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The origin of cynodonts, the group ancestral to and including mammals, is one of the major outstanding problems in therapsid evolution. One of the most troubling aspects of the cynodont fossil record is the lengthy Permian ghost lineage between the latest possible divergence from its sister group Therocephalia and the first appearance of definitive cynodonts in the late Permian. The absence of cynodonts and dominance of therocephalians in middle Permian strata has led some workers to argue that cynodonts evolved from within therocephalians, rendering the latter paraphyletic, but more recent analyses support the reciprocal monophyly of Cynodontia and Therocephalia. Furthermore, although a fundamental dichotomy in the derived subclade Eucynodontia is well-supported in cynodont phylogeny, the relationships of more stemward cynodonts from the late Permian and Early Triassic are unresolved. Here, we provide a re-evaluation of the phylogeny of Eutheriodontia (Cynodontia + Therocephalia) and an assessment of character evolution within the group. Using computed tomographic data derived from extensive sampling of the earliest known (late Permian and Early Triassic) cynodonts and selected exemplars of therocephalians and later (Middle Triassic onwards) cynodonts, we describe novel aspects of the endocranial anatomy of these animals. These data were incorporated into a new phylogenetic data set including a comprehensive sample of early cynodonts. Our phylogenetic analyses support some results previously recovered by other authors, but recover therocephalians as paraphyletic with regards to cynodonts, with cynodonts and eutherocephalians forming a clade to the exclusion of the "basal therocephalian" families Lycosuchidae and Scylacosauridae. Though both conservatism and homoplasy mark the endocranial anatomy of early non-mammalian cynodonts, we were able to identify several new endocranial synapomorphies for eutheriodont subclades and recovered generally better-supported topologies than previous analyses using primarily external craniodental characters., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

Evolutionary radiations generate most of Earth's biodiversity, but are there common ecomorphological traits among the progenitors of radiations? In Synapsida (the mammalian total group), 'small-bodied faunivore' has been hypothesized as the ancestral state of most major radiating clades, but this has not been quantitatively assessed across multiple radiations. To examine macroevolutionary patterns in a phylogenetic context, we generated a time-calibrated metaphylogeny ('metatree') comprising 1,888 synapsid species from the Carboniferous through the Eocene (305-34 Ma) based on 269 published character matrices. We used comparative methods to investigate body size and dietary evolution during successive synapsid radiations. Faunivory is the ancestral dietary regime of each major synapsid radiation, but relatively small body size is only established as the common ancestral state of radiations near the origin of Mammaliaformes in the Late Triassic. The faunivorous ancestors of synapsid radiations typically have numerous novel characters compared with their contemporaries, and these derived traits may have helped them to survive faunal turnover events and subsequently radiate., (© 2023. The Author(s), under exclusive licence to Springer Nature Limited.)

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., Competing Interests: The authors declare that they have no competing interests., (© 2023 Bendel et al.)

Catastrophic ecosystem disruption in the late Permian period resulted in the greatest loss of biodiversity in Earth's history, the Permian-Triassic mass extinction (PTME). 1 The dominant terrestrial vertebrates of the Permian (synapsids) suffered major losses at this time, leading to their replacement by reptiles in the Triassic. 2 The dominant late Permian predatory synapsids, gorgonopsians, were completely extirpated by the PTME. The largest African gorgonopsians, the Rubidgeinae, have traditionally been assumed to go extinct at the Permo-Triassic boundary (PTB). 3 , 4 , 5 However, this apparent persistence through the sustained extinction interval characterizing the continental PTME 6 is at odds with ecological theory indicating that top predators have high extinction risk. 7 Here, we report the youngest known large-bodied gorgonopsians, gigantic specimens from the PTB site of Nooitgedacht 68 in South Africa. These specimens are not rubidgeine, and instead are referable to Inostrancevia, a taxon previously thought to be a Russian endemic. 8 Based on comprehensive review of the South African gorgonopsian record, we show that rubidgeines were early victims of ecosystem disruption preceding the PTME and were replaced as top predators by Laurasian immigrant inostranceviines. The reign of this latter group was short-lived, however; by the PTB, gorgonopsians were extinct, and a different group (therocephalians) became the largest synapsid predators, before themselves going extinct. The extinction and replacement of top predators in rapid succession at the clade level underlines the extreme degree of ecosystem instability in the latest Permian and earliest Triassic, a phenomenon that was likely global in extent., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 North Carolina Department of Natural and Cultural Resources (DNCR). Published by Elsevier Inc. All rights reserved.)

