School of Geosciences (ETDs)

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Cranial orientation and the lateral semicircular canal in primates: implications for palaeobiological reconstructions and the evolution of locomotor repertoires
(University of the Witwatersrand, Johannesburg, 2023) Pestana, Christopher; Benoit, Julien; Beaudet, Amélie
The lateral semicircular canal and its predicted relation to head posture have been used in reconstructions of locomotion and posture of contemporary and extinct species, and in the evolution of bipedalism. Inferences of head posture in fossil species sometimes assume that the lateral semicircular canal is held near the earth’s horizontal when the head is at rest. Despite the physiological importance of the vestibular system, the relationship between head posture and lateral semicircular canal orientation in primates has not been explored on a statistically significant sample, using phylogenetically corrected methods. This study tests the hypothesised relationship between lateral semicircular canal orientation and head posture in primates, and investigates potential links to locomotor categories. This study finds that lateral canal orientation is not significantly correlated to positional repertoires. Significant differences in canal orientation are detected between terrestrial and arboreal species. Neutral head posture distinguishes several locomotor categories, and explains a moderate proportion of the variance in positional behaviour. Brain mass is found to correlate with positional behaviour when correcting for the effects of the phylogeny. The implications of the evolution of head posture in fossil species are discussed.
Reassessment of the Phylogeny of Basal Therapsida Using Micro-Tomography and Bayesian Phylogeny
(University of the Witwatersrand, Johannesburg, 2023) Duhamel, Alienor; Rubidge, Bruce; Benoit, Julien
This PhD research is focussed on the evolution and cranial anatomy of the earliest therapsids. The Therapsida is a paraphyletic group of land tetrapods that lived mostly during the Permian and Triassic Periods. They emerged quite suddenly in the fossil record during the middle Permian, already showing diversification into five distinct groups: Biarmosuchia, Dinocephalia, Anomodontia, Gorgonopsia and Therocephalia. A sixth group, the Cynodontia, appeared during the late Permian. Of these, the anomodonts, therocephalians, and cynodonts survived the devastating Permo-Triassic mass extinction and continued to be the dominant tetrapods during the Early and Middle Triassic Periods. Ultimately, the cynodonts would evolve into the earliest mammals. Chapter 1 introduces the scientific questions tackled in this thesis. Chapter 2 provides the necessary explanations about the material and methods used for this work. The use of CT-scanning technology allows for the detailed examination of cranial internal morphology, with the goal of incorporating inner cranial characters into a phylogenetic framework. Chapters 3 and 6 focus on the fossilisation gap at the early to middle Permian transition, which has led to debates over the timing of the early diversification of the Therapsida. In Chapter 3, the geologic position and anatomy of one of the oldest known therapsids, Raranimus dashankouensis is reviewed. The chapter concludes with a reassessment of the basal phylogenetic position of Raranimus and suggests a Roadian origin for the taxon. Chapter 6 presents a comprehensive analysis of cranial morphological characters based on the descriptions and findings from Chapters 3, 4, and 5. Using both traditional maximum parsimony and clock-based Bayesian inference of phylogeny for the first time on the Therapsida, the results suggest that therapsids originated around 280.5 million years ago and rapidly diversified into several distinct clades during the Kungurian and Roadian Epochs. The phylogenetic relationships of several taxa are reevaluated and the findings suggest that Biseridens is more closely related to the Dinocephalia and Biarmosuchia, rather than the Anomodontia. Furthermore, the results suggest that Sinophoneus may represent the basal-most dinocephalian, and that Therocephalia might be paraphyletic. Therapsids were characterised by a wide range of cranial morphologies, with the Biarmosuchia and Anomod ontia exhibiting distinct cranial features. Both groups are considered basal among therapsids. Chapter 4 of this PhD research focuses on the postnatal ontogenetic development of cranial ornamentation in Biarmosuchia and possible impact on phylogeny. The results suggests that cranial bosses and ridges are ontogenetic features and the parietal bone originates from multiple centres of ossification. Chapter 5 is a comprehensive cranial description of several basal anomodont specimens. The Chapter concludes with a review of the taxonomy of the genus Eodicynodon and proposes the creation of a new taxonomical unit for specimen NMQR 2913.
The Taxonomy and Phylogeny of Varanopidae from the Middle Permian of outh Africa
(University of the Witwatersrand, Johannesburg, 2024-08) Sibiya, Zoleka; Rubidge, Bruce; Benoit, Julien
Varanopidae are the only pelycosaur-grade tetrapods from the Abrahamskraal Formation of the Beaufort Group. Although four varanopid species are described from the middle Permian of South Africa (SA), their validity has been questionable. Moreover, Ford and Benson recently proposed that varanopids belong to the Diapsida rather than Synapsida. Given this taxonomic and phylogenetic turmoil, this study i) describes two newly discovered and well-preserved varanopid specimens from the middle Permian of SA using CT scanning and ii) re-assesses the validity of the four species from the Karoo Basin in the light of new data from these specimens. The phylogeny of the group is addressed by updating two pre-existing cladistics matrices. The two new specimens exhibit a mosaic of features that overlap the diagnoses of several species of SA varanopids. In addition, some diagnostic traits of the four SA varanopid genera and species are found to be variable across ontogeny. For instance, dorsal osteoderms and the extent of ornamentation on the angular bone co-vary with size. It is therefore proposed that all varanopid specimens currently described from the middle Permian of South Africa belong to a single valid species, Heleosaurus scholtzi, with differences between specimens being accounted for by ontogenetic changes. The phylogenetic analysis supports Ford and Benson’s hypothesis but contra the prediction of this hypothesis, no differences in cranial fenestration during ontogeny are observed. Morphological characters preserved in the manus of the newly discovered specimen BP/1/8499 suggest that Heleosaurus was arboreal.