Marc Jones | University of Adelaide (original) (raw)

Papers by Marc Jones

Journal of the Royal Society, Interface, 2017

The role of soft tissues in skull biomechanics remains poorly understood. Not least, the chondroc... more The role of soft tissues in skull biomechanics remains poorly understood. Not least, the chondrocranium, the portion of the braincase which persists as cartilage with varying degrees of mineralization. It also remains commonplace to overlook the biomechanical role of sutures despite evidence that they alter strain distribution. Here, we examine the role of both the sutures and the chondrocranium in the South American tegu lizardWe use multi-body dynamics analysis (MDA) to provide realistic loading conditions for anterior and posterior unilateral biting and a detailed finite element model to examine strain magnitude and distribution. We find that strains within the chondrocranium are greatest during anterior biting and are primarily tensile; also that strain within the cranium is not greatly reduced by the presence of the chondrocranium unless it is given the same material properties as bone. This result contradicts previous suggestions that the anterior portion (the nasal septum) ac...

Frontiers of Oral Biology, 2009

The Tuatara, Sphenodon, is a small reptile currently restricted to islands off the coast of New Z... more The Tuatara, Sphenodon, is a small reptile currently restricted to islands off the coast of New Zealand where it feeds mainly on arthropods. A widely held misconception is that 'Sphenodon does not have real teeth' and instead possesses 'serrations on the jaw bone'. One hatchling and one adult dentary were examined under SEM. Two longitudinal ground sections 100-microm thick were prepared through a lower canine tooth and its supporting tissues. There was clear evidence of aprismatic enamel (primless enamel) containing dentine tubules crossing the EDJ, dentine, cementum and a basal-bone attachment. Enamel increments averaged approximately 3 microm/day and extension rates were approximately 30 microm/day. The base of the tooth consisted of basal attachment bone that graded from few cell inclusions to lamella or even Haversian-like bone with evidence of remodeling. A string of sclerosed pulp-stone like structures filled the pulp chamber and were continuous with the bone of attachment. Bone beneath the large central nutrient mandibular (Meckel's) canal was quite unlike lamella bone and appeared to be fast growing and to contain wide alternating cell-rich and cell-free zones. Bone cells were rounded (never fusiform) and had few, if any, canaliculi. The dentine close to the EDJ formed at about the same rate as enamel but also contained longer period increments approximately 100 microm apart. These were spaced appropriately for monthly lunar growth bands, which would explain the basis of the banding pattern observed in the fast growing basal bone beneath the Meckel's canal.

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The patella (kneecap) is the largest and best-known of the sesamoid bones, postulated to confer b... more The patella (kneecap) is the largest and best-known of the sesamoid bones, postulated to confer biomechanical advantages including increasing joint leverage and reinforcing the tendon against compression. It has evolved several times independently in amniotes, but despite apparently widespread occurrence in lizards, the patella remains poorly characterised in this group and is, as yet, completely undescribed in their nearest extant relative Sphenodon (Rhynchocephalia). Through radiography, osteological and fossil studies we examined patellar presence in diverse lizard and lepidosauromorph taxa, and using computed tomography, dissection and histology we investigated in greater depth the anatomy and morphology of the patella in 16 lizard species and 19 Sphenodon specimens. We have found the first unambiguous evidence of a mineralised patella in Sphenodon, which appears similar to the patella of lizards and shares several gross and microscopic anatomical features. Although there may be a common mature morphology, the squamate patella exhibits a great deal of variability in development (whether from a cartilage anlage or not, and in the number of mineralised centres) and composition (bone, mineralised cartilage or fibrotendinous tissue). Unlike in mammals and birds, the patella in certain lizards and Sphenodon appears to be a polymorphic trait. We have also explored the evolution of the patella through ancestral state reconstruction, finding that the patella is ancestral for lizards and possibly Lepidosauria as a whole. Clear evidence of the patella in rhynchocephalian or stem lepidosaurian fossil taxa would clarify the evolutionary origin(s) of the patella, but due to the small size of this bone and the opportunity for degradation or loss we could not definitively conclude presence or absence in the fossils examined. The pattern of evolution in lepidosaurs is unclear but our data suggest that the emergence of this sesamoid may be related to the evolution of secondary ossification centres and/or changes in knee joint conformation, where enhancement of extensor muscle leverage would be more beneficial.

BMC Evolutionary Biology

Abstract Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologi... more Abstract 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 alllepidosaurs range between approximately 226 and 289 Mya whereas estimates for crowngroup 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 a...

