Anatomy and phylogenetic relationships of Tazoudasaurus naimi (Dinosauria, Sauropoda) from the late Early Jurassic of Morocco

The complete osteology of the basal sauropod Tazoudasaurus naimi from the late Early Jurassic Toundoute continental series of Ouarzazate Province, Morocco, is presented. The described material belongs to juvenile to adult individuals. The skeleton of Tazoudasaurus is virtually complete except for the skull and presents a combination of plesiomorphic and apomorphic sauropodomorph characters. Phylogenetic analysis indicates that Tazoudasaurus shares with Vulcanodon several derived features that include strongly transversely flattened tibial shaft and the marked dorsoventral flattening of the unguals of pedal digits II and III. Both taxa are placed within the Vulcanodontidae, at the base of a new clade named Gravisauria n. nom. Our analysis underscores the major morphological changes that occur among Gravisauria between the Vulcanodontidae and the Eusauropoda. The numerous remains of Tazoudasaurus were recovered from a bone-bed associated with a few remains of the basal abelisauroid Berberosaurus. A minimum of six individuals was buried at the site. Taphonomical data suggest that Tazoudasaurus had a gregarious behaviour. The different interpretations of the evolution of the configuration and posture of the manus in basal sauropods can be tested thanks to the discovery of a complete articulated manus of Tazoudasaurus. The latter is clearly digitigrade with a spreading configuration of the metacarpus. Early sauropod evolution is analyzed in a broad extinction/radiation perspective. Prosauropoda and Coelophysoidea extinction around the Pliensbachian-Toarcian boundary, followed by the late Early Jurassic to Middle Jurassic radiation of Gravisauria, Neoceratosauria, Tetanurae and Euornithopoda are linked to the Pliensbachian-Toarcian mass extinction event.