Early synapsids neurosensory diversity revealed by CT and synchrotron scanning.

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 wide range of unexpected NMS paleoneurological diversity. First, we describe adaptations to low-frequency hearing and low-light conditions in the non-mammalian cynodont Cistecynodon parvus, supporting adaptations to an obligatory fossorial lifestyle. Second, we describe the uniquely complex and three-dimensional maxillary canal morphology of the biarmosuchian Pachydectes elsi, which suggests that it may have used its cranial bosses for display or low-energy combat. Finally, we introduce a paleopathology found in the skull of Moschognathus whaitsi. Since the specimen was not fully grown, this condition suggests the possibility that this species might have engaged in playful fighting as juveniles-a behavior that is both social and structured. Additionally, this paper discusses other evidence that could indicate that tapinocephalid dinocephalians were social animals, living and interacting closely with one another. Altogether, these examples evidence the wide range of diversity of neurological structures and complex behavior in NMS.

The postcranial anatomy of Gorgonops torvus (Synapsida, Gorgonopsia) from the late Permian of South Africa.

Gorgonopsians are among the most recognizable groups of synapsids from the Permian period and have an extensive but mostly cranial fossil record. By contrast, relatively little is known about their postcranial anatomy. Here, we describe a nearly complete, semi-articulated skeleton of a gorgonopsian (identified as Gorgonops torvus) from the late Permian Endothiodon Assemblage Zone of the South African Karoo Basin and discuss its paleobiological implications. Known gorgonopsian postcrania indicate morphological conservatism in the group, but the skeletal anatomy of Gorgonops does differ from that of other gorgonopsians in some respects, such as in the triangular radiale and short terminal phalanges in the manus, and a weakly developed distinction between pubis and ischium in ventral aspect of the pelvic girdle. Similarities between the specimen described herein and a historically problematic specimen originally referred to "Scymnognathus cf. whaitsi" confirm referral of the latter specimen to Gorgonops. Since descriptions of gorgonopsian postcrania are rare, new interpretations of the lifestyle and ecology of Gorgonopsia can be drawn from our contribution. We conclude that gorgonopsians were likely ambush predators, able to chase their prey over short distances and pin them down with strong forelimbs before using their canines for the kill. This is evidenced by their different fore- and hindlimb morphology; the former stouter and more robust in comparison to the longer, more gracile, back legs. Furthermore, the completeness of the study specimen facilitates calculation of an estimated body mass of approximately 98 kg, similar to that of a modern lioness.

Revisiting the evolutionary trend toward the mammalian lower jaw in non-mammalian synapsids in a phylogenetic context.

The mammalian lower jaw comprises a single bone, the dentary, which is a unique feature among vertebrates. The lower jaws of extinct non-mammalian synapsids were composed of the dentary and several postdentary bones. Synapsid fossils exhibit variation in the dentary size relative to the overall lower jaw. An evolutionary trend toward dentary enlargement and postdentary reduction in non-mammalian synapsids has long been documented but has not been established using modern phylogenetic comparative methods. In this study, we examine the evolutionary pattern of dentary size relative to the lower jaw through phylogenetic analyses of measurements in a broad range of non-mammalian synapsid taxa. Our analyses revealed an evolutionary trend toward dentary area enlargement relative to the overall lower jaw in the lateral view across all non-mammalian synapsids. This trend is likely due to vertical expansion of the dentary given that the same trend is not evident when looking at anterior to posterior measurements of the dentary relative to the lower jaw as a whole in lateral view. Ancestral character reconstructions revealed that the evolution of the measurements was not unidirectional in non-mammalian synapsids. Our results provide no evidence of an evolutionary trend toward the dentary enlargement at the expense of postdentary bones across non-mammalian synapsids. This implies that the evolutionary origin of the mammalian lower jaw is not adequately explained by the evolutionary trend of dentary enlargement throughout non-mammalian synapsids. Instead, selection that occurred during the transition from non-mammalian cynodonts to early mammals may have produced the mammalian lower jaw.

At the root of the mammalian mind: The sensory organs, brain and behavior of pre-mammalian synapsids.

