Bizarre dermal armour suggests the first African ankylosaur.

Ankylosauria is a diverse clade of armoured dinosaurs whose members were important constituents of many Cretaceous faunas. Phylogenetic analyses imply that the clade diverged from its sister taxon, Stegosauria, during the late Early Jurassic, but the fossil records of both clades are sparse until the Late Jurassic (~150 million years ago). Moreover, Ankylosauria is almost entirely restricted to former Laurasian continents, with only a single valid Gondwanan taxon. Spicomellus afer gen. et sp. nov. appears to represent the earliest-known ankylosaur and the first to be named from Africa, from the Middle Jurassic (Bathonian-Callovian) of Morocco, filling an important gap in dinosaur evolution. The specimen consists of a rib with spiked dermal armour fused to its dorsal surface, an unprecedented morphology among extinct and extant vertebrates. The specimen reveals an unrealized morphological diversity of armoured dinosaurs during their early evolution, and implies the presence of an important but undiscovered Gondwanan fossil record.

The anatomy and palaeobiology of the early armoured dinosaur Scutellosaurus lawleri (Ornithischia: Thyreophora) from the Kayenta Formation (Lower Jurassic) of Arizona.

The armoured dinosaurs, Thyreophora, were a diverse clade of ornithischians known from the Early Jurassic to the end of the Cretaceous. During the Middle and Late Jurassic, the thyreophorans radiated to evolve large body size, quadrupedality, and complex chewing mechanisms, and members of the group include some of the most iconic dinosaurs, including the plated Stegosaurus and the club-tailed Ankylosaurus; however, the early stages of thyreophoran evolution are poorly understood due to a paucity of relatively complete remains from early diverging thyreophoran taxa. Scutellosaurus lawleri is generally reconstructed as the earliest-diverging thyreophoran and is known from over 70 specimens from the Lower Jurassic Kayenta Formation of Arizona, USA. Whereas Scutellosaurus lawleri is pivotal to our understanding of character-state changes at the base of Thyreophora that can shed light on the early evolution of the armoured dinosaurs, the taxon has received limited study. Herein, we provide a detailed account of the osteology of Scutellosaurus lawleri, figuring many elements for the first time. Scutellosaurus lawleri was the only definitive bipedal thyreophoran. Histological studies indicate that it grew slowly throughout its life, possessing lamellar-zonal tissue that was a consequence neither of its small size nor phylogenetic position, but may instead be autapomorphic, and supporting other studies that suggest thyreophorans had lower basal metabolic rates than other ornithischian dinosaurs. Faunal diversity of the Kayenta Formation in comparison with other well-known Early Jurassic-aged dinosaur-bearing formations indicates that there was considerable spatial and/or environmental variation in Early Jurassic dinosaur faunas.

Dinosaur diversification rates were not in decline prior to the K-Pg boundary.

Determining the tempo and mode of non-avian dinosaur extinction is one of the most contentious issues in palaeobiology. Extensive disagreements remain over whether their extinction was catastrophic and geologically instantaneous or the culmination of long-term evolutionary trends. These conflicts have arisen due to numerous hierarchical sampling biases in the fossil record and differences in analytical methodology, with some studies identifying long-term declines in dinosaur richness prior to the Cretaceous-Palaeogene (K-Pg) boundary and others proposing continued diversification. Here, we use Bayesian phylogenetic generalized linear mixed models to assess the fit of 12 dinosaur phylogenies to three speciation models (null, slowdown to asymptote, downturn). We do not find strong support for the downturn model in our analyses, which suggests that dinosaur speciation rates were not in terminal decline prior to the K-Pg boundary and that the clade was still capable of generating new taxa. Nevertheless, we advocate caution in interpreting the results of such models, as they may not accurately reflect the complexities of the underlying data. Indeed, current phylogenetic methods may not provide the best test for hypotheses of dinosaur extinction; the collection of more dinosaur occurrence data will be essential to test these ideas further.

The postcranial skeleton of the erythrosuchid archosauriform Garjainia prima from the Early Triassic of European Russia.

