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Digital specializations of geckos are widely associated with their climbing abilities. A recurring feature that has independently emerged within the sister families Gekkonidae and Phyllodactylidae is the presence of neomorphic paraphalanges (PPEs), usually paired, paraxial skeletal structures lying adjacent to interphalangeal and metapodial-phalangeal joints. The incorporation of PPEs into gekkotan autopodia has the potential to modify the modularity and integration of the ancestral limb pattern by affecting information flow among skeletal limb parts. Here we explore the influence of PPEs on limb organization using anatomical networks. We modeled the fore- and hindlimbs in species ancestrally devoid of PPEs (Iguana iguana and Gekko gecko) and paraphalanx-bearing species (Hemidactylus mabouia and Uroplatus fimbriatus). To further clarify the impact of PPEs we also expunged PPEs from paraphalanx-bearing network models. We found that PPEs significantly increase modularity, giving rise to tightly integrated sub-modules along the digits, suggesting functional specialization. Species-specific singularities also emerged, such as the trade-off between the presence of PPEs favoring modularity (along the proximodistal axis) and the interdigital webbing favoring integration (across the lateromedial axis) in the limbs of U. fimbriatus. The PPEs are characterized by low connectivity compared with other skeletal elements; nevertheless, this varies based on their specific location and seemingly reflects developmental constraints. Our results also highlight the importance of the fifth metatarsal in generating a shift in lepidosaurian hindlimb polarity that contrasts with the more symmetrical bauplan of tetrapods. Our findings support extensive modification of the autopodial system in association with the addition of the neomorphic and intriguing PPEs.

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Evolutionary tempo and mode summarize ancient and controversial subjects of theoretical biology such as gradualism, convergence, contingence, trends, and entrenchment. We employed an integrative methodological approach to explore the evolutionary tempo and mode of Lepidosaurian phalangeal formulae (PFs). This approach involves quantifying the frequencies of morphological changes along an evolutionary trajectory. The five meristic characters encoded by PFs are particularly valuable in revealing evolutionary patterns, owing to their discrete nature and extensive documentation in the literature. Based on a pre-existing dataset of PFs from 649 taxa (35 Lepidosauria families, including fossils), from which there exists a unique repertoire of 53 formulations, our approach simultaneously considers phenetic and phylogenetic data. This culminates in a diagram accounting for the phylogenetic dynamic of evolution traversing across different regions of morphospace. The method involves enumerating phenotypical options, reconstructing phenotypes across the phylogeny, projecting phenotypes onto a morphospace, and constructing a flow network from the frequency of evolutionary transitions between unique phenotypic conditions. This approach links Markovian chains and evolutionary trajectories to formally define parameters that describe the underlying transitions of morphological change. Among other results, we found that (a) PF evolution exhibits a clear trend towards reduction in the phalangeal count and that (b) evolutionary change tends to occur significantly between morphologically similar PFs. Notwithstanding, although minor but not trivial, transitions between distant formulas -jumps- occur. Our results support a pluralistic view including stasis, gradualism, and saltationism discriminating their prevalence in a target character evolution.

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Skeletal remains of a small lepidosaurian reptile from the Middle Triassic (Ladinian: Longobardian) Erfurt Formation, exposed in a commercial limestone quarry near Vellberg (Germany), represent the oldest rhynchocephalian known to date. The new taxon, Wirtembergia hauboldae, is diagnosed by the following combination of features: Premaxilla with four teeth, first being largest and decreasing in size from first to fourth. Jugal with tiny, spur-like posterior process. Lateral surface of dentary strongly convex dorsoventrally for much of length of bone, bearing distinct longitudinal ridge and sculpturing in large specimens. Coronoid eminence of dentary low, subrectangular, and with dorsoventrally concave lateral surface in larger specimens. Dentition with pleurodont anterior and acrodont posterior teeth. Posterior (=additional) teeth with (in side view) triangular, at mid-crown level labiolingually somewhat flattened crowns, and with oval bases. Phylogenetic analysis recovered the new rhynchocephalian as the earliest-diverging member of its clade known to date.