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 were unrefined compared to those of modern mammals. Recent works have, however, revealed that neurological, sensory, and behavioral traits previously considered typically mammalian, such as whiskers, enhanced olfaction, nocturnality, parental care, and complex social interactions evolved before the origin of Mammaliaformes, among the early-diverging "mammal-like reptiles." In contrast, an enlarged brain did not evolve immediately after the origin of mammaliaforms. As such, in terms of paleoneurology, the last "mammal-like reptiles" were not significantly different from the earliest mammaliaforms. The abundant data and literature published in the last 10 years no longer supports the "three pulses" scenario of synapsid brain evolution proposed by Rowe and colleagues in 2011, but supports the new "outside-in" model of Rodrigues and colleagues proposed in 2018, instead. As Mesozoic reptiles were becoming the dominant taxa within terrestrial ecosystems, synapsids gradually adapted to smaller body sizes and nocturnality. This resulted in a sensory revolution in synapsids as olfaction, audition, and somatosensation compensated for the loss of visual cues. This altered sensory input is aligned with changes in the brain, the most significant of which was an increase in relative brain size.

Computed Tomography elucidates ontogeny within the basal therapsid clade Biarmosuchia.

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 other therapsids with pachyostosed crania, which complicates burnetiamorph taxonomic distinction and thus it is essential to better understand cranial ontogeny of the Burnetiamorpha. Here, three new juvenile biarmosuchian skulls from the late Permian of South Africa are described using X-ray micro computed tomography (CT). We found that juvenile biarmosuchians are distinguished from adults by their relatively large orbits, open cranial sutures, and incomplete ossification of the braincase and bony labyrinth. Also, they manifest multiple centres of ossification within the parietal and preparietal bones. CT examination reveals that the holotype of Lemurosaurus pricei (BP/1/816), previously alleged to be a juvenile, shows no evidence of juvenility and is thus probably an adult. This suggests that the larger skull NMQR 1702, previously considered to be an adult L. pricei, may represent a new taxon. This study provides, for the first time, a list of characters by which to recognise juvenile biarmosuchians.

Cranial anatomy of Bolotridon frerensis, an enigmatic cynodont from the Middle Triassic of South Africa, and its phylogenetic significance.

The cynodont fauna of the Trirachodon-Kannemeyeria Subzone of the Middle Triassic Cynognathus Assemblage Zone (AZ) is almost exclusively represented by taxa belonging to the clade Eucynodontia. However, there is one basal (non-eucynodont) cynodont known to have survived into this assemblage: the enigmatic Bolotridon frerensis. BSPG 1934-VIII-7 represents by far the most extensive specimen of B. frerensis, consisting of a partial skull with occluded lower jaw. The specimen was initially described by Broili & Schröder (1934), but their description was limited to surface details of the skull and the dental morphology. Here, by using a computed tomographic (CT) reconstruction, we redescribe this specimen, providing novel information on its palatal and internal anatomy. New endocranial characters recognized for this taxon include ridges in the nasal cavity indicating the presence of cartilaginous respiratory turbinals. New data obtained from the CT scan were incorporated into the most recently published data matrix of early non-mammalian cynodonts to test the previously unstable phylogenetic position of Bolotridon. Our phylogenetic analyses recovered Bolotridon as the sister-taxon of Eucynodontia, a more crownward position than previously hypothesized.

The paleobiology and paleoecology of South African Lystrosaurus.

