Cranial anatomy of Emeroleter levis and the phylogeny of Nycteroleteridae.

Among the diverse basal reptile clade Parareptilia, the nycteroleters are among the most poorly understood. The interrelationships of nycteroleters are contentious, being recovered as both monophyletic and paraphyletic in different analyses, yet their anatomy has received little attention. We utilized x-ray computed tomography to investigate the skull of the nycteroleterid Emeroleter levis, revealing aspects of both the external and internal cranial anatomy that were previously unknown or undescribed, especially relating to the palate, braincase, and mandible. Our results reveal a greater diversity in nycteroleter cranial anatomy than was previously recognized, including variation in the contribution of the palatal elements to the orbitonasal ridge among nycteroleters. Of particular note are the unique dentition patterns in Emeroleter, including the presence of dentition on the ectopterygoid, an element which is typically edentulous in most parareptiles. We then incorporate the novel information gained from the computed tomography analysis into an updated phylogenetic analysis of parareptiles, producing a fully resolved Nycteroleteridae and further supporting previous suggestions that the genus 'Bashkyroleter' is paraphyletic.