Cranial mechanics and feeding in Tyrannosaurus rex

It has been suggested that the large theropod dinosaur Tyrannosaurus rex was capable of producing extremely powerful bite forces and resisting multi-directional loading generated during feeding. Contrary to this suggestion is the observation that the cranium is composed of often loosely articulated facial bones, although these bones may have performed a shock-absorption role. The structural analysis technique finite element analysis (FEA) is employed here to investigate the functional morphology and cranial mechanics of the T. rex skull. In particular, I test whether the skull is optimized for the resistance of large bi-directional feeding loads, whether mobile joints are adapted for the localized resistance of feeding-induced stress and strain, and whether mobile joints act to weaken or strengthen the skull overall. The results demonstrate that the cranium is equally adapted to resist biting or tearing forces and therefore the 'puncture-pull' feeding hypothesis is well supported. Finite-element-generated stress-strain patterns are consistent with T. rex cranial morphology: the maxilla-jugal suture provides a tensile shock-absorbing function that reduces localized tension yet 'weakens' the skull overall. Furthermore, peak compressive and shear stresses localize in the nasals rather than the fronto-parietal region as seen in Allosaurus, offering a reason why robusticity is commonplace in tyrannosaurid nasals.     

Correlations between lizard cranial shape and diet: a quantitative, phylogenetically informed analysis

Although the relationship between dietary and phenotypic specialization has been well documented for many vertebrate groups, it has been stated that few such general trends can be established for lizards. This is often thought to be due to the lack of dietary specialization in many lizards. For example, many species that are reported to be insectivorous may also consume a variety of plant materials, and the reverse is often true as well. In this study, we investigate whether a correlation exists between general cranial form and dietary niche in lizards. Additionally, we test previously proposed hypotheses suggesting that herbivorous lizards should be larger bodied than lizards with other diets. Our data indicate that lizards specializing in food items imposing different mechanical demands on the feeding system show clear patterns of morphological specialization in their cranial morphology. True herbivores (diet of fibrous and tough foliage) are clearly distinguished from omnivorous and carnivorous lizards by having taller skulls and shorter snouts, likely related to the need for high bite forces. This allows herbivores to mechanically reduce relatively less digestible foliage. Carnivores have relatively longer snouts and retroarticular processes, which may result in more efficient capture and processing of elusive prey. When analysed in an explicit phylogenetic context, only snout length and skull mass remained significantly different between dietary groups. The small number of differences in the phylogenetic analyses is likely the result of shared evolutionary history and the relative paucity of independent origins of herbivory and omnivory in our sample. Analyses of the relationship between diet and body size show that on average herbivores have a larger body size than carnivores, with omnivores intermediate between the two other dietary groups. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 433–466.     

Ecological and evolutionary factors in the morphological diversification of South American spiny rats

Understanding the processes underlying morphological diversification is a central goal in ecology and evolutionary biology and requires the integration of information about phylogenetic divergence and ecological niche diversity. In the present study, we use geometric morphometrics and comparative methods to investigate morphological diversification in Neotropical spiny rats of the family Echimyidae. Morphological diversification is studied as shape variation in the skull, comprising a structure composed of four distinct units: vault, base, orognathofacial complex, and mandible. We demonstrate association among patterns of variation in shape in different cranial units, levels of phylogenetic divergence, and ecological niche diversification. At the lower level of phylogenetic divergence, there is significant and positive concordance between patterns of phylogenetic divergence and cranial shape variation in all cranial units. This concordance may be attributable to the phylogenetic and shape distances being calculated between species that occupy the same niche. At higher phylogenetic levels of divergence and with ecological niche diversity, there is significant concordance between shape variation in all four cranial units and the ecological niches. In particular, the orognathofacial complex revealed the most significant association between shape variation and ecological niche diversity. This association may be explained by the great functional importance of the orognathofacial complex.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 646–660.     

