior part of the deep masseter arises entirely from the lateral side of an aponeurosis, i.e., superior zygomatic plate aponeurosis, whereas N. mexicana has an additional aponeurosis for this part of the muscle, and the fibers attach on both sides of the superior zygomatic plate aponeurosis. Although the structure of the temporalis muscle is nearly identical in the two genera, a clear aponeurosis of origin occurs only in N. mexicana. These characteristics allow fibrous tissues to be processed with a large occlusal force. The deep masseter, internal pterygoid, and external pterygoid muscles of N. mexicana incline more anterodorsally than those of P. maniculatus. The transverse force component of these muscles relative to whole muscle force is smaller in N. mexicana than in P. maniculatus, with the exception of the internal pterygoid. The anterior part of the temporalis muscle of N. mexicana is specialized to produce occlusal pressure. These findings suggest that in N. mexicana a large anterior force is required to move the heavy mandible, due to the hypsodont molars, against frictional force from food, and that the posterior pull of the temporalis, which adjusts the forward force by the other jaw adductor muscles to a suitable level, need not be large for the mandibular movement.     

Cranial functional morphology of fossil dogs and adaptation for durophagy in Borophagus and Epicyon (Carnivora,Mammalia)

Morphological specialization is a complex interplay of adaptation and constraint, as similarly specialized features often evolve convergently in unrelated species, indicating that there are universally adaptive aspects to these morphologies. The evolutionary history of carnivores offers outstanding examples of convergent specialization. Among larger predators, borophagine canids were highly abundant during the tertiary of North America and are regarded as the ecological vicars of Afro‐Eurasian hyenas. Borophaginae is an extinct group of 60+ species, the largest forms evolving robust skulls with prominently domed foreheads, short snouts, and hypertrophied fourth premolars. These specializations have been speculated to enhance bone cracking. To test the extent that the skulls of derived borophagines were adapted for producing large bite forces and withstanding the mechanical stresses associated with bone cracking relative to their nonrobust sister clades, we manipulated muscle forces in models of six canid skulls and analyzed their mechanical response using 3D finite element analysis. Performance measures of bite force production efficiency and deformation minimization showed that skulls of derived borophagines Borophagus secundus and Epicyon haydeni are particularly strong in the frontal region; maximum stresses are lower and more evenly distributed over the skull than in other canids. Frontal strength is potentially coupled with a temporalis‐driven bite to minimize cranial stress during biting in the two derived genera, as tensile stress incurred by contracting temporalis muscles is dissipated rostro‐ventrally across the forehead and face. Comparison of estimated masticatory muscle cross section areas suggests that the temporalis‐masseter ratio is not strongly associated with morphological adaptations for bone cracking in Borophagus and Epicyon; larger body size may explain relatively larger temporalis muscles in the latter. When compared with previous studies, the overall cranial mechanics of the derived borophagines is more similar to bone‐cracking hyaenids and percrocutids than to their canid relatives, indicating convergence in both morphological form and functional capability. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Quantitative genetic analysis of cranial morphology in the cotton-top (Saguinus oedipus) and saddle-back (S. fuscicollis) tamarins

Patterns of morphological variation play an important role in evolutionary diversification and are critical to an informed interpretation of interspecific differences. When patterns of genetic variation have not diverged substantially, it is possible to reconstruct the differences in selection which gave rise to morphological differences among extant species. Morphological variation patterns are compared between two tamarin species, the cotton-top tamarin (Saguinus oedipus) and the saddle-back tamarin (S. fuscicollis illigeri). Genetic, phenotypic, and environmental variance/covariance and correlation matrices were obtained for a series of 39 cranial characters in each species (cotton-top tamarin, N = 328; saddle-back tamarin, N = 209) and for the species combined using crania from individuals of known genealogical relationship. After accounting for the effects of estimation error on measures of matrix similarity, patterns of phenotypic, genetic, and environmental variation and correlation were found to be very similar across species and among the types of variance within species. Taking the saddle-back tamarins as the standard, cotton-top tamarins have been selected for an enlarged anterior temporalis attachment area and increased facial prognathism. In primates, an enlarged anterior temporalis muscle is associated with incisive food preparation, especially at wide gape.     

