An overview of the appendicular skeletal anatomy of South American titanosaurian sauropods, with definition of a newly recognized clade.

In the last two decades, the number of phylogenetically informative anatomical characters recognized in the appendicular skeleton of titanosaurian sauropod dinosaurs has increased dramatically with the discovery of new and comparatively complete specimens. Here we provide an overview of the appendicular skeletal morphology of South American titanosaurs and discuss its significance for phylogenetic reconstruction. The appendicular skeletal diversity of South American titanosaurs is substantially greater than was initially appreciated. Moreover, some regions of the appendicular skeleton, such as the pes, exhibit remarkable variability in form. Multiple synapomorphies of Titanosauria and the less inclusive clades Lithostrotia and Saltasauridae consist of characters of the girdles and limbs. Although the phylogenetic definitions of titanosaurian clades such as Saltasaurinae and Lognkosauria are stable, the taxonomic content of these clades has varied in recent analyses depending on the phylogenetic topology recovered. Within Titanosauria, the results of four recent, largely independent analyses support the existence of a derived titanosaurian lineage distinct from the 'Saltasaurinae line,' which is herein termed Colossosauria. At present, this clade is mainly comprised by taxa within Lognkosauria and Rinconsauria, and is useful in discussions of titanosaurian lower-level relationships.

Histological evidence for a supraspinous ligament in sauropod dinosaurs.

Supraspinous ossified rods have been reported in the sacra of some derived sauropod dinosaurs. Although different hypotheses have been proposed to explain the origin of this structure, histological evidence has never been provided to support or reject any of them. In order to establish its origin, we analyse and characterize the microstructure of the supraspinous rod of two sauropod dinosaurs from the Upper Cretaceous of Argentina. The supraspinous ossified rod is almost entirely formed by dense Haversian bone. Remains of primary bone consist entirely of an avascular tissue composed of two types of fibre-like structures, which are coarse and longitudinally (parallel to the main axis of the element) oriented. These structures are differentiated on the basis of their optical properties under polarized light. Very thin fibrous strands are also observed in some regions. These small fibres are all oriented parallel to one another but perpendicular to the element main axis. Histological features of the primary bone tissue indicate that the sacral supraspinous rod corresponds to an ossified supraspinous ligament. The formation of this structure appears to have been a non-pathological metaplastic ossification, possibly induced by the continuous tensile forces applied to the element.

A gigantic, exceptionally complete titanosaurian sauropod dinosaur from southern Patagonia, Argentina.

Titanosaurian sauropod dinosaurs were the most diverse and abundant large-bodied herbivores in the southern continents during the final 30 million years of the Mesozoic Era. Several titanosaur species are regarded as the most massive land-living animals yet discovered; nevertheless, nearly all of these giant titanosaurs are known only from very incomplete fossils, hindering a detailed understanding of their anatomy. Here we describe a new and gigantic titanosaur, Dreadnoughtus schrani, from Upper Cretaceous sediments in southern Patagonia, Argentina. Represented by approximately 70% of the postcranial skeleton, plus craniodental remains, Dreadnoughtus is the most complete giant titanosaur yet discovered, and provides new insight into the morphology and evolutionary history of these colossal animals. Furthermore, despite its estimated mass of about 59.3 metric tons, the bone histology of the Dreadnoughtus type specimen reveals that this individual was still growing at the time of death.

Lamniform shark teeth from the late cretaceous of southernmost South America (Santa Cruz province, Argentina).

Here we report multiple lamniform shark teeth recovered from fluvial sediments in the (Campanian-Maastrichtian) Cerro Fortaleza Formation, Santa Cruz Province, Argentina. This small tooth assemblage is compared to various lamniform sharks possessing similar dental morphologies, including Archaeolamna, Cretalamna, Dwardius, Dallasiella, and Cretodus. Although the teeth share numerous morphological features with the genus Archaeolamna, including a developed neck that maintains a relatively consistent width along the base of the crown, the small sample size and incomplete nature of these specimens precludes definitive taxonomic assignment. Regardless, the discovery of selachian teeth unique from those previously described for the region broadens the known diversity of Late Cretaceous South American sharks. Additionally, the discovery of the teeth in fluvial sandstone may indicate a euryhaline paleobiology in the lamniform taxon or taxa represented by this tooth assemblage.

A reappraisal of Notohypshilophodon comodorensis (Ornithischia: Ornithopoda) from the Late Cretaceous of Patagonia, Argentina.

The Bajo Barreal Formation (Cenomanian, Late Cretaceous; central Patagonia, Argentina) preserves an important and rich fossil record. Among the dinosaurs described, a small ornithischian, Notohypsilophon comodorensis, was named in the 1990s. This small-bodied dinosaur, the most complete representative of the group discovered in that formation, was described as a "hypsilophodontid" ornithopod based on close morphological affinities with other members of that group, which is currently regarded as paraphyletic. Within this context, we present a restudy of Notohypsilophodon. This dinosaur is considered a basal ornithopod, probably more basal than Gasparinisaura. Likewise, this analysis emends and provides additional unique anatomical characters that support its taxonomic validity and position. On the basis of weak evidence, Notohypsilophodon might share the presence of gastroliths with other basal ornithopods (Gasparinisaura and Haya), which could suggest a specific behavior in response to its food ingestion. Finally, this study increases our knowledge of the evolutionary dynamics of South American Cretaceous ornithopods, and therefore broadens our knowledge of the early Late Cretaceous continental vertebrate assemblages of central Patagonia and of the Southern Hemisphere in general.

The behavioral implications of a multi-individual bonebed of a small theropod dinosaur.

BACKGROUND: Central Patagonia, Argentina, preserves an abundant and rich fossil record. Among vertebrate fossils from the Upper Cretaceous Bajo Barreal Formation of Patagonia, five individuals of the small, non-avian theropod dinosaur Aniksosaurus darwini were recovered. Group behavior is an important aspect of dinosaur paleoecology, but it is not well-documented and is poorly understood among non-avian Theropoda. METHODS/PRINCIPAL FINDINGS: The taphonomic association of individuals from the Bajo Barreal Formation and aspects of their bone histology suggest gregarious behavior for Aniksosaurus, during at least a portion of the life history of this species. Histology indicates that the specimens were juvenile to sub-adult individuals. In addition, morphological differences between individuals, particularly proportions of the appendicular bones, are probably related to body-size dimorphism rather than ontogenetic stage. CONCLUSIONS/SIGNIFICANCE: Gregarious behaviour may have conferred a selective advantage on Aniksosaurus individuals, contributing to their successful exploitation of the Cretaceous paleoenvironment preserved in the Bajo Barreal Formation. The monospecific assemblage of Aniksosaurus specimens constitutes only the second body fossil association of small, coelurosaurian theropods in South America and adds valuable information about the paleoecologies of non-avian theropod dinosaurs, particularly in the early Late Cretaceous of Patagonia.