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Current hypotheses of early tetrapod evolution posit close ecological and biogeographic ties to the extensive coal-producing wetlands of the Carboniferous palaeoequator with rapid replacement of archaic tetrapod groups by relatives of modern amniotes and lissamphibians in the late Carboniferous (about 307 million years ago). These hypotheses draw on a tetrapod fossil record that is almost entirely restricted to palaeoequatorial Pangea (Laurussia)1,2. Here we describe a new giant stem tetrapod, Gaiasia jennyae, from high-palaeolatitude (about 55° S) early Permian-aged (about 280 million years ago) deposits in Namibia that challenges this scenario. Gaiasia is represented by several large, semi-articulated skeletons characterized by a weakly ossified skull with a loosely articulated palate dominated by a broad diamond-shaped parasphenoid, a posteriorly projecting occiput, and enlarged, interlocking dentary and coronoid fangs. Phylogenetic analysis resolves Gaiasia within the tetrapod stem group as the sister taxon of the Carboniferous Colosteidae from Euramerica. Gaiasia is larger than all previously described digited stem tetrapods and provides evidence that continental tetrapods were well established in the cold-temperate latitudes of Gondwana during the final phases of the Carboniferous-Permian deglaciation. This points to a more global distribution of continental tetrapods during the Carboniferous-Permian transition and indicates that previous hypotheses of global tetrapod faunal turnover and dispersal at this time2,3 must be reconsidered.

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Sauropodomorph dinosaurs dominated the herbivorous niches during the first 40 million years of dinosaur history (Late Triassic-Early Jurassic), yet palaeobiological factors that influenced their evolutionary success are not fully understood. For instance, knowledge on their behaviour is limited, although herding in sauropodomorphs has been well documented in derived sauropods from the Late Jurassic and Cretaceous. Here we report an exceptional fossil occurrence from Patagonia that includes over 100 eggs and skeletal specimens of 80 individuals of the early sauropodomorph Mussaurus patagonicus, ranging from embryos to fully-grown adults, with an Early Jurassic age as determined by high-precision U-Pb zircon geochronology. Most specimens were found in a restricted area and stratigraphic interval, with some articulated skeletons grouped in clusters of individuals of approximately the same age. Our new discoveries indicate the presence of social cohesion throughout life and age-segregation within a herd structure, in addition to colonial nesting behaviour. These findings provide the earliest evidence of complex social behaviour in Dinosauria, predating previous records by at least 40 My. The presence of sociality in different sauropodomorph lineages suggests a possible Triassic origin of this behaviour, which may have influenced their early success as large terrestrial herbivores.

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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.

RESUMEN

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.

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BACKGROUND: Temnospondyls are one of the earliest radiations of limbed vertebrates. Skeletal remains of more than 190 genera have been identified from late Paleozoic and early Mesozoic rocks. Paleozoic temnospondyls comprise mainly small to medium sized forms of diverse habits ranging from fully aquatic to fully terrestrial. Accordingly, their ichnological record includes tracks described from many Laurasian localities. Mesozoic temnospondyls, in contrast, include mostly medium to large aquatic or semi-aquatic forms. Exceedingly few fossil tracks or trackways have been attributed to Mesozoic temnospondyls, and as a consequence very little is known of their locomotor capabilities on land. METHODOLOGY/PRINCIPAL FINDINGS: We report a ca. 200 Ma trackway, Episcopopus ventrosus, from Lesotho, southern Africa that was made by a 3.5 m-long animal. This relatively long trackway records the trackmaker dragging its body along a wet substrate using only the tips of its digits, which in the manus left characteristic drag marks. Based on detailed mapping, casting, and laser scanning of the best-preserved part of the trackway, we identified synapomorphies (e.g., tetradactyl manus, pentadactyl pes) and symplesiomorphies (e.g., absence of claws) in the Episcopopus trackway that indicate a temnospondyl trackmaker. CONCLUSIONS/SIGNIFICANCE: Our analysis shows that the Episcopopus trackmaker progressed with a sprawling posture, using a lateral-sequence walk. Its forelimbs were the major propulsive elements and there was little lateral bending of the trunk. We suggest this locomotor style, which differs dramatically from the hindlimb-driven locomotion of salamanders and other extant terrestrial tetrapods can be explained by the forwardly shifted center of mass resulting from the relatively large heads and heavily pectoral girdles of temnospondyls.

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