Lystrosaurus was one of the few tetrapods to survive the end-Permian mass extinction (EPME), the most catastrophic biotic crisis in Phanerozoic history. The significant increased abundance of this genus during the post-extinction Early Triassic recovery period has made Lystrosaurus an iconic survivor taxon globally and ideal for studying changes in growth dynamics during a mass extinction. There is potential evidence of a Lilliput effect in Lystrosaurus in South Africa as the two Triassic species that became highly abundant after the EPME are relatively smaller than the two Permian species. In order to test this hypothesis a detailed examination of the body size and life history of Permo-Triassic Lystrosaurus is required. In this study, the basal skull length and growth patterns of the four South African Lystrosaurus species from the Karoo Basin, L. maccaigi, L. curvatus, L. murrayi and L. declivis, were examined using cranial measurements and bone histology. The basal skull length measurements show that the Triassic species are smaller than the Permian species and supports previous studies. The osteohistology examination of all four species reveal rapidly forming fibrolamellar bone tissues during early to mid-ontogeny. Growth marks are common in L. maccaigi and L. curvatus, but rare and inconsistent in the purely Triassic L. murrayi and L. declivis. The inconsistency of the growth marks in these latter two taxa suggests the presence of developmental plasticity. This feature may have been advantageous in allowing these species to alter their growth patterns in response to environmental cues in the post-extinction Early Triassic climate. An overall transition to slower forming parallel-fibered bone is observed in the largest individuals of L. maccaigi, but absent from the limb bones of the other species. The absence of such bone tissue or outer circumferential lamellae in L. curvatus, L. murrayi and L. declivis indicates that even the largest collected specimens do not represent fully grown individuals. Although L. murrayi and L. declivis are smaller in size, the lack of a growth asymptote in the largest specimens indicates that adult individuals would have been notably larger and may have been similar in size to large L. maccaigi and L. curvatus when fully grown. Thus, the previously described Lilliput effect, recognized by some authors in the Karoo fossil record (such as the therocephalian Moschorhinus kitchingi), may be a product of high juvenile excess mortality in the Triassic rather than a strict "dwarfing" of Lystrosaurus species. The lifestyle of Lystrosaurus was also re-examined. Although previous studies have proposed an aquatic lifestyle for the genus, the similar morphology and bone microanatomy to several other large terrestrial Permo-Triassic dicynodonts supports a fully terrestrial mode of life.

The original boneheads: histologic analysis of the pachyostotic skull roof in Permian burnetiamorphs (Therapsida: Biarmosuchia).

Thickened, pachyostotic skulls are best known in pachycephalosaur dinosaurs, but evolved convergently in Permian burnetiamorphs as well as in some other stem-mammal groups and Triassic archosauromorphs. Until now, only pachycephalosaur domes have been histologically sampled to reveal patterns of bone tissue microstructure and growth. Using computed tomography and osteohistology, we serially thin-sectioned one of the smallest burnetiamorph skull caps ever recovered (estimated skull length = 10 cm), as well as an individual nearly twice as large, and here report the first cranial histological data from this clade. We recognize four highly vascularized histological zones visible in coronal thin-sections, only one of which shares morphological similarities with the tripartite zonation previously reported in pachycephalosaur domes. Zone A forms the endocranial region of the skull cap and records disorganized primary osteons in a fibrolamellar complex. Zone B preserves a border of compact, avascular layers of parallel-fibered bone surrounding an interior of partially remodeled vascular canals. Interestingly, the outline of Zone B resembles the shape of an incipient skull roof. Zone C forms the thickest portion of the skull cap and is composed of fast-growing woven bone with minimal osteonal development. The superficial Zone D has a matrix of predominantly woven bone with narrower primary vascular canals than in deeper regions of the skull caps. Unlike in pachycephalosaurs, where primary vascular porosity is thought to decrease through ontogeny, both burnetiamorph skull caps preserve a thick Zone C of highly vascularized tissue. Additionally, the remnants of sutures are visible as radial struts that taper superficially, leaving no trace on the surface of the skull. Even in the smallest individual, the sutures are closed ectocranially, which is unusual, given that some large, presumably adult pachycephalosaur domes preserve open sutural gaps. Although pachycephalosaur and burnetiamorph skull domes are superficially similar, histological analysis reveals differences in their vascularity and construction that imply multiple evolutionary pathways to form an elaborate pachyostotic dome.

Antiquity of forelimb ecomorphological diversity in the mammalian stem lineage (Synapsida).