The Burgersdorp Formation of South Africa is a richly fossiliferous rock sequence at the top of the Permian-Triassic Beaufort Group and is known for its abundance of Early-Middle Triassic vertebrate remains, particularly cynodonts. Fossils from the Burgersdorp Formation are referred biostratigraphically to the Cynognathus Assemblage Zone (CAZ), which is further divided into three subzones: Langbergia - Garjainia, Trirachodon - Kannemeyeria , and Cricodon-Ufudocyclops . Each subzone is characterised by the presence of a distinct species of trirachodontid, a group of gomphodont cynodonts found relatively abundantly throughout the CAZ, with the lower two subzones characterised by the medium-sized trirachodontids Langbergia and Trirachodon . The uppermost part of the formation, the Cricodon-Ufudocyclops subzone, yields trirachodontids of larger size. The majority of these trirachodontid specimens have previously been referred to Cricodon metabolus , a taxon also known from the Manda Beds of Tanzania and the Ntawere Formation of Zambia. Here we identify one of the specimens (BP/1/5538) previously referred to Cricodon as a new taxon, Guttigomphus avilionis . Guttigomphus can be distinguished from other gomphodont cynodonts by features of the upper postcanine teeth, such as an asymmetric crown in occlusal view (crown narrower along the lingual margin than the labial). Our phylogenetic analysis recovers Guttigomphus as a basal member of Trirachodontidae, outside of the clade including Cricodon , Langbergia and Trirachodon ., Competing Interests: The authors declare that they have no competing interests., (© 2022 Rayner et al.)

Climate change-induced mass extinctions provide unique opportunities to explore the impacts of global environmental disturbances on organismal evolution. However, their influence on terrestrial ecosystems remains poorly understood. Here, we provide a new time tree for the early evolution of reptiles and their closest relatives to reconstruct how the Permian-Triassic climatic crises shaped their long-term evolutionary trajectory. By combining rates of phenotypic evolution, mode of selection, body size, and global temperature data, we reveal an intimate association between reptile evolutionary dynamics and climate change in the deep past. We show that the origin and phenotypic radiation of reptiles was not solely driven by ecological opportunity following the end-Permian extinction as previously thought but also the result of multiple adaptive responses to climatic shifts spanning 57 million years.

The sternum is a stabilizing element in the axial skeleton of most tetrapods, closely linked with the function of the pectoral girdle of the appendicular skeleton. Modern mammals have a distinctive sternum characterized by multiple ossified segments, the origins of which are poorly understood. Although the evolution of the pectoral girdle has been extensively studied in early members of the mammalian total group (Synapsida), only limited data exist for the sternum. Ancestrally, synapsids exhibit a single sternal element and previously the earliest report of a segmental sternum in non-mammalian synapsids was in the Middle Triassic cynodont Diademodon tetragonus. Here, we describe the well-preserved sternum of a gorgonopsian, a group of sabre-toothed synapsids from the Permian. It represents an ossified, multipartite element resembling the mammalian condition. This discovery pulls back the origin of the distinctive "mammalian" sternum to the base of Theriodontia, significantly extending the temporal range of this morphology. Through a review of sternal morphology across Synapsida, we reconstruct the evolutionary history of this structure. Furthermore, we explore its role in the evolution of mammalian posture, gait, and ventilation through progressive regionalization of the postcranium as well as the posteriorization of musculature associated with mammalian breathing., (© 2022. The Author(s).)

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., Competing Interests: The authors declare that they have no competing interests., (© 2021 Pusch et al.)

