We describe a new specimen of the aïstopod Oestocephalus from Five Points, Ohio, which preserves much of the posterior braincase. The specimen, the largest aïstopod skull described, preserves the postorbital region to the occiput. The posterior braincase has coossified the basioccipital, exoccipitals, and opisthotic. The parasphenoid is rostrally restricted, toothless, and highly vaulted along the cultriform process. The lateral walls of the cultriform process are further reinforced by large longitudinally running, ventral flanges from the parietals. Two large endochondral ventral projections from the basioccipital, previously interpreted as basal tuberosities for hypaxial muscle insertion, are here instead interpreted as articulations for the branchial skeleton. This interpretation is further supported by traces of vasculature that is consistent with what is seen in gill-bearing species. A model for the reorganisation of the basicranial region on the transition from hyomandibula to stapes is proposed, which suggests that gills, or gill-support skeletal elements, might be further distributed along the tetrapod stem than previously thought. These data further support the placement of aïstopods in the tetrapod stem group and require a reconsideration of our understanding of early tetrapod evolution.

The morphologies of the hyobranchial apparatus in early tetrapods are reviewed, based primarily on first-hand examination and supplemented by published descriptions. The basic arrangement of the “aquatic” hyobranchium, with four pairs of branchial arches and internal gills, was conserved to a remarkable degree across the fish–to–tetrapod transition and was retained in further evolution in adults of several tetrapod lineages. Thus, a fish-like hyobranchium in basal tetrapods does not necessarily represent a larval or paedomorphic character, respectively, as was often suggested in analogy to extant salamanders. Rather, it represents the plesiomorphic state of the adult hyobranchium in tetrapods. The changes in the hyobranchium during the fish–to–tetrapod transition include the reduction of the number of skeletal elements and their morphological simplification. In all three presently discussed scenarios of lissamphibian origin, the temnospondyl, lepospondyl and diphyly hypotheses, the internal gills were reduced independently within temnospondyls and on the amniote stem below seymouriamorphs. Evidence of remodelling into a true “terrestrial” hyobranchium, with reduction of the posterior branchial arches and modification to support terrestrial tongue feeding, is scarce in early tetrapods. It evolved within temnospondyls in zatracheids, amphibamids and lissamphibians, as well as once or several times in early amniotes or in their immediate stem-forms.