Erythrosuchidae were large-bodied, quadrupedal, predatory archosauriforms that dominated the hypercarnivorous niche in the aftermath of the Permo-Triassic mass extinction. Garjainia, one of the oldest members of the clade, is known from the late Olenekian of European Russia. The holotype of Garjainia prima comprises a well-preserved skull, but highly incomplete postcranium. Recent taxonomic reappraisal demonstrates that material from a bone bed found close to the type locality, previously referred to as 'Vjushkovia triplicostata', is referable to G. prima. At least, seven individuals comprising cranial remains and virtually the entire postcranium are represented, and we describe this material in detail for the first time. An updated phylogenetic analysis confirms previous results that a monophyletic Garjainia is the sister taxon to a clade containing Erythrosuchus, Shansisuchus and Chalishevia. Muscle scars on many limb elements are clear, allowing reconstruction of the proximal locomotor musculature. We calculate the body mass of G. prima to have been 147-248 kg, similar to that of an adult male lion. Large body size in erythrosuchids may have been attained as part of a trend of increasing body size after the Permo-Triassic mass extinction and allowed erythrosuchids to become the dominant carnivores of the Early and Middle Triassic.

Cranial anatomy and taxonomy of the erythrosuchid archosauriform 'Vjushkovia triplicostata' Huene, 1960, from the Early Triassic of European Russia.

Erythrosuchidae are a globally distributed and important group of apex predators that occupied Early and Middle Triassic terrestrial ecosystems following the Permo-Triassic mass extinction. The stratigraphically oldest known genus of Erythrosuchidae is Garjainia Ochev, 1958, which is known from the late Early Triassic (late Olenekian) of European Russia and South Africa. Two species of Garjainia have been reported from Russia: the type species, Garjainia prima Ochev, 1958, and 'Vjushkovia triplicostata' von Huene, 1960, which has been referred to Garjainia as either congeneric (Garjainia triplicostata) or conspecific (G. prima). The holotype of G. prima has received relatively extensive study, but little work has been conducted on type or referred material attributed to 'V. triplicostata'. However, this material includes well-preserved fossils representing all parts of the skeleton and comprises seven individuals. Here, we provide a comprehensive description and review of the cranial anatomy of material attributed to 'V. triplicostata', and draw comparisons with G. prima. We conclude that the two Russian taxa are indeed conspecific, and that minor differences between them result from a combination of preservation or intraspecific variation. Our reassessment therefore provides additional information on the cranial anatomy of G. prima. Moreover, we quantify relative head size in erythrosuchids and other early archosauromorphs in an explicit phylogenetic context for the first time. Our results show that erythrosuchids do indeed appear to have disproportionately large skulls, but that this is also true for other early archosauriforms (i.e. proterosuchids), and may reflect the invasion of hypercarnivorous niches by these groups following the Permo-Triassic extinction.

The systematic position of the enigmatic thyreophoran dinosaur Paranthodon africanus, and the use of basal exemplifiers in phylogenetic analysis.

The first African dinosaur to be discovered, Paranthodon africanus was found in 1845 in the Lower Cretaceous of South Africa. Taxonomically assigned to numerous groups since discovery, in 1981 it was described as a stegosaur, a group of armoured ornithischian dinosaurs characterised by bizarre plates and spines extending from the neck to the tail. This assignment has been subsequently accepted. The type material consists of a premaxilla, maxilla, a nasal, and a vertebra, and contains no synapomorphies of Stegosauria. Several features of the maxilla and dentition are reminiscent of Ankylosauria, the sister-taxon to Stegosauria, and the premaxilla appears superficially similar to that of some ornithopods. The vertebral material has never been described, and since the last description of the specimen, there have been numerous discoveries of thyreophoran material potentially pertinent to establishing the taxonomic assignment of the specimen. An investigation of the taxonomic and systematic position of Paranthodon is therefore warranted. This study provides a detailed re-description, including the first description of the vertebra. Numerous phylogenetic analyses demonstrate that the systematic position of Paranthodon is highly labile and subject to change depending on which exemplifier for the clade Stegosauria is used. The results indicate that the use of a basal exemplifier may not result in the correct phylogenetic position of a taxon being recovered if the taxon displays character states more derived than those of the basal exemplifier, and we recommend the use, minimally, of one basal and one derived exemplifier per clade. Paranthodon is most robustly recovered as a stegosaur in our analyses, meaning it is one of the youngest and southernmost stegosaurs.