Jaw-muscle fiber architecture in tufted capuchins favors generating relatively large muscle forces without compromising jaw gape

Tufted capuchins (sensu lato) are renowned for their dietary flexibility and capacity to exploit hard and tough objects. Cebus apella differs from other capuchins in displaying a suite of craniodental features that have been functionally and adaptively linked to their feeding behavior, particularly the generation and dissipation of relatively large jaw forces. We compared fiber architecture of the masseter and temporalis muscles between C. apella (n = 12) and two “untufted” capuchins (C. capucinus, n = 3; C. albifrons, n = 5). These three species share broadly similar diets, but tufted capuchins occasionally exploit mechanically challenging tissues. We tested the hypothesis that tufted capuchins exhibit architectural properties of their jaw muscles that facilitate relatively large forces including relatively greater physiologic cross-sectional areas (PCSA), more pinnate fibers, and lower ratios of mass to tetanic tension (Mass/P₀). Results show some evidence supporting these predictions, as C. apella has relatively greater superficial masseter and temporalis PCSAs, significantly so only for the temporalis following Bonferroni adjustment. Capuchins did not differ in pinnation angle or Mass/P₀. As an architectural trade-off between maximizing muscle force and muscle excursion/contraction velocity, we also tested the hypothesis that C. apella exhibits relatively shorter muscle fibers. Contrary to our prediction, there are no significant differences in relative fiber lengths between tufted and untufted capuchins. Therefore, we attribute the relatively greater PCSAs in tufted capuchins primarily to their larger muscle masses. These findings suggest that relatively large jaw-muscle PCSAs can be added to the suite of masticatory features that have been functionally linked to the exploitation of a more resistant diet by C. apella. By enlarging jaw-muscle mass to increase PCSA, rather than reducing fiber lengths and increasing pinnation, tufted capuchins appear to have increased jaw-muscle and bite forces without markedly compromising muscle excursion and contraction velocity. One performance advantage of this morphology is that it promotes relatively large bite forces at wide jaw gapes, which may be useful for processing large food items along the posterior dentition. We further hypothesize that this morphological pattern may have the ecological benefit of facilitating the dietary diversity seen in tufted capuchins. Lastly, the observed feeding on large objects, coupled with a jaw-muscle architecture that facilitates this behavior, raises concerns about utilizing C. apella as an extant behavioral model for hominins that might have specialized on small objects in their diets.     

Reconstruction of the cranial musculature and masticatory function of the Pleistocene panamerican ground sloth Eremotherium laurillardi (Mammalia,Xenarthra, Megatheriidae)

Cranial musculature, dental function and mandibular movement patterns in Eremotherium laurillardi were reconstructed from the examination of crania and dentitions. Size, shape and pattern of muscle divisions were reconstructed from the examination of bony rugosities indicating muscle attachments. Details of masticatory muscle structure and function were based on dissections of the tree sloths Bradypus and Choloepus. Among sloths, masticatory muscles in E. laurillardi demonstrate a different synergist–antagonist pattern, reflecting greater emphasis on mediolateral mandibular movements. Eight cranial character complexes (anterior facial, zygomatic arch, superficial masseter, deep masseter–zygomaticomandibularis, pterygoid, temporal, occipital and occlusal) determined by interrelated contributions of each component made to group functions were identified. An elongate anterior face and predental spout in E. laurillardi allowed protrusion of a long narrow tongue at small degrees of gape, reflecting a probably ancestral xenarthran condition. Gape minimisation, in conjunction with the mediolaterally directed masticatory stroke in E. laurillardi, was a unique solution to increase masticatory efficiency by permitting molariform tooth shearing surfaces to remain in or near occlusion for a greater percentage of each chewing cycle.     

Masticatory function in patients with xerostomia

The effects of reduced salivary output in patients suffering from xerostomia on masticatory function has not been previously studied. This study compares masticatory performance and kinematic activity of patients suffering from xerostomia with age-, sex-, and number of occluding pairs-matched healthy controls. Masticatory function was evaluated by assessment of chewing motion and muscle activity during chewing an artificial food (CutterSil®), chewing gum and swallowing a bolus of almond. Chewing motion was recorded with the Optotrak® computer system. Bilateral muscle activity of both masseter and anterior temporalis was recorded using surface electrodes. Results of this study revealed significant differences between patients and controls in their ability to process food and masticatory muscle activity. The majority of patients could not break down the artificial food, others had a larger median particle size than the controls. A significant difference was also observed in the number of chewing cycles required to swallow almonds, the patients required more than twice as many chews as the controls, P<0.001. The right masseter muscle displayed significantly less activity for the patient than the controls. These findings suggest that patients with xerostomia exhibit reduced ability to process food. The observed decline in masticatory performance is probably due to reduced activity of the muscles of mastication.     

