The upper Stormberg Group (Elliot and Clarens formations) of the main Karoo Basin is well-known for its fossil vertebrate fauna, comprising early branching members of lineages including mammals, dinosaurs, and testudinates. Despite 150 years of scientific study, the upper Stormberg Group lacks radioisotopic age constraints and remains coarsely dated via imprecise faunal correlations. Here we synthesise previous litho-and magnetostratigraphic studies, and present a comprehensive biostratigraphic review of upper Stormberg fauna. We also present the results of the first geochronological assessment of the unit across the basin, using U-Pb dates derived from detrital zircons obtained from tuffaceous sandstones and siltstones, the youngest of which are considered maximum depositional ages. Our results confirm that the Elliot Formation contains the Triassic-Jurassic boundary, making it one of the few fossiliferous continental units that records the effects of the end-Triassic Mass Extinction event. Our work suggests a mid-Norian-Rhaetian age for the lower Elliot Formation and a Hettangian-Sinemurian age for the upper Elliot Formation, although the precise stratigraphic position of the Triassic/Jurassic (Rhaetian/Hettangian) boundary remains somewhat uncertain. A mainly Pliensbachian age is obtained for the Clarens Formation. The new dates allow direct comparison with better-calibrated Triassic-Jurassic faunas of the Western Hemisphere (e.g., Chinle and Los Colorados formations). We show that sauropodomorph, but not ornithischian or theropod, dinosaurs were well-established in the main Karoo Basin ~220 million years ago, and that typical Norian faunas (e.g., aetosaurs, phytosaurs) are either rare or absent in the lower Elliot Formation, which is paucispecific compared to the upper Elliot. While this is unlikely the result of geographic sampling biases, it could be from historical sampling intensity differences.

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 Brasilichniumlike 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.

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.

A palaeosurface with one megatheropod trackway and several theropod tracks and trackways from the Lower Jurassic upper Elliot Formation (Stormberg Group, Karoo Supergroup) in western Lesotho is described. The majority of the theropod tracks are referable to either Eubrontes or Kayentapus based on their morphological characteristics. The larger megatheropod tracks are 57 cm long and have no Southern Hemisphere equivalent. Morphologically, they are more similar to the Early Jurassic Kayentapus, as well as the much younger Upper Cretaceous ichnogenus Irenesauripus, than to other contemporaneous ichnogenera in southern Africa. Herein they have been placed within the ichnogenus Kayentapus and described as a new ichnospecies (Kayentapus ambrokholohali). The tracks are preserved on ripple marked, very fine-grained sandstone of the Lower Jurassic upper Elliot Formation, and thus were made after the end-Triassic mass extinction event (ETE). This new megatheropod trackway site marks the first occurrence of very large carnivorous dinosaurs (estimated body length >8–9 meters) in the Early Jurassic of southern Gondwana, an evolutionary strategy that was repeatedly pursued and amplified in the following ~135 million years, until the next major biotic crisis at the end-Cretaceous.

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The end-Triassic mass extinction events mark a pivotal period in archosaur history, and have been proposed to contribute to the rise and dominance of dinosaurs throughout the Mesozoic. In southern Africa, the Triassic–Jurassic boundary is contained within the richly fossiliferous fluvio-lacustrine-aeolian deposits of the upper Stormberg Group in the main Karoo Basin. Due to an absence of high-resolution radioisotopic age constraints, the exact placement of the boundary remains difficult. The Stormberg Group theropod osteological record is limited to scarce, fragmentary material; therefore, the abundant Norian–Pliensbachian tridactyl tracks attributed to theropods are vital for unraveling theropod dinosaur evolutionary trends in southwestern Gondwana. This study considers over 200 upper Stormberg Group tridactyl tracks assigned to the Kayentapus-Grallator-Anchisauripus-Eubrontes (K-GAE) plexus, to quantify their morphological variation across a time span of ∼35 million years. Our findings show that within the upper Stormberg Group, and across the Triassic–Jurassic boundary, the younger tracks become larger, have a decreased mesaxony and a reduced digit III projection. This reduced emphasis of the medial digit is also observed across the K-GAE plexus, and for the individual ichnotaxa across time in the main Karoo Basin, e.g., Eubrontes tracks become less mesaxonic and have a reduced digit III projection higher up in the stratigraphy. This suggests that these morphological trends are not simply linked to size but may reflect a change in autopod morphology through time, which has implications for pedal functionality. Furthermore, being morphologically distinct from contemporaneous North American K-GAE tracks (e.g., reduced elongation and mesaxony, no correlation between digit divarication angles and size), these southern African footprints warrant further investigation.