Mammals with a long diastema typically also have dominant masseter and pterygoid muscles

A few orders of mammals contain many individuals with dominant masseter and pterygoid muscles that pull up and forward as they close the jaw. A dominant temporalis muscle that pulls the jaw up and to the rear is the more common condition in mammals. A long toothless region (diastema) is present in almost all mammals with a large masseter/pterygoid complex. The presence of a diastema, when few teeth have been lost and their size has not changed significantly over evolutionary time, implies that the jaws have lengthened, as in horses and selenodont artiodactyls. (A long jaw with a shorter diastema will also form if very long incisors develop as in rodents.) The sum of the forces of all the jaw muscles (represented by an arrow) typically divides the jaw into a posterior, toothless region and an anterior region where the teeth are located. In most mammals, the sum of all the bite forces at the teeth is maximized when the lengths of the projections of these two regions, onto a line perpendicular to the arrow, are in the ratio of 3 : 7. If the tooth-bearing region of the jaws becomes longer over evolutionary time this ratio will obviously be disturbed. A change in the location of some basic bony features of the jaw mechanism could maintain this ratio, but this requires major disruption of the skull and jaws. Alternatively, simply changing the masses of the muscles that close the jaw (smaller temporalis, larger masseter and/or pterygoid, or some combination), so that the lower jaw is pulled up and forward, rather than backward, also maintains the ratio. According to this view, if the jaw lengthens over evolutionary time, the relative sizes of the jaw muscles will change so that the masseter/pterygoid complex will become dominant.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 625–629.     

First complete restoration of the Oreopithecus skull

The crushed skull ofOreopithecus bambolii (IGF 11778) has been fully reconstructed in Florence, Italy and shows important differences with previous drawn reconstructions. The jaws are massive and projecting, such that the face is not as short as was believed. The incisors are small and do not project significantly forward beyond the position of the canines, and the anterior symphyseal surface of the mandible projects in front of the canines and well infront of the upper canines and incisors. There is a prominent sagittal crest, a high ascending mandibular ramus, and a nuchal plane which faces postero-inferiorly.     

Ecomorphological correlates of craniodental variation in bears and paleobiological implications for extinct taxa: an approach based on geometric morphometrics

Relative warp analyses of landmarks describing cranial and mandibular shape are used for investigating patterns of morphological variation among extant bears (Mammalia, Carnivora, Ursidae) indicative of diet and feeding behavior. These patterns are used for deriving inferences about the autecology of two extinct species previously assumed to have had different dietary preferences, the North American giant, short-faced bear Arctodus simus and the Eurasian cave bear Ursus spelaeus . Results reveal a set of shared craniodental traits among the herbivorous bears, including short and vaulted skulls with well-developed zygomatic arches, lateralized orbits and small canines, concave jaws with a highly positioned condyle, large moment arms for the temporalis and masseter muscles, and long cheek teeth. In contrast, those bears that consume animal resources have long skulls with small zygomatic arches, frontalized orbits and well-developed canines, and long jaws with a deep mandibular symphysis, low muscle leverages, a condyle situated at the level of the tooth row and reduced cheek teeth. The craniodental morphology of omnivorous bears is intermediate between those of faunivores and herbivores. This is also the case of the short-faced bear and the cave bear, which suggests that previous reconstructions of the feeding ecology of these extinct species (highly carnivorous for A. simus and herbivorous for U. spelaeus ) should be revised.     

Molar occlusion and jaw mechanics of the Eocene primate Adapis

Wear facets on molars of the Eocene primate Adapis magnus are described. Striations on these wear facets indicate three separate directions of mandibular movement during mastication. One direction corresponds to a first stage of mastication involving orthal retraction of the mandible. The remaining two directions correspond to buccal and lingual phases of a second stage of mastication involving a transverse movement of the mandible. The mechanics of jaw adduction are analysed for both the orthal retraction and transverse stages of mastication. During the orthal retraction stage the greatest component of bite force is provided by the temporalis muscles acting directly against the food with the mandible functioning as a link rather than as a lever. A geometrical argument suggests that during the transverse stage of mastication bite force is provided by the temporalis muscles of both sides, the ipsilateral medial and lateral pterygoid muscles, and the contralateral masseter muscle.     

