The generalized carnivore jaw

A model is presented of the jaw mechanism that relies on the geometrical similarities among mammalian carnivores with carnassial teeth. These similarities, together with estimates of the location of the resultant force of the jaw muscles, allow the model to predict that the mechanical advantage of the jaw lever system is the same in all carnivores with carnassials and, therefore, that the magnitude of the bite force is mainly determined by the absolute amount of jaw musculature.     

Correlation of carnassial tooth size and body weight in recent carnivores (mammalia)

The correlation between the lower carnassial crown area and the body weight is examined for modern carnivores. It is very high, but the ursids and felids show a relationship different from the other carnivore families. Some exceptions are discussed. The possibility of predicting body weight for fossils is given.     

Morphological and functional diversity of the mandible in suckermouth armored catfishes (Siluriformes: Loricariidae)

We examined the mandibles of 377 individuals representing 25 species, 12 genera, 5 tribes, and 2 subfamilies of the Loricariidae, a species‐rich radiation of detritivorous–herbivorous neotropical freshwater fishes distinguished by having a ventral oral disk and jaws specialized for surface attachment and benthic feeding. Loricariid mandibles are transversely oriented and bilaterally independent, each rotating predominantly around its long axis, although rotational axes likely vary with mandibular geometry. On each mandible, we measured three traditional and three novel morphological parameters chosen primarily for their functional relevance. Five parameters were linear distances and three of these were analogous to traditional teleost in‐ and out‐levers for mandibular adduction. The sixth parameter was insertion area of the combined adductor mandibulae muscle (AM_area), which correlated with adductor mandibulae volume across a subset of taxa and is interpreted as being proportional to maximum force deliverable to the mandible. Multivariate analysis revealed distributions of phylogenetically diagnosed taxonomic groupings in mandibular morphospace that are consistent with an evolutionary pattern of basal niche conservatism giving rise to multiple adaptive radiations within nested clades. Correspondence between mandibular geometry and function was explored using a 3D model of spatial relationships among measured parameters, potential forces, and axes of rotation. By combining the model with known loricariid jaw kinematics, we developed explicit hypotheses for how individual parameters might relate to each other during kinesis. We hypothesize that the ratio [AM_area/tooth row length²] predicts interspecific variation in the magnitude of force entering the mandible per unit of substrate contacted during feeding. Other newly proposed metrics are hypothesized to predict variation in aspects of mandibular mechanical advantage that may be specific to Loricariidae and perhaps shared with other herbivorous and detritivorous fishes. 2011. © 2011Wiley Periodicals, Inc.     

Using linkage models to explore skull kinematic diversity and functional convergence in arthrodire placoderms

Biomechanical models offer a powerful set of tools for quantifying the diversity of function across fossil taxa. A computer‐based four‐bar linkage model previously developed to describe the potential feeding kinematics of Dunkleosteus terrelli is applied here to several other arthrodire placoderm taxa from different lineages. Arthrodire placoderms are a group of basal gnathostomes showing one of the earliest diversifications of jaw structures. The linkage model allows biomechanical variation to be compared across taxa, identify trends in skull morphology among arthrodires that potentially influence function and explore the role of linkage systems in the early evolution of jaw structures. The linkage model calculates various kinematic metrics including gape angle, effective mechanical advantage, and kinematic transmission coefficients. Results indicate that the arthrodire feeding system may be more diverse and complex than previously thought. A range of potential kinematic profiles among arthrodire taxa illustrate a diversity of feeding function comparable with modern teleost fishes. Previous estimates of bite force in Dunkleosteus are revised based on new morphological data. High levels of kinematic transmission among arthrodires suggest the potential for rapid gape expansion and possible suction feeding. Morphological comparisons indicate that there were several morphological solutions for obtaining these fast kinematics, which allowed different taxa to achieve similar kinematic profiles while varying other aspects of the feeding apparatus. Mapping of key morphological components of the linkage system on a general placoderm phylogeny illustrates the potential importance of four‐bar systems to the early evolution of jaw structures. J. Morphol. 271:990–1005, 2010. © 2010 Wiley‐Liss, Inc.     

