New information on the pterosaur Tupandactylus imperator, with comments on the relationships of Tapejaridae

A new specimen of Tupandactylus imperator, comprising an incomplete skull with associated lower jaw, is described. The material is the best preserved specimen of this species known so far and provides new information on the anatomy of this pterodactyloid pterosaur, especially with respect to the morphology of the lower jaw, the first one formally described for the species. Also, the new specimen shows an extensive preservation of soft tissues such as the soft-tissue component of the headcrest, ramphoteca associated with the premaxillae and lower jaw, as well as probable pycnofibres. A phylogenetic analysis was performed in order to test the relationships of the taxon within Tapejaridae. The results of the analysis support Tapejaridae, as well as monophyly of Tapejarinae and Thalassodrominae.