Explore the vast range of titles available.

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.

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 samplestandardized 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.

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.

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.

Therapsids were a dominant component of middle–late Permian terrestrial ecosystems worldwide, eventually giving rise to mammals during the early Mesozoic. However, little is currently known about the time and place of origin of Therapsida. Here we describe a definitive therapsid from the lower–?middle Permian palaeotropics, a partial skeleton of a gorgonopsian from the island of Mallorca, western Mediterranean. This specimen represents, to our knowledge, the oldest gorgonopsian record worldwide, and possibly the oldest known therapsid. Using emerging relaxed clock models, we provide a quantitative timeline for the origin and early diversification of therapsids, indicating a long ghost lineage leading to the evolutionary radiation of all major therapsid clades within less than 10 Myr, in the aftermath of Olson’s Extinction. Our findings place this unambiguous early therapsid in an ancient summer wet biome of equatorial Pangaea, thus suggesting that the group originated in tropical rather than temperate regions. Gorgonopsians were sabre toothed therapsids, a group that is ancestral to mammals. The authors here describe a gorgonopsian from the early–middle Permian of Mallorca, suggesting that therapsids originated in tropical regions and diversified quickly over 10 million years.

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.

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.

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 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.

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.