Osteology, relationships and functional morphology of Weigeltisaurus jaekeli (Diapsida, Weigeltisauridae) based on a complete skeleton from the Upper Permian Kupferschiefer of Germany

Abstract

Background Weigeltisauridae is a clade of small-bodied diapsids characterized by a horned cranial frill, slender trunk and limbs, and a patagium supported by elongated bony rods. Partial skeletons and fragments are definitively known only from upper Permian (Lopingian) rocks in England, Germany, Madagascar and Russia. Despite these discoveries, there have been few detailed descriptions of weigeltisaurid skeletons, and the homologies of many skeletal elements—especially the rods supporting the patagium—remain the subject of controversy. Materials & Methods Here, we provide a detailed description of a nearly complete skeleton of Weigeltisaurus jaekeli from the upper Permian (Lopingian: Wuchiapingian) Kupferschiefer of Lower Saxony, Germany. Briefly addressed by past authors, the skeleton preserves a nearly complete skull, postcranial axial skeleton, appendicular skeleton, and patagial supports. Through comparisons with extant and fossil diapsids, we examine the hypotheses for the homologies of the patagial rods. To examine the phylogenetic position of Weigeltisauridae and characterize the morphology of the clade, we integrate the material and other weigeltisaurids into a parsimony-based phylogenetic analysis focused on Permo-Triassic non-saurian Diapsida and early Sauria (61 taxa, 339 characters). Results We recognize a number of intriguing anatomical features in the weigeltisaurid skeleton described here, including hollow horns on the post-temporal arch, lanceolate teeth in the posterior portion of the maxilla, the absence of a bony arch connecting the postorbital and squamosal bones, elongate and slender phalanges that resemble those of extant arboreal squamates, and patagial rods that are positioned superficial to the lateral one third of the gastral basket. Our phylogenetic study recovers a monophyletic Weigeltisauridae including Coelurosauravus elivensis, Weigeltisaurus jaekeli, and Rautiania spp. The clade is recovered as the sister taxon to Drepanosauromorpha outside of Sauria (=Lepidosauria + Archosauria). Conclusions Our anatomical observations and phylogenetic analysis show variety of plesiomorphic diapsid characters and apomorphies of Weigeltisauridae in the specimen described here. We corroborate the hypothesis that the patagial ossifications are dermal bones unrelated to the axial skeleton. The gliding apparatus of weigeltisaurids was constructed from dermal elements unknown in other known gliding diapsids. SMNK-PAL 2882 and other weigeltisaurid specimens highlight the high morphological disparity of Paleozoic diapsids already prior to their radiation in the early Mesozoic.