The evolution of cranial form and function in theropod dinosaurs: insights from geometric morphometrics

Theropod dinosaurs, an iconic clade of fossil species including Tyrannosaurus and Velociraptor, developed a great diversity of body size, skull form and feeding habits over their 160+ million year evolutionary history. Here, we utilize geometric morphometrics to study broad patterns in theropod skull shape variation and compare the distribution of taxa in cranial morphospace (form) to both phylogeny and quantitative metrics of biting behaviour (function). We find that theropod skulls primarily differ in relative anteroposterior length and snout depth and to a lesser extent in orbit size and depth of the cheek region, and oviraptorosaurs deviate most strongly from the "typical" and ancestral theropod morphologies. Noncarnivorous taxa generally fall out in distinct regions of morphospace and exhibit greater overall disparity than carnivorous taxa, whereas large-bodied carnivores independently converge on the same region of morphospace. The distribution of taxa in morphospace is strongly correlated with phylogeny but only weakly correlated with functional biting behaviour. These results imply that phylogeny, not biting function, was the major determinant of theropod skull shape.     

Is the morphological disparity of lemur skulls (Primates) controlled by phylogeny and/or environmental constraints?

The cranial morphology of Lemur catta and of the five species of the genus Eulemur is investigated here by landmark identification and Procrustes superimpositions. This geometrical morphometric method makes it possible to describe pure shape differences independent of size effects, and to quantify differences between specimens. The aim of this study is to determine whether the morphological disparity of lemur skulls is constrained by the environment and/or by the group's phylogenetic history. First, Procrustes residuals are analysed by principal component analysis and the scatter-plots interpreted against the geographical distributions of taxa to determine whether morphology is correlated with geography. Then, a morphological distance tree is computed and compared with various cladograms reported in the literature to test for any correlation between morphology and phylogeny. Morphological disparity is found to be closely correlated with geographical distribution but independent of phylogeny. This confirms that the morphological disparity of lemur skulls is associated with a high degree of homoplasy, probably as a result of ecological constraints.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 577–590.     

The relationship between skull morphology, biting performance and foraging mode in Kalahari lacertid lizards

Lizards are a diverse clade in which one radiation consists entirely of sit-and-wait foragers and another consists of wide foragers. Lizards utilizing these two foraging modes are known to differ in diet, but little is known about how feeding morphology relates to diet and/or foraging mode. This study tested the hypothesis that skull morphology and biting performance are related to diet preference, and consequently, coevolve with foraging mode. Four species of lacertid lizard were studied because they vary in foraging mode, their phylogenetic relationships are known and they are well studied ecologically. Using an 'ecomorphological' approach, skull morphology and biting performance were quantified and mapped on to the phylogeny for the species. The results indicate that sit-and-wait species have shorter, wider skulls than the wide foraging species, and that all are significantly different in overall head shape. The sit-and-wait species had similar values for biting performance; however, clear phylogenetic patterns of covariation were not present between sit-and-wait and wide foraging species for either biting performance or skull morphology. Thus, skull morphology and performance have little influence on diet and foraging mode in these species. Instead it is likely that other factors such as seasonal prey availability and/or life history strategy shape foraging mode decisions.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 403–416.     

The cranial skeleton of the Early Permian aquatic reptile Mesosaurus tenuidens: implications for relationships and palaeobiology

The cranial osteology of the aquatic reptile Mesosaurus tenuidens is redescribed on the basis of new and previously examined materials from the Lower Permian of both southern Africa and South America. Mesosaurus is distinguished from other mesosaurs in exhibiting an absolutely larger skull and possessing relatively longer marginal teeth. The teeth gradually angle outwards as one progresses anteriorly in the tooth row and become conspicuously procumbent at the tip of the snout. The suggestion that mesosaurs used their conspicuous dental apparatus as a straining device for filter feeding is based upon erroneous reconstruction of a high number of teeth in this mesosaur. Reinterpretation of the morphology and the organization of the marginal teeth of Mesosaurus suggests that they were used to capture individually small, nektonic prey. General morphological aspects of the skull support the idea that Mesosaurus was an aquatic predator and that the skull was well adapted for feeding in an aqueous environment. The anatomical review permits critical reappraisal of several cranial characters that have appeared in recent phylogenetic analyses of early amniotes. Emendation of problematic characters and reanalysis of amniote phylogeny using a slightly modified data matrix from the literature strengthens the hypothesis that mesosaurs form a clade with millerettids, procolophonoids and pareiasaurs within Reptilia.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 146 , 345–368.     