Comparative and developmental functional morphology of the jaws of living and fossil gars (Actinopterygii: Lepisosteidae)

The feeding mechanism of gars (Ginglymodi : Lepisosteidae) is characterized by cranial elevation and lower jaw rotation but minimal cranial kinesis. Gar jaws have numerous, sharply pointed, elongate teeth for capture of evasive prey. Their mandibles range from relatively short to extremely long depending on the species. Jaw length and lever dimensions were hypothesized to affect the biomechanics of force and motion during feeding, according to simple mechanical models of muscles exerting force through first- or third-order levers. A morphometric protocol was used to measure the jaw structure of seven living and five fossil species of gar and these data were used to calculate the mechanical advantage (a measure of force transmission) for both opening and closing of the mandible. Gars were found to possess low mechanical advantage (MA) and high transmission of motion, although gars occupy a range of biomechanical states across the continuum of force vs. velocity transmission. The long-nose gar, Lepisosteus osseus, has one of the lowest jaw closing MAs (0.05) ever measured in fishes. Intraspecific lever mechanics were also calculated for a developmental series (from feeding larvae to adults) of L. osseus and Atractosteus spatula. A characteristic ontogenetic curve in MA of the lower jaw was obtained, with a large decrease in MA between larva and juvenile, followed by a steady increase during adult growth. This curve correlates with a change in prey type, with the small, robust-jawed individuals feeding mainly on crustaceans and insects and the large, long-jawed individuals of all species becoming mainly piscivorous. Principal components analysis of functionally important morphometrics shows that several gar species occupy different regions of functional morphospace. Some fossil gar species are also placed within functional morphospace using this approach.     

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.     

A longirostrine Temnodontosaurus (Ichthyosauria) with comments on Early Jurassic ichthyosaur niche partitioning and disparity

Abstract: We describe an almost complete ichthyosaur skeleton from the middle Toarcian (Lower Jurassic) of the Beaujolais foothills near Lyon, France, and assign it to Temnodontosaurus azerguensis sp. nov. This new species exhibits cranial peculiarities such as a thin, elongated and possibly edentulous rostrum, as well as a reduced quadrate. These characters indicate dietary preferences that markedly differ from other species referred to Temnodontosaurus, a genus previously considered as the top predator of the Early Jurassic seas. Despite a conservative postcranial skeleton, we propose that Temnodontosaurus is one of the most ecologically disparate genera of ichthyosaurs, including apex predators and now a soft prey longirostrine hunter. Ammonites collected from the same stratigraphic level as the described specimen indicate that the new species is somewhat younger (bifrons ammonite zone) than the most known Toarcian ichthyosaurs and therefore slightly postdates the interval of severe environmental changes and marine invertebrate extinctions known as the Toarcian Oceanic Anoxic Event. The present study therefore raises the question of whether postcrisis recovery of vertebrate faunas, including the radiation of Temnodontosaurus into a new ecological niche, may have been a consequence of marine ecosystem reorganization across this event.     

Dry versus wet and gross: Comparisons between the dry skull method and gross dissection in estimations of jaw muscle cross-sectional area and bite forces in sea otters

Bite force is a measure of feeding performance used to elucidate links between animal morphology, ecology, and fitness. Obtaining live individuals for in vivo bite-force measurements or freshly deceased specimens for bite force modeling is challenging for many species. Thomason's dry skull method for mammals relies solely on osteological specimens and, therefore, presents an advantageous approach that enables researchers to estimate and compare bite forces across extant and even extinct species. However, how accurately the dry skull method estimates physiological cross-sectional area (PCSA) of the jaw adductor muscles and theoretical bite force has rarely been tested. Here, we use an ontogenetic series of southern sea otters (Enhydra lutris nereis) to test the hypothesis that skeletomuscular traits estimated from the dry skull method accurately predicts test traits derived from dissection-based biomechanical modeling. Although variables from these two methods exhibited strong positive relationships across ontogeny, we found that the dry skull method overestimates PCSA of the masseter and underestimates PCSA of the temporalis. Jaw adductor in-levers for both jaw muscles and overall bite force are overestimated. Surprisingly, we reveal that sexual dimorphism in craniomandibular shape affects temporalis PCSA estimations; the dry skull method predicted female temporalis PCSA well but underestimates male temporalis PCSA across ontogeny. These results highlight the importance of accounting for sexual dimorphism and other intraspecific variation when using the dry skull method. Together, we found the dry skull method provides an underestimation of bite force over ontogeny and that the underlying anatomical components driving bite force may be misrepresented.     