Mechanical advantage of area of origin for the external pterygoid muscle in two murid rodents, Apodemus speciosus and Clethrionomys rufocanus

The actions of masticatory muscles in relation to transverse grinding, associated with forward masticatory movement of the mandible, were investigated by using a mechanical model in the two murid rodents, the Japanese field mouse (Apodemus speciosus: subfamily Murinae) and the gray red-backed vole (Clethrionomys rufocanus: subfamily Arvicolinae). Furthermore, statics of the masticatory system on a sagittal plane while chewing is taking place were also analyzed in these rodents. The inward grinding movements of hemimandibles are generated by the posterior temporalis and internal and external pterygoids in both species. In addition to these muscles, the anterior temporalis also moves the hemimandibles lingually in Apodemus speciosus. The area of origin of the external pterygoid seems more advantageous for transverse grinding in A. speciosus than in Clethrionomys rufocanus. On the basis of the static analysis, the anterodorsal area of origin of the external pterygoid to the upper second and third molars in Clethrionomys rufocanus appears to be an adaptive character to prevent the jaw joints from dislocation during occlusion at a posterior point on the elongated row of cheek teeth.     

Association between the lack of teeth and the expression of myosins in masticatory muscles of microphthalmic mouse

The purposes of the present study were to elucidate the influences of the deficiency of teeth on masticatory muscles, such as the masseter, temporalis and digastric muscles and compare the influence among masticatory muscles. We analysed the expressions of myosin heavy chain (MyHC) isoform messenger RNA (mRNA) and protein in these muscles in the microphthalmic (mi/mi) mouse, whose teeth cannot erupt because of a mutation in the mitf gene locus. The expression levels of MyHC mRNA and protein in the masseter, temporalis, digastric, tibialis anterior and gastrocnemius muscles of +/+ and mi/mi mice were analysed with real‐time polymerase chain reaction and sodium dodecyl sulfate‐polyacrylamide gel electrophoresis, respectively. The mi/mi masseter muscle at 8 weeks of age expressed 4·1‐fold (p < 0·05) and 3.3‐fold (p < 0·01) more MyHC neonatal mRNA and protein than that in the +/+, respectively; the expression level of MyHC neonatal protein was 19% of the total MyHC protein in the masseter muscle of mi/mi mice. In the digastric muscle, the expression levels of MyHC I mRNA and protein in the mi/mi mice were 4·7‐fold (p < 0·05) and 5‐fold (p < 0·01) higher than those in the +/+ mice. In the temporalis, tibialis anterior and gastrocnemius muscles, there was no significant difference in the expression levels of any MyHC isoform mRNA and protein between +/+ and mi/mi mice. These results indicate associations between the lack of teeth and the expression of MyHC in the masseter and digastric muscles but not such associations in the temporalis muscle, suggesting that the influence of tooth deficiency varies among the masticatory muscles. Copyright © 2011 John Wiley & Sons, Ltd.     

Supraorbital ridge and masticatory apparatus I: Primates

The supraorbital ridge formation in apes is a result of the traction of the masticatory force exerted by the anterior part of the temporalis muscle. This force varies inversely with the ratio of the power arm to the load arm of the mandible. The smaller this ratio, the greater the anterior part of the temporalis muscle and the larger the supraorbital ridge. The size of the ridge is independent of the actual size of the skull. The orang has a larger skull but the smallest brow ridge of the three species studied: gorilla, chimpanzee and orang. Pearson’s correlation coefficient shows a positive correlation between the anterior temporalis muscle and the ridge as well as between the ridge/sinus and anterior/whole temporalis muscle. The correlation is negative between the power/load arms and anterior/whole muscle and also between the power/load arms and the ridge/sinus ratio.     

Contour analysis of masticatory jaw movements and muscle activity in Macaca mulatta

Frontal plane mandibular movements during mastication and the associated electromyographic (EMG) activity for left and right superficial masseter, posterior temporalis, anterior temporalis, and anterior belly of the digastric (ABD) were studied for two adult male Macaca mulatta by the new technique of “contour” analysis. Contour analysis allowed graphic and quantitative portrayal of multiple chew cycle patterns of mandibular movement and EMG activity during active mastication. A series of computer programs (ATS, ATSED, ATSXYZ) facilitated the collection, editing and definition, and finally processing of these masticatory data into contour plots. These preliminary data indicated the essential symmetry of mandibular movement patterns, high chew cycle variability inferior to occlusion, multiple centers of intense EMG activity for balancing-side superficial masseter, and no difference between working-side anterior and posterior temporalis EMG patterns. Maximum EMG amplitude was found in the area of buccal phase power stroke (BPS). Maximum EMG amplitude for ABD was located medial and inferior to occlusion; all other muscle maximum amplitudes were buccal and inferior to occlusion. The location of maximum EMG amplitudes for superficial masseter and ABD were closer to occlusion (more superior) during mastication of carrot than were maximum amplitudes during biscuit mastication. The absence of any detectable shift of EMG maximum amplitude location between biscuit and carrot for posterior and anterior temporalis suggested, along with the continuous EMG activity of working-side posterior temporalis, a secondary role for the temporalis (compensation for superficial masseter activity) during active mastication.     