Bite-force performance is an ecologically important measure of whole-organism performance that sh... more Bite-force performance is an ecologically important measure of whole-organism performance that shapes dietary breadth and feeding strategies and, in some taxa, determines reproductive success. It also is a metric that is crucial to testing and evaluating biomechanical models. We reviewed nearly 100 published studies of a range of taxa that incorporate direct in vivo measurements of bite force. Problematically, methods of data collection and processing vary considerably among studies. In particular, there is little consensus on the appropriate substrate to use on the biting surface of force transducers. In addition, the bite out-lever, defined as the distance from the fulcrum (i.e. jaw joint) to the position along the jawline at which the jaws engage the transducer, is rarely taken into account. We examined the effect of bite substrate and bite out-lever on bite-force estimates in a diverse sample of lizards. Results indicate that both variables have a significant impact on the accuracy of measurements. Maximum bite force is significantly greater using leather as the biting substrate compared with a metal substrate. Less-forceful bites on metal are likely due to inhibitory feedback from mechanoreceptors that prevent damage to the feeding apparatus. Standardization of bite out-lever affected which trial produced maximum performance for a given individual. Indeed, maximum bite force is usually underestimated without standardization because it is expected to be greatest at the minimum out-lever (i.e. back of the jaws), which in studies is rarely targeted with success. We assert that future studies should use a pliable substrate, such as leather, and use appropriate standardization for bite out-lever.

The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In rec... more The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In recent years, many new species have been described and understanding of the history and relationships of this fauna has been greatly advanced by molecular studies, but very little is known of the fossil history of frogs on the island. Beelzebufo ampinga, the first named pre-Holocene frog from Madagascar, was described in 2008 on the basis of numerous disarticulated cranial and postcranial elements from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. These specimens documented the presence of a hyperossified taxon that differed strikingly from extant Malagasy frogs in its large size and heavy coarse cranial exostosis. Here we describe and analyse new, articulated, and more complete material of the skull, vertebral column, and hind limb, as well as additional isolated elements discovered since 2008. μCT scans allow a detailed understanding of both internal and external morphology and permit a more accurate reconstruction. The new material shows Beelzebufo to have been even more bizarre than originally interpreted, with large posterolateral skull flanges and sculptured vertebral spine tables. The apparent absence of a tympanic membrane, the strong cranial exostosis, and vertebral morphology suggest it may have burrowed during seasonally arid conditions, which have been interpreted for the Maevarano Formation from independent sedimentological and taphonomic evidence. New phylogenetic analyses, incorporating both morphological and molecular data, continue to place Beelzebufo with hyloid rather than ranoid frogs. Within Hyloidea, Beelzebufo still groups with the South American Ceratophryidae thus continuing to pose difficulties with both biogeographic interpretations and prior molecular divergence dates.

Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologically ... more 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 at least 11 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 77 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.

Background: To date, the Lower Cretaceous Jehol Group of western Liaoning, China has yielded five... more Background: To date, the Lower Cretaceous Jehol Group of western Liaoning, China has yielded five monotypic genera of anurans, including Liaobatrachus grabaui, Callobatrachus sanyanensis, Mesophryne beipiaoensis, Dalianbatrachus mengi, and Yizhoubatrachus macilentus. However, the validity and distinctness of these taxa have been questioned.

The skull is composed of many bones that come together at sutures. These sutures are important si... more The skull is composed of many bones that come together at sutures. These sutures are important sites of growth, and as growth ceases some become fused while others remain patent. Their mechanical behaviour and how they interact with changing form and loadings to ensure balanced craniofacial development is still poorly understood. Early suture fusion often leads to disfiguring syndromes, thus is it imperative that we understand the function of sutures more clearly. By applying advanced engineering modelling techniques, we reveal for the first time that patent sutures generate a more widely distributed, high level of strain throughout the reptile skull. Without patent sutures, large regions of the skull are only subjected to infrequent low-level strains that could weaken the bone and result in abnormal development. Sutures are therefore not only sites of bone growth, but could also be essential for the modulation of strains necessary for normal growth and development in reptiles.

Abstract The Purbeck Limestone Group of England has yielded a rich assemblage of Late Jurassic to... more Abstract The Purbeck Limestone Group of England has yielded a rich assemblage of Late Jurassic to Early Cretaceous (Berriasian) vertebrate fossils, including one of the most diverse Early Cretaceous lizard assemblages on record. Here we describe the first articulated lizard skull from Purbeck. The specimen was rediscovered in the collections of the British Geological Survey, having been excavated at least a century ago.

