Phylogenetic relationships of the European trilophosaurids Tricuspisaurus thomasi and Variodens inopinatus

Trilophosauridae are an extinct group of archosauromorphs from the Upper Triassic of western North America and Nova Scotia. Possible trilophosaurids Tricuspisaurus thomasi and Variodens inopinatus have also been reported from Upper Triassic fissure fills in the southwest of the United Kingdom, based on their multi-cuspid dentitions. Here, we provide an updated description of V. inopinatus and add to the description of T. thomasi, including re-examining CT scans of its holotype dentary. We confirm that the apparent gap on the dentary of T. thomasi is pathological, with the dentine of the teeth on either side of the gap asymmetrically thick, and that heavy vascularization of the anterior part of the dentary confirms it likely had a keratinous rhamphotheca. Our cladistic analyses, to which we add nine new characters, confirm that T. thomasi and V. inopinatus are closely related taxa and the youngest known trilophosaurids. Further, we support a previous suggestion that the relatively small size of these two European trilophosaurids results from insular dwarfism.