Application and validation of a three-dimensional mathematical model of the human masticatory system in vivo.

A previously described three-dimensional mathematical model of the human masticatory system, predicting maximum possible bite forces in all directions and the recruitment patterns of the masticatory muscles necessary to generate these forces, was validated in in vivo experiments. The morphological input parameters to the model for individual subjects were collected using MRI scanning of the jaw system. Experimental measurements included recording of maximum voluntary bite force (magnitude and direction) and surface EMG from the temporalis and masseter muscles. For bite forces with an angle of 0, 10 and 20 degrees relative to the normal to the occlusal plane the predicted maximum possible bite forces were between 0.9 and 1.2 times the measured ones and the average ratio of measured to predicted maximum bite force was close to unity. The average measured and predicted muscle recruitment patterns showed no striking differences. Nevertheless, some systematic differences, dependent on the bite force direction, were found between the predicted and the measured maximum possible bite forces. In a second series of simulations the influence of the direction of the joint reaction forces on these errors was studied. The results suggest that they were caused primarily by an improper determination of the joint force directions.     

Pharyngeal teeth and masticatory process of the basioccipital bone in Japanese bitterlings (Cyprinidae)

The pharyngeal teeth and the masticatory process of the basioccipital bone were compared in fifteen species and subspecies of Japanese bitterlings.Acheilognathus lanceolata, A. limbata and two subspecies ofRhodeus ocellatus, which are characterized by the absence of serrations on the side of the pharyngeal teeth, have reduced occlusal grooves on the pharyngeal teeth. In the larvae of these species, the occlusal grooves are fairly developed. The occlusal grooves of the adult pharyngeal teeth in herbivorous species are more developed than those in omnivorous species. The occlusal grooves seem to have developed in relation to the feeding habits. On the basis of the combination of developmental degrees of the anterior part of the masticatory process, the occlusal grooves and the chewing area on the pharyngeal first tooth, Japanese bitterlings were classified into two groups and two types and/or five subtypes, suggesting phylogenetic relationships among them.     

Relationships between attrition and lingual tilting in human teeth

Associations between occlusal dental attrition and the lingual tilting of human teeth were investigated in two aboriginal California populations. A literature survey suggests lingual tilting is related geographically and temporally to the helicoidal occlusal plane in humans; if true, lingual tilting should be expected throughout the Homo osteological record. Buccal alveolar abscesses, exposed pulp chambers, extent of tooth attrition, angle of lingual tilt, and amount of overjet were observed for lingually tilted teeth. These attributes were analyzed statistically, leading to a conclusion that lingual tilting is slight and infrequent unless tooth attrition is pronounced. It is suggested that lingual tilting is due in part to masticatory stress. A feedback model considers lingual tilting as one conspicuous manifestation of a more complex “severe attrition syndrome,” the initial stimulus for which may derive from heavy occlusal tooth wear. Untested possibilities should be examined–e.g., age-related changes, the impact of the helicoidal occlusal plane, and the role of masticatory forces.     

Biomechanics of the masticatory apparatus of Pteropus giganteus (Megachiroptera)

Jaw mechanics in Pteropus were studied by means of a three-dimensional model. The model included several parameters of muscle architecture, combined with quantified movement and electromyographical data. Estimates of the nature of the applied forces that act upon the mandible during a chewing cycle, and subsequent estimates of reaction forces at the bite point and joints during the powerstroke, were thus obtained for different food consistencies. The resultant muscle force (relative to the palate) shifts from upward and slightly backward at large gapes to upward and markedly backward at the end of closing. The resultant simultaneously moves anteriorly. During the powerstroke it retains a constant position and orientation along the thickened anterior edge of the coronoid process. The early stages of opening are guided by the slope of the teeth and mandibular fossa; during the remaining part of opening the working line of the resultant crosses the skull behind the joint and thus acquires an opening moment. The bite force has downward and forward components, and a slight transverse component. For a given applied muscular force its magnitude is larger in more posteriorly positioned bite points. Both joints are loaded, the contralateral one more than the ipsilateral. Food consistency affects magnitude and orientation of the applied force, and hence, magnitude and orientation of the bite force and magnitude of the joint reaction forces. The magnitude of masseter activity relative to temporalis activity appears to be the key factor for the orientation of the bite force, and hence for the mechanical optimal position of the food. The adaptive value of the general topography of the masticatory muscles in Pteropus is discussed.     