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BACKGROUND: The shape of the semicircular canals of the inner ear of living squamate reptiles has been used to infer phylogenetic relationships, body size, and life habits. Often these inferences are made without controlling for the effects of the other ones. Here we examine the semicircular canals of 94 species of extant limbed lepidosaurs using three-dimensional landmark-based geometric morphometrics, and analyze them in phylogenetic context to evaluate the relative contributions of life habit, size, and phylogeny on canal shape. RESULTS: Life habit is not a strong predictor of semicircular canal shape across this broad sample. Instead, phylogeny plays a major role in predicting shape, with strong phylogenetic signal in shape as well as size. Allometry has a limited role in canal shape, but inner ear size and body mass are strongly correlated. CONCLUSIONS: Our wide sampling across limbed squamates suggests that semicircular canal shape and size are predominantly a factor of phylogenetic relatedness. Given the small proportion of variance in semicircular canal shape explained by life habit, it is unlikely that unknown life habit could be deduced from semicircular canal shape alone. Overall, semicircular canal size is a good estimator of body length and even better for body mass in limbed squamates. Semiaquatic taxa tend to be larger and heavier than non-aquatic taxa, but once body size and phylogeny are accounted for, they are hard to distinguish from their non-aquatic relatives based on bony labyrinth shape and morphology.

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The squamates (lizards, snakes, and relatives) today comprise more than 10,000 species, and yet their sister group, the Rhynchocephalia, is represented by a single species today, the tuatara. The explosion in squamate diversity has been tracked back to the Cretaceous Terrestrial Revolution, 100 million years ago (Ma), the time when flowering plants began their takeover of terrestrial ecosystems, associated with diversification of coevolving insects and insect-eating predators such as lizards, birds, and mammals. Squamates arose much earlier, but their long pre-Cretaceous history of some 150 million years (Myr) is documented by sparse fossils. Here, we provide evidence for an initial radiation of squamate morphology in the Middle and Late Jurassic (174-145 Ma), and show that they established their key ecological roles much earlier than had been assumed, and they have not changed them much since.

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Performance traits are tightly linked to the fitness of organisms. However, because studies of variation in performance traits generally focus on just one or several closely related species, we are unable to draw broader conclusions about how and why these traits vary across clades. One important performance trait related to many aspects of an animal's life history is bite-force. Here, we use a clade-wide phylogenetic comparative approach to investigate relationships between size, head dimensions and bite-force among lizards and tuatara (lepidosaurs), using the largest bite-force dataset collated to date for any taxonomic group. We test four predictions: that bite-force will be greater in larger species, and for a given body size, bite-force will be greatest in species with acrodont tooth attachment, herbivorous diets, and non-burrowing habits. We show that bite-force is strongly related to body and head size across lepidosaurs and, as predicted, larger species have the greatest bite-forces. Contrary to our other predictions, tooth attachment, diet and habit have little predictive power when accounting for size. Herbivores bite more forcefully simply because they are larger. Our results also highlight priorities for future sampling to further enhance our understanding of broader evolutionary patterns.

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The ability to repair injuries among reptiles, i.e., ectothermic amniotes, is similar to that of mammals with some noteworthy exceptions. While large wounds in turtles and crocodilians are repaired through scarring, the reparative capacity involving the tail derives from a combined process of wound healing and somatic growth, the latter being continuous in reptiles. When the tail is injured in juvenile crocodilians, turtles and tortoises as well as the tuatara (Rhynchocephalia: Sphenodon punctatus, Gray 1842), the wound is repaired in these reptiles and some muscle and connective tissue and large amounts of cartilage are regenerated during normal growth. This process, here indicated as "regengrow", can take years to produce tails with similar lengths of the originals and results in only apparently regenerated replacements. These new tails contain a cartilaginous axis and very small (turtle and crocodilians) to substantial (e.g., in tuatara) muscle mass, while most of the tail is formed by an irregular dense connective tissue containing numerous fat cells and sparse nerves. Tail regengrow in the tuatara is a long process that initially resembles that of lizards (the latter being part of the sister group Squamata within the Lepidosauria) with the formation of an axial ependymal tube isolated within a cartilaginous cylinder and surrounded by an irregular fat-rich connective tissue, some muscle bundles, and neogenic scales. Cell proliferation is active in the apical regenerative blastema, but much reduced cell proliferation continues in older regenerated tails, where it occurs mostly in the axial cartilage and scale epidermis of the new tail, but less commonly in the regenerated spinal cord, muscles, and connective tissues. The higher tissue regeneration of Sphenodon and other lepidosaurians provides useful information for attempts to improve organ regeneration in endothermic amniotes.