Mammals and their closest fossil relatives are unique among tetrapods in expressing a high degree of pectoral girdle and forelimb functional diversity associated with fully pelagic, cursorial, subterranean, volant, and other lifestyles. However, the earliest members of the mammalian stem lineage, the "pelycosaur"-grade synapsids, present a far more limited range of morphologies and inferred functions. The more crownward nonmammaliaform therapsids display novel forelimb morphologies that have been linked to expanded functional diversity, suggesting that the roots of this quintessentially mammalian phenotype can be traced to the pelycosaur-therapsid transition in the Permian period. We quantified morphological disparity of the humerus in pelycosaur-grade synapsids and therapsids using geometric morphometrics. We found that disparity begins to increase concurrently with the emergence of Therapsida, and that it continues to rise until the Permo-Triassic mass extinction. Further, therapsid exploration of new regions of morphospace is correlated with the evolution of novel ecomorphologies, some of which are characterized by changes to overall limb morphology. This evolutionary pattern confirms that nonmammaliaform therapsid forelimbs underwent ecomorphological diversification throughout the Permian, with functional elaboration initially being more strongly expressed in the proximal end of the humerus than the distal end. The role of the forelimbs in the functional diversification of therapsids foreshadows the deployment of forelimb morphofunctional diversity in the evolutionary radiation of mammals.

A new therocephalian (Gorynychus masyutinae gen. et sp. nov.) from the Permian Kotelnich locality, Kirov Region, Russia.

A new therocephalian taxon (Gorynychus masyutinae gen. et sp. nov.) is described based on a nearly complete skull and partial postcranium from the Permian Kotelnich locality of Russia. Gorynychus displays an unusual mixture of primitive ("pristerosaurian") and derived (eutherocephalian) characters. Primitive features of Gorynychus include extensive dentition on the palatal boss and transverse process of the pterygoid, paired vomers, and a prominent dentary angle; derived features include the absence of the postfrontal. Gorynychus can be distinguished from all other therocephalians by its autapomorphic dental morphology, with roughly denticulated incisors and postcanines. Phylogenetic analysis recovers Gorynychus as a non-lycosuchid, non-scylacosaurid therocephalian situated as sister-taxon to Eutherocephalia. The identification of Gorynychus as the largest predator from Kotelnich indicates that therocephalians acted as apex predators in middle-late Permian transition ecosystems in Russia, corroborating a pattern observed in South African faunas. However, other aspects of the Kotelnich fauna, and Permian Russian tetrapod faunas in general, differ markedly from those of South Africa and suggest that Karoo faunas are not necessarily representative of global patterns.

Gorgonopsian therapsids (Nochnitsa gen. nov. and Viatkogorgon) from the Permian Kotelnich locality of Russia.

The early evolution of gorgonopsians is poorly understood. New material from the Kotelnich locality in Russia expands our knowledge of middle/earliest late Permian gorgonopsians from Laurasia. Two gorgonopsian taxa are recognized from Kotelnich: Viatkogorgon ivakhnenkoi Tatarinov, 1999 and Nochnitsa geminidens gen. et sp. nov. Nochnitsa can be distinguished from all known gorgonopsians by its unique upper postcanine tooth row, composed of pairs of teeth (a small anterior and larger posterior) separated by diastemata. Both Viatkogorgon and Nochnitsa are relatively small gorgonopsians, comparable in size to the South African middle Permian taxon Eriphostoma. Inclusion of Viatkogorgon and Nochnitsa in a phylogenetic analysis of gorgonopsians recovers them in basal positions, with Nochnitsa representing the earliest-diverging gorgonopsian genus. All other sampled gorgonopsians fall into two major subclades: one made up entirely of Russian taxa (Inostrancevia, Pravoslavlevia, Sauroctonus, and Suchogorgon) and the other containing only African gorgonopsians. The high degree of endemism indicated in this analysis for gorgonopsians is remarkable, especially given the extensive intercontinental dispersal inferred for coeval therapsid groups.

Endocranial Casts of Pre-Mammalian Therapsids Reveal an Unexpected Neurological Diversity at the Deep Evolutionary Root of Mammals.