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.     

Jaw muscles of new world squirrels

The jaw, suprahyoid, and extrinsic tongue muscles are described for eight species of New World squirrels, spanning more than an order of magnitude in body mass. Anatomical differences are discussed in the light of body size, natural history, and phylogeny. The relative sizes of different muscles, their orientations, and the shapes and positions of their areas of attachment vary but show few trends in relation to body size. The anatomical differences are likewise not readily explained by the mechanical requirements of the animals' diets, which are similar. The most marked anatomical differences occur in Sciurillus (the pygmy tree squirrel), as well as those genera—Glaucomys (the flying squirrel) and Tamias (the chipmunk)—that are taxonomically most distinct from the tree squirrels. sciurillus is noteworthy for its unusually small temporalis and an anterior deep masseter that is oriented to assist in retraction of the jaw. Tamias has a more vertically oriented temporalis and greater inclination in the anterior masseter muscles than the other squirrels, features that may be associated with its large diastema and relatively posteriorly situated cheek teeth, which in turn may relate to its having cheek pouches. Our results form a valuable database of information to be used in further studies of functional morphology and phylogeny. © 1995 Wiley-Liss, Inc.     

The functional relationship between feeding type and jaw and cranial morphology in ungulates

The relationship between jaw and skull morphology and feeding type (grazer, mixed feeder, browser, frugivorous, omnivorous) was analysed in 94 species of extant ungulates. A total of 21 morphological traits of the jaw and skull (17 and 4, respectively) were analysed using analysis of covariance, with body mass as covariate. To take into account the phylogenetic effect, simulations were generated under the Brownian motion model of character evolution. Analysis of covariance was applied to these simulations and the simulated F-ratios were used to assess the signification of the F-ratios for the real values of the traits. The feeding types had a weak effect on ungulate cranial and jaw morphology in comparison with the phylogenetic effect, since, before phylogeny correction, the analysis of covariance showed statistically significant differences associated with feeding type in 15 out of the 21 traits analysed. After controlling for phylogeny, only 2 significant traits remained, the length of the coronoid process and the occipital height. Omnivorous species had shorter coronoid processes than grazers or mixed feeders, and the occipital height was greater in the omnivorous species than in the grazers, mixed feeders or browsers. The coronoid process is involved in the generation of bite force, being the effective moment arm of the temporalis muscle, and occipital height is positively related to the force exerted by the temporalis muscle. This result matches the hypothesis that species with a toughness diet should show higher bite force (“toughness” describes the resistance of a material to being mechanically broken down). When the omnivorous species were excluded from the analysis, no differences in jaw and skull morphology were detected between the rest of the feeding types. Received: 1 September 1998 / Accepted: 2 November 1998     

Modulatory complexity of the feeding repertoire in scincid lizards

The kinematics of jaws and tongue, and jaw muscle activity patterns were investigated in the omnivorous lizard Tiliqua rugosa, and the herbivorous Corucia zebrata (Scincidae) during feeding. Small metal markers were inserted into different parts of the skull, the jaws, and the tongue. Video and cineradiographic images were digitized and displacements of the head, jaws, and tongue were quantified. Additionally, muscle activity patterns were recorded, digitized and several variables were determined quantitatively. The effect of food type on the jaw and hyolingual movement patterns and the jaw muscle activity patterns was investigated for both species. The kinematic data indicate that distinct aspects of gape and tongue cycles are modulated in response to the food characteristics. Similarly, in both species, muscle activity patterns are altered in response to the type of food eaten. A comparison of kinematic and electromyographic patterns during intraoral transport cycles for both species shows that these can be related to food characteristics such as toughness and mobility. Differences between both species in the response to changes in food characteristics are minor. Clearly both species are able to fine tune the activation of the jaw muscles, resulting in the appropriate movement patterns for the type of food eaten. Accepted: 30 January 1999     

