The oldest fossil bird-like footprints from the upper Triassic of southern Africa.

Footprint morphology reflects the anatomy of the trackmaker's foot and is direct evidence for the animal's behaviour. Consequently, fossil tracks can be used to infer ancient diversity, ethology, and evolutionary trends. This is particularly useful for deep-time intervals during which the early history of an animal group is reliant upon limited fossil skeletal material. Fossil tracks of early birds and theropods, the co-existing dinosaurian ancestors of birds, co-occur in the rock record since the Early Cretaceous. However, the evolutionary transition from dinosaur to bird and the timing of the birds' origin are still contested. Skeletal remains of the basal-most birds Aurornis, Anchiornis, Archaeopteryx and Xiaotingia are Middle to Late Jurassic, while tracks with tentative bird affinities, attributed to dinosaurs, are known from as early as the Late Triassic. Here, we present numerous, well-provenanced, Late Triassic and Early Jurassic tridactyl tracks from southern Africa, with demonstrable bird-like affinities, predating basal bird body fossils by c. 60 million years.

Basal sauropodomorph locomotion: ichnological lessons from the Late Triassic trackways of bipeds and quadrupeds (Elliot Formation, main Karoo Basin).

Using modern ichnological and stratigraphic tools, we reinvestigate two iconic sauropodomorph-attributed tetradactyl ichnogenera, Pseudotetrasauropus and Tetrasauropus, and their stratigraphic occurrences in the middle Upper Triassic of Lesotho. These tracks have been reaffirmed and are stratigraphically well-constrained to the lower Elliot Formation (Stormberg Group, Karoo Basin) with a maximum depositional age range of <219-209 Ma (Norian). This represents the earliest record of basal sauropodomorph trackways in Gondwana, if not globally. Track and trackway morphology, the sedimentary context of the tracks, and unique features (e.g., drag traces) have enabled us to discuss the likely limb postures and gaits of the trackmakers. Pseudotetrasauropus has bipedal (P. bipedoida) and quadrupedal (P. jaquesi) trackway states, with the oldest quadrupedal Pseudotetrasauropus track and trackway parameters suggestive of a columnar, graviportal limb posture in the trackmaker. Moreover, an irregularity in the intermanus distance and manus orientation and morphology, in combination with drag traces, is indicative of a non-uniform locomotory suite or facultative quadrupedality. Contrastingly, Tetrasauropus, the youngest trackway, has distinctive medially deflected, robust pedal and manual claw traces and a wide and uniform intermanus distance relative to the interpedal. These traits suggest a quadrupedal trackmaker with clawed and fleshy feet and forelimbs held in a wide, flexed posture. Altogether, these trackways pinpoint the start of the southern African ichnological record of basal sauropodomorphs with bipedal and quadrupedal locomotory habits to, at least, c. 215 Ma in the middle Late Triassic.

A new Heterodontosaurus specimen elucidates the unique ventilatory macroevolution of ornithischian dinosaurs.

Ornithischian dinosaurs were ecologically prominent herbivores of the Mesozoic Era that achieved a global distribution by the onset of the Cretaceous. The ornithischian body plan is aberrant relative to other ornithodiran clades, and crucial details of their early evolution remain obscure. We present a new, fully articulated skeleton of the early branching ornithischian Heterodontosaurus tucki. Phase-contrast enhanced synchrotron data of this new specimen reveal a suite of novel postcranial features unknown in any other ornithischian, with implications for the early evolution of the group. These features include a large, anteriorly projecting sternum; bizarre, paddle-shaped sternal ribs; and a full gastral basket - the first recovered in Ornithischia. These unusual anatomical traits provide key information on the evolution of the ornithischian body plan and suggest functional shifts in the ventilatory apparatus occurred close to the base of the clade. We complement these anatomical data with a quantitative analysis of ornithischian pelvic architecture, which allows us to make a specific, stepwise hypothesis for their ventilatory evolution.

