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The first unambiguous evidence of the presence of osteoderms in sauropod dinosaurs came from the discovery of Saltasaurus loricatus, a titanosaur from the Upper Cretaceous of Argentina. The dermal armor of Saltasaurus is composed of bony plates and small dermal ossicles. Here, we analyze the bone microstructure of these elements and provide information regarding its origin and development. The bony plates are composed almost entirely of reconstructed cancellous bone. Remains of primary bone consist of coarse bundles of mineralized collagenous fibers towards the external surface. Also, woven fibered bone tissue appears in the basal and lateral regions. Dermal ossicles lack secondary remodeling, and their matrix is formed by three orthogonal systems of collagenous fiber bundles. Growth lines are present in both bony plates and ossicles. Bone histology reveals that osteoderms mainly originated through direct mineralization (metaplasia) of the dermis, although other mechanisms are also involved (at least in the origin of dermal plates). The common features of development and integumental location of the osteoderms of Saltasaurus and other non−related vertebrates (e.g., lepidosaurs, crocodylomorphs) are linked to the intrinsic skeletogenic properties of the dermis.
The Late Cretaceous South American sauropods Neuquensaurus australis and Saltasaurus loricatus are represented by well−preserved and abundant material that has been integral to our understanding of titanosaur anatomy for decades. Although the hypodigms for these species span most of the skeleton, holotypic materials are limited to a few bones that do not overlap between the two taxa. In this contribution, we augment the holotype of Neuquensaurus australis with a partial sacrum that was preserved in articulation with one of the caudal vertebrae from its original description, but not recognised as such at the time. We document this field association via the presence of a broken piece of matrix on the sixth sacral vertebral centrum that has a snap−fit to matrix on the rim of the anterior condyle of the holotypic biconvex vertebra. Based on comparisons with a more complete sacrum and ilium of a referred specimen of Neuquensaurus australis, we interpret this biconvex vertebra to be the seventh sacral vertebra. This raises the possibility that the biconvex “first caudal” vertebra of some other titanosaurs may be part of the sacrum as well. Augmentation of the Neuquensaurus australis holotype to include a sacrum makes it directly comparable to the holotype of Saltasaurus loricatus. Morphological differences in the number, shape, and proportion of sacral vertebrae allow discrimination between Neuquensaurus and Saltasaurus, confirming their generic separation. The El Brete quarry, which preserves the holotypic sacrum and abundant referred specimens of Saltasaurus loricatus, also preserves a sacrum consisting of seven vertebrae that bears autapomorphies of Neuquensaurus australis, indicating that these two saltasaurines coexisted.
Lirainosaurus is the only titanosaurian sauropod described to date from the Late Cretaceous of the Iberian Peninsula. The type of Lirainosaurus astibiae consists of both cranial and postcranial remains that were found as disarticulated elements in the Laño quarry (Treviño, northern Spain). This taxon was diagnosed originally on the basis of vertebral and appendicular autapomorphic traits. The study of a paratypic skull fragment and a second referred specimen provides information about its braincase morphology. Lirainosaurus is regarded as a derived titanosaur on the basis of the complete fusion between the prootic and the exoccipital−opisthotic complex, the position of the cranial foramina, and the shape and orientation of the occipital condyle. The braincase of L. astibiae appears to be diagnostic in the presence of a foramen distally on each basal tubera. The absence of median subcondylar foramina in the basioccipital may be an autopomorphic trait or be due to ontogenetic growth. A comparison with other partial skulls known in Europe suggests a high diversity during the Campanian/Maastrichtian, with at least three different titanosaurian species living in the Ibero−Armorican Island.
A new sauropod titanosaur from the Upper Cretaceous Anacleto Formation is described. Narambuenatitan palomoi gen. et sp. nov., is diagnosed by cranial and axial autapomorphies. The holotype, which represent a subadult individual, consists of the left premaxilla and maxilla, braincase, both quadrates, one cervical vertebrae, one dorsal vertebra, fragments of cervical and dorsal ribs, seventeen caudal vertebrae, caudal transverse processes, fragments of haemal arches, left sternal plate, right coracoid, left humerus, left ulnae, both pubes, iliac pedicel, proximal fragment of right ischia, and an incomplete left femur. The phylogenetic analysis indicates that Narambuenatitan is a non−eutitanosaurian lithostrotian, and that it shares with Epachthosaurus a neural spine in middle caudal vertebrae which are laminar and posteriorly elongated.
Knowledge of titanosaurian cranial anatomy has improved substantially in the last decade because several skulls have come to light or were restudied. The discovery of Bonitasaura salgadoi, a partial titanosaurian skeleton including cranial bones, permitted the definitive recognition of square jaws in a titanosaurian sauropod as well as a peculiar skull morphology that increases the morphological diversity of the group. Here we present a full description and illustration of the skull material of B. salgadoi. Among cranial bones, the lacrimal, quadrate, and dentary exhibit apomorphic differences from those of other titanosaurians. Conversely, the frontal and parietal are more conservative. A phylogenetic analysis recovers B. salgadoi as a member of the Titanosauria, related to mid−sized to large titanosauroids from the Turonian–Campanian of South America, in contrast to a previous hypothesis that suggested a nemegtosaurid affinity. The skull reconstruction presented here shows that the skull of B. salgadoi is anteroposteriorly short and dorsoventrally high, contrasting with the elongate skull of Rapetosaurus krausei.
The dinosaur record of the Salitral Moreno locality (Río Negro Province, Argentina) is characterized by a high diversity of herbivore taxa, among them hadrosaurs, ankylosaurs, and titanosaur sauropods, but carnivores are rare, consisting of only a few fragmentary bones of small forms. Titanosaurs are represented by Rocasaurus muniozi and Aeolosaurus sp., and at least four other taxa, represented by fragmentary material. The elements preserved include a cervical, dorsal and caudal vertebrae, chevron, humerii, ulnae, radii, metacarpal, femora, tibiae, metatarsal, ischia, pubis, and ilium. The Allen Formation is thought to be correlated with the Marília Formation in Brazil, and their faunas have certain elements in common such as aeolosaurines, but saltasaurines and hadrosaurs, are known exclusively from the Allen Formation. These absences, and particularly that of the saltasaurines, may be because those sauropods originated late in the Cretaceous, probably in southern South America (Northern Patagonia?), and they did not have time to disperse to northern South America.
Exceptionally well−preserved embryonic skulls of Upper Cretaceous (Campanian Anacleto Formation) sauropods from Auca Mahuevo (Neuquén Province, Argentina) provide important insights into the ontogeny and evolution of titanosaurian neosauropods. The most important cranial modifications occurring during titanosaurian ontogeny appear to be centered on the infraorbital and narial regions, which exhibit a substantial degree of “mosaic” evolution. On one hand, the Auca Mahuevo embryos show a large jugal that forms part of the lower margin of the skull and unretracted external nares, as indicated by the position and orientation of the lacrimals as well as the anterior extension of the frontals. Both of these features are ancestral for neosauropods, being present in prosauropods. On the other hand, the embryonic skull exhibits a large ventral notch, tentatively interpreted as homologous to the neosauropod preantorbital fenestra, that opens ventral to the jugal and between the maxilla and the quadratojugal, and a temporal region that closely resembles the adult neosauropod condition. This mosaic of character states indicates that different regions of the skull of titanosaurian neosauropods acquired their characteristic morphology at substantially different rates during their ontogenetic development.
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