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.     

Craniodental biomechanics and dietary toughness in the genus Cebus

The tufted capuchin (Cebus apella) has been used in a number of comparative studies to represent a primate with craniofacial morphology indicative of hard-object feeding. Researchers have specifically referred to the tufted capuchin as a seed predator. Craniofacial features exhibited by the tufted capuchin, such as thick cortical bone in the mandibular corpus and symphysis, and a broad face associated with large masticatory muscles, permit the production and dissipation of relatively high masticatory forces. These morphologies, however, cannot distinguish between the tufted capuchin's propensity to exert higher forces when opening food with its anterior dentition or with its cheek teeth. It is also unclear whether these are adaptations for biting or chewing foods. This study uses a constrained lever model to compare the masticatory adaptations of C. apella to other cebids and atelids. Results show that the temporalis and masseter muscles in C. apella and C. olivaceus are more anteriorly positioned relative to nine other platyrrhine taxa. This condition, which appears to be ancestral among the Cebinae, increases force production at the incisors and canines while compromising third molar function. Cebus apella, has exaggerated this pattern. Field data on dietary toughness show that both capuchins typically select foods of low toughness, but on occasion, C. apella ingests food items of exceptional toughness. Thus, C. apella appears to maintain these biomechanical relationships by producing particularly high but relatively infrequent bite forces, particularly at the incisors and canines. However, adaptations for anterior dental use do not tightly constrain the diet of Cebus apella. This approach can be used to clarify the dietary adaptations of fossil taxa.     

A functional analysis of the primate masticatory system and the origin of the anthropoid post-orbital septum

Study of the dry weights of primate and non-primate masticatory musculature reveals a significant relationship between the Anterior Temporalis/Masseter ratio and the relative development of the anterior dentition. Available dietary information demonstrates that species emphasizing incisal preparation of food have a high AT/M index; species emphasizing molar occlusion have a low AT/M index. Utilizing this information, a model is presented of the origin of the anthropoid post-orbital septum. Frugivory or extensive incisal preparation of food is causally related to the development of the post-orbital septum, because diet can then create selection pressures for an increasingly tendinous and enlarging anterior segment of the temporalis muscle which requires additional bony areas of origin in the anterior temporal fossa. Cenozoic climatic oscillations leading to increasing seasonality may have been the triggering element in this model, because seasonality creates periods in which the availability of fruit is relatively predictable.     

Feeding mechanics and dietary implications in the fossil sloth Neocnus (Mammalia: Xenarthra: Megalonychidae) from haiti

Haitian species of the extinct ground sloth genus Neocnus (Mammalia: Pilosa: Megalonychidae) have previously been hypothesized to have a much reduced jugal bone and a correspondingly reduced masseter musculature but a paucity of specimens has prevented further investigation of this hypothesis. Recent discovery of jugal bones belonging to Haitian specimens of Neocnus within the University of Florida Museum collections enables the element to be more accurately described. The discovery also makes it possible to explore mastication in these sloths. Osteological characters related to feeding were examined, along with comparative estimations of bite force with the extant tree sloths, Bradypus and Choloepus, and their known dietary habits as a means to infer aspects of the paleodiet of Neocnus. There is a significant difference in moment arm calculations for m. masseter between predicted and actual jugals, but the overall significance for bite force is lost and hampered by small sample size. Neocnus demonstrates a variety of characters that are similar to those of Bradypus and not to Choloepus, which is a close phylogenetic relative. The masticatory musculature of Neocnus enabled a chewing cycle emphasizing a grinding combination of mesiodistal and linguobuccal movements of the molariform dentition. The orientations of m. masseter and m. temporalis are estimated to produce relatively high bite force ratios that imply a masticatory system with stronger versus faster components. Because of the similarity of bite forces and jaw mechanics to those of Bradypus, in addition to a number of osteological adaptations indicative of herbivorous grazers (elevated mandibular condyle, large and complex masseter, and robust angular process), the Haitian forms of Neocnus are considered to have been selective feeders with a folivorous diet. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.