A survey of skeletal elements in the orbit of Pseudosuchia and the origin of the crocodylian palpebral

Ossified skeletal elements within the orbit, such as scleral ossicles, palpebrals, supraorbitals and sesamoids, are widespread across Reptilia, including extant members of Crocodylia, Aves, Squamata and Chelonia. Extant crocodylians lack scleral ossicles, but have a unique palpebral that has recently been shown, through developmental studies, to be an osteoderm in the upper eyelid. Here, we examine the diversity and disparity of the crocodylian palpebral in extant members of Crocodylia and, through the fossil record, trace the origin and evolutionary history of the element throughout Pseudosuchia (all archosaurs more closely related to crocodylians than avians). We show that the crocodylian palpebral originated outside of Crocodylomorpha in early pseudosuchian groups (e.g. Aetosauria, Loricata) and that scleral ossicles are lost in nearly all groups of pseudosuchians, but appear in poposauroids, at least one early crocodylomorph, and some partially or fully marine Crocodyliformes (e.g. thalattosuchians). The morphology and number of palpebrals differs across Crocodyliformes; the presence of two palpebrals is plesiomorphic, but this is reduced to one by Crocodylia. We further recommend the restriction of the term palpebral to the structure in crocodylians and their homologues, but not to the structure in ornithischian dinosaurs.