Post-weaning maternal effects and the evolution of female dominance in the spotted hyena

Mammalian societies in which females dominate males are rare, and the factors favouring the evolution of female dominance have yet to be clearly identified. We propose a new hypothesis for the evolution of female dominance and test its predictions with empirical data from the spotted hyena (Crocuta crocuta), a well-studied species characterized by female dominance. We suggest that constraints imposed by the development of a feeding apparatus specialized for bone cracking, in combination with the intensive feeding competition characteristic of spotted hyenas, led to the evolution of female dominance. Specifically, we propose that protracted development of the feeding apparatus in young hyenas led to selection for increased aggressiveness in females as a compensatory mechanism for mothers to secure food access for their young after weaning. Our analyses yielded results consistent with this hypothesis. Morphological and behavioural measurements indicate that skull development is indeed protracted in this species; spotted hyenas do not achieve adult skull size or feeding performance capabilities until after sexual maturity. The period between weaning and completed skull development is particularly challenging, as indicated by high mortality. Finally, maternal presence between weaning and full skull maturity, as well as the relative ability of females to aggressively displace conspecifics from food, are important determinants of offspring survival.     

Ancient mitochondrial genomes from Chinese cave hyenas provide insights into the evolutionary history of the genus Crocuta

Cave hyenas (genus Crocuta) are extinct bone-cracking carnivores from the family Hyaenidae and are generally split into two taxa that correspond to a European/Eurasian and an (East) Asian lineage. They are close relatives of the extant African spotted hyenas, the only extant member of the genus Crocuta. Cave hyenas inhabited a wide range across Eurasia during the Pleistocene, but became extinct at the end of the Late Pleistocene. Using genetic and genomic datasets, previous studies have proposed different scenarios about the evolutionary history of Crocuta. However, causes of the extinction of cave hyenas are widely speculative and samples from China are severely understudied. In this study, we assembled near-complete mitochondrial genomes from two cave hyenas from northeastern China dating to 20 240 and 20 253 calBP, representing the youngest directly dated fossils of Crocuta in Asia. Phylogenetic analyses suggest a monophyletic clade of these two samples within a deeply diverging mitochondrial haplogroup of Crocuta. Bayesian analyses suggest that the split of this Asian cave hyena mitochondrial lineage from their European and African relatives occurred approximately 1.85 Ma (95% CI 1.62–2.09 Ma), which is broadly concordant with the earliest Eurasian Crocuta fossil dating to approximately 2 Ma. Comparisons of mean genetic distance indicate that cave hyenas harboured higher genetic diversity than extant spotted hyenas, brown hyenas and aardwolves, but this is probably at least partially due to the fact that their mitochondrial lineages do not represent a monophyletic group, although this is also true for extant spotted hyenas. Moreover, the joint female effective population size of Crocuta (both cave hyenas and extant spotted hyenas) has sustained two declines during the Late Pleistocene. Combining this mitochondrial phylogeny, previous nuclear findings and fossil records, we discuss the possible relationship of fossil Crocuta in China and the extinction of cave hyenas.     

The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave,Germany: specialised cave bear predators of Europe

Remains of 13 individuals with 3/1 male/female ratio of the extinct Upper Pleistocene lion Panthera leo spelaea (Goldfuss, 1810) from the Zoolithen Cave near Burggeilenreuth (Bavaria, Germany) include the holotype skull and all paratype material. The highest mortality rate for the Zoolithen Cave lions is in their reproductive adult ages. Bite marks on lion bones or skulls are results of hyena activities, or rare cannibalism of lions under stress situations. Lions were possibly also killed in battles with cave bears during predation on hibernating bears in winter times. This cave bear hunt specialisation in caves overlaps with the ecological behaviour of cave bear feeding by Ice Age-spotted hyenas. Both largest Ice Age predators, lions and hyenas, had to specialise on feeding herbivorous cave bears in boreal forest mountainous cave rich regions, where the mammoth steppe megafauna prey was absent. This cave bear hunt by felids, and scavenging by hyenas and other large carnivores such as leopards and wolves explains why cave bears hibernated deep in to the European caves, for protection reasons against predators. Within such lion–cave bear and even lion–hyena conflicts in the caves lions must have been killed sometimes, explaining mainly the skeleton occurrences in different European caves.     

