Evolution of cranial shape in a continental‐scale evolutionary radiation of Australian lizards

A defining character of adaptive radiations is the evolution of a diversity of morphological forms that are associated with the use of different habitats, following the invasion of vacant niches. Island adaptive radiations have been thoroughly investigated but continental scale radiations are more poorly understood. Here, we use 52 species of Australian agamid lizards and their Asian relatives as a model group, and employ three‐dimensional geometric morphometrics to characterize cranial morphology and investigate whether variation in cranial shape reflects patterns expected from the ecological process of adaptive radiation. Phylogenetic affinity, evolutionary allometry, and ecological life habit all play major roles in the evolution of cranial shape in the sampled lizards. We find a significant association between cranial shapes and life habit. Our results are in line with the expectations of an adaptive radiation, and this is the first time detailed geometric morphometric analyses have been used to understand the selective forces that drove an adaptive radiation at a continental scale

Visualising muscle anatomy using three-dimensional computer models - an example using the head and neck muscles of Sphenodon

We demonstrate how the computer-based technique of multi-body dynamics analysis (MDA) can be used to create schematic, but informative three-dimensional (3D) representations of complex muscle anatomy. As an example we provide an overview of the head and neck muscles present in Sphenodon (Diapsida: Lepidosauria: Rhynchocephalia). First a computer model based on micro-computed tomography datasets provides a detailed and anatomically correct three-dimensional (3D) framework to work from. Secondly, muscles are represented by groups of cylinders that can be colour coded as desired. This allows muscle positions, attachment areas, and 3D orientation to be visualised clearly. This method has advantages over imaging techniques such as two-dimensional drawings and permits the form and function of the muscles to be understood in a way that is not always possible with more classical visualisation techniques. Copyright: Palaeontological Association December 2009

The head and neck muscles associated with feeding in sphenodon (Reptilia: Lepidosauria: Rhynchocephalia)

Feeding in Sphenodon, the tuatara of New Zealand, is of interest for several rea-sons. First, the modern animal is threatened by extinction, and some populations are in competition for food with Pacific rats. Second, Sphenodon demonstrates a feeding apparatus that is unique to living amniotes: an enlarged palatine tooth row, acrodont dentition, enlarged incisor-like teeth on the premaxilla, a posterior extension of the dentary and an elongate articular surtace that permits prooral shearing. Third, Spheno-don has a skull with two complete lateral temporal bars and is therefore structurally analogous to the configuration hypothesised for the ancestral diapsid reptile. Further-more, the fossil relatives of Sphenodon demonstrate considerable variation in terms of feeding apparatus and skull shape. Lastly, as Sphenodon is the only extant rhyn-chocephalian it represents a potentially useful reference taxon for both muscle recon-struction in extinct reptile taxa and determination of muscle homology in extant taxa. Here we provide an up-to-date consensus view of osteology and musculature in Sphenodon that is relevant to feeding. Discrepancies within previous descriptions are evaluated and synthesised with new observations. This paper displays the complex muscle arrangement using a range of different imaging techniques and a variety of different angles. This includes photographs, illustrations, schematic diagrams, and microcomputed tomography (micro-CT) slice images. © Palaeontological Association August 2009

Ontogenetic allometry underlies trophic diversity in sea turtles (Chelonioidea)

Despite only comprising seven species, extant sea turtles (Cheloniidae and Dermochelyidae) display great ecological diversity, with most species inhabiting a unique dietary niche as adults. This adult diversity is remarkable given that all species share the same dietary niche as juveniles. These ontogenetic shifts in diet, as well as a dramatic increase in body size, make sea turtles an excellent group to examine how morphological diversity arises by allometric processes and life habit specialisation. Using three-dimensional geometric morphometrics, we characterise ontogenetic allometry in the skulls of all seven species and evaluate variation in the context of phylogenetic history and diet. Among the sample, the olive ridley (Lepidochelys olivacea) has a seemingly average sea turtle skull shape and generalised diet, whereas the green (Chelonia mydas) and hawksbill (Eretmochelys imbricata) show different extremes of snout shape associated with their modes of food gathering (grazing vs. grasping, respectively). Our ontogenetic findings corroborate previous suggestions that the skull of the leatherback (Dermochelys coriacea) is paedomorphic, having similar skull proportions to hatchlings of other sea turtle species and retaining a hatchling-like diet of relatively soft bodied organisms. The flatback sea turtle (Natator depressus) shows a similar but less extreme pattern. By contrast, the loggerhead sea turtle (Caretta caretta) shows a peramorphic signal associated with increased jaw muscle volumes that allow predation on hard shelled prey. The Kemp’s ridley (Lepidochelys kempii) has a peramorphic skull shape compared to its sister species the olive ridley, and a diet that includes harder prey items such as crabs. We suggest that diet may be a significant factor in driving skull shape differences among species. Although the small number of species limits statistical power, differences among skull shape, size, and diet are consistent with the hypothesis that shifts in allometric trajectory facilitated diversification in skull shape as observed in an increasing number of vertebrate groups

A new rhynchocephalian from the late jurassic of Germany with a dentition that is unique amongst tetrapods.