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Whether scales reduce cutaneous evaporative water loss in lepidosaur reptiles (Superorder Lepidosauria) such as lizards and snakes has been a contentious issue for nearly half a century. Furthermore, while many studies have looked at whether dehydration affects thermal preference in lepidosaurs, far fewer have examined whether normally hydrated lepidosaurs can assess their instantaneous rate of evaporative water loss and adjust their thermal preference to compensate in an adaptive manner. We tested both of these hypotheses using three captive-bred phenotypes of bearded dragon (Pogona vitticeps) sourced from the pet trade: 'wild-types' with normal scalation, 'leatherbacks' exhibiting scales of reduced prominence, and scaleless bearded dragons referred to as 'silkbacks'. Silkbacks on average lost water evaporatively at about twice the rate that wild-types did. Leatherbacks on average were closer in their rates of evaporative water loss to silkbacks than they were to wild-types. Additionally, very small (at most ∼1°C) differences in thermal preference existed between the three phenotypes that were not statistically significant. This suggests a lack of plasticity in thermal preference in response to an increase in the rate of evaporative water loss, and may be reflective of a thermal 'strategy' as employed by thermoregulating bearded dragons that prioritises immediate thermal benefits over the threat of future dehydration. The results of this study bolster an often-discounted hypothesis regarding the present adaptive function of scales and have implications for the applied fields of animal welfare and conservation.

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Cranial morphology in lepidosaurs is highly disparate and characterised by the frequent loss or reduction of bony elements. In varanids and geckos, the loss of the postorbital bar is associated with changes in skull shape, but the mechanical principles underlying this variation remain poorly understood. Here, we sought to determine how the overall cranial architecture and the presence of the postorbital bar relate to the loading and deformation of the cranial bones during biting in lepidosaurs. Using computer-based simulation techniques, we compared cranial biomechanics in the varanid Varanus niloticus and the teiid Salvator merianae, two large, active foragers. The overall strain magnitude and distribution across the cranium were similar in the two species, despite lower strain gradients in V. niloticus In S. merianae, the postorbital bar is important for resistance of the cranium to feeding loads. The postorbital ligament, which in varanids partially replaces the postorbital bar, does not affect bone strain. Our results suggest that the reduction of the postorbital bar impaired neither biting performance nor the structural resistance of the cranium to feeding loads in V. niloticus Differences in bone strain between the two species might reflect demands imposed by feeding and non-feeding functions on cranial shape. Beyond variation in cranial bone strain related to species-specific morphological differences, our results reveal that similar mechanical behaviour is shared by lizards with distinct cranial shapes. Contrary to the situation in mammals, the morphology of the circumorbital region, calvaria and palate appears to be important for withstanding high feeding loads in these lizards.

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Archosaurs displayed an evolutionary trend toward increasing bipedalism in their evolutionary history, that is, forelimbs tend to be reduced in contrast to the development of hindlimbs becoming major weight-bearing and locomotor appendages. The archosaurian locomotion has been extensively discussed based on their limb morphology because the latter reflects their locomotor modes very well. However, despite some attempts of reconstructing the hindlimb musculature in Archosauria, that of the most distal portion, the pes, has often been neglected. In order to rectify this trend, detailed homologies of pedal muscles among sauropsids were established based on dissections and literature reviews of adult conditions. As a result, homologies of some pedal muscles between non-avian sauropsids and avians were revised, challenging classical hypotheses. The present new hypothesis postulates that the avian m. tibialis cranialis and non-avian m. extensor digitorum longus, as well as the avian m. extensor digitorum longus and non-avian m. tibialis anterior, are homologous with each other, respectively. This is more plausible because it requires no drastical change in the attachment sites between the avian and non-avian homologues unlike the classical hypothesis. Many interosseous muscles in non-archosaurian sauropsids that have long been regarded as a part of short digital extensors or flexors are also divided into multiple distinct muscles so that they can be homologized with short pedal muscles among all sauropsids. In addition, osteological correlates of attachments are identified for most of the pedal muscles, contributing to future attempts of reconstruction of this muscle system in fossil archosaurs.

