The skull of the titanosaur Tapuiasaurus macedoi (Dinosauria: Sauropoda), a basal titanosaur from the Lower Cretaceous of Brazil

Abstract

Although Titanosauria is the most diverse and late-surviving sauropod lineage, cranial elements are known for just over 24 of its 70+ genera – the vast majority of which are fairly fragmentary and restricted to the Late Cretaceous. Only three complete titanosaur skulls have been described to date; two of these are from the latest Cretaceous (Nemegtosaurus, Rapetosaurus), and the third, Tapuiasaurus, is from the Early Cretaceous (Aptian). In this contribution, we build on the initial treatment of the taxon by providing a complete description of the cranial elements that benefits from additional preparation and computed tomography imaging. We identify six additional features diagnosing Tapuiasaurus macedoi, including a jugal with an elongate lacrimal process forming much of the posteroventral border of the antorbital fenestra, a lateral temporal fenestra divided by a second squamosal–postorbital contact, and upper jaw teeth with labial wear facets. We directed the new morphological data in Tapuiasaurus as well as other observations towards a re-analysis of its phylogenetic position within Titanosauria. Our analysis yielded 34 most parsimonious trees, most of which recovered Tapuiasaurus in a basal position adjacent to the Early Cretaceous taxa Malawisaurus and Tangvayosaurus, but two trees recovered it within Late Cretaceous nemegtosaurids. We explored the effects of missing data and missing stratigraphic ranges on our results, concluding that (1) when missing data levels are high, resolution of even small amounts of that missing data can have dramatic effects on topology, (2) taxa that are mostly scored for characters that cannot be scored in other taxa may be topologically unstable and (3) there were several suboptimal trees that had greatly improved stratigraphic fit with relatively little compromise in terms of tree length.