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.     

A quantitative approach to the cranial ontogeny of <Emphasis Type="Italic">Lycalopex culpaeus</Emphasis> (Carnivora: Canidae)

The study of cranial ontogeny is important for understanding the relationship between form and function in developmental, ecological, and evolutionary contexts. The transition from lactation to the diet of adult carnivores must be accompanied by pronounced modifications in skull morphology and feeding behavior. Our goal was to study relative growth and development in the skull ontogeny of the canid Lycalopex culpaeus, and interpret our findings in a functional context, thereby exploring the relationship between changes in shape and size with dietary habits and age stages. We performed quantitative analyses, including multivariate allometry and geometric morphometrics. Our results indicate that shape changes are related to functional improvements of the jaw mechanics related for food catching/processing. Estimates of full muscle size, mechanical advantage, and adult cranial shape are reached after sexual maturity, while adult mandible and skull size are reached after weaning, which is related to diet change (incorporation of meat and other food items). The ontogenetic pattern observed in L. culpaeus is similar to those observed in Canis familiaris and C. latrans. However, the magnitude of change seen in L. culpaeus is smaller than those seen in the felid Puma concolor and considerably smaller than those seen in the bone cracker hyaenid Crocuta crocuta. These patterns are associated with dietary habits and specializations in skull anatomy, as L. culpaeus, domestic dog and coyote are generalist species compared with hypercarnivores such as C. crocuta and P. concolor.     

Feeding biomechanics of five demersal Antarctic fishes

There is scant information available on the ecomorphology of Antarctic fishes, and especially on their feeding capabilities. We measured interspecific variation in mechanical advantage (MA), force-producing capability, and suction index for the jaws of the five dominant taxa of high-Antarctic fishes: the nototheniid Trematomus bernacchii; the zoarcids Pachycara brachycephalum, Lycodichthys dearborni, and Ophthalmolycus amberensis; and the liparid Paraliparis devriesi. Analysis of variance indicated significant differences in jaw metrics, and ordinations of morphological traits identified three loosely defined groups reflecting their family-level taxonomy. Principal component analyses showed distinct segregation between the nototheniid and the liparid, indicating that they are at the extremes of the feeding performance continuum. The zoarcids fell in the middle, suggesting that they utilize a combination of feeding modes to capture prey. The liparid had the lowest MA and bite force, but a large epaxialis implied a ram-suction-feeding mode. The large adductor mandibulae in the zoarcids P. brachycephalum and L. dearborni suggest that they are capable of grasping mobile prey and manipulating sedentary, hard-shelled macroinvertebrates. The zoarcids had a smaller epaxialis than the liparid and may not be as efficient as suction-feeders. Values for mechanical advantage ratios and suction indices in Antarctic fishes were within the range known for non-Antarctic fishes. The five Antarctic species do not possess dentition specialized for durophagous feeding; however, the high mechanical advantage ratio in the nototheniid and, to a lesser extent, in the zoarcids, suggests that durophagy may be possible.     

SKULL SHAPE EVOLUTION IN DUROPHAGOUS CARNIVORANS

In this article, we investigate convergent evolution toward durophagy in carnivoran skull shape using geometric morphometrics in a sample of living and extinct species. Principal components analysis indicate that, in spite of the different dietary resources consumed by durophages—that is, bone‐crackers and bamboo‐feeders—both groups of carnivorans share portions of skull phenotypic spaces. We identify by discriminant analyses a shared set of adaptations toward durophagy in the skull of carnivores. However, ancestral states indicate that although durophages reached similar phenotypes, the evolutionary pathways that they followed are different depending upon the family to which they belong. Furthermore, while the carnivoran cranium more closely reflects the nature of the resources consumed—that is, soft or hard and tough items—the mandible shows particular feeding adaptations—that is, bamboo or bone. This finding supports the interpretation that the mandible has more evolutionary plasticity than the cranium, which is more limited to evolve toward a particular feeding adaptation. However, we find that the shapes of the cranium and the mandible are highly integrated for the whole order Carnivora. Published studies of teratological cats and dogs indicate that the role of internal constraints in shaping this pattern of integration is absent or weak and malleable by selection.     

