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The cranial anatomy of the Lower Jurassic ornithischian dinosaur Heterodontosaurus tucki Crompton & Charig, 1962 is described in detail for the first time on the basis of two principal specimens: the holotype (SAM-PK-K337) and referred skull (SAM-PK-K1332). In addition several other specimens that have a bearing on the interpretation of the anatomy and biology of Heterodontosaurus are described. The skull and lower jaw of Heterodontosaurus are compact and robust but perhaps most notable for the heterodont dentition that merited the generic name. Details of the cranial anatomy are revealed and show that the skull is unexpectedly specialized in such an early representative of the Ornithischia, including: the closely packed, hypsodont crowns and 'warping' of the occlusal surfaces (created by progressive variation in the angulation of wear on successive crowns) seen in the cheek dentition; the unusual sutural relationships between the bones along the dorsal edge of the lower jaw; the very narrow, deeply vaulted palate and associated structures on the side wall of the braincase; and the indications of cranial pneumatism (more commonly seen in basal archosaurs and saurischian dinosaurs). Evidence for tooth replacement (which has long been recognized, despite frequent statements to the contrary) is suggestive of an episodic, rather than continuous, style of tooth replacement that is, yet again, unusual in diapsids generally and particularly so amongst ornithischian dinosaurs. Cranial musculature has been reconstructed and seems to conform to that typically seen in diapsids, with the exception of the encroachment of M. adductor mandibulae externus superficialis across the lateral surface of the temporal region and external surface of the lower jaw. Indications, taken from the unusual shape of the occlusal surfaces of the cheek dentition and jaw musculature, are suggestive of a novel form of jaw action in this dinosaur. The taxonomy of currently known late Karoo-aged heterodontosaurids from southern Africa is reviewed. Although complicated by the inadequate nature of much of the known material, it is concluded that two taxa may be readily recognized: H. tucki and Abrictosaurus consors. At least one additional taxon is recognized within the taxa presently named Lanasaurus and Lycorhinus; however, both remain taxonomically problematic and their status needs to be further tested and may only be resolved by future discoveries. The only other named taxon, Geranosaurus atavus, represents an invalid name. The recognition of at least four distinct taxa indicates that the heterodontosaurids were speciose within the late Karoo ecosystem. The systematics of Heterodontosaurus and its congeners has been analysed, using a restricted sample of taxa. A basal (nongenasaurian) position within Ornithischia is re-affirmed. There are at least four competing hypotheses 182 concerning the phylogenetic placement of the Heterodontosauridae, so the evidence in support of the various hypotheses is reviewed in some detail. At present the best-suppor...

Sauria is the crown-group of Diapsida and is subdivided into Lepidosauromorpha and Archosauromorpha, comprising a high percentage of the diversity of living and fossil tetrapods. The split between lepidosauromorphs and archosauromorphs (the crocodile-lizard, or bird-lizard, divergence) is considered one of the key calibration points for molecular analyses of tetrapod phylogeny. Saurians have a very rich Mesozoic and Cenozoic fossil record, but their late Paleozoic (Permian) record is problematic. Several Permian specimens have been referred to Sauria, but the phylogenetic affinity of some of these records remains questionable. We reexamine and review all of these specimens here, providing new data on early saurian evolution including osteohistology, and present a new morphological phylogenetic dataset. We support previous studies that find that no valid Permian record for Lepidosauromorpha, and we also reject some of the previous referrals of Permian specimens to Archosauromorpha. The most informative Permian archosauromorph is Protorosaurus speneri from the middle Late Permian of Western Europe. A historically problematic specimen from the Late Permian of Tanzania is redescribed and reidentified as a new genus and species of basal archosauromorph: Aenigmastropheus parringtoni. The supposed protorosaur Eorasaurus olsoni from the Late Permian of Russia is recovered among Archosauriformes and may be the oldest known member of the group but the phylogenetic support for this position is low. The assignment of Archosaurus rossicus from the latest Permian of Russia to the archosauromorph clade Proterosuchidae is supported. Our revision suggests a minimum fossil calibration date for the crocodile-lizard split of 254.7 Ma. The occurrences of basal archosauromorphs in the northern (30°N) and southern (55°S) parts of Pangea imply a wider paleobiogeographic distribution for the group during the Late Permian than previously appreciated. Early archosauromorph growth strategies appear to be more diverse than previously suggested based on new data on the osteohistology of Aenigmastropheus.

The fossil record of crocodylians and their relatives (pseudosuchians) reveals a rich evolutionary history, prompting questions about causes of long-term decline to their present-day low biodiversity. We analyse climatic drivers of subsampled pseudosuchian biodiversity over their 250 million year history, using a comprehensive new data set. Biodiversity and environmental changes correlate strongly, with long-term decline of terrestrial taxa driven by decreasing temperatures in northern temperate regions, and biodiversity decreases at lower latitudes matching patterns of increasing aridification. However, there is no relationship between temperature and biodiversity for marine pseudosuchians, with sea-level change and post-extinction opportunism demonstrated to be more important drivers. A ‘modern-type' latitudinal biodiversity gradient might have existed throughout pseudosuchian history, and range expansion towards the poles occurred during warm intervals. Although their fossil record suggests that current global warming might promote long-term increases in crocodylian biodiversity and geographic range, the 'balancing forces' of anthropogenic environmental degradation complicate future predictions.

