noting that spermaceti is more likely to be used during aggression or for sonar. They agreed that the idea of the crest having an in-flight function was tempting and sideways movement of the head would have helped it change direction, but biomechanical and flight-mechanical studies of the crest would have to be conducted to determine the animal's aerodynamics. Witton also expressed hope for further analysis of thalassodromid locomotion. He noted that since their limb proportions were similar to those of the better-studied azhdarchids, the shape of their wings and style of flight might have been similar. Thalassodromids might also have been adapted for inland flight; their wings were short and broad (unlike the long, narrow wings of marine soarers), and were more manoeuvrable and less likely to snag on obstacles. Their lower shoulder muscles appear to have been enlarged, which would have helped with powerful (or frequent) wing downstrokes and takeoff ability. Although it may have had to compensate for its large crest during flight, its development late in growth indicates that it did not develop primarily for aerodynamics. Witton suggested that the proportional similarity between the limbs of thalassodromids and azhdarchids also indicates that their terrestrial abilities would have been comparable. Their limbs would have been capable of long strides, and their short, compact feet would have made these mechanics efficient. The enlarged shoulder muscles may have allowed them to accelerate quickly when running, and they may have been as adapted for movement on the ground as has been suggested for azhdarchids; Witton cautioned that more analysis of thalassodromids was needed to determine this. Palaeoecology Thalassodromeus is known from the Romualdo Formation, which dates to the Albian stage of the Early Cretaceous period (about 110million years ago). The formation is part of the Santana Group and, at the time Thalassodromeus was described, was thought to be a member of what was then considered the Santana Formation. The Romualdo Formation is a Lagerstätte (a sedimentary deposit that preserves fossils in excellent condition) consisting of lagoonal limestone concretions embedded in shales, and overlies the Crato Formation. It is well known for preserving fossils three-dimensionally in calcareous concretions, including many pterosaur fossils. As well as muscle fibres of pterosaurs and dinosaurs, fish preserving gills, digestive tracts, and hearts have been found there. The formation's tropical climate largely corresponded to today's Brazilian climate. Most of its flora were xerophytic (adapted to dry environments).