Feeding mechanisms of the sauropod dinosaurs Brachiosaurus,Camarasaurus, Diplodocus and Dicraeosaurus

Skull morphologies and dental wear patterns have been examined in four sauropod genera to evaluate their probable feeding mechanisms. Wear facets on teeth are generally confined to their apices in Brachiosaurus and Dicraeosaurus and they are sometimes also present on the mesial and distal carinae. Skull morphology and dental wear patterns in Diplodocus and Dicraeosaurus are consistent with a raking motion of the jaws during feeding. Diplodocus became mechanically adapted to feed in this way by evolving anteriorly directed teeth in the premaxilla and mesial parts of the maxilla, and by changing the direction of jaw adduction relative to the long axis of the skull. Similar features are present in the few known skulls of Apatosaurus and they may also have been present in Dicraeosaurus. In Brachiosaurus dental wear patterns also imply a raking motion of the jaws, although the more robust skull and teeth and the more vertically directed action of the jaw adductor muscles have led some to suggest the possibility of isognathous occlusion. Camarasaurus employed a powerful bite in its feeding, possibly with slight propaliny of the lower jaw, and its skull was modified to cope with increased stresses arising from mastication. Archaic sauropods appear largely to have employed isognathic occlusion in chopping off vegetation. The raking motion employed by diplodocids and dicraeosaurids was an advanced mode of cropping and stripping, linked evolutionarily to their highly apomorphic cranial morphology.     

Three-dimensional computer simulations of feeding behaviour in red and giant pandas relate skull biomechanics with dietary niche partitioning

The red (Ailurus fulgens) and giant (Ailuropoda melanoleuca) pandas are mammalian carnivores convergently adapted to a bamboo feeding diet. However, whereas Ailurus forages almost entirely on younger leaves, fruits and tender trunks, Ailuropoda relies more on trunks and stems. Such difference in foraging mode is considered a strategy for resource partitioning where they are sympatric. Here, we use finite-element analysis to test for mechanical differences and similarities in skull performance between Ailurus and Ailuropoda related to diet. Feeding simulations suggest that the two panda species have similar ranges of mechanical efficiency and strain energy profiles across the dentition, reflecting their durophagous diet. However, the stress distributions and peaks in the skulls of Ailurus and Ailuropoda are remarkably different for biting at all tooth locations. Although the skull of Ailuropoda is capable of resisting higher stresses than the skull of Ailurus, the latter is able to distribute stresses more evenly throughout the skull. These differences in skull biomechanics reflect their distinct bamboo feeding preferences. Ailurus uses repetitive chewing in an extended mastication to feed on soft leaves, and Ailuropoda exhibits shorter and more discrete periods of chomp-and-swallow feeding to break down hard bamboo trunks.     

A clan of Late Pleistocene hyenas,Crocuta crocuta spelaea (Goldfuss 1823), from the Rösenbeck Cave (Germany) and a contribution to cranial shape variability

The remains of a large population of Late Pleistocene Ice Age spotted hyenas (Crocuta crocuta spelaea Goldfuss 1823) are described from the Rösenbeck Cave in the Sauerland Karst of Germany. They include four skulls and 79 other skeletal parts, mainly from adult to senile animals, making this an important Late Pleistocene hyena cave‐den site in Europe. The skulls have been compared with 30 other hyena skull specimens from open air and cave‐den sites in central Europe (Germany, Austria, the Czech Republic and Romania) in order to achieve an understanding of sexual dimorphism in the crania of Ice Age spotted hyenas from the Upper Pleistocene cold period (Weichselian/Wuermian), and the types of injuries that they acquired during their lifetimes. Three different types of cranial shape have been distinguished, one of which appears to have been a consequence of pathologies that developed in response to injuries caused by bites received during the animal's lifetime, as a result of either intraspecies fights or fights with lions. Although cave bears penetrated to great depths within the Rösenbeck Cave for hibernation purposes, hyenas appear to have utilized only a short section of the cave that branched off directly from the entrance area. Hyena cub material is scarce, suggesting that this area was used as a communal den rather than for cub rearing. Bones exhibiting gnaw marks, representing prey imported by hyenas, are also rare but include horse (Equus caballus przewalskii) and reindeer (Rangifer tarandus) remains. The scarcity of bones from hyena prey suggests that this cave was not used as a food storage site. Some Ursus spelaeus cave bear remains, including skulls, show evidence of having been gnawed, chewed and cracked by hyenas, indicating that the hyenas periodically fed on cave bear carcasses in a specialization response to the mammoth steppe megafauna absence of the boreal mountain forest regions. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 191–220.     

