Julien Benoit

University of the Witwatersrand | wits · Evolutionary Studies Institute

The Horned Serpent panel at La Belle France (Free State Province, South Africa) was painted by the San at least two hundred years ago. It pictures, among many other elements, a tusked animal with a head that resembles that of a dicynodont, the fossils of which are abundant and conspicuous in the Karoo Basin. This picture also seemingly relates to a...

Mammalian origin is often traced back to the Therapsida, a clade of diverse synapsids that emerged during the early-middle Permian from “pelycosaur” grade ancestors. The six major therapsid groups (Biarmosuchia, Dinocephalia, Anomodontia, Gorgonopsia, Therocephalia, and Cynodontia) evolved rapidly between the early and middle Permian. This fast rad...

Therocephalians represent one of the major lineages of Permian-Triassic non-mammalian therapsids, surviving the end-Permian mass extinction. Among Therapsida, they are most closely related to Cynodontia, the clade including mammals, which is derived from within Therocephalia, rendering the latter paraphyletic. Despite their importance, abundance, b...

Non‐mammaliaform synapsids (NMS) represent the closest relatives of today's mammals among the early amniotes. Exploring their brain and nervous system is key to understanding how mammals evolved. Here, using CT and Synchrotron scanning, we document for the first time three extreme cases of neurosensory and behavioral adaptations that probe into the...

The Dicynodontia (Therapsida: Anomodontia) is one of the most successful Permo-Triassic terrestrial tetrapod clades and the oldest specimens are recorded from the middle Permian Eodicynodon Assemblage Zone of South Africa. Their fossil record is abundant and species-rich across Pangea. By contrast, the fossil record of the basal-most anomodonts, wh...

Diarthrognathus broomi is a transitional taxon between non-mammaliaform cynodonts and Mammaliaformes that occurred during the Late Triassic to Early Jurassic. All known specimens of Diarthrognathus represent juveniles, and the postcrania have not been thoroughly described. The palatal, basicranial and postcranial elements of the referred specimen N...

Therocephalians were an ecomorphologically varied group of therapsids with global distribution during the late Permian and earliest Triassic periods. Here, we redescribe the holotype of the therocephalian Olivierosuchus parringtoni (BP/1/3849) from the Early Triassic Lystrosaurus declivis Assemblage Zone in the main Karoo Basin of South Africa. The...

Compared to other continents, the study of indigenous (non-western) knowledge of palaeontology in Africa is a relatively new field. The literature reviewed here nevertheless suggests a long-lasting string of traditions, geomyths and folklore related to fossilised items on the whole continent and encompassing many African cultures. It is often diffi...

The maxillary canal of the titanosuchid dinocephalian Jonkeria is described based on digitised serial sections. We highlight that its morphology is more like that of the tapinocephalid Moschognathus than that of Anteosaurus. This is unexpected given the similarities between the dentition of Jonkeria and Anteosaurus (i.e., presence of a canine) and...

Mammals are diagnosed by more than 30 osteological characters (e.g. squamosal- dentary jaw joint, three inner ear ossicles, etc.) that are readily preserved in the fossil record. However, it is the suite of physiological, soft tissue and behavioural characters (e.g. endothermy, hair, lactation, isocortex and parental care), the evolutionary origins...

All modern mammals are descendants of the paraphyletic non-mammaliaform Synapsida, colloquially referred to as the “mammal-like reptiles.” It has long been assumed that these mammalian ancestors were essentially reptile-like in their morphology, biology, and behavior, i.e., they had a small brain, displayed simple behavior, and their sensory organs...

The elephant brain is famous for its higher than average encephalization quotient, memory capacities, large cerebellum, large facial and trigeminal nerves, and the extensive repertoire of complex behaviors and social interactions it produces, the last of which being supported by infrasonic communication. The evolutionary history of Proboscidea is a...