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 non-mammaliaform Therapsida (NMT). This is because the majority of the NMT have a non-ossified braincase, making the study of their endocranial cast (sometimes called the "fossil brain") problematic. Thus, descriptions of the morphology and size of NMT endocranial casts have been based largely on approximations rather than reliable determination. Accordingly, here we use micro-CT scans of the skulls of 1 Dinocephalia and 3 Biarmosuchia, which are NMT with a fully ossified braincase and thus a complete endocast. For the first time, our work enables the accurate determination of endocranial shape and size in NMT. This study suggests that NMT brain size falls in the upper range of the reptilian and amphibian variation. Brain size in the dicynodont Kawingasaurus is equivalent to that of early Mammaliaformes, whereas the Dinocephalia show evidence of a secondary reduction of brain size. In addition, unlike other NMT in which the endocast has a tubular shape and its parts are arranged in a linear manner, the biarmosuchian endocast is strongly flexed at the level of the midbrain, creating a near right angle between the fore- and hindbrain. These data highlight an unexpected diversity of endocranial size and morphology in NMT, features that are usually considered conservative in this group.

What did the "Unossified zone" of the non-mammalian therapsid braincase house?

Most nonmammalian synapsids possess a mid-dorsal depression in the brain cavity known as the "unossified zone." It remains obscure which structures this zone contained, and, as candidates, the vermis of the cerebellum, the superior sagittal sinus, a junction of several blood vessels, the pineal gland or other midbrain structures were considered. Neutron tomography of a skull of Diictodon feliceps (Therapsida, Anomodontia) revealed some clear impressions of canals in this region of the brain cavity. Furthermore, the prootic sinus probably ran on the internal surface of the pila antotica and had a similar course in anomodonts as it has been proposed for cynodonts and Mesozoic mammals. Comparisons with the vascular systems of nonmammalian synapsids and mammals suggest that the unossified zone is best interpreted as a terminal chamber of the anterior segment of the medial head vein, which housed the junction of the superior sagittal sinus and the transverse sinuses. Consequently, the system of cranial vessels in Diictodon reveals a partial division of the medial head vein system into an anterior and a posterior segment at an early stage of synapsid evolution, which is consistent with the well-known common pattern of early ontogenetic development in amniotes. J. Morphol., 2017. © 2017 Wiley Periodicals, Inc.

Evidence for convergent evolution of a neocortex-like structure in a late Permian therapsid.

The special sensory, motor, and cognitive capabilities of mammals mainly depend upon the neocortex, which is the six-layered cover of the mammalian forebrain. The origin of the neocortex is still controversial and the current view is that larger brains with neocortex first evolved in late Triassic Mammaliaformes. Here, we report the earliest evidence of a structure analogous to the mammalian neocortex in a forerunner of mammals, the fossorial anomodont Kawingasaurus fossilis from the late Permian of Tanzania. The endocranial cavity of Kawingasaurus is almost completely ossified, which allowed a less hypothetical virtual reconstruction of the brain endocast to be generated. A parietal foramen is absent. A small pit between the cerebral hemispheres is interpreted as a pineal body. The inflated cerebral hemispheres are demarcated from each other by a median sulcus and by a possible rhinal fissure from the rest of the endocast. The encephalization quotient estimated by using the method of Eisenberg is 0.52, which is 2-3 times larger than in other nonmammalian synapsids. Another remarkable feature are the extremely ramified infraorbital canals in the snout. The shape of the brain endocast, the extremely ramified maxillary canals as well as the small frontally placed eyes suggest that special sensory adaptations to the subterranean habitat such as a well developed sense of touch and binocular vision may have driven the parallel evolution of an equivalent of the mammalian neocortex and a mammal-like lemnothalamic visual system in Kawingasaurus. The gross anatomy of the brain endocast of Kawingasaurus supports the Outgroup Hypothesis, according to which the neocortex evolved from the dorsal pallium of an amphibian-like ancestor, which receives sensory projections from the lemnothalamic pathway. The enlarged brain as well as the absence of a parietal foramen may be an indication for a higher metabolic rate of Kawingasaurus compared to other nonmammalian synapsids.

