Evaluation of tridactyl theropod tracks in southern Africa: quantitative morphometric analysis across the Triassic–Jurassic boundary

In the Mesozoic, dinosaur abundance and diversity steadily increased from the Carnian to the Triassic booming soon after the end Triassic Mass-Extinction event (ETE), marking a key period in archosaur history. In southern Africa, the Triassic–Jurassic Boundary (TJB) is contained in the richly fossiliferous, fluvio-lacustrine-aeolian Upper Triassic to Lower Jurassic strata of the Stormberg Group. More specifically, the middle Norian – Pliensbachian Elliot and Clarens formations (upper Stormberg Group) of main Karoo Basin, host a diverse tetrapod osteological and ichnite record. Due to an absence of high resolution radioisotopic age determinations, the exact stratigraphic placement of the TJB remains unknown. Although diverse ichnofossils attributed to Saurischians and Ornithischians are preserved in the Stormberg Group, the record is dominated by isolated tridactyl tracks that can be assigned to common ichnogenera like Grallator, Eubrontes and Kayentapus. Ideally, these track morphologies would reflect the trackmaker's autopod morphology, but complex interactions between the trackmaker and tracking substrate may affect the final footprint shape. Tracks with a high morphological preservation grade may be used to infer information (e.g., body length, hip height, weight) about the trackmaker, which is especially useful when skeletal remains are scarce, as is the case with theropod body fossils in the Elliot and Clarens formations. Herein, we present the findings of an extensive southern African field-based study to quantify the morphological variation of Grallator, Eubrontes and Kayentapus tracks across the TJB in the upper Stormberg Group. Furthermore, this study produced the first detrital zircon Uranium–Lead (U-Pb) LA-ICPMS ages of the major ichnosites from this region, and confirmed that the TJB is within the Elliot Formation, near the boundary of the lower and upper Elliot Formation (i.e., near the contact of the informal lEF and uEF). Across this contact, the considered tridactyl tracks become more abundant, larger and have a less pronounced medial digit projection. These morphological changes are gradational, with tracks from the Clarens Formation being distinct to tracks from the lEF, while the uEF tracks being intermediate between the lEF and Clarens Formation. A decrease in the mesaxony (Dp/TS ratio) and a decrease in medial digit projection relative to track length can be detected in both small and large tridactyl tracks. These apparent trends in the upper Stormberg Group are consistent with global tridactyl trends, which suggest an overall increase in theropod abundance and body size across the Jurassic. Moreover, the reason for the less prominent medial pedal digit is linked to a better weight distribution across the autopod in the increasingly larger theropods. Last but not least, Grallator, Eubrontes and Kayentapus ichnogenera which may be attributed to at least three different groups of theropod-like trackmakers, suggest a higher palaeo-diversity and abundance of tridactyl dinosaurs in southern Africa than is known from the osteological record

Quantification of morphological variation in Late Triassic to Early Jurassic theropod tracks of southern Africa

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Re-evaluating the wealth of the Late Triassic Phuthiatsana ichnosite, Maseru District, Lesotho

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The Late Sinemurian ichnofossils from Mampoboleng (Upper Moyeni), Lesotho

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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

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

Large tridactyl dinosaur tracks from the Early Jurassic of southern Gondwana : uppermost Elliot Formation, Upper Moyeni, Lesotho

A new ichnosite in southwest Lesotho (Upper Moyeni; Quthing District) is located within the uppermost part of the highly fossiliferous Elliot Formation, ~35 m below the conformably overlying Clarens Formation and ~65 m above the world-renowned Lower Moyeni ichnosite. While the Lower Moyeni site preserves diverse Early Jurassic ichnofossils, the ichnites at the Upper Moyeni comprise one vertebrate burrow and ~50 tridactyl tracks with footprint lengths between 15 and 51 cm. Many of the tracks preserve digital pad impressions, claw marks and displacement rims, all related to substrate conditions. The morphometric parameters of the Upper Moyeni tracks are consistent with Grallator, Eubrontes and Kayentapus. Several larger tracks with footprint lengths > 40 cm are Kayentapus-like and Eubrontes-like, and are comparable to previously described very large theropods tracks with lengths > 50 cm from the uppermost Elliot and Clarens formations. On the basis of sedimentological and ichnological evidence, the Upper Moyeni ichnofossils were formed in a palaeolandscape with small rivers and shallow lakes by burrowing tetrapods and a variety of bipedal dinosaurs (theropods), some of which were up to 7–8 m in body length. The Upper Moyeni tracks, together with the other very large tracks from coeval locations in southern Africa, collectively highlight the tendency towards increasing diversity in size of tridactyl tracks and by extension theropod trackmaker body size body size, which runs in tandem with the increasing diversity of non-sauropod, sauropodomorph body fossils in the Sinemurian-Pliensbachian of southern Gondwana

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

Scampering, trotting, walking tridactyl bipedal dinosaurs in southern Africa: ichnological account of a Lower Jurassic palaeosurface (upper Elliot Formation, Roma Valley) in Lesotho

<p>In Gondwana, Early Jurassic dinosaur track sites are especially concentrated in Lesotho. Despite intensive investigations during the third quarter of the twentieth century, a limited number of vertebrate track sites of this country have been studied with rigorous ichnological and sedimentological methodology. Here, we present a previously mentioned, but undescribed track site in the upper Elliot Formation (Hettangian?) of Lesotho, located near Roma (at Lephoto dam). Two tridactyl ichnite morphologies, made by bipedal vertebrate trackmakers are recognised. The first can be identified as <i>Grallator</i>-like, an ichnotaxon common in the Lower Jurassic of both Laurasia and Gondwana that can be attributed to small and medium-size theropod dinosaurs. In contrast, the second ichnite type is reminiscent of <i>Trisauropodiscus</i>, which is a rare ichnotaxon that resembles tracks of small birds and is known with certainty in Lesotho from only a few places. We suggest that at our upper Elliot Formation study site, <i>Trisauropodiscus</i> was potentially made by a heterodontosaurid ornithischian dinosaur. Our work provides further evidence that the ichnological record of the Stormberg Group of southern Africa is in a unique position to shed light not only on Early Jurassic biostratigraphy and palaeoenvironments but also on the biodiversity and palaeobiology of early dinosaurs.</p