Lumkuia fuzzi is a small non-mammalian cynodont from the Middle Triassic of South Africa. It has traditionally been phylogenetically identified as a basal Probainognathia, but some studies place it more basally as the sister-group to Eucynodontia. Lumkuia is known from a single specimen comprising a skull and partial skeleton. Only the skull has be...

Endothermy underpins the ecological dominance of mammals and birds in diverse environmental settings1,2. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most of the fossil evidence is ambiguous3–17. Here we show that this key evolutionary transition can be investigated using the morphology of the e...

Emydopoidea is one of the major dicynodont subclades and includes some purported fossorial taxa. Various cranial and postcranial adaptations for fossoriality have long been recognized in cistecephalid emydopoids, but anatomical variation of their braincases remains poorly understood. Here, using laboratory and synchrotron X-ray tomography, we provi...

Pareiasaurs were relatively abundant, globally distributed, herbivorous parareptiles of the mid to late Permian. The basal-most forms, all members of the Bradysauria, are restricted to the Guadalupian (mid-Permian) of South Africa and went extinct in the late Capitanian near the top of the Tapinocephalus Assemblage Zone. Currently four species are...

Biarmosuchia is a clade of basal therapsids that includes forms possessing plesiomorphic ‘pelycosaurian’ cranial characters as well as the highly derived Burnetiamorpha which are characterised by cranial pachyostosis and a variety of cranial bosses. Potential ontogenetic variation in these structures has been suggested based on growth series of oth...

Several amniote lineages independently evolved multiple rows of marginal teeth in response to the challenge of processing high fiber plant matter. Multiple tooth rows develop via alterations to tooth replacement in captorhinid reptiles and ornithischian dinosaurs, but the specific changes that produce this morphology differ, reflecting differences...

A partial skull (BP/1/7976) of a very large cynodont from the Middle Triassic Cynognathus Assemblage Zone (Cricodon-Ufudocyclops subzone) of South Africa is described. The specimen represents a new gomphodont taxon, Impidens hancoxi, gen. et sp. nov., diagnosed by five sectorial teeth constituting just over half of the length of the upper postcanin...

Despite their significance for paleobiological interpretations, bite marks have been rarely reported in non-mammalian therapsids (NMT). Here we describe, for the first time, the occurrence of a tooth embedded in the snout of a gorgonopsian. The tooth is surrounded by a bony callus, which demonstrates that the animal was still alive after the attack...

The non-mammalian therapsids comprise a paraphyletic assemblage of Permian-Jurassic synapsids closely related to mammals that includes six major clades of largely unresolved phylogenetic affinity. Understanding the early evolutionary radiation of therapsids is complicated by a gap in the fossil record during the Roadian (middle Permian) known as Ol...

Lanthanostegus is an unusual dicynodont known from only two partial skulls from a single locality near Jansenville in the Eastern Cape Province of South Africa. Although these specimens can be constrained to near the base of the late middle Permian (Guadalupian) Abrahamskraal Formation, their precise age is uncertain as a result of diachroneity of...

Endothermy (“warm-bloodedness”) underpins the ecological dominance of mammals and birds in diverse environmental settings1-3. However, it is unclear when this crucial feature emerged during mammalian evolutionary history, as most fossil evidence is ambiguous4-25. Here, we show that new information on this key evolutionary transition can be obtained...

Postcranial morphology of the South African middle Permian pareiasaurs from the Karoo Basin of South Africa.

Pareiasaurs were globally distributed, abundant, herbivorous parareptiles with the basal-most members found only in the mid-Permian of South Africa. These basal forms form a monophyletic group and were locally abundant and became extinct at the top of the Tapinocephalus Assemblage Zone at the end of the Guadalupian. Four species of basal pareiasaur...

Dinocephalians (Therapsida), some of the earliest amniotes to have evolved large body size, include the carnivorous Anteosauria and mostly herbivorous Tapinocephalia. Whilst the palaeoneurology of the Tapinocephalia has been investigated in Moschognathus whaitsi, that of the Anteosauria remains completely unknown. Here we used X-ray micro-Computed...