The fossilised skeletons of long extinct dinosaurs are more than just stones. By comparing these remains to their living relatives such as birds and crocodiles, palaeontologists can reveal how dinosaurs grew, moved, ate and socialised. Previous research indicates that dinosaurs were likely warm-blooded and also more active than modern reptiles. This means they would have required breathing mechanisms capable of supplying enough oxygen to allow these elevated activity levels. So far, much of our insight into dinosaur breathing biology has been biased towards dinosaur species more closely related to modern birds, such as Tyrannosaurus rex, as well as the long-necked sauropods. The group of herbivorous dinosaurs known as ornithischians, which include animals with head ornamentation, spikes and heavy body armour, like that found in Triceratops and Stegosaurus, have often been overlooked. As a result, there are still significant gaps in ornithischian biology, especially in understanding how they breathed. Radermacher et al. used high-powered X-rays to study a new specimen of the most primitive ornithischian dinosaur, Heterodontosaurus tucki, and discovered that this South African dinosaur has bones researchers did not know existed in this species. These include bones that are part of the breathing system of extant reptiles and birds, including toothpick-shaped bones called gastralia, paired sternal bones and sternal ribs shaped like tennis rackets. Together, these new pieces of anatomy form a complicated chest skeleton with a large range of motion that would have allowed the body to expand during breathing cycles. But this increased motion of the chest was only possible in more primitive ornithischians. More advanced species lost much of the anatomy that made this motion possible. Radermacher et al. show that while the chest was simpler in advanced species, their pelvis was more specialised and likely played a role in breathing as it does in modern crocodiles. This new discovery could inform the work of biologists who study the respiratory diversity of both living and extinct species. Differences in breathing strategies might be one of the underlying reasons that some lineages of animals go extinct. It could explain why some species do better than others under stressful conditions, like when the climate is warmer or has less oxygen.

Tracking the Pliensbachian-Toarcian Karoo firewalkers: Trackways of quadruped and biped dinosaurs and mammaliaforms.

The Karoo igneous rocks represent one of the largest continental flood basalt events (by volume) on Earth, and are not normally associated with fossils remains. However, these Pliensbachian-Toarcian lava flows contain sandstone interbeds that are particularly common in the lower part of the volcanic succession and are occasionally fossiliferous. On a sandstone interbed in the northern main Karoo Basin, we discovered twenty-five tridactyl and tetradactyl vertebrate tracks comprising five trackways. The tracks are preserved among desiccation cracks and low-amplitude, asymmetrical ripple marks, implying deposition in low energy, shallow, ephemeral water currents. Based on footprint lengths of 2-14 cm and trackway patterns, the trackmakers were both bipedal and quadrupedal animals of assorted sizes with walking and running gaits. We describe the larger tridactyl tracks as "grallatorid" and attribute them to bipedal theropod dinosaurs, like Coelophysis, a genus common in the Early Jurassic of southern Africa. The smallest tracks are tentatively interpreted as Brasilichnium-like tracks, which are linked to synapsid trackmakers, a common attribution of similar tracks from the Lower to Middle Jurassic record of southern and southwestern Gondwana. The trackway of an intermediate-sized quadruped reveals strong similarities in morphometric parameters to a post-Karoo Zimbabwean trackway from Chewore. These trackways are classified here as a new ichnogenus attributable to small ornithischian dinosaurs as yet without a body fossil record in southern Africa. These tracks not only suggest that dinosaurs and therapsids survived the onset of the Drakensberg volcanism, but also that theropods, ornithischians and synapsids were among the last vertebrates that inhabited the main Karoo Basin some 183 Ma ago. Although these vertebrates survived the first Karoo volcanic eruptions, their rapidly dwindling habitat was turned into a land of fire as it was covered by the outpouring lavas during one of the most dramatic geological episodes in southern Africa.

A micro X-ray computed tomography dataset of fossil echinoderms in an ancient obrution bed: a robust method for taphonomic and palaeoecologic analyses.

BACKGROUND: Taphonomic and palaeoecologic studies of obrution beds often employ conventional methods of investigation such as physical removal and extraction of fossils from their host rock (matrix) by mechanical preparation. This often-destructive method is not suitable for studying mold fossils, which are voids left in host rocks due to dissolution of the original organism in post-depositional processes. FINDINGS: Microcomputed tomography (µCT) scan data of 24 fossiliferous rock samples revealed thousands of Paleozoic echinoderms. Digitally "stitching" together individually µCT scanned rock samples within three-dimensional (3D) space allows for quantifiable taphonomic data on a fossil echinoderm-rich obrution deposit from the Devonian (Emsian) of South Africa. Here, we provide a brief step-by-step guide on creating, segmenting, and ultimately combining sections of richly fossiliferous beds to create virtual models suited for the quantitative and qualitative taphonomic analyses of fossil invertebrate assemblages. CONCLUSIONS: Visualizing the internal features of fossiliferous beds in 3D is an invaluable taphonomic tool for analyzing delicate fossils, accounting for all specimens irrespective of their preservation stages and with minimal damage. This technique is particularly useful for analyzing fossiliferous deposits with mold fossils that prove to be difficult to study with traditional methods, because the method relies on the large density contrast between the mold and host rock.

