Cranial functional morphology of the pseudosuchian Effigia and implications for its ecological role in the Triassic

Abstract

Pseudosuchians, archosaurian reptiles more closely related to crocodylians than to birds, exhibited high morphological diversity during the Triassic with numerous examples of morphological convergence described between Triassic pseudosuchians and post-Triassic dinosaurs. One example is the shuvosaurid Effigia okeeffeae which exhibits an "ostrich-like" bauplan comprising a gracile skeleton with edentulous jaws and large orbits, similar to ornithomimid dinosaurs and extant palaeognaths. This bauplan is regarded as an adaptation for herbivory, but this hypothesis assumes morphological convergence confers functional convergence, and has received little explicit testing. Here, we restore the skull morphology of Effigia, perform myological reconstructions, and apply finite element analysis to quantitatively investigate skull function. We also perform finite element analysis on the crania of the ornithomimid dinosaur Ornithomimus edmontonicus, the extant palaeognath Struthio camelus and the extant pseudosuchian Alligator mississippiensis to assess the degree of functional convergence with a taxon that exhibit "ostrich-like" bauplans and its closest extant relatives. We find that Effigia possesses a mosaic of mechanically strong and weak features, including a weak mandible that likely restricted feeding to the anterior portion of the jaws. We find limited functional convergence with Ornithomimus and Struthio and limited evidence of phylogenetic constraints with extant pseudosuchians. We infer that Effigia was a specialist herbivore that likely fed on softer plant material, a niche unique among the study taxa and potentially among contemporaneous Triassic herbivores. This study increases the known functional diversity of pseudosuchians and highlights that superficial morphological similarity between unrelated taxa does not always imply functional and ecological convergence.

Keywords: Effigia; Ornithomimus; Struthio; Triassic; convergence; functional morphology; herbivory; pseudosuchian.