What is a tuatara? The tuatara is a modern reptile found only in New Zealand (Figure 1). Although... more What is a tuatara? The tuatara is a modern reptile found only in New Zealand (Figure 1). Although it superficially looks like a medium-sized lizard it is in fact the only living member of the Rhynchocephalia, a group of animals that were successful, globally distributed, and ecologically diverse during the Mesozoic, alongside dinosaurs. A wealth of molecular and fossil evidence shows that Rhynchocephalia are the closest living relatives of Squamata (lizards and snakes), and that the two groups diverged about 250 million years ago. Therefore, humans are more closely related to kangaroos, for example, than the tuatara is to a lizard.

Background: Sea turtles (Chelonoidea) are a charismatic group of marine reptiles that occupy a r... more Background:
Sea turtles (Chelonoidea) are a charismatic group of marine reptiles that occupy a range of important ecological roles. However, the diversity and evolution of their feeding anatomy remain incompletely known.

Methodology/Principal Findings:
Using computed tomography and classical comparative anatomy we describe the cranial anatomy in two sea turtles, the loggerhead (Caretta caretta) and Kemp’s ridley (Lepidochelys kempii), for a better understanding of sea turtle functional anatomy and morphological variation. In both taxa the temporal region of the skull is enclosed by bone and the jaw joint structure and muscle arrangement indicate that palinal jaw movement is possible. The tongue is relatively small, and the hyoid apparatus is not as conspicuous as in some freshwater aquatic turtles. We find several similarities between the muscles of C. caretta and L. kempii, but comparison with other turtles suggests only one of these characters may be derived: connection of the m. adductor mandibulae internus into the Pars intramandibularis via the Zwischensehne. The large fleshy origin of the m. adductor mandibulae externus Pars superficialis from the jugal seems to be a characteristic feature of sea turtles.

Conclusions/Significance:
In C. caretta and L. kempii the ability to suction feed does not seem to be as well developed as that found in some freshwater aquatic turtles. Instead both have skulls suited to forceful biting. This is consistent with the observation that both taxa tend to feed on relatively slow moving but sometimes armoured prey. The broad fleshy origin of the m. adductor mandibulae externus Pars superficialis may be linked to thecheek region being almost fully enclosed in bone but the relationship is complex.

The tuatara (Sphenodon punctatus) is the only living representative of Rhynchocephalia, a group o... more The tuatara (Sphenodon punctatus) is the only living representative of Rhynchocephalia, a group of small vertebrates that originated about 250 million years ago. The tuatara has been referred to as a living fossil; however, the group to which it belongs included a much greater diversity of forms in the Mesozoic. We explore the morphological diversity of Rhynchocephalia and stem lepidosaur relatives (Sphenodon plus 13 fossil relatives) by employing a combination of geometric morphometrics and comparative methods. Geometric morphometrics is used to explore cranium size and shape at interspecific scale, while comparative methods are employed to test association between skull shape and size and tooth number after taking phylogeny into account. Two phylogenetic topologies have been considered to generate a phylomorphospace and quantify the phylogenetic signal in skull shape data, the ancestral state reconstruction as well as morphological disparity using disparity through time plots (DTT). Rhynchocephalia exhibit a significant phylogenetic signal in skull shape that compares well with that computed for other extinct vertebrate groups. A consistent form of allometry has little impact on skull shape evolution while the number of teeth significantly correlates with skull shape also after taking phylogeny into account. The ancestral state reconstruction demonstrates a dramatic shape difference between the skull of Sphenodon and its much larger Cretaceous relative Priosphenodon. Additionally, DTT demonstrates that skull shape disparity is higher between rather than within clades while the opposite applies to skull size and number of teeth. These results were not altered by the use of competing phylogenic hypotheses. Rhynchocephalia evolved as a morphologically diverse group with a dramatic radiation in the Late Triassic and Early Jurassic about 200 million years ago. Differences in size are not marked between species whereas changes in number of teeth are associated with co-ordinated shape changes in the skull to accommodate larger masticatory muscles. These results show that the tuatara is not the product of evolutionary stasis but that it represents the only survivor of a diverse Mesozoic radiation whose subsequent decline remains to be explained.