Are Cranial Biomechanical Simulation Data Linked to Known Diets in Extant Taxa? A Method for Applying Diet-Biomechanics Linkage Models to Infer Feeding Capability of Extinct Species

Performance of the masticatory system directly influences feeding and survival, so adaptive hypotheses often are proposed to explain craniodental evolution via functional morphology changes. However, the prevalence of “many-to-one” association of cranial forms and functions in vertebrates suggests a complex interplay of ecological and evolutionary histories, resulting in redundant morphology-diet linkages. Here we examine the link between cranial biomechanical properties for taxa with different dietary preferences in crown clade Carnivora, the most diverse clade of carnivorous mammals. We test whether hypercarnivores and generalists can be distinguished based on cranial mechanical simulation models, and how such diet-biomechanics linkages relate to morphology. Comparative finite element and geometric morphometrics analyses document that predicted bite force is positively allometric relative to skull strain energy; this is achieved in part by increased stiffness in larger skull models and shape changes that resist deformation and displacement. Size-standardized strain energy levels do not reflect feeding preferences; instead, caniform models have higher strain energy than feliform models. This caniform-feliform split is reinforced by a sensitivity analysis using published models for six additional taxa. Nevertheless, combined bite force-strain energy curves distinguish hypercarnivorous versus generalist feeders. These findings indicate that the link between cranial biomechanical properties and carnivoran feeding preference can be clearly defined and characterized, despite phylogenetic and allometric effects. Application of this diet-biomechanics linkage model to an analysis of an extinct stem carnivoramorphan and an outgroup creodont species provides biomechanical evidence for the evolution of taxa into distinct hypercarnivorous and generalist feeding styles prior to the appearance of crown carnivoran clades with similar feeding preferences.     

Cranial anatomy and variation in Prosaurolophus maximus (Dinosauria: Hadrosauridae)

Prosaurolophus maximus Brown is a saurolophine hadrosaurid known from numerous complete, articulated skulls from the Dinosaur Park Formation (Campanian, Alberta, Canada) that range in size by approximately half a metre in total skull length. Therefore, it is an important taxon for understanding patterns of growth and variation in saurolophines. This study describes the cranial anatomy of P. maximus from the type locality of Dinosaur Provincial Park (Dinosaur Park Formation: Campanian) on the basis of ten articulated skulls, quantitatively examines its range of osteological variation, and provides the first hypothesized ontogenetic series for this taxon. A second species, Prosaurolophus blackfeetensis Horner, was named on the basis of geologically younger material from Montana (Two Medicine Formation: Campanian) that is diagnosed by putative morphological differences in the nasal crest. However, considerable nasal crest variation in the sample from the Dinosaur Park Formation does not permit quantitative differentiation of P. blackfeetensis from P. maximus. Furthermore, a species‐level phylogenetic analysis of saurolophines that includes both P. maximus and P. blackfeetensis as originally defined recovers them as sister taxa that do not differ morphologically in the character matrix. Based on both the morphometric and phylogenetic data, this study supports the hypothesis that P. blackfeetensis is a junior synonym of P. maximus, thereby substantially increasing its temporal range to 1.6 million years, and a concomitant period of morphological stasis in this taxon. © 2013 The Linnean Society of London     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

Postnatal growth of skull linear measurements of Cape Hare Lepus capensis in northern China: an analysis in an adaptive context