The tooth of perfection: functional and spatial constraints on mammalian tooth shape

This paper addresses the question of how close mammalian teeth are to ideal functional forms. An 'ideal' form is a morphology predicted to be the best functional shape according to information of the relationships between shape and function. Deviations from an ideal form are likely to indicate the presence of developmental or genetic constraints on form. Model tools were constructed to conform to functional principles from engineering and dental studies. The final model shapes are very similar to several mammalian tooth forms (carnassial teeth and tribosphenic-like cusps), suggesting that these tooth forms very closely approach ideal functional forms. Further evidence that these tooth forms are close to ideal comes from the conservation over 140 million years, the independent derivation and/or the occurrence over a size range of several orders of magnitude of these basic tooth forms. One of the main functional shapes derived here is the 'protoconoid', a fundamental design for double-bladed tools that fits a large number of functional parameters. This shape occurs in tooth forms such as tribosphenic, dilambdodont and zalambdodont. This study extends our understanding of constraints on tooth shape in terms of geometry (how space influences tooth shape) and function (how teeth divide food).  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 173–191.     

Basicranial axis length v. skull length in analysis of carnivore skull shape

Skull length is the measurement most commonly used as a standard against which other aspects of cranial morphology are compared to derive an index of relative size or proportions. However, skull length is composed of two different functional components, facial skull and cerebral skull, which vary independently and have different scaling relationships with body size. An analysis of carnivore skull shape with measurements standardized against basicranium length produced very different results than an analysis using skull length as the standard. For example, expressions of relative size of cranial measurements were reduced by 13% in mustelids and increased by 20% in canids, reflecting removal of jaw length (short in mustelids and long in canids) from the comparative standard (basicranial axis length). Cranial measurements scale with higher allometric exponents against basicranial axis length than against skull length.     

A new psittacosaur from Inner Mongolia and the parrot-like structure and function of the psittacosaur skull

We describe a new species of psittacosaur, Psittacosaurus gobiensis, from the Lower Cretaceous of Inner Mongolia and outline a hypothesis of chewing function in psittacosaurs that in many respects parallels that in psittaciform birds. Cranial features that accommodate increased bite force in psittacosaurs include an akinetic skull (both cranium and lower jaws) and differentiation of adductor muscle attachments comparable to that in psittaciform birds. These and other features, along with the presence of numerous large gastroliths, suggest that psittacosaurs may have had a high-fibre, nucivorous (nut-eating) diet.Psittacosaurs, alone among ornithischians, generate oblique wear facets from tooth-to-tooth occlusion without kinesis in either the upper or lower jaws. This is accomplished with a novel isognathous jaw mechanism that combines aspects of arcilineal (vertical) and propalinal (horizontal) jaw movement. Here termed clinolineal (inclined) jaw movement, the mechanism uses posteriorly divergent tooth rows, rather than kinesis, to gain the added width for oblique occlusion. As the lower tooth rows are drawn posterodorsally into occlusion, the increasing width between the upper tooth rows accommodates oblique shear. With this jaw mechanism, psittacosaurs were able to maintain oblique shearing occlusion in an akinetic skull designed to resist high bite forces.     

A functional analysis of how frogs jump out of water

Euphlyctis cyanophlyctis and E. hexadactylus are two common frog species from south-east Asia. Both species have the remarkable ability to leap from a floating position out of the water. Rana esculenta is a European species from the same family that barely manages to do so. The species' morphology, however, looks roughly the same. We studied the unique abilities of the Euphlyctis species by videotaping the three species mentioned above whilst they jumped out of the water. These recordings enabled us to study the hind leg movements and to test several hypotheses. Differences between the species only became apparent when details of the behaviour were analysed. The three species mainly differed in their slip factor at the start of the propulsive phase, resulting in a difference in slip distance. This difference could be traced back to the difference in web shape by means of a physical model. The Euphlyctis species can prevent their feet from slipping because of their rounded and convex webbing, resulting in a larger take-off velocity and thereby a successful emergence out of the water.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 413–420.     