In the late 1980's the discovery of late Permian helical burrow casts containing articulated skeletons of the small herbivorous therapsid Diictodon feliceps led to conjecture that they may have been used for oviposition/parturition and shelter for infants. Here we present new fossil evidence in support of this interpretation and discuss the possibi...

Synchrotron-based X-ray analytical techniques are gaining in prominence as a crucial yet highly specialized tool for advancing knowledge within the broad field of the Earth Sciences. In terms of global scientific competitiveness, African scientists are somewhat disadvantaged by not having access to their own synchrotron on the African continent. Ho...

For over a century, researchers have assumed that the plane of the lateral semicircular canal of the inner ear lies parallel to the horizon when the head is at rest, and used this assumption to reconstruct head posture in extinct species. Although this hypothesis has been repeatedly questioned, it has never been tested on a large sample size and at...

In mammals, the infraorbital canal provides a passage for the infraorbital ramus of the maxillary branch of the trigeminal nerve. The infraorbital nerve ensures tactile sensitivity of the upper teeth and face between the eye and upper lip and, more significantly, the innervation of mystacial vibrissae (whiskers). In contrast, most non-mammalian syn...

Styracocephalus platyrhynchus is an unusual dinocephalian therapsid, known only from a handful of specimens from the Tapinocephalus Assemblage Zone of South Africa. It has had a chequered taxonomic history, largely because it is characterized by cranial pachyostosis and the presence of horn-like structures that project posteriorly from the temporal...

Cistecephalids are among the most distinctive Permian dicynodonts because of their highly derived skulls and postcrania, which indicate a fossorial ecology. Four cistecephalid species have been described from India, South Africa, and Tanzania; a fifth putative species has been reported from the Luangwa Basin of Zambia but never formally described....

As the largest and among the most behaviourally complex extant terrestrial mammals, proboscideans (elephants and their extinct relatives) are iconic representatives of the modern megafauna. the timing of the evolution of large brain size and above average encephalization quotient remains poorly understood due to the paucity of described endocranial...

Compared with other parts of the world, the study of geomythology in southern Africa, and the associated documentation of non-western awareness of palaeontological and geological phenomena, is in an early phase. We focus on examples of rocks and fossils as items of special interest and curiosity, and we search for evidence of an indigenous palaeont...

Bipedalism is a hallmark of being human and the human foot is modified to reflect this unique form of locomotion. Leonardo da Vinci is credited with calling the human foot “a masterpiece of engineering and a work of art.” However, a scientific approach to human origins has revealed that our feet are products of a long, evolutionary history in which...

The Beaufort Group of the South African Karoo Supergroup is internationally celebrated as one of the richest sources of vertebrate fossils chronicling the Permo-Triassic evolutionary radiation of Therapsida. This clade includes modern mammals as well as their non-mammalian ancestors and relatives. Biarmosuchia is the most basal clade of the Theraps...

Biarmosuchia is a very poor therapsid group from Middle Permian deposits. Most of the genera are monospecific and represented by only one cranial specimen. It result that we have no idea of the intraspecific variation among the group (ontogeny/dimorphism). During that presentation, we presented two small biarmosuchians specimens from the South Afri...

Anomodontia was the most successful herbivorous clade of the mammalian stem lineage (non-mammalian synapsids) during the late Permian and Early Triassic. Among anomodonts, Dicynodontia stands apart because of the presence of an osseous beak that shows evidence of the insertion of a cornified sheath, the ramphotheca. In this study, fourteen anomodon...

The origin and evolution of the mammalian brain has long been the focus of scientific enquiry. Conversely, little research has focused on the palaeoneurology of the stem group of Mammaliaformes, the Permian and Triassic nonmammaliaform Therapsida (NMT). This is because the majority of the NMT have a non-ossified braincase, making the study of their...