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Among amniote vertebrates, nonavian reptiles (chelonians, crocodilians, and lepidosaurs) are regarded as using vocal signals rarely (compared to birds and mammals). In all three reptilian clades, however, certain taxa emit distress calls and advertisement calls using modifications of regions of the upper respiratory tract. There is no central tendency in either acoustic mechanisms or the structure of the vocal apparatus, and many taxa that vocalize emit only relatively simple sounds. Available evidence indicates multiple origins of true vocal abilities within these lineages. Reptiles thus provide opportunities for studying the early evolutionary stages of vocalization. The early literature on the diversity of form of the laryngotracheal apparatus of reptiles boded well for the study of form-function relationships, but this potential was not extensively explored. Emphasis shifted away from anatomy, however, and centered instead on acoustic analysis of the sounds that are produced. New investigative techniques have provided novel ways of studying the form-function aspects of the structures involved in phonation and have brought anatomical investigation to the forefront again. In this review we summarize what is known about hearing in reptiles in order to contextualize the vocal signals they generate and the sound-producing mechanisms responsible for them. The diversity of form of the sound producing apparatus and the increasing evidence that reptiles are more dependent upon vocalization as a communication medium than previously thought indicates that they have a significant role to play in the understanding of the evolution of vocalization in amniotes.

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Extant and extinct reptiles exhibit numerous combinations of tooth implantation and attachment. Tooth implantation ranges from those possessing roots and lying within a socket (thecodonty), to teeth lying against the lingual wall of the jawbone (pleurodonty), to teeth without roots or sockets that are attached to the apex of the marginal jawbones (acrodonty). Attachment may be ligamentous (gomphosis) or via fusion (ankylosis). Generally speaking, adaptative reasonings are proposed as an underlying driver for evolutionary changes in some forms of tooth implantation and attachment. However, a substantiated adaptive hypothesis is lacking for the state of acrodont ankylosis that is seen in several lineages of Lepidosauria, a clade that is plesiomorphically pleurodont. The convergent evolution of acrodont ankylosis in several clades of lepidosaurs suggests a selective pressure shaped the evolution of the trait. We hypothesize that acrodont ankylosis as seen in Acrodonta and Sphenodon punctatus, is an adaptation either resulting from or allowing for a stronger bite force. We analyzed bite force data gathered from the literature to show that those taxa possessing acrodont dentition possess a stronger bite force on average than those taxa with pleurodont dentition. Dietary specialists with pleurodont dentition may also possess relatively high bite forces, though body size may also play a role in their ability to bite hard. Furthermore, our results have implications for the evolution of acrodont ankylosis and potential behaviors related to strong bite force that influenced the evolution of acrodonty within Acrodonta and Rhynchocephalia.