Functional relationship between skull form and feeding mechanics in Sphenodon, and implications for diapsid skull development

The vertebrate skull evolved to protect the brain and sense organs, but with the appearance of jaws and associated forces there was a remarkable structural diversification. This suggests that the evolution of skull form may be linked to these forces, but an important area of debate is whether bone in the skull is minimised with respect to these forces, or whether skulls are mechanically "over-designed" and constrained by phylogeny and development. Mechanical analysis of diapsid reptile skulls could shed light on this longstanding debate. Compared to those of mammals, the skulls of many extant and extinct diapsids comprise an open framework of fenestrae (window-like openings) separated by bony struts (e.g., lizards, tuatara, dinosaurs and crocodiles), a cranial form thought to be strongly linked to feeding forces. We investigated this link by utilising the powerful engineering approach of multibody dynamics analysis to predict the physiological forces acting on the skull of the diapsid reptile Sphenodon. We then ran a series of structural finite element analyses to assess the correlation between bone strain and skull form. With comprehensive loading we found that the distribution of peak von Mises strains was particularly uniform throughout the skull, although specific regions were dominated by tensile strains while others were dominated by compressive strains. Our analyses suggest that the frame-like skulls of diapsid reptiles are probably optimally formed (mechanically ideal: sufficient strength with the minimal amount of bone) with respect to functional forces; they are efficient in terms of having minimal bone volume, minimal weight, and also minimal energy demands in maintenance.     

Convergence and functional evolution of longirostry in crocodylomorphs

During the Mesozoic, Crocodylomorpha had a much higher taxonomic and morphological diversity than today. Members of one particularly successful clade, Thalattosuchia, are well‐known for being longirostrine: having long, slender snouts. It has generally been assumed that Thalattosuchia owed their success in part to the evolution of longirostry, leading to a feeding ecology similar to that of the living Indian gharial, Gavialis. Here, we compare form and function of the skulls of the thalattosuchian Pelagosaurus and Gavialis using digital reconstructions of the skull musculoskeletal anatomy and finite element models to show that they had different jaw muscle arrangements and biomechanical behaviour. Additionally, the relevance of feeding‐related mandibular traits linked to longirostry in the radiation of crocodylomorph clades was investigated by conducting an evolutionary rates analysis under the variable rates model. We find that, even though Pelagosaurus and Gavialis share similar patterns of stress distribution in their skulls, the former had lower mechanical resistance. This suggests that compared to Gavialis, Pelagosaurus was unable to process large, mechanically less tractable prey, instead operating as a specialized piscivore that fed on softer and smaller prey. Secondly, innovation of feeding strategies was achieved by rate acceleration of functional characters of the mandible, a key mechanism for the diversification of certain clades like thalattosuchians and eusuchians. Different rates of functional evolution suggest divergent diversification dynamics between teleosaurids and metriorhynchids in the Jurassic.     

Ontogenetic relationships between cranium and mandible in coyotes and hyenas

Developing animals must resolve the conflicting demands of survival and growth, ensuring that they can function as infants or juveniles while developing toward their adult form. In the case of the mammalian skull, the cranium and mandible must maintain functional integrity to meet the feeding needs of a juvenile even as the relationship between parts must change to meet the demands imposed on adults. We examine growth and development of the cranium and mandible, using a unique ontogenetic series of known‐age coyotes (Canis latrans), analyzing ontogenetic changes in the shapes of each part, and the relationship between them, relative to key life‐history events. Both cranial and mandibular development conform to general mammalian patterns, but each also exhibits temporally and spatially localized maturational transformations, yielding a complex relationship between growth and development of each part as well as complex patterns of synchronous growth and asynchronous development between parts. One major difference between cranium and mandible is that the cranium changes dramatically in both size and shape over ontogeny, whereas the mandible undergoes only modest shape change. Cranium and mandible are synchronous in growth, reaching adult size at the same life‐history stage; growth and development are synchronous for the cranium but not for the mandible. This synchrony of growth between cranium and mandible, and asynchrony of mandibular development, is also characteristic of a highly specialized carnivore, the spotted hyena (Crocuta crocuta), but coyotes have a much less protracted development, being handicapped relative to adults for a much shorter time. Morphological development does not predict life‐history events in these two carnivores, which is contrary to what has been reported for two rodent species. The changes seen in skull shape in successive life‐history stages suggest that adult functional demands cannot be satisfied by the morphology characterizing earlier life‐history stages. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Functional morphogenesis from embryos to adults: Late development shapes trophic niche in coral reef damselfishes