"The New Zealand tuatara, Sphenodon, has a specialized feeding system in which the teeth of the l... more "The New Zealand tuatara, Sphenodon, has a specialized feeding system in which the teeth of the lower jaw close between two upper tooth rows before sliding forward to slice food apart like a draw cut saw. This shearing action is unique amongst living amniotes but has been compared with the chewing power stroke of mammals. We investigated details of the jaw movement using multibody dynamics analysis of an anatomically accurate three-dimensional computer model constructed from computed tomography scans. The model predicts that a flexible symphysis is necessary for changes in the intermandibular angle that permits prooral (=proal) movement. Models with the greatest symphysial flexibility allow the articulation surface of the articular to follow the quadrate cotyle with the least restriction, and suggest that shearing is accompanied by a long axis rotation of the lower jaws. This promotes precise point loading between the cutting edges of particular teeth, enhancing the effectiveness of the shearing action. Given that Sphenodon is a relatively inactive reptile, we suggest that the link between oral food processing and endothermy has been overstated. Food processing improves feeding efficiency, a consideration
of particular importance when food availability is unpredictable.
Although this feeding mechanism is today limited to Sphenodon, a survey
of fossil rhynchocephalians suggests that it was once more widespread."

The vertebrate skull evolved to protect the brain and sense organs, but with the appearance of ja... more 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.

Rhynchocephalia achieved a global distribution during the Mesozoic but the history of sphenodonti... more Rhynchocephalia achieved a global distribution during the Mesozoic but the history of sphenodontines, the clade containing the extant genus Sphenodon (the New Zealand tuatara), remains poorly understood. Here, we describe a partial maxilla from the Late Cretaceous of Argentina bearing teeth that closely resemble those of modern Sphenodon. This material helps to fill in a notable gap in the fossil history of lepidosaurs because it represents the first evidence of a sphenodontine from South America and increases the number of known Late Cretaceous rhynchocephalian taxa from that region. The morphological disparity encompassed by these records is consistent with suggestions that rhynchocephalians remained diverse in the Late Cretaceous of South America despite a concurrent disappearance from Laurasia. Moreover, the new record supports the hypothesis that sphenodontines were once found widely throughout Gondwana, before its constituent landmasses began to separate about 80 million years ago. The extant Sphenodon probably represents a biogeographic remnant of this distribution, but whether its relatively large size and its ability to remain active at cold temperatures reflects a high latitude ancestry requires further examination.

Although diplodocoid sauropods from Africa and the Americas are well known, their European record... more Although diplodocoid sauropods from Africa and the Americas are well known, their European record remains largely neglected. Here we redescribe Dinheirosaurus lourinhanensis from the Late Jurassic of Portugal. The holotype comprises two posterior cervical vertebrae, the dorsal series and a caudal centrum. Redescription demonstrates its validity on the basis of three autapomorphies: (1) posteriorly restricted ventral keel on posterior cervical vertebrae; (2) three small subcircular fossae posterior to the lateral coel on posterior cervical neural spines; (3) accessory lamina linking the hyposphene with base of the posterior centrodiapophyseal lamina in middle-posterior dorsal vertebrae. Phylogenetic analysis places Dinheirosaurus as the sister taxon to Supersaurus, and this clade forms the sister taxon to other diplodocines. However, this position should be treated with caution as Dinheirosaurus displays several plesiomorphic features absent in other diplodocids (including unbifurcated presacral neural spines, and dorsolaterally projecting diapophyses on dorsal vertebrae) and only four additional steps are required to place Dinheirosaurus outside of Flagellicaudata. We identify Amazonsaurus as the basal-most rebbachisaurid and recover Zapalasaurus outside of the South American Limaysaurinae, suggesting the biogeographic history of rebbachisaurids is more complex than previously proposed. Review of the European diplodocoid record reveals evidence for the earliest known diplodocid, as well as additional diplodocid remains from the Late Jurassic of Spain. A Portuguese specimen, previously referred to Dinheirosaurus, displays strong similarities to Apatosaurus from the contemporaneous Morrison Formation of North America, indicating the presence of a second Late Jurassic Portuguese diplodocid taxon. Along with Dinheirosaurus, these Portuguese remains provide further evidence for a Late Jurassic palaeobiogeographic connection between Europe and North America. No dicraeosaurids are currently known from Europe, but rebbachisaurids are present in the Early Cretaceous, with weak evidence for the earliest known representative from the Late Jurassic of Spain; however, more complete material is required to recognize early members of this clade.

