Cranial functional specialisation for strength precedes morphological evolution in Oviraptorosauria.

Oviraptorosaurians were a theropod dinosaur group that reached high diversity in the Late Cretaceous. Within oviraptorosaurians, the later diverging oviraptorids evolved distinctive crania which were extensively pneumatised, short and tall, and had a robust toothless beak, interpreted as providing a powerful bite for their herbivorous to omnivorous diet. The present study explores the ability of oviraptorid crania to resist large mechanical stresses compared with other theropods and where this adaptation originated within oviraptorosaurians. Digital 3D cranial models were constructed for the earliest diverging oviraptorosaurian, Incisivosaurus gauthieri, and three oviraptorids, Citipati osmolskae, Conchoraptor gracilis, and Khaan mckennai. Finite element analyses indicate oviraptorosaurian crania were stronger than those of other herbivorous theropods (Erlikosaurus and Ornithomimus) and were more comparable to the large, carnivorous Allosaurus. The cranial biomechanics of Incisivosaurus align with oviraptorids, indicating an early establishment of distinctive strengthened cranial biomechanics in Oviraptorosauria, even before the highly modified oviraptorid cranial morphology. Bite modelling, using estimated muscle forces, suggests oviraptorid crania may have functioned closer to structural safety limits. Low mechanical stresses around the beaks of oviraptorids suggest a convergently evolved, functionally distinct rhamphotheca, serving as a cropping/feeding tool rather than for stress reduction, when compared with other herbivorous theropods.

Hwiccewyrm trispiculum gen. et sp. nov., a new leptopleuronine procolophonid from the Late Triassic of southwest England.

The fissure fill localities of southwest England and South Wales are well-known for preserving rich assemblages of predominantly small-bodied Late Triassic to Early Jurassic tetrapods, but many aspects of these assemblages remain contentious. The age of the Late Triassic fissures is disputed, with some lines of argument suggesting a latest Triassic (Rhaetian) age, whereas other evidence suggests they may be as old as Carnian. The fissures have been hypothesized by some workers to have formed on an archipelago, with island effects invoked to explain aspects of the assemblages such as the abundance of small-bodied species. Procolophonids were a successful group of Triassic parareptiles, best known from Early to early Late Triassic assemblages, but have only recently been described from one of the fissure fill sites (Ruthin) based upon fragmentary remains. Here, we describe new procolophonid specimens from another fissure (Cromhall) that represent at least six individuals of different sizes, with much of the skeleton represented including well-preserved skull material. The Cromhall procolophonid shows strong similarities to Late Triassic procolophonids from Scotland, Brazil and North America, but both autapomorphies and a unique character combination demonstrate that it represents a new species, which we name as Hwiccewyrm trispiculum gen. et sp. nov. Phylogenetic analysis places Hwiccewyrm in a derived clade within Leptopleuroninae, together with Leptopleuron, Hypsognathus, and Soturnia. The largest specimens of Hwiccewyrm demonstrate a body size that is similar to Leptopleuron and Hypsognathus, supporting other recent work that has questioned the insular dwarfism hypothesis for the fissure fill assemblages.

A toolbox for the retrodeformation and muscle reconstruction of fossil specimens in Blender.

Accurate muscle reconstructions can offer new information on the anatomy of fossil organisms and are also important for biomechanical analysis (multibody dynamics and finite-element analysis (FEA)). For the sake of simplicity, muscles are often modelled as point-to-point strands or frustra (cut-off cones) in biomechanical models. However, there are cases in which it is useful to model the muscle morphology in three dimensions, to better examine the effects of muscle shape and size. This is especially important for fossil analyses, where muscle force is estimated from the reconstructed muscle morphology (rather than based on data collected in vivo). The two main aims of this paper are as follows. First, we created a new interactive tool in the free open access software Blender to enable interactive three-dimensional modelling of muscles. This approach can be applied to both palaeontological and human biomechanics research to generate muscle force magnitudes and lines of action for FEA. Second, we provide a guide on how to use existing Blender tools to reconstruct distorted or incomplete specimens. This guide is aimed at palaeontologists but can also be used by anatomists working with damaged specimens or to test functional implication of hypothetical morphologies.

