The Presence of Ankylosaur Tracks in the Guará Formation (Brazil) and Remarks on the Spatial and Temporal Distribution of Late Jurassic Dinosaurs

The Guara Formation (Parana Basin, southern Brazil) is an Upper Jurassic unit that yielded a dinosaur ichnoassemblage composed of theropod, ornithopod, and sauropod tracks. A new set of footprints is described herein and its major features are heteropody, paraxony, and both manual and pedal tetradactyly, among others. Using ichnological, osteological, and stratigraphic approaches, we interpret these tracks as produced by an ankylosaur dinosaur. The record of these armored dinosaurs in South America is scarce and restricted to the Cretaceous units of Argentina, Bolivia, and Brazil. Therefore, the presence of these tracks in the Guara Formation provides the oldest evidence of ankylosaurs in western Gondwana and the first uncontroversial record of this group in Brazil. In addition, a comparison between the Guara Formation fossil record and other Kimmeridgian-Tithonian dinosaur-bearing units worldwide indicates that more efforts are needed to better understand the geographical distribution of Late Jurassic dinosaurs.