Journal of the Royal Society, Interface, 2017

The role of soft tissues in skull biomechanics remains poorly understood. Not least, the chondroc... more The role of soft tissues in skull biomechanics remains poorly understood. Not least, the chondrocranium, the portion of the braincase which persists as cartilage with varying degrees of mineralization. It also remains commonplace to overlook the biomechanical role of sutures despite evidence that they alter strain distribution. Here, we examine the role of both the sutures and the chondrocranium in the South American tegu lizardWe use multi-body dynamics analysis (MDA) to provide realistic loading conditions for anterior and posterior unilateral biting and a detailed finite element model to examine strain magnitude and distribution. We find that strains within the chondrocranium are greatest during anterior biting and are primarily tensile; also that strain within the cranium is not greatly reduced by the presence of the chondrocranium unless it is given the same material properties as bone. This result contradicts previous suggestions that the anterior portion (the nasal septum) ac...

Frontiers of Oral Biology, 2009

The Tuatara, Sphenodon, is a small reptile currently restricted to islands off the coast of New Z... more The Tuatara, Sphenodon, is a small reptile currently restricted to islands off the coast of New Zealand where it feeds mainly on arthropods. A widely held misconception is that 'Sphenodon does not have real teeth' and instead possesses 'serrations on the jaw bone'. One hatchling and one adult dentary were examined under SEM. Two longitudinal ground sections 100-microm thick were prepared through a lower canine tooth and its supporting tissues. There was clear evidence of aprismatic enamel (primless enamel) containing dentine tubules crossing the EDJ, dentine, cementum and a basal-bone attachment. Enamel increments averaged approximately 3 microm/day and extension rates were approximately 30 microm/day. The base of the tooth consisted of basal attachment bone that graded from few cell inclusions to lamella or even Haversian-like bone with evidence of remodeling. A string of sclerosed pulp-stone like structures filled the pulp chamber and were continuous with the bone of attachment. Bone beneath the large central nutrient mandibular (Meckel's) canal was quite unlike lamella bone and appeared to be fast growing and to contain wide alternating cell-rich and cell-free zones. Bone cells were rounded (never fusiform) and had few, if any, canaliculi. The dentine close to the EDJ formed at about the same rate as enamel but also contained longer period increments approximately 100 microm apart. These were spaced appropriately for monthly lunar growth bands, which would explain the basis of the banding pattern observed in the fast growing basal bone beneath the Meckel's canal.

UCL logo UCL Discovery. ...

The patella (kneecap) is the largest and best-known of the sesamoid bones, postulated to confer b... more The patella (kneecap) is the largest and best-known of the sesamoid bones, postulated to confer biomechanical advantages including increasing joint leverage and reinforcing the tendon against compression. It has evolved several times independently in amniotes, but despite apparently widespread occurrence in lizards, the patella remains poorly characterised in this group and is, as yet, completely undescribed in their nearest extant relative Sphenodon (Rhynchocephalia). Through radiography, osteological and fossil studies we examined patellar presence in diverse lizard and lepidosauromorph taxa, and using computed tomography, dissection and histology we investigated in greater depth the anatomy and morphology of the patella in 16 lizard species and 19 Sphenodon specimens. We have found the first unambiguous evidence of a mineralised patella in Sphenodon, which appears similar to the patella of lizards and shares several gross and microscopic anatomical features. Although there may be a common mature morphology, the squamate patella exhibits a great deal of variability in development (whether from a cartilage anlage or not, and in the number of mineralised centres) and composition (bone, mineralised cartilage or fibrotendinous tissue). Unlike in mammals and birds, the patella in certain lizards and Sphenodon appears to be a polymorphic trait. We have also explored the evolution of the patella through ancestral state reconstruction, finding that the patella is ancestral for lizards and possibly Lepidosauria as a whole. Clear evidence of the patella in rhynchocephalian or stem lepidosaurian fossil taxa would clarify the evolutionary origin(s) of the patella, but due to the small size of this bone and the opportunity for degradation or loss we could not definitively conclude presence or absence in the fossils examined. The pattern of evolution in lepidosaurs is unclear but our data suggest that the emergence of this sesamoid may be related to the evolution of secondary ossification centres and/or changes in knee joint conformation, where enhancement of extensor muscle leverage would be more beneficial.

BMC Evolutionary Biology

Abstract Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologi... more Abstract 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 alllepidosaurs range between approximately 226 and 289 Mya whereas estimates for crowngroup 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 a...