The orientation of the force of the jaw muscles and the length of the mandible in mammals

Ungulates generally have large masseter and pterygoid muscles and a necessarily large angular process provides attachment surface on the mandible. The temporalis muscle tends to be small. It has been suggested that this is an adaptation for enhanced control of the lower jaw and reduction of forces at the jaw joint. I suggest an additional reason: because of the geometry of the jaw, the length of that segment of the lower jaw that spans the distance from the jaw joint to the most posterior tooth is significantly reduced when the masseler and pterygoid are the dominant muscles; this region is necessarily much longer when the temporalis is large.     

The relationships among jaw‐muscle fiber architecture,jaw morphology,and feeding behavior in extant apes and modern humans

The jaw‐closing muscles are responsible for generating many of the forces and movements associated with feeding. Muscle physiologic cross‐sectional area (PCSA) and fiber length are two architectural parameters that heavily influence muscle function. While there have been numerous comparative studies of hominoid and hominin craniodental and mandibular morphology, little is known about hominoid jaw‐muscle fiber architecture. We present novel data on masseter and temporalis internal muscle architecture for small‐ and large‐bodied hominoids. Hominoid scaling patterns are evaluated and compared with representative New‐ (Cebus) and Old‐World (Macaca) monkeys. Variation in hominoid jaw‐muscle fiber architecture is related to both absolute size and allometry. PCSAs scale close to isometry relative to jaw length in anthropoids, but likely with positive allometry in hominoids. Thus, large‐bodied apes may be capable of generating both absolutely and relatively greater muscle forces compared with smaller‐bodied apes and monkeys. Compared with extant apes, modern humans exhibit a reduction in masseter PCSA relative to condyle‐M₁ length but retain relatively long fibers, suggesting humans may have sacrificed relative masseter muscle force during chewing without appreciably altering muscle excursion/contraction velocity. Lastly, craniometric estimates of PCSAs underestimate hominoid masseter and temporalis PCSAs by more than 50% in gorillas, and overestimate masseter PCSA by as much as 30% in humans. These findings underscore the difficulty of accurately estimating jaw‐muscle fiber architecture from craniometric measures and suggest models of fossil hominin and hominoid bite forces will be improved by incorporating architectural data in estimating jaw‐muscle forces. Am J Phys Anthropol 151:120–134, 2013. © 2013 Wiley Periodicals, Inc.     

Shape analysis of the jaws between two minnow species over ontogeny

This study compares sand shiner (Notropis stramineus ) and silverjaw (Ericymba buccata ) minnows, in terms of the morphological shape changes of the upper, lower, and pharyngeal jaws over ontogeny. These two species of minnows initially feed on midge larvae and undergo an ontogenic prey shift. The traditional morphometrics measured—total length, snout‐to‐vent length, eye diameter, premaxilla length, lower jaw length, gape—were regressed onto total length to test for allometry. Digital pictures were processed with tpsDig and further analyzed with MorphoJ utilizing a regular geometric morphometrics procedure using principle component analyses. We examined gut contents for 16 fish of each species. For the silverjaw minnows, we found all jaw variables to exhibit positive allometric growth with increasing total length, while most of the jaw variables for the sand shiner exhibited negative allometric growth with increasing total length. This correlates with an ontogenic prey shift for both species. Sand shiner minnows have been found to be more omnivorous, feeding on algae later in life, while silverjaw minnows undergo a prey shift to larger invertebrates. These species lack oral dentition causing an increased reliance on the pharyngeal apparatus. Principle component analyses revealed elongation of pharyngeal jaw elements in the silverjaw minnows and a relative shortening and bulking of pharyngeal jaws in the sand shiner minnows. The ontogenic dietary shifts observed in these two species provide possible explanation for the morphological changes over ontogeny in jaw elements, which are likely enabling these species to occupy the same habitat with little niche overlap.     