304 skulls of Cape hare (Lepus capensis) were collected from two climatically distinct localities in northern China. With eye lens weight as a continuous age variable, postnatal growth patterns of 25 cranial linear measurements in relation to sex, growth season and region were analysed to understand the morphological basis of life history adaptation. In almost all the skull measurements, no significant differences were found between either sex or growth seasons. Principal component analysis revealed that facial elements accounted for the greatest proportion of skull morphological variation. Von Bertalanffy function was applied to describe growth trajectories of the skull elements. Based on this model, the growth rates of skull elements and the age at which they reached a certain proportion (95%) of asymptotes were compared. The results showed that skull growth exhibited an allometric pattern, with neural components attaining their final size more rapidly (at about 2–3 months old in tympanic bulla and 4–6 months old in others) than did the facial, which continued to grow well into postnatal life (at 6–10 months old). The earlier establishment of neurocranial morphology was associated with a fully developed central nervous system, which may play a key role in improving the survival of animals during the early phase of life. There was a regional difference in developmental rate of the hare skull. For all the skull parameters, northern hares had a more rapid rate of cranial growth compared to the southern, i.e. skull elements of juveniles from northern population were relatively larger at comparable ages and achieved adult size 0.5–4.0 months earlier than those from the south. In adult hares, however, no significant regional differences in any of the skull parameters were present. Adaptive explanations for the regional difference in ontogenetic pattern of skull morphology include age‐specific thermoregulation constraint, season‐related food availability and age‐dependent predation pressure. Based on the findings of this study, it is suggested that the postnatal growing period represents a crucial time of life, and that improvement of survivorship when young by growth adaptation forms an important aspect of the hare's life history strategies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 343–353.     

Morphology of the endocranial cavities of Campinasuchus dinizi (Crocodyliformes: Baurusuchidae) from the Upper Cretaceous of Brazil

Two specimens of Campinasuchus dinizi (CPPLIP 1319 and CPPLIP 1360) belonging to Baurusuchidae (Crocodyliformes, Notosuchia) from the Upper Cretaceous Bauru Group of Minas Gerais state (Brazil) were scanned in a Toshiba Aquilion 64 CT machine. Based on these data, it was possible to identify and reconstruct the paranasal sinuses, the nasal cavity proper, the nasopharyngeal duct, the encephalon, the paratympanic sinuses, and the semicircular canals of the inner ear. The paranasal sinuses present similar morphology to those of other mesoeucrocodylians, especially eusuchians. The nasal cavity proper occupies the entire rostral region, with an expansion in the olfactory region. The expansion in the nasal cavity is present in other notosuchians and theropod dinosaurs (e.g., Tyrannosaurus rex Osborn, 1905), but less developed in aquatic crocodilians, which may indicate an olfactory acuity related to terrestrial habits. The encephalon is similar in shape to that of other mesoeucrocodylians. The rostral semicircular canal is smaller than the caudal one, differing from most mesoeucrocodylians. The paratympanic sinuses are more developed in C. dinizi than in eusuchians, being more similar to Tyrannosaurus rex. Campinasuchus dinizi presents few variations in the internal structures of the skull in relation to taxa with different ecological niches, probably indicating that ecological factors do not strongly influence the morphology of these structures.     

Phylogenetic implications of the morphology of the braincase of caecilian amphibians (Gymnophiona)

Caecilian morphology is strongly modified in association with their fossorial mode of life. Currently phylogenetic analyses of characters drawn from the morphology of caecilians lack resolution, as well as complementarity, with results of phylogenetic analyses that employ molecular data. Stemming from the hypothesis derived from the mammal literature that the braincase has the greatest potential (in comparison to other cranial units) to yield phylogenetic information, the braincase and intimately associated stapes of 27 species (23 genera) of extant caecilians were examined using images assembled via microcomputed tomography. Thirty‐four new morphological characters pertaining to the braincase and stapes were identified and tested for congruence with previously recognized morphological characters. The results reveal that when added to previous character matrices, characters of the braincase and stapes resolve generic‐level relationships in a way that is largely congruent with the results of molecular analyses. Analysis of a combined data set of molecular and morphological data provides a framework for conducting ancestral character state reconstructions, which resulted in the identification of 95 new synapomorphies for various clades and taxa, 27 of which appear to be unique for the taxa that possess them. Together these data demonstrate the utility of the application of characters of the braincase and stapes for resolving phylogenetic relationships for a group whose morphology is largely confounded by functional modifications. In addition this study provides evidence of the utility of the braincase in resolving problematic morphology‐based phylogeny outside of Amniota, in an amphibian group. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 160–201.     