Dinosaurs were remarkably successful during the Mesozoic and one subgroup, birds, remain an important component of modern ecosystems. Although the extinction of non-avian dinosaurs at the end of the Cretaceous has been the subject of intense debate, comparatively little attention has been given to the origin and early evolution of dinosaurs during the Late Triassic and Early Jurassic, one of the most important evolutionary radiations in earth history. Our understanding of this keystone event has dramatically changed over the past 25 years, thanks to an influx of new fossil discoveries, reinterpretations of long-ignored specimens, and quantitative macroevolutionary analyses that synthesize anatomical and geological data. Here we provide an overview of the first 50 million years of dinosaur history, with a focus on the large-scale patterns that characterize the ascent of dinosaurs from a small, almost marginal group of reptiles in the Late Triassic to the preeminent terrestrial vertebrates of the Jurassic and Cretaceous. We provide both a biological and geological background for early dinosaur history. Dinosaurs are deeply nested among the archosaurian reptiles, diagnosed by only a small number of characters, and are subdivided into a number of major lineages. The first unequivocal dinosaurs are known from the late Carnian of South America, but the presence of their sister group in the Middle Triassic implies that dinosaurs possibly originated much earlier. The three major dinosaur lineages, theropods, sauropodomorphs, and ornithischians, are all known from the Triassic, when continents were joined into the supercontinent Pangaea and global climates were hot and arid. Although many researchers have long suggested that dinosaurs outcompeted other reptile groups during the Triassic, we argue that the ascent of dinosaurs was more of a matter of contingency and opportunism. Dinosaurs were overshadowed in most Late Triassic ecosystems by crocodile-line archosaurs and showed no signs of outcompeting their rivals. Instead, the rise of dinosaurs was a two-stage process, as dinosaurs expanded in taxonomic diversity, morphological disparity, and absolute faunal abundance only after the extinction of most crocodile-line reptiles and other groups.

The fossil record is our only direct means for evaluating shifts in biodiversity through Earth's history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic -Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent 'Lagerstätten effects'. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous -Palaeogene extinction.

Pneumatic (air-filled) postcranial bones are unique to birds among extant tetrapods. Unambiguous skeletal correlates of postcranial pneumaticity first appeared in the Late Triassic (approximately 210 million years ago), when they evolved independently in several groups of bird-line archosaurs (ornithodirans). These include the theropod dinosaurs (of which birds are extant representatives), the pterosaurs, and sauropodomorph dinosaurs. Postulated functions of skeletal pneumatisation include weight reduction in large-bodied or flying taxa, and density reduction resulting in energetic savings during foraging and locomotion. However, the influence of these hypotheses on the early evolution of pneumaticity has not been studied in detail previously. We review recent work on the significance of pneumaticity for understanding the biology of extinct ornithodirans, and present detailed new data on the proportion of the skeleton that was pneumatised in 131 non-avian theropods and Archaeopteryx. This includes all taxa known from significant postcranial remains. Pneumaticity of the cervical and anterior dorsal vertebrae occurred early in theropod evolution. This 'common pattern' was conserved on the line leading to birds, and is likely present in Archaeopteryx. Increases in skeletal pneumaticity occurred independently in as many as 12 lineages, highlighting a remarkably high number of parallel acquisitions of a bird-like feature among non-avian theropods. Using a quantitative comparative framework, we show that evolutionary increases in skeletal pneumaticity are significantly concentrated in lineages with large body size, suggesting that mass reduction in response to gravitational constraints at large body sizes influenced the early evolution of pneumaticity. However, the body size threshold for extensive pneumatisation is lower in theropod lineages more closely related to birds (maniraptorans). Thus, relaxation of the relationship between body size and pneumatisation preceded the origin of birds and cannot be explained as an adaptation for flight. We hypothesise that skeletal density modulation in small, non-volant, maniraptorans resulted in energetic savings as part of a multi-system response to increased metabolic demands. Acquisition of extensive postcranial pneumaticity in small-bodied maniraptorans may indicate avian-like high-performance endothermy.

The relationship between dinosaurs and other reptiles is well established, but the sequence of acquisition of dinosaurian features has been obscured by the scarcity of fossils with transitional morphologies. The closest extinct relatives of dinosaurs either have highly derived morphologies or are known from poorly preserved or incomplete material. Here we describe one of the stratigraphically lowest and phylogenetically earliest members of the avian stem lineage (Avemetatarsalia), Teleocrater rhadinus gen. et sp. nov., from the Middle Triassic epoch. The anatomy of T. rhadinus provides key information that unites several enigmatic taxa from across Pangaea into a previously unrecognized clade, Aphanosauria. This clade is the sister taxon of Ornithodira (pterosaurs and birds) and shortens the ghost lineage inferred at the base of Avemetatarsalia. We demonstrate that several anatomical features long thought to characterize Dinosauria and dinosauriforms evolved much earlier, soon after the bird-crocodylian split, and that the earliest avemetatarsalians retained the crocodylian-like ankle morphology and hindlimb proportions of stem archosaurs and early pseudosuchians. Early avemetatarsalians were substantially more species-rich, widely geographically distributed and morphologically diverse than previously recognized. Moreover, several early dinosauromorphs that were previously used as models to understand dinosaur origins may represent specialized forms rather than the ancestral avemetatarsalian morphology.