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A mid-Permian (Guadalupian epoch) extinction event at approximately 260 Ma has been mooted for two decades. This is based primarily on invertebrate biostratigraphy of Guadalupian–Lopingian marine carbonate platforms in southern China, which are temporally constrained by correlation to the associated Emeishan Large Igneous Province (LIP). Despite attempts to identify a similar biodiversity crisis in the terrestrial realm, the low resolution of mid-Permian tetrapod biostratigraphy and a lack of robust geochronological constraints have until now hampered both the correlation and quantification of terrestrial extinctions. Here we present an extensive compilation of tetrapod-stratigraphic data analysed by the constrained optimization (CONOP) algorithm that reveals a significant extinction event among tetrapods within the lower Beaufort Group of the Karoo Basin, South Africa, in the latest Capitanian. Our fossil dataset reveals a 74–80% loss of generic richness between the upper Tapinocephalus Assemblage Zone (AZ) and the mid-Pristerognathus AZ that is temporally constrained by a U–Pb zircon date (CA-TIMS method) of 260.259 ± 0.081 Ma from a tuff near the top of the Tapinocephalus AZ. This strengthens the biochronology of the Permian Beaufort Group and supports the existence of a mid-Permian mass extinction event on land near the end of the Guadalupian. Our results permit a temporal association between the extinction of dinocephalian therapsids and the LIP volcanism at Emeishan, as well as the marine end-Guadalupian extinctions.

Sixteen specimens of the Early Triassic cynodont Galesaurus planiceps (including eight that were scanned using micro-computed tomography) representing different ontogenetic stages were assembled to study the dental replacement in the species. The growth series shows that the incisors and postcanines continue to develop and replace, even in the largest (presumably oldest) specimen. In contrast, replacement of the canines ceased with the attainment of skeletal maturity, at a basal skull length of ~90 mm, suggesting that Galesaurus had a finite number of canine replacement cycles. Additionally, the functional canine root morphology of these larger specimens showed a tendency to be open-rooted, a condition not previously reported in Mesozoic theriodonts. An alternating pattern of tooth replacement was documented in the maxillary and mandibular postcanine series. Both postcanine series increased in tooth number as the skull lengthened, with the mandibular postcanine series containing more teeth than the maxillary series. In the maxilla, the first postcanine is consistently the smallest tooth, showing a proportional reduction in size as skull length increased. The longer retention of a tooth in this first locus is a key difference between Galesaurus and Thrinaxodon , in which the mesial-most postcanines are lost after replacement. This difference has contributed to the lengthening of the postcanine series in Galesaurus , as teeth continued to be added to the distal end of the tooth row through ontogeny. Overall, there are considerable differences between Galesaurus and Thrinaxodon relating to the replacement and development of their teeth.

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 synapsids display a more “reptilian-like” ancestral condition in which a long and ramified maxillary canal completely enclosed the infraorbital nerve along with other branches of the trigeminal nerve. The phylogenetic transition from the ancestral “reptilian-like” to the derived “mammal-like” condition has been hypothesized to occur at the base of the Probainognathia clade. Using μCT and synchrotron scanning, this study aims to document this transition in detail by focusing on a sample of non-mammalian probainognathian cynodonts and early mammaliaforms. We find that the mammalian condition is the result of a gradual shortening of the maxillary canal, which enabled the infraorbital nerve to ramify within the soft tissues of the face. Mobile whiskers became possible only after the mammalian infraorbital nerve had evolved, which suggest that these structures appeared in Probainognathus and more derived cynodonts. Finally a foramen located on the ventral margin of the lacrimal bone, which has been often homologized with the infraorbital foramen of derived Probainognathia and early Mammaliaformes, is most probably homologous to the mammalian zygomaticofacial foramen.

Large-bodied temnospondyl amphibians were the dominant predators in non-marine aquatic ecosystems from the Carboniferous to the Middle Triassic. In the Permian-aged lower Beaufort Group of the main Karoo Basin, South Africa, temnospondyls are represented exclusively by the family Rhinesuchidae and are well represented by body fossils, whereas trace fossils are scarce. Accordingly, most interpretations of the behaviour of this family are based on skeletal morphology and histological data. Here we document the sedimentology and palaeontology of a late Permian palaeosurface situated immediately below the palaeoshoreline of the Ecca Sea (transition from the Ecca Group to the Beaufort Group) near the town of Estcourt in KwaZulu-Natal Province. The surface preserves numerous ichnofossils, including tetrapod footprints and fish swim-trails, but most striking are seven body impressions and associated swim trails that we attribute to a medium-sized (~1.9 m long) rhinesuchid temnospondyl. These provide valuable insight into the behaviour of these animals. The sinuous shape of some of the traces suggest that the tracemaker swam with continuous sub-undulatory propulsion of the tail.

