Subaqueous foraging among carnivorous dinosaurs.

Secondary aquatic adaptations evolved independently more than 30 times from terrestrial vertebrate ancestors1,2. For decades, non-avian dinosaurs were believed to be an exception to this pattern. Only a few species have been hypothesized to be partly or predominantly aquatic3-11. However, these hypotheses remain controversial12,13, largely owing to the difficulty of identifying unambiguous anatomical adaptations for aquatic habits in extinct animals. Here we demonstrate that the relationship between bone density and aquatic ecologies across extant amniotes provides a reliable inference of aquatic habits in extinct species. We use this approach to evaluate the distribution of aquatic adaptations among non-avian dinosaurs. We find strong support for aquatic habits in spinosaurids, associated with a marked increase in bone density, which precedes the evolution of more conspicuous anatomical modifications, a pattern also observed in other aquatic reptiles and mammals14-16. Spinosaurids are revealed to be aquatic specialists with surprising ecological disparity, including subaqueous foraging behaviour in Spinosaurus and Baryonyx, and non-diving habits in Suchomimus. Adaptation to aquatic environments appeared in spinosaurids during the Early Cretaceous, following their divergence from other tetanuran theropods during the Early Jurassic17.

Tail-propelled aquatic locomotion in a theropod dinosaur.

In recent decades, intensive research on non-avian dinosaurs has strongly suggested that these animals were restricted to terrestrial environments1. Historical proposals that some groups, such as sauropods and hadrosaurs, lived in aquatic environments2,3 were abandoned decades ago4-6. It has recently been argued that at least some of the spinosaurids-an unusual group of large-bodied theropods of the Cretaceous era-were semi-aquatic7,8, but this idea has been challenged on anatomical, biomechanical and taphonomic grounds, and remains controversial9-11. Here we present unambiguous evidence for an aquatic propulsive structure in a dinosaur, the giant theropod Spinosaurus aegyptiacus7,12. This dinosaur has a tail with an unexpected and unique shape that consists of extremely tall neural spines and elongate chevrons, which forms a large, flexible fin-like organ capable of extensive lateral excursion. Using a robotic flapping apparatus to measure undulatory forces in physical models of different tail shapes, we show that the tail shape of Spinosaurus produces greater thrust and efficiency in water than the tail shapes of terrestrial dinosaurs and that these measures of performance are more comparable to those of extant aquatic vertebrates that use vertically expanded tails to generate forward propulsion while swimming. These results are consistent with the suite of adaptations for an aquatic lifestyle and piscivorous diet that have previously been documented for Spinosaurus7,13,14. Although developed to a lesser degree, aquatic adaptations are also found in other members of the spinosaurid clade15,16, which had a near-global distribution and a stratigraphic range of more than 50 million years14, pointing to a substantial invasion of aquatic environments by dinosaurs.

New Information on Tataouinea hannibalis from the Early Cretaceous of Tunisia and Implications for the Tempo and Mode of Rebbachisaurid Sauropod Evolution.

The rebbachisaurid sauropod Tataouinea hannibalis represents the first articulated dinosaur skeleton from Tunisia and one of the best preserved in northern Africa. The type specimen was collected from the lower Albian, fluvio-estuarine deposits of the Ain el Guettar Formation (southern Tunisia). We present detailed analyses on the sedimentology and facies distribution at the main quarry and a revision of the vertebrate fauna associated with the skeleton. Data provide information on a complex ecosystem dominated by crocodilian and other brackish water taxa. Taphonomic interpretations indicate a multi-event, pre-burial history with a combination of rapid segregation in high sediment supply conditions and partial subaerial exposure of the carcass. After the collection in 2011 of the articulated sacrum and proximalmost caudal vertebrae, all showing a complex pattern of pneumatization, newly discovered material of the type specimen allows a detailed osteological description of Tataouinea. The sacrum, the complete and articulated caudal vertebrae 1-17, both ilia and ischia display asymmetrical pneumatization, with the left side of vertebrae and the left ischium showing a more extensive invasion by pneumatic features than their right counterparts. A pneumatic hiatus is present in caudal centra 7 to 13, whereas caudal centra 14-16 are pneumatised by shallow fossae. Bayesian inference analyses integrating morphological, stratigraphic and paleogeographic data support a flagellicaudatan-rebbachisaurid divergence at about 163 Ma and a South American ancestral range for rebbachisaurids. Results presented here suggest an exclusively South American Limaysaurinae and a more widely distributed Rebbachisaurinae lineage, the latter including the South American taxon Katepensaurus and a clade including African and European taxa, with Tataouinea as sister taxon of Rebbachisaurus. This scenario would indicate that South America was not affected by the end-Jurassic extinction of diplodocoids, and was most likely the centre of the rapid radiation of rebbachisaurids to Africa and Europe between 135 and 130 Ma.