Earth's largest biotic crisis occurred during the Permo-Triassic Transition (PTT). On land, this event witnessed a turnover from synapsid- to archosauromorph-dominated assemblages and a restructuring of terrestrial ecosystems. However, understanding extinction patterns has been limited by a lack of high-precision fossil occurrence data to resolve events on submillion-year timescales. We analyzed a unique database of 588 fossil tetrapod specimens from South Africa's Karoo Basin, spanning ∼4 My, and 13 stratigraphic bin intervals averaging 300,000 y each. Using sample-standardized methods, we characterized faunal assemblage dynamics during the PTT. High regional extinction rates occurred through a protracted interval of ∼1 Ma, initially co-occurring with low origination rates. This resulted in declining diversity up to the acme of extinction near the Daptocephalus - Lystrosaurus declivis Assemblage Zone boundary. Regional origination rates increased abruptly above this boundary, co-occurring with high extinction rates to drive rapid turnover and an assemblage of short-lived species symptomatic of ecosystem instability. The "disaster taxon" Lystrosaurus shows a long-term trend of increasing abundance initiated in the latest Permian. Lystrosaurus comprised 54% of all specimens by the onset of mass extinction and 70% in the extinction aftermath. This early Lystrosaurus abundance suggests its expansion was facilitated by environmental changes rather than by ecological opportunity following the extinctions of other species as commonly assumed for disaster taxa. Our findings conservatively place the Karoo extinction interval closer in time, but not coeval with, the more rapid marine event and reveal key differences between the PTT extinctions on land and in the oceans., Competing Interests: The authors declare no competing interest.

Taphonomic deformation, the distortion of fossils as a result of geological processes, poses problems for the use of geometric morphometrics in addressing paleobiological questions. Signal from biological variation, such as ontogenetic trends and sexual dimorphism, may be lost if variation from deformation is too high. Here, we investigate the effects of taphonomic deformation on geometric morphometric analyses of the abundant, well known Permian therapsid Diictodon feliceps . Distorted Diictodon crania can be categorized into seven typical styles of deformation: lateral compression, dorsoventral compression, anteroposterior compression, "saddle-shape" deformation (localized collapse at cranial mid-length), anterodorsal shear, anteroventral shear, and right/left shear. In simulated morphometric datasets incorporating known "biological" signals and subjected to uniform shear, deformation was typically the main source of variance but accurate "biological" information could be recovered in most cases. However, in empirical datasets, not only was deformation the dominant source of variance, but little structure associated with allometry and sexual dimorphism was apparent, suggesting that the more varied deformation styles suffered by actual fossils overprint biological variation. In a principal component analysis of all anomodont therapsids, deformed Diictodon specimens exhibit significant dispersion around the "true" position of this taxon in morphospace based on undistorted specimens. The overall variance associated with deformation for Anomodontia as a whole is minor, and the major axes of variation in the study sample show a strong phylogenetic signal instead. Although extremely problematic for studying variation in fossil taxa at lower taxonomic levels, the cumulative effects of deformation in this study are shown to be random, and inclusion of deformed specimens in higher-level analyses of morphological disparity are warranted. Mean morphologies of distorted specimens are found to approximate the morphology of undistorted specimens, so we recommend use of species-level means in higher-level analyses when possible., Competing Interests: The authors declare that they have no competing interests., (© 2020 Kammerer et al.)

Early members of the dinosaur-pterosaur clade Ornithodira are very rare in the fossil record, obscuring our understanding of the origins of this important group. Here, we describe an early ornithodiran ( Kongonaphon kely gen. et sp. nov.) from the Mid-to-Upper Triassic of Madagascar that represents one of the smallest nonavian ornithodirans. Although dinosaurs and gigantism are practically synonymous, an analysis of body size evolution in dinosaurs and other archosaurs in the context of this taxon and related forms demonstrates that the earliest-diverging members of the group may have been smaller than previously thought, and that a profound miniaturization event occurred near the base of the avian stem lineage. In phylogenetic analysis, Kongonaphon is recovered as a member of the Triassic ornithodiran clade Lagerpetidae, expanding the range of this group into Africa and providing data on the craniodental morphology of lagerpetids. The conical teeth of Kongonaphon exhibit pitted microwear consistent with a diet of hard-shelled insects, indicating a shift in trophic ecology to insectivory associated with diminutive body size. Small ancestral body size suggests that the extreme rarity of early ornithodirans in the fossil record owes more to taphonomic artifact than true reflection of the group's evolutionary history., Competing Interests: The authors declare no competing interest.