The damselfishes are one of the dominant coral reef fish lineages. Their ecological diversification has involved repeated transitions between pelagic feeding using fast bites and benthic feeding using forceful bites. A highly‐integrative approach that combined gene expression assays, shape analyses, and high‐speed video analyses was used to examine the development of trophic morphology in embryonic, larval, juvenile, and adult damselfishes. The anatomical characters that distinguish pelagic‐feeding and benthic‐feeding species do not appear until after larval development. Neither patterns of embryonic jaw morphogenesis, larval skull shapes nor larval bite mechanics significantly distinguished damselfishes from different adult trophic guilds. Analyses of skull shape and feeding performance identified two important transitions in the trophic development of a single species (the orange clownfish; Amphiprion percula): (a) a pronounced transformation in feeding mechanics during metamorphosis; and (b) more protracted cranial remodeling over the course of juvenile development. The results of this study indicate that changes in postlarval morphogenesis have played an important role in damselfish evolution. This is likely to be true for other fish lineages, particularly if they consist of marine species, the majority of which have planktonic larvae with different functional requirements for feeding in comparison to their adult forms.     

Dispersal Status Influences Hormones and Behavior in the Male Spotted Hyena

Male spotted hyenas (Crocuta crocuta) reach puberty at 24 months of age and then invariably emigrate from their natal clans 1 to 38 months later. Thus there are two classes of reproductively mature males in everyCrocutaclan: adult natal males born in the clan and adult immigrant males born elsewhere. In one free-living hyena population in Kenya, these two groups of males were compared with respect to measures of aggression, social dominance, sexual behavior, and circulating hormone levels. Adult natal males engaged in higher hourly rates of aggression than did immigrants, won all fights with immigrants, and were socially dominant to immigrants. In addition, adult natal males engaged in far lower hourly rates of sexual behavior with resident females than did immigrants, and natal males were never observed to copulate with natal females. Mean basal plasma cortisol values did not differ between the two groups of adult males, but cortisol concentrations in immigrants were positively correlated with tenure in the clan and with immigrant male social rank. Adult natal males had plasma testosterone levels significantly lower than those of immigrants. Social rank and plasma testosterone values were positively correlated among immigrant males. Thus two different relationships appear to exist between circulating testosterone and social rank in maleCrocuta:one apparent in immigrants and the other in natal adult males. Our results suggest that dispersal might disinhibit testosterone secretion in postpubertal male hyenas.     

The role of cranial kinesis in birds.

In birds, the ability to move the upper beak relative to the braincase has been the subject of many functional morphological investigations, but in many instances the adaptive significance of cranial kinesis remains unclear. Alternatively, cranial kinesis may be considered a consequence of the general design of the skull, rather than an adaptive trait as such. The present study reviews some results related to the mechanism and functional significance of cranial kinesis in birds. Quantitative three-dimensional X-ray has shown that in skulls morphologically as divers as paleognaths and neognaths the mechanism for elevation of the upper beak is very similar. One of the mechanisms proposed for avian jaw movement is a mechanical coupling of the upper and the lower jaw movement by the postorbital ligament. Such a mechanical coupling would necessitate upper beak elevation. However, independent control of upper and lower jaw has been shown to occur during beak movements in birds. Moreover, kinematic modeling and force measurements suggests that the maximum extensibility of collagen, in combination with the short distance of the insertion of the postorbital ligament to the quadrato-mandibular articulation do not constitute a block to lower jaw depression. The lower jaw ligaments serve to limit the maximal extension of the mandibula. It is suggested here that cranial kinesis in avian feeding may have evolved as a consequence of an increase in eye size. This increase in size led to a reduction of bony bars in the lateral aspect of the skull enabling the transfer of quadrate movement to the upper jaw. The selective forces favoring the development of a kinetic upper beak in birds may be subtle and act in different ecological contexts. Simultaneous movement of the upper and lower jaw not only increases the velocity of beak movements, but with elevated upper beak also less force is required to open the lower jaw. However, the penalty of increased mobility of elements in a lightweight skull and a large eye is potential instability of skull elements during biting, smaller bite forces and limitations on joint reaction forces. Such a lightly built, kinetic skull may have evolved in animals that feed on small plant material or insects. This type of food does not require the resistance of large external forces on the jaws as in carnivores eating large prey.     