Bite-force performance is an ecologically important measure of whole-organism performance that sh... more Bite-force performance is an ecologically important measure of whole-organism performance that shapes dietary breadth and feeding strategies and, in some taxa, determines reproductive success. It also is a metric that is crucial to testing and evaluating biomechanical models. We reviewed nearly 100 published studies of a range of taxa that incorporate direct in vivo measurements of bite force. Problematically, methods of data collection and processing vary considerably among studies. In particular, there is little consensus on the appropriate substrate to use on the biting surface of force transducers. In addition, the bite out-lever, defined as the distance from the fulcrum (i.e. jaw joint) to the position along the jawline at which the jaws engage the transducer, is rarely taken into account. We examined the effect of bite substrate and bite out-lever on bite-force estimates in a diverse sample of lizards. Results indicate that both variables have a significant impact on the accuracy of measurements. Maximum bite force is significantly greater using leather as the biting substrate compared with a metal substrate. Less-forceful bites on metal are likely due to inhibitory feedback from mechanoreceptors that prevent damage to the feeding apparatus. Standardization of bite out-lever affected which trial produced maximum performance for a given individual. Indeed, maximum bite force is usually underestimated without standardization because it is expected to be greatest at the minimum out-lever (i.e. back of the jaws), which in studies is rarely targeted with success. We assert that future studies should use a pliable substrate, such as leather, and use appropriate standardization for bite out-lever.

The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In rec... more The extant anuran fauna of Madagascar is exceptionally rich and almost completely endemic. In recent years, many new species have been described and understanding of the history and relationships of this fauna has been greatly advanced by molecular studies, but very little is known of the fossil history of frogs on the island. Beelzebufo ampinga, the first named pre-Holocene frog from Madagascar, was described in 2008 on the basis of numerous disarticulated cranial and postcranial elements from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. These specimens documented the presence of a hyperossified taxon that differed strikingly from extant Malagasy frogs in its large size and heavy coarse cranial exostosis. Here we describe and analyse new, articulated, and more complete material of the skull, vertebral column, and hind limb, as well as additional isolated elements discovered since 2008. μCT scans allow a detailed understanding of both internal and external morphology and permit a more accurate reconstruction. The new material shows Beelzebufo to have been even more bizarre than originally interpreted, with large posterolateral skull flanges and sculptured vertebral spine tables. The apparent absence of a tympanic membrane, the strong cranial exostosis, and vertebral morphology suggest it may have burrowed during seasonally arid conditions, which have been interpreted for the Maevarano Formation from independent sedimentological and taphonomic evidence. New phylogenetic analyses, incorporating both morphological and molecular data, continue to place Beelzebufo with hyloid rather than ranoid frogs. Within Hyloidea, Beelzebufo still groups with the South American Ceratophryidae thus continuing to pose difficulties with both biogeographic interpretations and prior molecular divergence dates.

Background Lepidosauria (lizards, snakes, tuatara) is a globally distributed and ecologically ... more 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 at least 11 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 77 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.

Background: To date, the Lower Cretaceous Jehol Group of western Liaoning, China has yielded five... more Background: To date, the Lower Cretaceous Jehol Group of western Liaoning, China has yielded five monotypic genera of anurans, including Liaobatrachus grabaui, Callobatrachus sanyanensis, Mesophryne beipiaoensis, Dalianbatrachus mengi, and Yizhoubatrachus macilentus. However, the validity and distinctness of these taxa have been questioned.

The skull is composed of many bones that come together at sutures. These sutures are important si... more The skull is composed of many bones that come together at sutures. These sutures are important sites of growth, and as growth ceases some become fused while others remain patent. Their mechanical behaviour and how they interact with changing form and loadings to ensure balanced craniofacial development is still poorly understood. Early suture fusion often leads to disfiguring syndromes, thus is it imperative that we understand the function of sutures more clearly. By applying advanced engineering modelling techniques, we reveal for the first time that patent sutures generate a more widely distributed, high level of strain throughout the reptile skull. Without patent sutures, large regions of the skull are only subjected to infrequent low-level strains that could weaken the bone and result in abnormal development. Sutures are therefore not only sites of bone growth, but could also be essential for the modulation of strains necessary for normal growth and development in reptiles.

Abstract The Purbeck Limestone Group of England has yielded a rich assemblage of Late Jurassic to... more Abstract The Purbeck Limestone Group of England has yielded a rich assemblage of Late Jurassic to Early Cretaceous (Berriasian) vertebrate fossils, including one of the most diverse Early Cretaceous lizard assemblages on record. Here we describe the first articulated lizard skull from Purbeck. The specimen was rediscovered in the collections of the British Geological Survey, having been excavated at least a century ago.