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Mammals and reptiles have evolved divergent adaptations for processing abrasive foods. Mammals have occluding, diphyodont dentitions with taller teeth (hypsodonty), more complex occlusal surfaces, continuous tooth eruption, and forms of prismatic enamel that prolong the functional life of each tooth [1, 2]. The evolution of prismatic enamel in particular was a key innovation that made individual teeth more resilient to abrasion in early mammals [2-4]. In contrast, reptiles typically have thin, non-prismatic enamel, and shearing, polyphyodont dentitions with multi-cusped or serrated tooth crowns, multiple tooth rows, rapid tooth replacement rates, or batteries made of hundreds of teeth [5-9]. However, there are rare cases where reptiles have evolved alternative solutions to cope with abrasive diets. Here, we show that the combined effects of herbivory and an ancestral loss of tooth replacement in a lineage of extinct herbivorous sphenodontians, distant relatives of the modern tuatara (Sphenodon punctatus) [10], are associated with the evolution of wear-resistant and highly complex teeth. Priosphenodon avelasi, an extinct sphenodontian from the Cretaceous of Argentina, possesses a unique cone-in-cone dentition with overlapping generations of teeth forming a densely packed tooth file. Each tooth is anchored to its predecessor via a rearrangement of dental tissues that results in a novel enamel-to-bone tooth attachment. Furthermore, the compound occlusal surfaces, thickened enamel, and the first report of prismatic enamel in a sphenodontian are convergent strategies with those in some mammals, challenging the perceived simplicity of acrodont dentitions [11-15] and showcasing the reptilian capacity to produce complex and unusual dentitions.

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Direct indicators of diet and predator-prey relationships are exceedingly rare in the fossil record [1, 2]. However, it is through such traces that we can best understand trophic interactions in ancient ecosystems [3], confirm dietary inferences derived from skeletal morphologies [4], and clarify behavioral and ecological interpretations [5]. Here, we identify a previously unrecognized lizard species in the abdomen of a specimen of Microraptor zhaoianus, a small, volant dromaeosaurid (Paraves) with asymmetrical flight feathers on both its forelimbs and hindlimbs from the Early Cretaceous Jehol Biota [6-8]. The lizard is largely complete and articulated, confirming the current perception of Microraptor as an agile opportunistic predator that, like extant reptiles, including raptorial birds, ingested small prey whole and head first [9]. The lizard can be readily distinguished from previously recognized Early Cretaceous species based on its unusual widely spaced and brachydont dentition. Phylogenetic analysis suggests Indrasaurus wangi gen. et sp. nov. is a basal scleroglossan closely related to the slightly older Liushusaurus [10]. Comparison of ingested remains preserved across Paraves suggests that dromaeosaurids retained the plesiomorphic condition in which ingested prey were fully digested, rather than egested, as has been demonstrated was the case in the probable troodontid Anchiornis [11]. This supports a closer relationship between Aves and Anchiornis [12, 13] and suggests that flight did not precipitate the evolution of pellet egestion in Paraves and that the evolution of the "modern avian" digestive system in paravians was highly homoplastic [14]. A preliminary Jehol food web is reconstructed from current data.

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In vivo bone strain data provide direct evidence of strain patterns in the cranium during biting. Compared with those in mammals, in vivo bone strains in lizard skulls are poorly documented. This paper presents strain data from the skulls of Anolis equestris, Gekko gecko, Iguana iguana and Salvator merianae during transducer biting. Analysis of variance was used to investigate effects of bite force, bite point, diet, cranial morphology and cranial kinesis on strain magnitude. Within individuals, the most consistent determinants of variance in bone strain magnitude were gauge location and bite point, with the importance of bite force varying between individuals. Inter-site variance in strain magnitude - strain gradient - was present in all individuals and varied with bite point. Between individuals within species, variance in strain magnitude was driven primarily by variation in bite force, not gauge location or bite point, suggesting that inter-individual variation in patterns of strain magnitude is minimal. Between species, variation in strain magnitude was significantly impacted by bite force and species membership, as well as by interactions between gauge location, species and bite point. Independent of bite force, species differences in cranial strain magnitude may reflect selection for different cranial morphology in relation to feeding function, but what these performance criteria are is not clear. The relatively low strain magnitudes in Iguana and Uromastyx compared with those in other lizards may be related to their herbivorous diet. Cranial kinesis and the presence or absence of postorbital and supratemporal bars are not important determinants of inter-specific variation in strain magnitude.

