Lacertoid tracks from the Botucatu Formation (Lower Cretaceous) with different locomotor behaviors: A new trackmaker with novel paleoecological implications

The Botucatu Formation (Paraná Basin) is an eolian deposit that represents an extensive paleoerg that existed in the south-central Gondwana during the Early Cretaceous. The fossil record of the unit is almost exclusively constituted of ichnofossils, including numerous tracks and burrows of vertebrates and invertebrates. The greatest trace fossil diversity of the Botucatu Formation is reported at the Ouro ichnosite, located in the Araraquara municipality region, São Paulo State, Brazil. This site comprises a faunal-rich setting of greater humidity in the ancient desert. In this work, a new footprint morphotype is reported for the Ouro ichnosite. Five trackways including tracks with an elongated anteroposterior axis associated with tail drag marks are described. The morphology of the autopodia and geometry of the trackways indicate an affinity with a lizard-like trackmaker. Furthermore, at least three different modes of locomotion were recognized: a typical walking gait, a pause-walking gait, and sideways drifting. The current report suggests Lepidosauria may have inhabited the ancient Botucatu desert, a group never reported before in this geological context. This finding expands the knowledge about the Botucatu Formation paleofauna and adds more complexity to its ecological network. Squamata and Sphenodontia are plausible candidates to have produced the described fossil tracks.