The origin of herbivory in the Carboniferous was a landmark event in the evolution of terrestrial ecosystems, increasing ecological diversity in animals but also giving them greater influence on the evolution of land plants. We evaluate the effect of early vertebrate herbivory on plant evolution by comparing local species richness of plant palaeofloras with that of vertebrate herbivores and herbivore body size. Vertebrate herbivores became diverse and achieved a much greater range of body sizes across the Carboniferous-Permian transition interval. This coincides with an abrupt reduction in local plant richness that persists throughout the Permian. Time-series regression analysis supports a negative relationship of plant richness with herbivore richness but a positive relationship of plant richness with minimum herbivore body size. This is consistent with studies of present-day ecosystems in which increased diversity of smaller, more selective herbivores places greater predation pressures on plants, while a prevalence of larger bodied, less selective herbivores reduces the dominance of a few highly tolerant plant species, thereby promoting greater local richness. The diversification of herbivores across the Carboniferous-Permian boundary, along with the appearance of smaller, more selective herbivores like bolosaurid parareptiles, constrained plant diversity throughout the Permian. These findings demonstrate that the establishment of widespread vertebrate herbivory has structured plant communities since the late Palaeozoic, as expected from examination of modern ecosystems, and illustrates the potential for fossil datasets in testing palaeoecological hypotheses.

The Pedra de Fogo Formation in the Parnaíba Basin of northeastern Brazil hosts a recently discovered lacustrine fauna and provides the only known record of the Captorhinidae in South America. Here, new captorhinid remains from this unit are described. Two partial mandibles, including one formerly ascribed to the genus Captorhinus , are here referred to Captorhinikos sp. a genus previously described from North America. The natural mould of a large mandible probably represents a new taxon within the captorhinid subclade Moradisaurinae, and a small skull roof is regarded as Captorhinidae indet. Captorhinids are generally considered to have been herbivores or omnivores. The Pedra de Fogo captorhinids likely played an important ecological role as primary consumers in the palaeoenvironment of this geological unit, which is also known for its extensive record of petrified forests. The new finds reinforce the close relationships between the continental faunas of palaeotropical western Gondwana and palaeoequatorial North America during the Cisuralian., Competing Interests: Claudia A. Marsicano is an Academic Editor for PeerJ., (© 2020 Cisneros et al.)

A single species of the dicynodontoid dicynodont Dicynodon is currently recognized from the late Permian Usili Formation of Tanzania: Dicynodon huenei Haughton, 1932. Restudy of the known Tanzanian materials of D . huenei demonstrates that they represent two distinct morphotypes, here considered separate taxa. The holotype of D . huenei is not referable to Dicynodon and instead is transferred to the genus Daptocephalus (but retained as a valid species, Daptocephalus huenei comb. nov.). A number of published dicynodontoid specimens from the Usili Formation, however, are referable to Dicynodon , and are here recognized as a new species ( Dicynodon angielczyki sp. nov.) Dicynodon angielczyki can be distinguished from its South African congener Dicynodon lacerticeps by the presence of an expansion of the squamosal and jugal beneath the postorbital bar and a curved, posterolateral expansion of the squamosal behind the temporal fenestra. Inclusion of Dicynodon angielczyki and D . huenei in a phylogenetic analysis supports their referral to Dicynodon and Daptocephalus (respectively). These results indicate higher basinal endemism in large late Permian dicynodonts than previously thought, a sharp contrast to the cosmopolitanism in the group in the earliest Triassic., Competing Interests: The author declares that they have no competing interests.