What is a tuatara? The tuatara is a modern reptile found only in New Zealand (Figure 1). Although... more What is a tuatara? The tuatara is a modern reptile found only in New Zealand (Figure 1). Although it superficially looks like a medium-sized lizard it is in fact the only living member of the Rhynchocephalia, a group of animals that were successful, globally distributed, and ecologically diverse during the Mesozoic, alongside dinosaurs. A wealth of molecular and fossil evidence shows that Rhynchocephalia are the closest living relatives of Squamata (lizards and snakes), and that the two groups diverged about 250 million years ago. Therefore, humans are more closely related to kangaroos, for example, than the tuatara is to a lizard.

Background: Sea turtles (Chelonoidea) are a charismatic group of marine reptiles that occupy a r... more Background:
Sea turtles (Chelonoidea) are a charismatic group of marine reptiles that occupy a range of important ecological roles. However, the diversity and evolution of their feeding anatomy remain incompletely known.

Methodology/Principal Findings:
Using computed tomography and classical comparative anatomy we describe the cranial anatomy in two sea turtles, the loggerhead (Caretta caretta) and Kemp’s ridley (Lepidochelys kempii), for a better understanding of sea turtle functional anatomy and morphological variation. In both taxa the temporal region of the skull is enclosed by bone and the jaw joint structure and muscle arrangement indicate that palinal jaw movement is possible. The tongue is relatively small, and the hyoid apparatus is not as conspicuous as in some freshwater aquatic turtles. We find several similarities between the muscles of C. caretta and L. kempii, but comparison with other turtles suggests only one of these characters may be derived: connection of the m. adductor mandibulae internus into the Pars intramandibularis via the Zwischensehne. The large fleshy origin of the m. adductor mandibulae externus Pars superficialis from the jugal seems to be a characteristic feature of sea turtles.

Conclusions/Significance:
In C. caretta and L. kempii the ability to suction feed does not seem to be as well developed as that found in some freshwater aquatic turtles. Instead both have skulls suited to forceful biting. This is consistent with the observation that both taxa tend to feed on relatively slow moving but sometimes armoured prey. The broad fleshy origin of the m. adductor mandibulae externus Pars superficialis may be linked to thecheek region being almost fully enclosed in bone but the relationship is complex.

The tuatara (Sphenodon punctatus) is the only living representative of Rhynchocephalia, a group o... more The tuatara (Sphenodon punctatus) is the only living representative of Rhynchocephalia, a group of small vertebrates that originated about 250 million years ago. The tuatara has been referred to as a living fossil; however, the group to which it belongs included a much greater diversity of forms in the Mesozoic. We explore the morphological diversity of Rhynchocephalia and stem lepidosaur relatives (Sphenodon plus 13 fossil relatives) by employing a combination of geometric morphometrics and comparative methods. Geometric morphometrics is used to explore cranium size and shape at interspecific scale, while comparative methods are employed to test association between skull shape and size and tooth number after taking phylogeny into account. Two phylogenetic topologies have been considered to generate a phylomorphospace and quantify the phylogenetic signal in skull shape data, the ancestral state reconstruction as well as morphological disparity using disparity through time plots (DTT). Rhynchocephalia exhibit a significant phylogenetic signal in skull shape that compares well with that computed for other extinct vertebrate groups. A consistent form of allometry has little impact on skull shape evolution while the number of teeth significantly correlates with skull shape also after taking phylogeny into account. The ancestral state reconstruction demonstrates a dramatic shape difference between the skull of Sphenodon and its much larger Cretaceous relative Priosphenodon. Additionally, DTT demonstrates that skull shape disparity is higher between rather than within clades while the opposite applies to skull size and number of teeth. These results were not altered by the use of competing phylogenic hypotheses. Rhynchocephalia evolved as a morphologically diverse group with a dramatic radiation in the Late Triassic and Early Jurassic about 200 million years ago. Differences in size are not marked between species whereas changes in number of teeth are associated with co-ordinated shape changes in the skull to accommodate larger masticatory muscles. These results show that the tuatara is not the product of evolutionary stasis but that it represents the only survivor of a diverse Mesozoic radiation whose subsequent decline remains to be explained.