Morphology of the Jaw-Closing Musculature in the Common Wombat (Vombatus ursinus) Using Digital Dissection and Magnetic Resonance Imaging

Wombats are unique among marsupials in having one pair of upper incisors, and hypsodont molars for processing tough, abrasive vegetation. Of the three extant species, the most abundant, the common wombat (Vombatus ursinus), has had the least attention in terms of masticatory muscle morphology, and has never been thoroughly described. Using MRI and digital dissection to compliment traditional gross dissections, the major jaw adductor muscles, the masseter, temporalis and pterygoids, were described. The masseter and medial pterygoid muscles are greatly enlarged compared to other marsupials. This, in combination with the distinctive form and function of the dentition, most likely facilitates processing a tough, abrasive diet. The broad, flat skull and large masticatory muscles are well suited to generate a very high bite force. MRI scans allow more detail of the muscle morphology to be observed and the technique of digital dissections greatly enhances the knowledge obtained from gross dissections.     

Biplanar X-ray motion analysis of the lower jaw movement during incisor interaction and mastication in the beaver (Castor fiber L. 1758)

The first biplanar X-ray motion analysis of mastication and food processing for Castor fiber is presented. While particles are chipped off interaction of incisors involves variable movements of the lower mandible and thus incisors. After jaw opening the tip of the lower incisors can reach different positions anteriorly of the upper incisors. Then the mandible moves upwards and backwards and brings the tips of the incisors into contact. The lower incisors slide along the wear facet of the upper to the ledge when the cheek teeth occlude. The glenoid fossa and lower jaw condyle are in close contact during incisor contact and no transverse movements are observed. Mastication involves interaction of the cheek teeth with no contact of the incisors. When the cheek teeth are in occlusal contact the mandible is moved forward and transverse, or mediolateral. In consecutive power strokes the jaw is moved alternately to the right and left side. When the jaw opens it is brought into a more central but not totally centred position. During mastication the condyles are positioned posteriorly to the glenoid allowing lateral movement of the mandible. The lateral movement is particularly noticeable in the anterior part of the mandible. With the lateral movements of the incisors one glenoid has to move posteriorly, the other anteriorly.     

Finite Element Analysis of the Cingulata Jaw: An Ecomorphological Approach to Armadillo’s Diets

Finite element analyses (FEA) were applied to assess the lower jaw biomechanics of cingulate xenarthrans: 14 species of armadillos as well as one Pleistocene pampathere (11 extant taxa and the extinct forms Vassallia, Eutatus and Macroeuphractus). The principal goal of this work is to comparatively assess the biomechanical capabilities of the mandible based on FEA and to relate the obtained stress patterns with diet preferences and variability, in extant and extinct species through an ecomorphology approach. The results of FEA showed that omnivorous species have stronger mandibles than insectivorous species. Moreover, this latter group of species showed high variability, including some similar biomechanical features of the insectivorous Tolypeutes matacus and Chlamyphorus truncatus to those of omnivorous species, in agreement with reported diets that include items other than insects. It remains unclear the reasons behind the stronger than expected lower jaw of Dasypus kappleri. On the other hand, the very strong mandible of the fossil taxon Vassallia maxima agrees well with the proposed herbivorous diet. Moreover, Eutatus seguini yielded a stress pattern similar to Vassalia in the posterior part of the lower jaw, but resembling that of the stoutly built Macroeuphractus outesi in the anterior part. The results highlight the need for more detailed studies on the natural history of extant armadillos. FEA proved a powerful tool for biomechanical studies in a comparative framework.     

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.     