An early geikiid dicynodont from the Tropidostoma Assemblage Zone (late Permian) of South Africa.

Based on specimens previously identified as Tropidostoma, a new taxon of dicynodont (Bulbasaurus phylloxyron gen. et sp. nov.) from the Karoo Basin of South Africa is described. Bulbasaurus is a medium-sized dicynodont (maximum dorsal skull length 16.0 cm) restricted to the Tropidostoma Assemblage Zone (early Lopingian) of the Beaufort Group. Bulbasaurus can be distinguished from Tropidostoma by an array of characters including the presence of a tall, sharp premaxillary ridge, large, rugose, nearly-confluent nasal bosses, a nasofrontal ridge, massive tusks, robust pterygoids, prominently twisted subtemporal bar, and absence of a distinct postfrontal. Inclusion of Bulbasaurus in a phylogenetic analysis of anomodont therapsids recovers it as a member of Geikiidae, a clade of otherwise later Permian dicynodonts such as Aulacephalodon and Pelanomodon. Bulbasaurus exhibits many of the characters typical of adult Aulacephalodon, but at substantially smaller skull size (these characters are absent in comparably-sized Aulacephalodon juveniles), suggesting that the evolution of typical geikiid morphology preceded gigantism in the clade. Bulbasaurus is the earliest known geikiid and the only member of the group known from the Tropidostoma Assemblage Zone; discovery of this taxon shortens a perplexing ghost lineage and indicates that abundant clades from the later Permian of South Africa (e.g., Geikiidae, Dicynodontoidea) may have originated as rare components of earlier Karoo assemblage zones.

Cranial Ontogeny of the Early Triassic Basal Cynodont Galesaurus planiceps.

Ontogenetic changes in the skull and mandible of thirty-one specimens of Galesaurus planiceps, a basal non-mammaliaform cynodont from the Early Triassic of South Africa, are documented. The qualitative survey indicated eight changes in the craniomandibular apparatus occurred during growth, dividing the sample into three ontogenetic stages: juvenile, subadult, and adult. Changes in the temporal region, zygomatic arch, occiput, and mandible occurred during the transition from the subadult to adult stage at a basal skull length of 90 mm. At least four morphological and allometric differences divided the adult specimens into two morphs, indicating the presence of sexual dimorphism in Galesaurus. Differences include extensive lateral flaring of the zygomatic arches in the "male" morph resulting in a more anterior orientation of the orbits, and a narrower snout in the "female". This is the first record of sexual dimorphism in a basal cynodont, and the first time it is quantitatively documented in a non-mammaliaform cynodont. An ontogenetic comparison between Galesaurus and the more derived basal cynodont Thrinaxodon revealed differences in the timing and extent of sagittal crest development. In Galesaurus, the posterior sagittal crest, located behind the parietal foramen, developed relatively later in ontogeny, and the anterior sagittal crest rarely formed suggesting the anterior fibres of the temporalis were less developed than in Thrinaxodon. In contrast, craniomandibular features related to the masseters became more developed during the ontogeny of Galesaurus. The development of the adductor musculature appears to be one of the main factors influencing skull growth in these basal non-mammaliaform cynodonts. Anat Rec, 300:353-381, 2017. © 2016 Wiley Periodicals, Inc.

Systematics of the Rubidgeinae (Therapsida: Gorgonopsia).