Cranial shape variation and molecular phylogenetic structure of crested newts (Triturus cristatus superspecies: Caudata,Salamandridae) in the Balkans

In the present study, we investigated the degree of congruence between phylogeny, as inferred from mitochondrial (mt)DNA sequences, and cranium shape variation of crested newts (Triturus cristatus superspecies) in the Balkans. These newts belong to four phylogenetic clades defined by mtDNA analysis, and significantly differed in cranial shape. Allometry explained a high percentage of shape variation in crested newts. The clade‐specific allometric slopes significantly diverged for both the ventral cranium and dorsal cranium, indicating that differences in shape between clades could not be a simple consequence of their difference in size. The analysis of hierarchical and spatial variation showed similarity in the patterns of global and spatially localized hierarchical variation of cranial shape. We also found significant congruence between the pattern of cranial shape variation and molecular phylogeny. The differences in morphology of Triturus dobrogicus in comparison to other crested newt clades, including marked differences in cranium shape, is discussed in the context of the evolution and ecology of crested newts. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 348–360.     

Turtle carotid circulation: a character analysis case study

The bony canals permeating the turtle skull associated with the cranial circulation have long been considered integral to an understanding of extinct and extant turtle systematics. Recent phylogenetic analyses, employing a variety of data sets, suggest alternatives to the traditional arrangement of crown turtles. Of particular note is the recent failure of investigations employing molecular techniques to retrieve a monophyletic Trionychoidea, a clade supported largely by shared circulatory features. These alternative phylogenies may also have implications for extinct forms. The turtle cranial arterial circulation therefore represents an ideal system upon which to conduct a detailed, multipronged analysis of a systematically influential character. In the present study, the theoretical underpinnings of character analysis are critically evaluated, and a new interpretation of circulatory variation in turtles is offered, including a revised phylogenetic character suite. This assessment indicates an autapomorphic circulatory pattern in Trionychia, whereas Trionychoidea is not supported.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 239–256.     

Does nasal echolocation influence the modularity of the mammal skull?

In vertebrates, changes in cranial modularity can evolve rapidly in response to selection. However, mammals have apparently maintained their pattern of cranial integration throughout their evolutionary history and across tremendous morphological and ecological diversity. Here, we use phylogenetic, geometric morphometric and comparative analyses to test the hypothesis that the modularity of the mammalian skull has been remodelled in rhinolophid bats due to the novel and critical function of the nasal cavity in echolocation. We predicted that nasal echolocation has resulted in the evolution of a third cranial module, the ‘nasal dome’, in addition to the braincase and rostrum modules, which are conserved across mammals. We also test for similarities in the evolution of skull shape in relation to habitat across rhinolophids. We find that, despite broad variation in the shape of the nasal dome, the integration of the rhinolophid skull is highly consistent with conserved patterns of modularity found in other mammals. Across their broad geographical distribution, cranial shape in rhinolophids follows two major divisions that could reflect adaptations to dietary and environmental differences in African versus South Asian distributions. Our results highlight the potential of a relatively simple modular template to generate broad morphological and functional variation in mammals.     

The first record of a nyctiphruretid parareptile from the Early Permian of North America,with a discussion of parareptilian temporal fenestration

The Richards Spur Locality of Oklahoma, USA, long known for its highly diverse Early Permian terrestrial tetrapod assemblage, is particularly interesting for the presence of many endemic taxa. The parareptilian component of the assemblage, rare members of other Early Permian communities, is especially diverse at Richards Spur, consisting of six species. The newest parareptile, A byssomedon williamsi gen. et sp. nov. , consists of an articulated left jaw and various disarticulated cranial and postcranial elements. A new phylogenetic analysis of parareptiles, based on an updated modified data matrix revealed that Ab . williamsi is a member of the small clade Nyctiphruretidae. This makes Ab . williamsi the first and oldest nyctiphruretid, a clade of parareptiles otherwise known from the Middle and Late Permian of Russia, extending the age of the clade back into the Early Permian. This discovery also raises the possibility that nyctiphruretids may have dispersed from western Laurasia to eastern Laurasia. The characteristic jugal morphology of Ab . williamsi shows that it would have possessed a slender, deep, temporal emargination. The current topology of Parareptilia indicates that there was considerable variability in the patterns of lateral temporal openings amongst the various members of this clade, suggesting that there may have been multiple, independent modifications of this region of the skull. © 2014 The Linnean Society of London