Morphology and evolution of the ectethmoid-mandibular articulation in the meliphagidae (Aves)

The ectethmoid-mandibular articulation in Melithreptus and Manorina (Meliphagidae: Aves) consists of the dorsal mandibular process fitting into and abutting against the ventral ectethmoid fossa; it forms a brace for the mandible. This articulation in Melithreptus is a typical diarthrosis with long folded capsular walls. The mandible, thus, has two separate articulations, each with a different axis of rotation. No other genus of Meliphagidae (except Ptiloprora) or any other avian family possesses a similar feature. The jaw and tongue musculature of Melithreptus are described. The two muscles opening the jaws are well developed, while those closing the jaws are small. The tongue muscles show no special developments. A large maxillary gland, presumably muscus secreting, covers the ventral surface of the jaw muscles. Its duct opens into the oral cavity just behind the tip of the upper jaw. The frilled tip of the tongue rests against the duct opening. The ectethmoid-mandibular articulation braces the adducted mandible against dorsoposteriorly directed forces. The mandible can be held closed without a compression force exerted by the mandible on the quadrate, permitting the bird to raise its upper jaw with greater ease and less loss of force. The tongue can be protruded through the slight gap between the jaws, moving against the duct opening and thus be coated with mucus. Presumably, these birds capture insects with their sticky tongue. Hence, the ectethmoid-mandibular articulation is an adaptation for this feeding method; it evolved independently in three genera of the Meliphagidae. The ectethmoid-mandibular articulation demonstrates that a bone can have two articulations with different axes of rotation, that the two articular halves can separate widely, and that articular cartilages can be flat and remain in contact over a large area. Its function suggests that the basitemporal articulation of the mandible found in many other birds has a similar function. And it demonstrates that in the evolution of the mammalian dentary-squamosal articulation, the new hinge did not have to lie on the same rotational axis as the existing quadrate-articular hinge.     

Masticatory motor patterns in ungulates: a quantitative assessment of jaw-muscle coordination in goats, alpacas and horses

We investigated patterns of jaw-muscle coordination during rhythmic mastication in three species of ungulates displaying the marked transverse jaw movements typical of many large mammalian herbivores. In order to quantify consistent motor patterns during chewing, electromyograms were recorded from the superficial masseter, deep masseter, posterior temporalis and medial pterygoid muscles of goats, alpacas and horses. Timing differences between muscle pairs were evaluated in the context of an evolutionary model of jaw-muscle function. In this model, the closing and food reduction phases of mastication are primarily controlled by two distinct muscle groups, triplet I (balancing-side superficial masseter and medial pterygoid and working-side posterior temporalis) and triplet II (working-side superficial masseter and medial pterygoid and balancing-side posterior temporalis), and the asynchronous activity of the working- and balancing-side deep masseters. The three species differ in the extent to which the jaw muscles are coordinated as triplet I and triplet II. Alpacas, and to a lesser extent, goats, exhibit the triplet pattern whereas horses do not. In contrast, all three species show marked asynchrony of the working-side and balancing-side deep masseters, with jaw closing initiated by the working-side muscle and the balancing-side muscle firing much later during closing. However, goats differ from alpacas and horses in the timing of the balancing-side deep masseter relative to the triplet II muscles. This study highlights interspecific differences in the coordination of jaw muscles to influence transverse jaw movements and the production of bite force in herbivorous ungulates.     