The cranial anatomy of the early non-mammalian cynodont Galesaurus planiceps from the South African Karoo Basin is redescribed on the basis of a computed tomographic reconstruction of the skull. Previously, little was known about internal skull morphology and the nervous and sensory system of this taxon. The endocranial anatomy of various cynodonts has been intensively studied in recent years to understand the origin of mammalian characters in the nasal capsule, brain and ear. However, these studies have focused on only a few taxa, the earliest of which is another Early Triassic cynodont, Thrinaxodon liorhinus. Galesaurus is phylogenetically stemward of Thrinaxodon and thus provides a useful test of whether the mammal-like features observed in Thrinaxodon were present even more basally in cynodont evolution. The cranial anatomy of G. planiceps is characterized by an intriguing mosaic of primitive and derived features within cynodonts. In contrast to the very similar internal nasal and braincase morphology of Galesaurus and Thrinaxodon, parts of the skull that seem to be fairly conservative in non-prozostrodont cynodonts, the morphology of the maxillary canal differs markedly between these taxa. Unusually, the maxillary canal of Galesaurus has relatively few ramifications, more similar to those of probainognathian cynodonts than that of Thrinaxodon. However, its caudal section is very short, a primitive feature shared with gorgonopsians and therocephalians. The otic labyrinth of Galesaurus is generally similar to that of Thrinaxodon, but differs in some notable features (e.g. proportional size of the anterior semicircular canal). An extremely large, protruding paraflocculus of the brain and a distinct medioventrally located notch on the anterior surface of the tabular, which forms the dorsal border of the large parafloccular lobe, are unique to Galesaurus among therapsids with reconstructed endocasts. These features may represent autapomorphies of Galesaurus, but additional sampling is needed at the base of Cynodontia to test this., (© 2019 Anatomical Society.)

Gorgonopsia is one of the major clades of non-mammalian synapsids, and includes an array of large-bodied carnivores that were the top terrestrial predators of the late Permian. Most research on the clade has focused on these largest members; small-bodied gorgonopsians are relatively little-studied. Here, we redescribe a small gorgonopsian skull (MB.R.999) from the late Permian (Tropidostoma Assemblage Zone) of South Africa on the basis of neutron and synchrotron CT reconstructions, which yield new data on internal cranial morphology in Gorgonopsia. Because of the largely undistorted nature of MB.R.999, we were also able to reconstruct unossified areas such as the brain endocast and the otic labyrinth. MB.R.999 can be referred to the taxon Cynariops robustus based on its general skull proportions, postcanine tooth count, preparietal morphology, and vomerine morphology. We refer additional small gorgonopsian specimens from the Victoria West area to Cynariops robustus, and consider Cynarioides grimbeeki and Cynarioides laticeps to be synonymous with C. robustus. Inclusion of Cynariops in a phylogenetic analysis of Gorgonopsia recovers it within a large clade of African taxa, more closely related to Lycaenops and rubidgeines than Eriphostoma or Gorgonops., Competing Interests: The authors have declared that no competing interests exist.

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., Competing Interests: The authors declare that they have no competing interests.

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., Competing Interests: The authors declare that they have no competing interests.

In this contribution we report the first occurrence of the enigmatic African probainognathian genus Aleodon in the Middle-early Late Triassic of several localities from the state of Rio Grande do Sul in southern Brazil. Aleodon is unusual among early probainognathians in having transversely-expanded postcanine teeth, similar to those of gomphodont cynognathians. This genus was previously known from the Manda Beds of Tanzania and the upper Omingonde Formation of Namibia. The Brazilian record of this genus is based upon multiple specimens representing different ontogenetic stages, including three that were previously referred to the sectorial-toothed probainognathian Chiniquodon theotonicus. We propose a new species of Aleodon (A. cromptoni sp. nov.) based on the specimens from Brazil. Additionally, we tentatively refer one specimen from the upper Omingonde Formation of Namibia to this new taxon, strengthening biostratigraphic correlations between these strata. Inclusion of A. cromptoni in a phylogenetic analysis of eucynodonts recovers it as the sister-taxon of A. brachyrhamphus within the family Chiniquodontidae. The discovery of numerous specimens of Aleodon among the supposedly monospecific Chiniquodon samples of Brazil raises concerns about chiniquodontid alpha taxonomy, particularly given the extremely broad geographic distribution of Chiniquodon. The discovery of Brazilian Aleodon and new records of the traversodontid Luangwa supports the hypothesis that at least two subzones can be recognized in the Dinodontosaurus Assemblage Zone.

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., Competing Interests: The authors declare there are no competing interests.