Rhynchocephalians, the sister group of squamates (lizards and snakes), are only represented by the single genus Sphenodon today. This taxon is often considered to represent a very conservative lineage. However, rhynchocephalians were common during the late Triassic to latest Jurassic periods, but rapidly declined afterwards, which is generally attributed to their supposedly adaptive inferiority to squamates and/or Mesozoic mammals, which radiated at that time. New finds of Mesozoic rhynchocephalians can thus provide important new information on the evolutionary history of the group. A new fossil relative of Sphenodon from the latest Jurassic of southern Germany, Oenosaurus muehlheimensis gen. et sp. nov., presents a dentition that is unique amongst tetrapods. The dentition of this taxon consists of massive, continuously growing tooth plates, probably indicating a crushing dentition, thus representing a previously unknown trophic adaptation in rhynchocephalians. The evolution of the extraordinary dentition of Oenosaurus from the already highly specialized Zahnanlage generally present in derived rhynchocephalians demonstrates an unexpected evolutionary plasticity of these animals. Together with other lines of evidence, this seriously casts doubts on the assumption that rhynchocephalians are a conservative and adaptively inferior lineage. Furthermore, the new taxon underlines the high morphological and ecological diversity of rhynchocephalians in the latest Jurassic of Europe, just before the decline of this lineage on this continent. Thus, selection pressure by radiating squamates or Mesozoic mammals alone might not be sufficient to explain the demise of the clade in the Late Mesozoic, and climate change in the course of the fragmentation of the supercontinent of Pangaea might have played a major role

Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards, snakes, and tuatara)

Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologically important group of over 9,000 reptile species. The earliest fossil records are currently restricted to the Late Triassic and often dated to 227 million years ago (Mya). As these early records include taxa that are relatively derived in their morphology (e.g. Brachyrhinodon), an earlier unknown history of Lepidosauria is implied. However, molecular age estimates for Lepidosauria have been problematic; dates for the most recent common ancestor of all lepidosaurs range between approximately 226 and 289 Mya whereas estimates for crown-group Squamata (lizards and snakes) vary more dramatically: 179 to 294 Mya. This uncertainty restricts inferences regarding the patterns of diversification and evolution of Lepidosauria as a whole. Results Here we report on a rhynchocephalian fossil from the Middle Triassic of Germany (Vellberg) that represents the oldest known record of a lepidosaur from anywhere in the world. Reliably dated to 238–240 Mya, this material is about 12 million years older than previously known lepidosaur records and is older than some but not all molecular clock estimates for the origin of lepidosaurs. Using RAG1 sequence data from 76 extant taxa and the new fossil specimens two of several calibrations, we estimate that the most recent common ancestor of Lepidosauria lived at least 242 Mya (238–249.5), and crown-group Squamata originated around 193 Mya (176–213). Conclusion A Early/Middle Triassic date for the origin of Lepidosauria disagrees with previous estimates deep within the Permian and suggests the group evolved as part of the faunal recovery after the end-Permain mass extinction as the climate became more humid. Our origin time for crown-group Squamata coincides with shifts towards warmer climates and dramatic changes in fauna and flora. Most major subclades within Squamata originated in the Cretaceous postdating major continental fragmentation. The Vellberg fossil locality is expected to become an important resource for providing a more balanced picture of the Triassic and for bridging gaps in the fossil record of several other major vertebrate groups

Neutron scanning reveals unexpected complexity in the enamel thickness of an herbivorous Jurassic reptile

Eilenodontines are one of the oldest radiation of herbivorous lepidosaurs (snakes, lizards and tuatara) characterized by batteries of wide teeth with thick enamel that bear mammal-like wear facets. Unlike most reptiles, eilenodontines have limited tooth replacement, making dental longevity particularly important to them. We use both X-ray and neutron computed tomography to examine a fossil tooth from the eilenodontine Eilenodon (Late Jurassic, USA). Of the two approaches, neutron tomography was more successful and facilitated measurements of enamel thickness and distribution. We find the enamel thickness to be regionally variable, thin near the cusp tip (0.10 mm) but thicker around the base (0.15–0.30 mm) and notably greater than that of other rhynchocephalians such as the extant Sphenodon (0.08–0.14 mm). The thick enamel in Eilenodon would permit greater loading, extend tooth lifespan and facilitate the establishment of wear facets that have sharp edges for orally processing plant material such as horsetails (Equisetum). The shape of the enamel dentine junction indicates that tooth development in Eilenodon and Sphenodon involved similar folding of the epithelium but different ameloblast activity

The impact of Stieltjes' work on continued fractions and orthogonal polynomials

Stieltjes' work on continued fractions and the orthogonal polynomials related to continued fraction expansions is summarized and an attempt is made to describe the influence of Stieltjes' ideas and work in research done after his death, with an emphasis on the theory of orthogonal polynomials

Financial and monetary policy responses to oil price shocks: evidence from oil-importing and oil-exporting countries

In this study, we investigate the financial and monetary policy responses to oil price shocks using a Structural VAR framework. We distinguish between net oil-importing and net oil-exporting countries. Since the 80s, a significant number of empirical studies have been published investigating the effect of oil prices on macroeconomic and financial variables. Most of these studies though, do not make a distinction between oil-importing and oil-exporting economies. Overall, our results indicate that the level of inflation in both net oil-exporting and net oil-importing countries is significantly affected by oil price innovations. Furthermore, we find that the response of interest rates to an oil price shock depends heavily on the monetary policy regime of each country. Finally, stock markets operating in net oil-importing countries exhibit a negative response to increased oil prices. The reverse is true for the stock market of the net oil-exporting countries. We find evidence that the magnitude of stock market responses to oil price shocks is higher for the newly established and/or less liquid stock market