Fossorialism is a beneficial adaptation for brooding, predator avoidance and protection from extreme climate. The abundance of fossilised burrow casts from the Early Triassic of southern Africa is viewed as a behavioural response by many tetrapods to the harsh conditions following the Permo-Triassic mass-extinction event. However, scarcity of vertebrate remains associated with these burrows leaves many ecological questions unanswered. Synchrotron scanning of a lithified burrow cast from the Early Triassic of the Karoo unveiled a unique mixed-species association: an injured temnospondyl amphibian (Broomistega) that sheltered in a burrow occupied by an aestivating therapsid (Thrinaxodon). The discovery of this rare rhinesuchid represents the first occurrence in the fossil record of a temnospondyl in a burrow. The amphibian skeleton shows signs of a crushing trauma with partially healed fractures on several consecutive ribs. The presence of a relatively large intruder in what is interpreted to be a Thrinaxodon burrow implies that the therapsid tolerated the amphibian's presence. Among possible explanations for such unlikely cohabitation, Thrinaxodon aestivation is most plausible, an interpretation supported by the numerous Thrinaxodon specimens fossilised in curled-up postures. Recent advances in synchrotron imaging have enabled visualization of the contents of burrow casts, thus providing a novel tool to elucidate not only anatomy but also ecology and biology of ancient tetrapods.

Out of the shadows Lampreys and hagfish are the only remaining jawless vertebrates and are commonly used as surrogate ancestors for comparative research on living jawed vertebrates. Until recently little was known of the evolutionary history of lampreys as the only known fossils were enigmatic examples from the Carboniferous period, around 300 million years ago. Then earlier this year Nature published a report of a fine specimen from the Cretaceous of China that looked very close to modern forms. This is now joined by a well preserved fossil from the Devonian of South Africa, which at about 360 million years old is the oldest known lamprey. It looks slightly different from modern lampreys, but is the same in essentials and differs from the various now-extinct armoured fishes with which it shared the Devonian world. Lampreys are the most scientifically accessible of the remaining jawless vertebrates, but their evolutionary history is obscure. In contrast to the rich fossil record of armoured jawless fishes, all of which date from the Devonian period and earlier 1 , 2 , 3 , only two Palaeozoic lampreys have been recorded, both from the Carboniferous period 1 . In addition to these, the recent report of an exquisitely preserved Lower Cretaceous example 4 demonstrates that anatomically modern lampreys were present by the late Mesozoic era. Here we report a marine/estuarine fossil lamprey from the Famennian (Late Devonian) of South Africa 5 , 6 , the identity of which is established easily because many of the key specializations of modern forms are already in place. These specializations include the first evidence of a large oral disc, the first direct evidence of circumoral teeth and a well preserved branchial basket. This small agnathan, Priscomyzon riniensis gen. et sp. nov., is not only more conventionally lamprey-like than other Palaeozoic examples 7 , 8 , but is also some 35 million years older. This finding is evidence that agnathans close to modern lampreys had evolved before the end of the Devonian period. In this light, lampreys as a whole appear all the more remarkable: ancient specialists that have persisted as such and survived a subsequent 360 million years.

Choerosaurus dejageri, a non-mammalian eutheriodont therapsid from the South African late Permian (~259 Ma), has conspicuous hemispheric cranial bosses on the maxilla and the mandible. These bosses, the earliest of this nature in a eutheriodont, potentially make C. dejageri a key species for understanding the evolutionary origins of sexually selective behaviours (intraspecific competition, ritualized sexual and intimidation displays) associated with cranial outgrowths at the root of the clade that eventually led to extant mammals. Comparison with the tapinocephalid dinocephalian Moschops capensis, a therapsid in which head butting is strongly supported, shows that the delicate structure of the cranial bosses and the gracile structure of the skull of Choerosaurus would be more suitable for display and low energy combat than vigorous head butting. Thus, despite the fact that Choerosaurus is represented by only one skull (which makes it impossible to address the question of sexual dimorphism), its cranial bosses are better interpreted as structures involved in intraspecific selection, i.e. low-energy fighting or display. Display structures, such as enlarged canines and cranial bosses, are widespread among basal therapsid clades and are also present in the putative basal therapsid Tetraceratops insignis. This suggests that sexual selection may have played a more important role in the distant origin and evolution of mammals earlier than previously thought. Sexual selection may explain the subsequent independent evolution of cranial outgrowths and pachyostosis in different therapsid lineages (Biarmosuchia, Dinocephalia, Gorgonopsia and Dicynodontia).

Euchambersia mirabilis is an iconic species of Permo-Triassic therapsid because of its unusually large external maxillary fossa linked through a sulcus to a ridged canine. This anatomy led to the commonly accepted conclusion that the large fossa accommodated a venom gland. However, this hypothesis remains untested so far. Here, we conducted a μCT scan assisted reappraisal of the envenoming capacity of Euchambersia, with a special focus on the anatomy of the maxillary fossa and canines. This study shows that the fossa, presumably for the venom-producing gland, is directly linked to the maxillary canal, which carries the trigeminal nerve (responsible for the sensitivity of the face). The peculiar anatomy of the maxillary canal suggests important reorganisation in the somatosensory system and that a ganglion could possibly have been present in the maxillary fossa instead of a venom gland. Nevertheless, the venom gland hypothesis is still preferred since we describe, for the first time, the complete crown morphology of the incisiform teeth of Euchambersia, which strongly suggests that the complete dentition was ridged. Therefore Euchambersia manifests evidence of all characteristics of venomous animals: a venom gland (in the maxillary fossa), a mechanism to deliver the venom (the maxillary canal and/or the sulcus located ventrally to the fossa); and an apparatus with which to inflict a wound for venom delivery (the ridged dentition).