Feeding mechanics in Triassic stem-group sauropterygians: the anatomy of a successful invasion of Mesozoic seas

The jaw adductor musculature in Triassic stem-group sauropterygians is reconstructed on the basis of a paradigmatic model of muscle architecture (functional equivalence of sarcomeres) and using invariant traits of the anatomy of the trigeminal jaw adductor muscles in extant reptiles. The reconstructed jaw adductor musculature predicts trophic specializations in stem-group sauropterygians. Suction feeding is a component in prey capture for some benthic feeding, as well as for some pelagic feeding taxa. The differentiation of 'pincer' jaws is correlated with the potential for rapid, snapping bites. There is some evidence for habitat partitioning among Triassic stem-group sauropterygians with respect to trophic specialization. © 2002 The Linnean Society of London. Zoological Journal of the Linnean Society , 2002, 135 , 33–63.     

Microsatellite analysis of kinkajou social organization

Kinkajou social groups generally consist of one adult female, two males, one subadult and one juvenile. Based on analysis of variation in 11 microsatellite loci, we assess the degree of kinship within and between four social groups totaling 25 kinkajous. We use exclusion and likelihood analyses to assign parents for seven of the eight offspring sampled, five with >/= 95% certainty, and two with >/= 80% certainty. Five of six identified sires of group offspring came from the same social group as the mother and pup. Adult males and females within a group were unrelated and subadults and juveniles were offspring of the group adults, suggesting a family structure. All five identified paternities within a social group were by the dominant male of the group. However, this copulation asymmetry does not necessarily reflect cooperation due to kinship ties between the two adult males within a group as one of two adult male pairs sampled was unrelated. Neighbouring male kinkajous were more closely related to each other than neighbouring female kinkajous, suggesting that females disperse more often or farther than males.     

Ontogeny of postcanine tooth form in the ferret,Mustela putorius (Carnivora: Mammalia), and the evolution of dental diversity within the mustelidae

This study describes dental development within the ferret, Mustela putorius, through study of the form of the carnassial teeth and the upper first molar at progressive growth stages. Primordial teeth were serially sectioned in sagittal and transverse planes and three-dimensional reconstructions of tooth primordia were generated using MacReco software. Regional growth of the crown and asynchronous maturation of the dental tissues were observed in each tooth. The upper carnassial blade develops early and the tooth increases in length rapidly. Lingual growth of the upper carnassial is less pronounced and the protocone and its surrounding region mature late. The lower carnassial blade develops early and the talonid is late to mature. Development of the upper first molar differs from carnassial development in the early emphasis upon transverse growth and reduced lengthwise expansion. The early development of the carnassial blades in the ferret is shared with other carnivores, and may reflect the functional significance of this feature. Later stages of tooth ontogeny differ among carnivoran taxa and the specialized morphology of ferret teeth results from an apparently truncated period of late tooth ontogeny. This suggests that carnivoran species may share a common path of early development that specifies the ontogeny of homologous tooth features and that in later stages developmental differences result in species-specific tooth forms. J. Morphol. 237:69–90, 1998. © 1998 Wiley-Liss, Inc.     

Jaw lever analysis of Hawaiian gobioid stream fishes: A simulation study of morphological diversity and functional performance

Differences in feeding behavior and performance among the five native Hawaiian gobioid stream fishes (Sicyopterus stimpsoni, Lentipes concolor, Awaous guamensis, Stenogobius hawaiiensis, and Eleotris sandwicensis) have been proposed based on the skeletal anatomy of their jaws and dietary specialization. However, performance of the feeding apparatus likely depends on the proportions and configurations of the jaw muscles and the arrangement of the jaw skeleton. We used a published mathematical model of muscle function to evaluate potential differences in jaw closing performance and their correlations with morphology among these species. For example, high output force calculated for the adductor mandibulae muscles (A2 and A3) of both A. guamensis and E. sandwicensis matched expectations based on the morphology of these species because these muscles are larger than in the other species. In contrast, Stenogobius hawaiiensis exhibited an alternative morphological strategy for achieving high relative output forces of both A2 and A3, in which the placement and configuration of the muscles conveyed high mechanical advantage despite only moderate cross‐sectional areas. These differing anatomical pathways to similar functional performance suggest a pattern of many‐to‐one mapping of morphology to performance. In addition, a functional differentiation between A2 and A3 was evident for all species, in which A2 was better suited for producing forceful jaw closing and A3 for rapid jaw closing. Thus, the diversity of feeding performance of Hawaiian stream gobies seems to reflect a maintenance of functional breadth through the retention of some primitive traits in combination with novel functional capacities in several species. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.