Dicynodonts were a highly successful group of herbivorous therapsids that inhabited terrestrial ecosystems from the Middle Permian through the end of the Triassic periods. Permian dicynodonts are extremely abundant in African deposits, but are comparatively poorly known from the other regions of Gondwana. Here we describe a new South American dicynodont, Rastodon procurvidens gen. et sp. nov., from the Boqueirão farm site of the Rio do Rasto Formation, Paraná Basin, Guadalupian/Lopingian of Brazil. Diagnostic features of R. procurvidens include uniquely anteriorly-curved maxillary tusks, well-developed ridges extending from the crista oesophagea anteriorly along the pterygoid rami, strong posterior angulation of the posterior pterygoid rami, and a bulbous, well-developed retroarticular process of the articular. Phylogenetic analysis indicates that R. procurvidens is the earliest and most basal member of Bidentalia, a cosmopolitan clade that includes Permian and Triassic dicynodonts whose dentition is usually reduced to a pair of maxillary tusks.

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.

Terrestrial vertebrates are first known to colonize high-latitude regions during the middle Permian (Guadalupian) about 270 million years ago, following the Pennsylvanian Gondwanan continental glaciation. However, despite over 150 years of study in these areas, the biogeographic origins of these rich communities of land-dwelling vertebrates remain obscure. Here we report on a new early Permian continental tetrapod fauna from South America in tropical Western Gondwana that sheds new light on patterns of tetrapod distribution. Northeastern Brazil hosted an extensive lacustrine system inhabited by a unique community of temnospondyl amphibians and reptiles that considerably expand the known temporal and geographic ranges of key subgroups. Our findings demonstrate that tetrapod groups common in later Permian and Triassic temperate communities were already present in tropical Gondwana by the early Permian (Cisuralian). This new fauna constitutes a new biogeographic province with North American affinities and clearly demonstrates that tetrapod dispersal into Gondwana was already underway at the beginning of the Permian.

The large dicynodont Eubrachiosaurus browni from the Upper Triassic Popo Agie Formation of Wyoming is redescribed. Eubrachiosaurus is a valid taxon that differs from Placerias hesternus, with which it was previously synonymized, by greater anteroposterior expansion of the scapula dorsally and a very large, nearly rectangular humeral ectepicondyle with a broad supinator process. Inclusion of Eubrachiosaurus in a revised phylogenetic analysis of anomodont therapsids indicates that it is a stahleckeriid closely related to the South American genera Ischigualastia and Jachaleria. The recognition of Eubrachiosaurus as a distinct lineage of North American dicynodonts, combined with other recent discoveries in the eastern USA and Europe, alters our perception of Late Triassic dicynodont diversity in the northern hemisphere. Rather than being isolated relicts in previously therapsid-dominated regions, Late Triassic stahleckeriid dicynodonts were continuing to disperse and diversify, even in areas like western North America that were otherwise uninhabited by coeval therapsids (i.e., cynodonts).

The feeding mechanism of gars (Ginglymodi : Lepisosteidae) is characterized by cranial elevation and lower jaw rotation but minimal cranial kinesis. Gar jaws have numerous, sharply pointed, elongate teeth for capture of evasive prey. Their mandibles range from relatively short to extremely long depending on the species. Jaw length and lever dimensions were hypothesized to affect the biomechanics of force and motion during feeding, according to simple mechanical models of muscles exerting force through first- or third-order levers. A morphometric protocol was used to measure the jaw structure of seven living and five fossil species of gar and these data were used to calculate the mechanical advantage (a measure of force transmission) for both opening and closing of the mandible. Gars were found to possess low mechanical advantage (MA) and high transmission of motion, although gars occupy a range of biomechanical states across the continuum of force vs. velocity transmission. The long-nose gar, Lepisosteus osseus, has one of the lowest jaw closing MAs (0.05) ever measured in fishes. Intraspecific lever mechanics were also calculated for a developmental series (from feeding larvae to adults) of L. osseus and Atractosteus spatula. A characteristic ontogenetic curve in MA of the lower jaw was obtained, with a large decrease in MA between larva and juvenile, followed by a steady increase during adult growth. This curve correlates with a change in prey type, with the small, robust-jawed individuals feeding mainly on crustaceans and insects and the large, long-jawed individuals of all species becoming mainly piscivorous. Principal components analysis of functionally important morphometrics shows that several gar species occupy different regions of functional morphospace. Some fossil gar species are also placed within functional morphospace using this approach., ((c) 2004 Wiley-Liss, Inc.)