Testing hypotheses of convergence with multivariate data: morphological and functional convergence among herbivorous lizards

Abstract Despite its importance to evolutionary theory, convergence remains an understudied phenomenon and is usually investigated using qualitative data. This paper advances a new, multidimensional view of convergence. Three patterns indicative of convergence are discussed, and techniques to discover and test convergent patterns in a quantitative framework are developed. These concepts and methods are applied to a dataset of digitized coordinates on 1554 lizard skulls and 1292 lower jaws to test hypotheses of convergence among herbivorous lizards. Encompassing seven independent acquisitions of herbivory, this lizard sample provides an ideal natural experiment for exploring ideas of convergence among different systems (here, morphological and functional). Three related questions are addressed: (1) Do herbivorous lizards show evidence of convergence in skull and lower jaw morphology? (2) What, if any, is the morphospace pattern associated with this convergence? (3) Is it possible to predict the direction of convergence using functional models? Relative warp analysis and permutation tests reveal that the skulls and lower jaws of herbivorous lizards do show evidence of convergence. Herbivore skulls deviate from their carnivorous or omnivorous sister groups toward the same area of morphospace. Without a phylogenetic perspective, this pattern would not be recognizable. Lower jaws of herbivores are not convergent in morphology but are convergent in function: herbivores deviate away from their carnivorous sister groups toward higher values of mechanical advantage. These results illustrate the desirability of quantitative methods, informed by phylogenetic information, in the study of convergence.     

Form and function of damselfish skulls: rapid and repeated evolution into a limited number of trophic niches

Background  

Damselfishes (Perciformes, Pomacentridae) are a major component of coral reef communities, and the functional diversity of their trophic anatomy is an important constituent of the ecological morphology of these systems. Using shape analyses, biomechanical modelling, and phylogenetically based comparative methods, we examined the anatomy of damselfish feeding among all genera and trophic groups. Coordinate based shape analyses of anatomical landmarks were used to describe patterns of morphological diversity and determine positions of functional groups in a skull morphospace. These landmarks define the lever and linkage structures of the damselfish feeding system, and biomechanical analyses of this data were performed using the software program JawsModel4 in order to calculate the simple mechanical advantage (MA) employed by different skull elements during feeding, and to compute kinematic transmission coefficients (KT) that describe the efficiency with which angular motion is transferred through the complex linkages of damselfish skulls.     

Mandibular morphology and diet in the genusCebus

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

Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace

Rhynchocephalians were a successful, globally distributed group of diapsid reptiles that thrived in the Mesozoic. Multiple species of Clevosaurus existed worldwide in the Late Triassic and Early Jurassic, characterized by shearing bladelike teeth perhaps functionally analogous to the carnassial teeth of mammals. Morphometric analysis shows that the dentary morphospace of clevosaurs differs significantly from that of other rhynchocephalians. Five Clevosaurus species occupied islands in the Bristol Channel archipelago of the UK, but generally not those occupied by mammaliaforms, suggesting dietary character displacement. Identifying the diet of such ancient, small tetrapods has been difficult. To identify the nature of their feeding mechanics and ecology, we apply finite element analysis to two near complete three-dimensional skulls of the species Clevosaurus hudsoni and Clevosaurus cambrica to estimate bite force, resistance to bending and torsion and the distribution of stresses in the jaws during biting. Both species had bite forces and tooth pressures sufficient to break apart chitin indicating that, like early Mesozoic mammaliaforms, clevosaurs could feed on tough-shelled beetles and possibly small vertebrates. In addition, the mechanical advantage of the jaws falls within the range of early mammaliaforms, so though we cannot demonstrate niche partitioning between members of the two clades, it raises the prospect that they may have been functionally similar.