A redescription of Carrolla craddocki (Lepospondyli: Brachystelechidae) based on high-resolution CT, and the impacts of miniaturization and fossoriality on morphology

Abstract

Some recent morphological analyses have brought into question the monophyly of Lissamphibia (frogs, salamanders, and caecilians). In these analyses, brachystelechid "microsaurs" are found to be sister group to caecilians. To test this hypothesis, the holotype specimen of the brachystelechid Carrolla craddocki was submitted to high-resolution X-ray computed tomography to gain insight into the nature of the morphology supporting the potential relationship between brachystelechids and caecilians. This analysis enabled us to conduct a detailed description of the internal anatomy such as the braincase and otic capsule endocast (the first of its kind for a lepospondyl), and new information regarding the architecture of the skull. Our results suggest brachystelechid cranial morphology is strongly influenced by miniaturization (enlarged sensory organs, anterior placement of the jaw articulation, and combination of both reduced- and hyper-ossifications) and burrowing habits (co-ossified braincase with broad, sloping occipital surface, overlapping joints between skull roof bones, and well-ossified anterior braincase). Characteristics of brachystelechids that appear unrelated to size-reduction and burrowing are the diamond-shaped skull and possible pedicellate dentition. We provide a revised diagnosis for Carrolla and identify possible new characters within the anatomy of the braincase and inner ear. Several characters currently uniting caecilians and "microsaurs" are among those associated with either miniaturization or burrowing, demonstrating that future efforts should continue to focus on fine details of anatomy minimally affected by these influences to contribute to the resolution of the question of the origin of caecilians.