Temporalis and masseter muscle function during incision in macaques and humans

Most previously published electromyographic (EMG) studies have indicated that the temporalis muscles in humans become almost electrically quiet during incisai biting. These data have led various workers to conclude that these muscles may contribute little to the incisai bite force. The feeding behavior and comparative anatomy of the incisors and temporalis muscles of certain catarrhine primates, however, suggest that the temporalis muscle is an important and powerful contributor to the bite force during incision. One purpose of this study is to analyze the EMG activity of the masseter and temporalis muscles in both humans and macaques with the intention of focusing on the conflict between published EMG data on humans and inferences of muscle function based on the comparative anatomy and behavior of catarrhine primates. The EMG data collected from humans in the present study indicate that, in five of seven subjects, the masseter,anterior temporalis, and posterior temporalis muscles are very active during apple incision (i.e., relative to EMG activity levels during apple and almond mastication). In the other two human subjects the EMG levels of these muscles are lower during incision than during mastication, but in no instance are these muscles ever close to becoming electrically quiet. The EMG data on macaques indicate that, in all six subjects, the masseter, anterior temporalis, and posterior temporalis muscles are very active during incision. These data are in general agreement with inferences on muscle function that have been drawn from the comparative anatomy and behavior of primates, but they do not agree with previous experimental data. The reason for this disagreement is probably due to differences in the experimental procedure. In previous studies subjects simply bit isometrically on their incisors and the resulting EMG pattern was compared to the pattern associated with powerful clenching in centric occlusion. In the present study the subjects incised into actual food objects, and the resulting EMG pattern was compared to the pattern associated with mastication of various foods. It is not surprising that these two procedures result in markedly different EMG patterns, which in turn result in markedly different interpretations of jaw-muscle function. In an attempt to explain the evolution of the postorbital septum in anthropoids, it has been suggested that the anterior temporalis is more active than the masseter during incision (Cachel, 1979). The human and macaque EMG data do not support this hypothesis; during incision, the two muscles show no consistent differences in humans and the masseter appears to be in fact more active than the anterior temporalis in macaques.     

Comparative functional morphology of mandibular forward movement during mastication of two murid rodents,Apodemus speciosus (Murinae) and Clethrionomys rufocanus (Arvicolinae)

The anatomy of the masticatory apparatus, the direction in which masticatory muscles act during mastication, and jaw muscle forces as estimated by muscle dry weight are compared between two murid rodents, the Japanese field mouse (Apodemus speciosus; subfamily Murinae) and the gray red-backed vole (Clethrionomys rufocanus; subfamily Arvicolinae). The occlusal forces exerted by the deep masseter and the anterior temporalis are large in C. rufocanus. Furthermore, in this species, the angle between the sagittal plane and the occlusal plane of the cheek teeth is larger than in A. speciosus. Therefore, a relatively large occlusal force can be generated in C. rufocanus. The estimated line of action of the anterior temporalis differs markedly between these two species. The functional significance of this difference is discussed relative to the adaptive dental characteristics for food processing, the forces required to masticate different types of food, and the forces that control mandibular forward movement. J Morphol 231:131–141, 1997. © 1997 Wiley-Liss, Inc.     

Jaw Function in Smilodon fatalis: A Reevaluation of the Canine Shear-Bite and a Proposal for a New Forelimb-Powered Class 1 Lever Model

The jaw function of Smilodon fatalis has long been a source of debate. Although modern-day lions subdue large prey through the use of a suffocating throat bite, the dramatically elongated maxillary canines of S. fatalis suggest an alternative bite mechanism. The current literature favors a “canine shear-bite,” in which the depression of the cranium by the ventral neck flexors assists the mandibular adductors in closing the jaws. Although the model makes intuitive sense and appears to be supported by scientific data, the mechanical feasibility of “neck-powered” biting has not been experimentally demonstrated. In the present study, the computer-assisted manipulation of digitized images of a high-quality replica of an S. fatalis neck and skull shows that a rotation of the cranium by the ventral neck flexors will not result in jaw closure. Instead, the cranium and mandible rotate ventrally together (at the atlantooccipital joint), and the jaws remain in an open configuration. The only manner by which rotation of the cranium can simultaneously result in jaw closure is by an anterior rotation at the temporomandibular joint. Based on this finding, the author proposes a new Class 1 lever mechanism for S. fatalis jaw function. In this model, the mandible is immobilized against the neck of the prey and a dorsally directed force from the extension of the forelimbs rotates the cranium anteriorly at the temporomandibular joint. The maxillary canines pierce the prey’s neck and assist in clamping the ventral neck structures. The model is based on a maximum gape angle of approximately 90° and incorporates a secondary virtual point of rotation located slightly anteroventral to the temporomandibular joint. The Class 1 Lever Model is mechanically feasible, consistent with current data on S. fatalis anatomy and ecology, and may provide a basis for similar studies on other fossil taxa.