Dinocephalian therapsids are renowned for their massive, pachyostotic and ornamented skulls adapted for head-to-head fighting during intraspecific combat. Synchrotron scanning of the tapinocephalid Moschops capensis reveals, for the first time, numerous anatomical adaptations of the central nervous system related to this combative behaviour. Many neural structures (such as the brain, inner ear and ophthalmic branch of the trigeminal nerve) were completely enclosed and protected by bones, which is unusual for non-mammaliaform therapsids. The nearly complete ossification of the braincase enables precise determination of the brain cavity volume and encephalization quotient, which appears greater than expected for such a large and early herbivore. The practice of head butting is often associated with complex social behaviours and gregariousness in extant species, which are known to influence brain size evolution. Additionally, the plane of the lateral (horizontal) semicircular canal of the bony labyrinth is oriented nearly vertically if the skull is held horizontally, which suggests that the natural position of the head was inclined about 60–65°to the horizontal. This is consistent with the fighting position inferred from osteology, as well as ground-level browsing. Finally, the unusually large parietal tube may have been filled with thick conjunctive tissue to protect the delicate pineal eye from injury sustained during head butting.

The only true living endothermic vertebrates are birds and mammals, which produce and regulate their internal temperature quite independently from their surroundings. For mammal ancestors, anatomical clues suggest that endothermy originated during the Permian or Triassic. Here we investigate the origin of mammalian thermoregulation by analysing apatite stable oxygen isotope compositions (δ18Op) of some of their Permo-Triassic therapsid relatives. Comparing of the δ18Op values of therapsid bone and tooth apatites to those of co-existing non-therapsid tetrapods, demonstrates different body temperatures and thermoregulatory strategies. It is proposed that cynodonts and dicynodonts independently acquired constant elevated thermometabolism, respectively within the Eucynodontia and Lystrosauridae + Kannemeyeriiformes clades. We conclude that mammalian endothermy originated in the Epicynodontia during the middle-late Permian. Major global climatic and environmental fluctuations were the most likely selective pressures on the success of such elevated thermometabolism. School textbooks often refer to “cold-blooded” and “warm-blooded” animals, but these terms are misleading. Rather than being cold, animals like reptiles have body temperatures that are mostly determined by their external environment and can actually achieve high body temperatures, for example, by basking in the sun. By contrast, “warm-blooded” mammals produce their own heat and typically maintain a body temperature that is warmer than their environment. As such, so-called warm-blooded animals are more accurately referred to as “endotherms” and cold-blooded animals as “ectotherms”. Endothermic animals share several characteristics, including insulating layers – like fur or feathers – that keep the body warm, and a secondary palate that separates the mouth and nose for continuous breathing, even while eating. Many of these traits are seen in fossils belonging to a group of animals called the therapsids. Also known as the “mammal-like reptiles”, these animals are descended from ectothermic reptiles but are the ancestors of the endothermic mammals. They dominated the land between 270 and 220 million years ago, during periods of time called the Permian and the Triassic. They also survived two major mass extinction events, including the most devastating mass extinction in all of Earth’s history. However, when the ancestors of mammals became truly endothermic remains an open question. Previous studies that have tried to determine this by focusing on the physical characteristics of therapsids have not yet given a consistent date. Rey et al. took a new approach to answer when endothermy first evolved in the mammal-like reptiles, and instead looked at the chemical makeup of minerals in over 100 fossils. Oxygen can exist in different forms called stable isotopes: oxygen-16 and the rarer and heavier oxygen-18. The ratio of these two isotopes in a fossil will depend on, among other things, where the animal lived and, importantly, its body temperature. Therefore, Rey et al. compared oxygen-containing minerals in the bones and teeth of therapsids to those of other animals that lived alongside them to look for signatures that indicated differences in body temperature and how it was regulated. It appears that two different branches of the therapsid’s family tree independently became endothermic. One branch includes the mammals and their direct ancestors, while the second is more distantly related to mammals. Both became endothermic towards the end of the Permian Period, between about 259 and 252 million years ago. Based on these findings, Rey et al. suggest that endothermy allowed these animals to better cope with fluctuating climates, which helped them to be among the few species that survived the mass extinction event at the end of the Permian. Going forward, these new findings can help scientists to understand which physical characteristics were necessary for endothermy to first develop and which helped to optimize it afterwards. Furthermore, they also suggest that endothermic animals are more able to survive fluctuations in climate, which could guide efforts to protect modern-day endangered species that are most at risk from the ongoing effects of climate change.