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Lepidosauria is a speciose clade with a long evolutionary history, but there have been few attempts to explore its taxon richness through time. Here we estimate patterns of terrestrial lepidosaur genus diversity for the Triassic-Palaeogene (252-23 Ma), and compare observed and sampling-corrected richness curves generated using Shareholder Quorum Subsampling and classical rarefaction. Generalized least-squares regression (GLS) is used to investigate the relationships between richness, sampling and environmental proxies. We found low levels of richness from the Triassic until the Late Cretaceous (except in the Kimmeridgian-Tithonian of Europe). High richness is recovered for the Late Cretaceous of North America, which declined across the K-Pg boundary but remained relatively high throughout the Palaeogene. Richness decreased following the Eocene-Oligocene Grande Coupure in North America and Europe, but remained high in North America and very high in Europe compared to the Late Cretaceous; elsewhere data are lacking. GLS analyses indicate that sampling biases (particularly, the number of fossil collections per interval) are the best explanation for long-term face-value genus richness trends. The lepidosaur fossil record presents many problems when attempting to reconstruct past diversity, with geographical sampling biases being of particular concern, especially in the Southern Hemisphere.

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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.

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We show-in contrast to the traditional textbook contention-that the first amniote lungs were complex, multichambered organs and that the single-chambered lungs of lizards and snakes represent a secondarily simplified rather than the plesiomorphic condition. We combine comparative anatomical and embryological data and show that shared structural principles of multichamberedness are recognizable in amniotes including all lepidosaurian taxa. Sequential intrapulmonary branching observed during early organogenesis becomes obscured during subsequent growth, resulting in a secondarily simplified, functionally single-chambered lung in lepidosaurian adults. Simplification of pulmonary structure maximized the size of the smallest air spaces and eliminated biophysically compelling surface tension problems that were associated with miniaturization evident among stem lepidosaurmorphs. The remaining amniotes, however, retained the multichambered lungs, which allowed both large surface area and high pulmonary compliance, thus initially providing a strong selective advantage for efficient respiration in terrestrial environments. Branched, multichambered lungs instead of simple, sac-like organs were part and parcel of the respiratory apparatus of the first amniotes and pivotal for their success on dry land, with the sky literally as the limit.

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The course of the medial ridge in the lizard jugal shows considerable morphological variation. There are four basic configurations: (1) the medial ridge is located ventral to mid-height on the suborbital process and anterior to mid-length on the postorbital process; (2) the medial ridge is located ventrally on the suborbital process (as above), but posteriorly on the postorbital process; (3) the medial ridge is located dorsally on the suborbital process and anteriorly on the postorbital process; and (4) the medial ridge is centrally located along the entire length of the jugal. Ancestral character state reconstruction shows that type 1 is plesiomorphic for Squamata regardless of the broad-scale phylogenetic topology. Type 3 is present in chamaeleonids and convergently in Anolis barbatus. Type 3 is a synapomorphy of the chamaeleonids. Type 2 is considered plesiomorphic for Anguidae, Heloderma and Xenosaurus, although it is independently modified in some extant members. These taxa form a clade in molecular phylogenies of Squamata, and the course of the medial ridge of the jugal therefore provides some measure of morphological support for this arrangement. The course of the medial ridge may be best explained by the position of the eye and by the angle of the jugal; its relations with other bony orbital structures (supraocular osteoderms, palpebral, supraorbital flanges) and the posterior extent of the maxilla are also discussed.

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Sphenodontians were a successful group of rhynchocephalian reptiles that dominated the fossil record of Lepidosauria during the Triassic and Jurassic. Although evidence of extinction is seen at the end of the Laurasian Early Cretaceous, they appeared to remain numerically abundant in South America until the end of the period. Most of the known Late Cretaceous record in South America is composed of opisthodontians, the herbivorous branch of Sphenodontia, whose oldest members were until recently reported to be from the Kimmeridgian-Tithonian (Late Jurassic). Here, we report a new sphenodontian, Sphenotitan leyesi gen. et sp. nov., collected from the Upper Triassic Quebrada del Barro Formation of northwestern Argentina. Phylogenetic analysis identifies Sphenotitan as a basal member of Opisthodontia, extending the known record of opisthodontians and the origin of herbivory in this group by 50 Myr.

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