The subfamily Rubidgeinae, containing the largest known African gorgonopsians, is thoroughly revised. Rubidgeinae is diagnosed by the absence of a blade-like parasphenoid rostrum and reduction or absence of the preparietal. Seven rubidgeine species from the Karoo Basin of South Africa are recognized as valid: Aelurognathus tigriceps, Clelandina rubidgei, Dinogorgon rubidgei, Leontosaurus vanderhorsti, Rubidgea atrox, Smilesaurus ferox, and Sycosaurus laticeps. Rubidgeines are also present in other African basins: A. tigriceps and S. laticeps occur in the Upper Madumabisa Mudstone Formation of Zambia, and D. rubidgei, R. atrox, and the endemic species Ruhuhucerberus haughtoni comb. nov. and Sycosaurus nowaki comb. nov. occur in the Usili Formation of Tanzania. Aelurognathus nyasaensis from the Chiweta Beds of Malawi also represents a rubidgeine, but of uncertain generic referral pending further preparation. No rubidgeine material is known outside of Africa: the purported Russian rubidgeine Leogorgon klimovensis is not clearly referable to this group and may not be diagnosable. Phylogenetic analysis of rubidgeines reveals strong support for a clade (Rubidgeini) of advanced rubidgeines including Clelandina, Dinogorgon, Leontosaurus, and Rubidgea. Support for Smilesaurus as a rubidgeine is weak; it may, as previous authors have suggested, represent an independent evolution of large body size from an Arctops-like ancestor. Temporally, rubidgeines are restricted to the Late Permian, first appearing in the Tropidostoma Assemblage Zone and reaching highest diversity in the Cistecephalus and Daptocephalus assemblage zones of the Beaufort Group.

Tiarajudens eccentricus and Anomocephalus africanus, two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana.

Anomodontia was a highly successful tetrapod clade during the Permian and the Triassic. New morphological information regarding two bizarre basal anomodonts is provided and their palaeoecological significance is explored. The osteology of the recently discovered Tiarajudens eccentricus Cisneros et al. 2011, from the Brazilian Permian, is described in detail. The taxon exhibits unusual postcranial features, including the presence of gastralia. Additional preparation and computed tomography scans of the holotype of Anomocephalus africanus Modesto et al. 1999 discovered in the Karoo Basin of South Africa allow a reappraisal of this genus. Anomocephalus is similar to Tiarajudens with regard to several traits, including a battery of large, transversally expanded, palatal teeth. Molariform teeth are present in the mandible of the African taxon, providing additional insight into the function of the earliest tooth-occlusion mechanism known in therapsids. At least two waves of tooth replacement can be recognized in the palate of Anomocephalus. The outsized, blade-like caniniforms of the herbivorous Tiarajudens allow several non-exclusive ecological interpretations, among which we favour intraspecific display or combat. This behaviour was an alternative to the head-butting practised by the contemporary dinocephalians. Combat specializations that are considered typical of Cenozoic herbivores likely evolved during the Middle Permian, at the time the first communities with diverse, abundant tetrapod herbivores were being assembled.

Physiological implications of the abnormal absence of the parietal foramen in a late Permian cynodont (Therapsida).

The third eye (pineal eye), an organ responsible for regulating exposure to sunlight in extant ectotherms, is located in an opening on the dorsal surface of the skull, the parietal foramen. The parietal foramen is absent in extant mammals but often observed in basal therapsids, the stem-group to true mammals. Here, we report the absence of the parietal foramen in a specimen of Cynosaurus suppostus, a Late Permian cynodont from South Africa (SA). Comparison with Procynosuchus delaharpeae, a contemporaneous non-mammalian cynodont from SA, demonstrates that the absence of this foramen is an abnormal condition for such a basal species. Because seasonality was marked during the Late Permian in SA, it is proposed that the third eye was functionally redundant in Cynosaurus, possibly due to the acquisition of better thermoregulation or the evolution of specialized cells in the lateral eyes to compensate for the role of the third eye.

Virtual reconstruction and description of the cranial endocast of Pristerodon mackayi (Therapsida, Anomodontia).

The cranial endocast of Pristerodon mackayi is described, which has been virtually reconstructed on the basis of neutron tomographic data. The gross anatomy of the cranial endocast of Pristerodon resembles other nonmammalian synapsids such as Thrinaxodon liorhinus in having a narrow, tubular forebrain, well developed olfactory bulbs, a large parietal foramen and unossified zone. As it is the case in cynodonts the hindbrain of Pristerodon is broader than the mid- and forebrain. Large paraflocculi are developed. The medulla oblongata can be well distinguished from the pons. The pons is divided by a median ridge into two portions. There is no evidence for a neocortex, which seems to be also reflected in the low encephalization quotient of 0.18 estimated according to the method of Eisenberg.