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Upper Permian rocks of the former supercontinent Gondwana record climax late Paleozoic terrestrial vertebrate faunas that were dominated numerically and ecologically by therapsid synapsids. Older faunal elements of earlier Paleozoic faunas, such as captorhinid reptiles, are rare and scattered components of the first amniote faunas to inhabit high-latitude regions. Here we describe a new genus and species of moradisaurine captorhinid that represents an archaic faunal element of the high-fibre herbivore fauna of the late Permian of what is now peninsular India. The presence of a relatively broad parietal and three rows of conical teeth on the maxilla and the dentary distinguishes Indosauriscus kuttyi gen. et sp. nov. from other moradisaurines. The hypodigm of I. kuttyi comprises skulls that range in length from 39 mm to 54 mm, and high ossification of the braincase elements and well developed skull-roof sutures, indicate that I. kuttyi adults were smaller than those of most moradisaurines. Results of our phylogenetic analyses suggest that moradisaurines, despite appearing first in the paleotropics, dispersed into temperate, high-latitude regions of Pangea early in their evolutionary history. Moradisaurines in dicynodont-dominated faunas, viz. Indosauriscus kuttyi and Gansurhinus naobaogouensis , were the smallest high-fibre herbivores in their respective faunas. This suggests that small body size may have evolved in these moradisaurines as they co-evolved with the more numerous and diverse dicynodont therapsids.

The initial stages of evolution of Diapsida (the large clade that includes not only snakes, lizards, crocodiles and birds, but also dinosaurs and numerous other extinct taxa) is clouded by an exceedingly poor Palaeozoic fossil record. Previous studies had indicated a 38 Myr gap between the first appearance of the oldest diapsid clade (Araeoscelidia), ca 304 million years ago (Ma), and that of its sister group in the Middle Permian (ca 266 Ma). Two new reptile skulls from the Richards Spur locality, Lower Permian of Oklahoma, represent a new diapsid reptile: Orovenator mayorum n. gen. et sp. A phylogenetic analysis identifies O. mayorum as the oldest and most basal member of the araeoscelidian sister group. As Richards Spur has recently been dated to 289 Ma, the new diapsid neatly spans the above gap by appearing 15 Myr after the origin of Diapsida. The presence of O. mayorum at Richards Spur, which records a diverse upland fauna, suggests that initial stages in the evolution of non-araeoscelidian diapsids may have been tied to upland environments. This hypothesis is consonant with the overall scant record for non-araeoscelidian diapsids during the Permian Period, when the well-known terrestrial vertebrate communities are preserved almost exclusively in lowland deltaic, flood plain and lacustrine sedimentary rocks.

Living species of mammals, crocodiles and most species of birds exhibit parental care, but evidence of this behaviour is extremely rare in the fossil record. Here, we present a new specimen of varanopid 'pelycosaur' from the Middle Permian of South Africa. The specimen is an aggregation, consisting of five articulated individuals preserved in undisturbed, close, lifelike, dorsal-up, subparallel positions, indicating burial in 'life position'. Two size classes are represented. One is 50% larger than the others, is well ossified, has fused neurocentral sutures and is distinguished by a coat of dermal ossifications that covers the neck and shoulder regions. We regard this individual to be an adult. The remaining four skeletons are considered to be juveniles as they are approximately the same size, are poorly ossified, have open neurocentral sutures and lack dermal ossifications. Aggregates of juvenile amniotes are usually siblings. Extant analogues of adult and juvenile groupings suggest that the adult is one of the parents, leading us to regard the aggregation as a family group. The Late Middle Permian age of the varanopid family predates the previously known oldest fossil evidence of parental care in terrestrial vertebrates by 140 Myr.

Recent morphological and molecular studies have challenged conventional hypotheses concerning the affinities of turtles, and this has led to unexpected and undocumented changes in the composition of the well-known taxon Reptilia, with additional ramifications for the nomenclature of some of its included taxa. Modesto and Anderson examine the consequences of the application of priority to the nomen Reptilia as their understanding of early amniote interrelationships has progressed over the past two decades, and offer a new definition that brings the phylogenetic concept of this taxon name into line with both currently accepted conventions of Phylogenetic Nomenclature and historical usage.

A group of early terrestrial vertebrates called varanopids, long regarded as mammal-line amniotes, is placed wholesale with reptiles in a new analysis of early amniote relationships. Meanwhile, a new species of varanopid from Canada provides the oldest evidence for extended parental care in terrestrial vertebrates.

Amniotes, tetrapods that evolved the cleidoic egg and thus independence from aquatic larval stages, appeared ca 314 Ma during the Coal Age. The rapid diversification of amniotes and other tetrapods over the course of the Late Carboniferous period was recently attributed to the fragmentation of coal-swamp rainforests ca 307 Ma. However, the amniote fossil record during the Carboniferous is relatively sparse, with ca 33% of the diversity represented by single specimens for each species. We describe here a new species of reptilian amniote that was collected from uppermost Carboniferous rocks of Prince Edward Island, Canada. Erpetonyx arsenaultorum gen. et sp. nov. is a new parareptile distinguished by 29 presacral vertebrae and autapomorphies of the carpus. Phylogenetic analyses of parareptiles reveal E. arsenaultorum as the closest relative of bolosaurids. Stratigraphic calibration of our results indicates that parareptiles began their evolutionary radiation before the close of the Carboniferous Period, and that the diversity of end-Carboniferous reptiles is 80% greater than suggested by previous work. Latest Carboniferous reptiles were still half as diverse as synapsid amniotes, a disparity that may be attributable to preservational biases, to collecting biases, to the origin of herbivory in tetrapods or any combination of these factors.

