A redescription of Brouffia orientalis Carroll & Baird, 1972 from the Upper Carboniferous of the Czech Republic and the status and affinities of protorothyridid amniotes

Abstract

The Upper Carboniferous protorothyridid amniote Brouffia orientalis from Czech Republic is redescribed. Photogrammetric scanning of the holotype and only known specimen yields considerable new information on the skull and postcranium of this tetrapod and allows us to amend previous morphological descriptions to a substantial degree. A virtual 3D model built from photogrammetry scan data is used as the basis for a new reconstruction of the skull in dorsal, lateral, and ventral aspects and the lower jaw in lateral aspect. We expand and refine the diagnosis of Brouffia and compare it with other protorothyridids. We discuss the affinities of this taxon by coding it in a recently published data matrix of early amniotes, which we subject to maximum parsimony and Bayesian fossilized birth–death analyses. Brouffia emerges as the sister taxon to Coelostegus in all analyses, but the position of these two taxa within amniotes varies. In a parsimony analysis with unweighted characters, the (Brouffia + Coelostegus) clade forms the sister group to Synapsida. In various experiments of character reweighting, that clade is placed crownward of Captorhinidae on the stem-group of Reptilia, but anticrownward of remaining protorothyridids. The latter constitute either a paraphyletic array relative to Diapsida or their monophyletic sister group. The Bayesian analysis retrieves (Brouffia + Coelostegus) as the most basal plesion on the stem-group of Reptilia.