Anatomy of the basal titanosaur (Dinosauria, Sauropoda) Andesaurus delgadoi from the mid-Cretaceous (Albian-early Cenomanian) Río Limay Formation, Neuquén Province, Argentina: implications for titanosaur systematics

Titanosauria is a taxonomically and morphologically diverse clade of sauropod dinosaurs that appeared in the Middle Jurassic and radiated in the mid-Late Cretaceous; however, its intrarelationships are poorly understood. The mid-Cretaceous Argentinean sauropod Andesaurus delgadoi has repeatedly been recovered at the base of Titanosauria, and thus represents a crucial taxon for determining the evolutionary history of this clade; yet it has only received a brief description. Here, we re-describe the holotype, comprising dorsal, sacral, and caudal vertebrae, as well as limb and pelvic elements. Detailed comparisons are made with a global array of titanosauriforms. Andesaurus is a valid genus and can be diagnosed by five autapomorphies: (1) posterior dorsal neural spine height greater than twice centrum height (autapomorphic within Macronaria); (2) square-shaped anterior-middle caudal centra in lateral view; (3) anteroposteriorly elongate fossa present on the anterodorsal corner of the lateral surface of middle-posterior caudal centra; (4) ridge along the midshaft of the ventral surface of metacarpal I, close to the ventromedial margin; (5) prominent ventromedial ridge along the distal half of metacarpal V. Other remains previously attributed to Andesaurus cannot be referred to this genus. Sixteen putative titanosaur synapomorphies can be recognized in Andesaurus, including: (1) lateral pneumatic foramina in dorsal vertebrae situated within fossae; (2) anterior-middle caudal vertebrae with ventrolateral ridges either side of a ventral midline hollow; and