A Giant Dinosaur from the Earliest Jurassic of South Africa and the Transition to Quadrupedality in Early Sauropodomorphs.

Sauropod dinosaurs were dominant, bulk-browsing herbivores for 130 million years of the Mesozoic, attaining gigantic body masses in excess of 60 metric tons [1, 2]. A columnar-limbed, quadrupedal posture enabled these giant body sizes [3], but the nature of the transition from bipedal sauropodomorph ancestors to derived quadrupeds remains contentious [4-6]. We describe a gigantic, new sauropodomorph from the earliest Jurassic of South Africa weighing 12 metric tons and representing a phylogenetically independent origin of sauropod-like body size in a non-sauropod. Osteohistological evidence shows that this specimen was an adult of maximum size and approximately 14 years old at death. Ledumahadi mafube gen. et sp. nov. shows that gigantic body sizes were possible in early sauropodomorphs, which were habitual quadrupeds but lacked the derived, columnar limb postures of sauropods. We use data from this new taxon and a discriminant analysis of tetrapod limb measurements to study postural evolution in sauropodomorphs. Our results show that quadrupedality appeared by the mid-Late Triassic (Norian), well outside of Sauropoda. Secondary reversion to bipedality occurred in some lineages phylogenetically close to Sauropoda, indicating early experimentation in locomotory styles. Morphofunctional observations support the hypothesis that partially flexed (rather than columnar) limbs characterized Ledumahadi and other early-branching quadrupedal sauropodomorphs. Patterns of locomotory and body-size evolution show that quadrupedality allowed Triassic sauropodomorphs to achieve body sizes of at least 3.8 metric tons. Ledumahadi's Early Jurassic age shows that maximum body mass in sauropodomorph dinosaurs was either unaffected or rapidly rebounded after the end-Triassic extinction event.

Digital reconstruction of the mandible of an adult Lesothosaurus diagnosticus with insight into the tooth replacement process and diet.

Fragmentary caudal ends of the left and right mandible assigned to Lesothosaurus diagnosticus, an early ornithischian, was recently discovered in the continental red bed succession of the upper Elliot Formation (Lower Jurassic) at Likhoele Mountain (Mafeteng District) in Lesotho. Using micro-CT scanning, this mandible could be digitally reconstructed in 3D. The replacement teeth within the better preserved (left) dentary were visualised. The computed tomography dataset suggests asynchronous tooth replacement in an individual identified as an adult on the basis of bone histology. Clear evidence for systematic wear facets created by attrition is lacking. The two most heavily worn teeth are only apically truncated. Our observations of this specimen as well as others do not support the high level of dental wear expected from the semi-arid palaeoenvironment in which Lesothosaurus diagnosticus lived. Accordingly, a facultative omnivorous lifestyle, where seasonality determined the availability, quality, and abundance of food is suggested. This would have allowed for adaptability to episodes of increased environmental stress.

Sedimentology and ichnology of the Mafube dinosaur track site (Lower Jurassic, eastern Free State, South Africa): a report on footprint preservation and palaeoenvironment.

Footprint morphology (e.g., outline shape, depth of impression) is one of the key diagnostic features used in the interpretation of ancient vertebrate tracks. Over 80 tridactyl tracks, confined to the same bedding surface in the Lower Jurassic Elliot Formation at Mafube (eastern Free State, South Africa), show large shape variability over the length of the study site. These morphological differences are considered here to be mainly due to variations in the substrate rheology as opposed to differences in the trackmaker's foot anatomy, foot kinematics or recent weathering of the bedding surface. The sedimentary structures (e.g., desiccation cracks, ripple marks) preserved in association with and within some of the Mafube tracks suggest that the imprints were produced essentially contemporaneous and are true dinosaur tracks rather than undertracks or erosional remnants. They are therefore valuable not only for the interpretation of the ancient environment (i.e., seasonally dry river channels) but also for taxonomic assessments as some of them closely resemble the original anatomy of the trackmaker's foot. The tracks are grouped, based on size, into two morphotypes that can be identified as Eubrontes-like and Grallator-like ichnogenera. The Mafube morphotypes are tentatively attributable to large and small tridactyl theropod trackmakers, possibly to Dracovenator and Coelophysis based on the following criteria: (a) lack of manus impressions indicative of obligate bipeds; (b) long, slender-digits that are asymmetrical and taper; (c) often end in a claw impression or point; and (d) the tracks that are longer than broad. To enable high-resolution preservation, curation and subsequent remote studying of the morphological variations of and the secondary features in the tracks, low viscosity silicone rubber was used to generate casts of the Mafube tracks.