Cranial muscle reconstructions quantify adaptation for high bite forces in Oviraptorosauria.

Oviraptorosaurians are an unusual and probably herbivorous group of theropod dinosaurs that evolved pneumatised crania with robust, toothless jaws, apparently adapted for producing a strong bite. Using 3D retrodeformed skull models of oviraptorid oviraptorosaurians Citipati, Khaan, and Conchoraptor, along with the earliest diverging oviraptorosaurian, Incisivosaurus, we digitally reconstruct jaw adductor musculature and estimate bite force to investigate cranial function in each species. We model muscle length change during jaw opening to constrain optimal and maximum gape angles. Results demonstrate oviraptorids were capable of much stronger bite forces than herbivorous theropods among Ornithomimosauria and Therizinosauria, relative to body mass and absolutely. Increased bite forces in oviraptorid oviraptorosaurians compared to the earliest diverging oviraptorosaurian result from expanded muscular space and different cranial geometry, not changes in muscular arrangement. Estimated optimal and maximum possible gapes are much smaller than published estimates for carnivorous theropods, being more similar to the herbivorous therizinosaurian theropod Erlikosaurus and modern birds. Restrictive gape and high bite force may represent adaptation towards exploiting tough vegetation, suggesting cranial function and dietary habits differed between oviraptorids and other herbivorous theropods. Differences in the relative strength of jaw adductor muscles between co-occurring oviraptorids may be a factor in niche partitioning, alongside body size.

Reconstructing development of the earliest seed integuments raises a new hypothesis for the evolution of ancestral seed-bearing structures.

How plant seeds originated remains unresolved, in part due to disconnects between fossil intermediates and developmental genetics in extant species. The Carboniferous fossil Genomosperma is considered among the most primitive known seeds, with highly lobed integument and exposed nucellus. We have used this key fossil taxon to investigate the evolutionary origins of seed development. We examined sectioned Genomosperma specimens using modern digital 3D reconstruction techniques and established population-level measurements of Genomosperma ovules for quantitative analysis. Genomosperma ovules show significant variation in integumentary lobe fusion and curvature. Our analysis suggests that this variation represents a single species with significant variations in lobe number and fusion, reminiscent of floral development in extant species. We conclude that changes in lobe flexure occurred late in development, consistent with a previously hypothesized function in pollen guidance/retention. We also identify seeds of Genomosperma within cupules for the first time. The presence of a cupule adds evidence towards the plesiomorphy of cupules within seed plants. Together with the similarities identified between the Genomosperma lobed integument and floral organs, we propose that the cupule, integument and nucellus together developed in a shoot-like fashion, potentially ancestral to extant seed plant reproductive shoots.

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.

A revision of tetrapod footprints from the late Carboniferous of the West Midlands, UK.

A series of sandstone slabs from Hamstead, Birmingham (West Midlands, UK), preserve an assemblage of tetrapod trackways and individual tracks from the Enville Member of the Salop Formation (late Carboniferous: late Moscovian-Kasimovian). This material has received limited previous study, despite being one of the few British sites to preserve Carboniferous tetrapod footprints. Here, we restudy and revise the taxonomy of this material, and document it using 3D models produced using photogrammetry. The assemblage is dominated by large tracks assigned to Limnopus isp., which were made by early amphibians (temnospondyls). A number of similar but smaller tracks are assigned to Batrachichnus salamandroides (also made by temnospondyls). Dimetropus leisnerianus (made by early synapsids) and Dromopus lacertoides (made by lizard-like sauropsids such as araeoscelids) are also present. This ichnofauna contrasts with a slightly stratigraphically older, more extensive and better-studied assemblage from Alveley (Shropshire), which is dominated by small amphibians with relatively rare reptiliomorphs, but which lacks Dromopus tracks. The presence of Dromopus lacertoides at Hamstead is consistent with the trend towards increasing aridity through the late Carboniferous. It is possible that the assemblage is the stratigraphically oldest occurrence of this important amniote ichnotaxon.