Captorhinikos valensis is a poorly known, multiple-tooth-rowed captorhinid reptile from the Lower Permian Vale Formation of Texas. Our reappraisal of C. valensis reveals it to be a small moradisaurine, exhibiting a maximum of five rows of bullet-shaped teeth in the multiple-rowed region of both the maxilla and the dentary. The slightly radiating organization of the tooth rows distinguishes C. valensis from the parallel arrangement of the tooth rows exhibited by all other moradisaurines. Captorhinikos valensis is also distinguishable from the coeval moradisaurine Labidosaurikos meachami by a more conspicuously denticulated, broader, ‘U’-shaped transverse flange of the pterygoid, a plesiomorphic morphology shared with the large, single-rowed captorhinid Labidosaurus hamatus. Postcranial information is limited to two short series of presacral vertebrae not associated with the cranial materials; open neurocentral sutures are present in one specimen, indicating immaturity at death. We investigated the relationships of C. valensis to other captorhinids by adding it to the data matrix of a previously published analysis that included several moradisaurine captorhinids. A branch-and-bound PAUP analysis discovered a single optimal tree. Whereas a previous analysis of captorhinid interrelationships found the (undifferentiated) genus Captorhinikos to fall outside of a clade composed of L. hamatus and the large moradisaurines, our analysis recovered C. valensis in a clade with the genera Labidosaurikos, Gansurhinus, Moradisaurus, and Rothianiscus (i.e., Moradisaurinae sensu stricto), and Captorhinikos chozaensis as the sister species of a clade that includes L. hamatus and Moradisaurinae s.s.; Captorhinikos chozaensis is no longer classifiable as a moradisaurine (according to our phylogenetic definition for the group), and should be assigned to a new genus. Stratigraphic calibration of our captorhinid phylogeny indicates that moradisaurines evolved by the middle Kungurian (middle Leonardian).

Description of a new species of the parareptile genus Delorhynchus is based on a well-preserved partial subadult skeleton, an isolated adult skull, and disarticulated elements recently collected from the Lower Permian Richards Spur locality of Oklahoma, U.S.A. Delorhynchus cifellii, sp. nov., is distinguished from Delorhynchus priscus by the lack of an accessory articulating anterodorsal flange of the maxilla. The hypodigm of Delorhynchus cifellii reveals that Delorhynchus is distinguished from other parareptiles by cranial dermal sculpture consisting of a system of low, smooth tuberosities and a pattern of diffuse shallow, circular dimples. In a phylogenetic analysis of parareptiles, Delorhynchus cifellii is positioned as the sister species of Lanthanosuchoidea. Recognition of Delorhynchus cifellii. sp. nov., and its phylogenetic position among parareptiles highlights the significance of the Richards Spur locality in our understanding of the early evolutionary history of reptiles.

We report on a large burrow cast with skeletal contents from Lower Triassic strata of the Palingkloof Member of the Balfour Formation, which forms the lowermost portion of the Lystrosaurus Assemblage Zone (LAZ) of South Africa. The burrow cast is similar to large burrow casts previously described from the LAZ that were identified as large-scale Scoyenia domichnia. It is the first large burrow cast from the LAZ found to contain diagnostic fossil bone. The burrow cast is a relatively straight, subhorizontal (inclined ∼12°), dorsoventrally compressed tube consisting of an entry ramp and living chamber; the entrance to the burrow is not preserved and there is no evidence that the ramp formed a spiral section. The skeletal material comprises a single, partial, disarticulated skeleton of a juvenile animal that can be assigned with confidence to the dicynodont genus Lystrosaurus. Whereas similar large-diameter burrow casts from strata slightly higher in the LAZ have been attributed to Lystrosaurus, we present an alternative hypothesis that a carnivorous tetrapod constructed the burrow. Our preferred hypothesis is supported by the observation that the interred Lystrosaurus skeleton is too small to be the maker of this particular burrow, by the general observation that carnivorous tetrapods construct relatively straight burrows, and by the partial, disarticulated state of the skeleton, which we interpret as the remains of larded prey. We suggest that akidnognathid theriodonts of the genera Moschorhinus or Olivierosuchus, the most conspicuous large predators of the LAZ, were the constructors of large-diameter, subhorizontal burrows.

Several new specimens of the Triassic therocephalian Tetracynodon darti have become available in recent years, allowing substantial corrections and expansions to previous descriptions. We here analyze T. darti in the context of therocephalian relationships and biology, using computed tomographic (CT) scanning to reveal details of the skull. Histological sections, as well as the degree of ossification of individual elements, both suggest that the available specimens are subadults (rather than rapidly growing neonates), and probably yearlings. A maxillary shelf bearing a long suture with the vomer similar to that of Lycideops is present, and T. darti also shares a nasal-lacrimal contact with that taxon. There is no evidence for the presence of maxilloturbinal bones. The postcranial skeleton is slender and similar to that of regisaurids and other small baurioids. The skull bones show clear indentations demarcating the major divisions of the brain, allowing one of the most detailed reconstructions of a non-mammalian therapsid brain published to date. The brain is strikingly plesiomorphic in many features, particularly in the retention of large dorsally positioned optic lobes of the tectum, but generally shows features that are intermediate between gorgonopsians and cynodonts, thus clarifying this stage of the evolution of the synapsid brain. A cladistic analysis of 33 therapsid taxa and 131 morphological characters supports the monophyly of Therocephalia, and there is evidence for a monophyletic Lycideopidae within Baurioidea. Lycideopidae contains species with long snouts, nasal-lacrimal contacts, and incipient secondary palates, including Lycideops, Choerosaurus, T. tenuis, and T. darti.