The origin and early radiation of archosaurs and closely related taxa (Archosauriformes) during the Triassic was a critical event in the evolutionary history of tetrapods. This radiation led to the dinosaur-dominated ecosystems of the Jurassic and Cretaceous, and the high present-day archosaur diversity that includes around 10,000 bird and crocodylian species. The timing and dynamics of this evolutionary radiation are currently obscured by the poorly constrained phylogenetic positions of several key early archosauriform taxa, including several species from the Middle Triassic of Argentina (Gracilisuchus stipanicicorum) and China (Turfanosuchus dabanensis, Yonghesuchus sangbiensis). These species act as unstable ‘wildcards’ in morphological phylogenetic analyses, reducing phylogenetic resolution.

A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252–201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian–Middle Jurassic. Maximum-likelihood analyses indicate that Cope's rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope's rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle–early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely-accepted ‘rule’ that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.

Archosauromorpha originated in the middle–late Permian, radiated during the Triassic, and gave rise to the crown group Archosauria, a highly successful clade of reptiles in terrestrial ecosystems over the last 250 million years. However, scientific attention has mainly focused on the diversification of archosaurs, while their stem lineage (i.e. non-archosaurian archosauromorphs) has often been overlooked in discussions of the evolutionary success of Archosauria. Here, we analyse the cranial disparity of late Permian to Early Jurassic archosauromorphs and make comparisons between non-archosaurian archosauromorphs and archosaurs (including Pseudosuchia and Ornithodira) on the basis of two-dimensional geometric morphometrics. Our analysis recovers previously unappreciated high morphological disparity for non-archosaurian archosauromorphs, especially during the Middle Triassic, which abruptly declined during the early Late Triassic (Carnian). By contrast, cranial disparity of archosaurs increased from the Middle Triassic into the Late Triassic, declined during the end-Triassic extinction, but re-expanded towards the end of the Early Jurassic. Our study indicates that non-archosaurian archosauromorphs were highly diverse components of terrestrial ecosystems prior to the major radiation of archosaurs, including dinosaurs, while disparity patterns of the Ladinian and Carnian indicate a gradual faunal replacement of stem archosaurs by the crown group, including a short interval of partial overlap in morphospace during the Ladinian.

The lower jaw of an unidentified Pennsylvanian (Late Carboniferous) tetrapod from Nova Scotia – the “Parrsboro jaw”- is redescribed in the light of recent tetrapod discoveries and work on evolution of tetrapod mandibular morphology and placed for the first time in a numerical cladistics analysis. All phylogenetic analyses place the jaw in a crownward polytomy of baphetids, temnospondyls, and embolomeres. Several features resemble baphetids and temnospondyls including dermal ornamentation, absence of coronoid teeth, and presence of coronoid shagreen. Dentary dentition is most similar to Baphetes. An adsymphysial toothplate may not preclude temnospondyl affinity. An apparent large exomeckelian fenestra, with the dorsal foraminal margins formed by an unossified element, echoes the morphology of the stem tetrapod Sigournea and is unusually primitive given the other features of the jaw. The jaw may thus provide an example of an intermediate stage in Meckelian element evolution.

The South African species Euparkeria capensis is of great importance for understanding the early radiation of archosauromorphs (including archosaurs) following the Permo–Triassic mass extinction, as most phylogenetic analyses place it as the sister taxon to crown group Archosauria within the clade Archosauriformes. Although a number of species from Lower–Middle Triassic deposits worldwide have been referred to the putative clade Euparkeriidae, the monophyly of Euparkeriidae is controversial and has yet to be demonstrated by quantitative phylogenetic analysis. Three Chinese taxa have been recently suggested to be euparkeriids: Halazhaisuchus qiaoensis, ‘Turfanosuchus shageduensis’, and Wangisuchus tzeyii, all three of which were collected from the Middle Triassic Ermaying Formation of northern China. Here, we reassess the taxonomy and systematics of these taxa. We regard Wangisuchus tzeyii as a nomen dubium, because the holotype is undiagnostic and there is no convincing evidence that the previously referred additional specimens represent the same taxon as the holotype. We also regard ‘Turfanosuchus shageduensis’ as a nomen dubium as we are unable to identify any diagnostic features. We refer the holotype to Archosauriformes, and more tentatively to Euparkeriidae. Halazhaisuchus qiaoensis and the holotype of ‘Turfanosuchus shageduensis’ are resolved as sister taxa in a phylogenetic analysis, and are in turn the sister taxon to Euparkeria capensis, forming a monophyletic Euparkeriidae that is the sister to Archosauria+Phytosauria. This is the first quantitative phylogenetic analysis to recover a non-monospecific, monophyletic Euparkeriidae, but euparkeriid monophyly is only weakly supported and will require additional examination. Given their similar sizes, stratigraphic positions and phylogenetic placement, the holotype of ‘Turfanosuchus shageduensis’ may represent a second individual of Halazhaisuchus qiaoensis, but no apomorphies or unique character combination can be identified to unambiguously unite the two. Our results have important implications for understanding the species richness and palaeobiogeographical distribution of early archosauriforms.

Since its discovery,Euparkeria capensishas been a key taxon for understanding the early evolution of archosaurs. The braincase ofEuparkeriawas described based on a single specimen, but much uncertainty remained. For the first time, all available braincase material ofEuparkeriais re-examined using micro-computed tomography scanning. Contrary to previous work, the parabasisphenoid does not form the posterior border of the fenestra ovalis in lateral view, but it does bear a dorsal projection that forms the anteroventral half of the fenestra. No bone pneumatization was found, but the lateral depression of the parabasisphenoid may have been pneumatic. We propose that the lateral depression likely corresponds to the anterior tympanic recess present in crown archosaurs. The presence of a laterosphenoid is confirmed forEuparkeria. It largely conforms to the crocodilian condition, but shows some features which make it more similar to the avemetatarsalian laterosphenoid. The cochlea ofEuparkeriais elongated, forming a deep cochlear recess. In comparison with other basal archosauromorphs, the metotic foramen is much enlarged and regionalized into vagus and recessus scalae tympani areas, indicating an increase in its pressure-relief mechanism. The anterior semicircular canal is extended and corresponds to an enlarged floccular fossa. These aspects of the braincase morphology may be related to the development of a more upright posture and active lifestyle. They also indicate further adaptations of the hearing system ofEuparkeriato terrestriality.