"The New Zealand tuatara, Sphenodon, has a specialized feeding system in which the teeth of the l... more "The New Zealand tuatara, Sphenodon, has a specialized feeding system in which the teeth of the lower jaw close between two upper tooth rows before sliding forward to slice food apart like a draw cut saw. This shearing action is unique amongst living amniotes but has been compared with the chewing power stroke of mammals. We investigated details of the jaw movement using multibody dynamics analysis of an anatomically accurate three-dimensional computer model constructed from computed tomography scans. The model predicts that a flexible symphysis is necessary for changes in the intermandibular angle that permits prooral (=proal) movement. Models with the greatest symphysial flexibility allow the articulation surface of the articular to follow the quadrate cotyle with the least restriction, and suggest that shearing is accompanied by a long axis rotation of the lower jaws. This promotes precise point loading between the cutting edges of particular teeth, enhancing the effectiveness of the shearing action. Given that Sphenodon is a relatively inactive reptile, we suggest that the link between oral food processing and endothermy has been overstated. Food processing improves feeding efficiency, a consideration
of particular importance when food availability is unpredictable.
Although this feeding mechanism is today limited to Sphenodon, a survey
of fossil rhynchocephalians suggests that it was once more widespread."

The vertebrate skull evolved to protect the brain and sense organs, but with the appearance of ja... more 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.

Rhynchocephalia achieved a global distribution during the Mesozoic but the history of sphenodonti... more Rhynchocephalia achieved a global distribution during the Mesozoic but the history of sphenodontines, the clade containing the extant genus Sphenodon (the New Zealand tuatara), remains poorly understood. Here, we describe a partial maxilla from the Late Cretaceous of Argentina bearing teeth that closely resemble those of modern Sphenodon. This material helps to fill in a notable gap in the fossil history of lepidosaurs because it represents the first evidence of a sphenodontine from South America and increases the number of known Late Cretaceous rhynchocephalian taxa from that region. The morphological disparity encompassed by these records is consistent with suggestions that rhynchocephalians remained diverse in the Late Cretaceous of South America despite a concurrent disappearance from Laurasia. Moreover, the new record supports the hypothesis that sphenodontines were once found widely throughout Gondwana, before its constituent landmasses began to separate about 80 million years ago. The extant Sphenodon probably represents a biogeographic remnant of this distribution, but whether its relatively large size and its ability to remain active at cold temperatures reflects a high latitude ancestry requires further examination.

Although diplodocoid sauropods from Africa and the Americas are well known, their European record... more Although diplodocoid sauropods from Africa and the Americas are well known, their European record remains largely neglected. Here we redescribe Dinheirosaurus lourinhanensis from the Late Jurassic of Portugal. The holotype comprises two posterior cervical vertebrae, the dorsal series and a caudal centrum. Redescription demonstrates its validity on the basis of three autapomorphies: (1) posteriorly restricted ventral keel on posterior cervical vertebrae; (2) three small subcircular fossae posterior to the lateral coel on posterior cervical neural spines; (3) accessory lamina linking the hyposphene with base of the posterior centrodiapophyseal lamina in middle-posterior dorsal vertebrae. Phylogenetic analysis places Dinheirosaurus as the sister taxon to Supersaurus, and this clade forms the sister taxon to other diplodocines. However, this position should be treated with caution as Dinheirosaurus displays several plesiomorphic features absent in other diplodocids (including unbifurcated presacral neural spines, and dorsolaterally projecting diapophyses on dorsal vertebrae) and only four additional steps are required to place Dinheirosaurus outside of Flagellicaudata. We identify Amazonsaurus as the basal-most rebbachisaurid and recover Zapalasaurus outside of the South American Limaysaurinae, suggesting the biogeographic history of rebbachisaurids is more complex than previously proposed. Review of the European diplodocoid record reveals evidence for the earliest known diplodocid, as well as additional diplodocid remains from the Late Jurassic of Spain. A Portuguese specimen, previously referred to Dinheirosaurus, displays strong similarities to Apatosaurus from the contemporaneous Morrison Formation of North America, indicating the presence of a second Late Jurassic Portuguese diplodocid taxon. Along with Dinheirosaurus, these Portuguese remains provide further evidence for a Late Jurassic palaeobiogeographic connection between Europe and North America. No dicraeosaurids are currently known from Europe, but rebbachisaurids are present in the Early Cretaceous, with weak evidence for the earliest known representative from the Late Jurassic of Spain; however, more complete material is required to recognize early members of this clade.