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.

The scant fossil record of caecilians has obscured the origin and evolution of this lissamphibian group. Eocaecilia micropodia from the Lower Jurassic of North America remains the only stem-group caecilian with an almost complete skull preserved. However, this taxon has been controversial, engendering re-evaluation of traits considered to be plesiomorphic for extant caecilians. Both the validity of the placement of E. micropodia as a stem caecilian and estimates of the plesiomorphic condition of extant caecilians have been questioned. In order to address these issues, the braincase of E. micropodia was examined via micro-computed tomography. The braincase is considered to be a more reliable phylogenetic indicator than peripheral regions of the skull. These data reveal significant new information, including the possession of an ossified nasal septum, ossified anterior wall of the sphenethmoid, long anterolateral processes on the sphenethmoid, and paired olfactory nerve foramina, which are known only to occur in extant caecilians; the latter are possibly related to the evolution of the tentacle, a caecilian autapomorphy. A phylogenetic analysis that included 64 non-amniote taxa and 308 characters represents the first extensive test of the phylogenetic affinities of E. micropodia. The results place E. micropodia securely on the stem of extant caecilians, representing a clade within Temnospondyli that is the sister taxon to batrachians plus Gerobatrachus. Ancestral character state reconstruction confirms the braincase of E. micropodia to be largely representative of the plesiomorphic condition of extant caecilians. Additionally, the results refine the context within which the evolution of the caecilian form can be evaluated. The robust construction and pattern of the dermal skull of E. micropodia is interpreted as symplesiomorphic with advanced dissorophoid temnospondyls, rather than being autapomorphic in its robust construction. Together these data increase confidence in incorporating E. micropodia into discussions of caecilian evolution.

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Caecilian morphology is strongly modified in association with their fossorial mode of life. Currently phylogenetic analyses of characters drawn from the morphology of caecilians lack resolution, as well as complementarity, with results of phylogenetic analyses that employ molecular data. Stemming from the hypothesis derived from the mammal literature that the braincase has the greatest potential (in comparison to other cranial units) to yield phylogenetic information, the braincase and intimately associated stapes of 27 species (23 genera) of extant caecilians were examined using images assembled via microcomputed tomography. Thirty-four new morphological characters pertaining to the braincase and stapes were identified and tested for congruence with previously recognized morphological characters. The results reveal that when added to previous character matrices, characters of the braincase and stapes resolve generic-level relationships in a way that is largely congruent with the results of molecular analyses. Analysis of a combined data set of molecular and morphological data provides a framework for conducting ancestral character state reconstructions, which resulted in the identification of 95 new synapomorphies for various clades and taxa, 27 of which appear to be unique for the taxa that possess them. Together these data demonstrate the utility of the application of characters of the braincase and stapes for resolving phylogenetic relationships for a group whose morphology is largely confounded by functional modifications. In addition this study provides evidence of the utility of the braincase in resolving problematic morphologybased phylogeny outside of Amniota, in an amphibian group.

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Bones of the cranial vault appear to be highly conserved among tetrapod vertebrates. Moreover, bones identified with the same name are assumed to be evolutionarily homologous. However, recent developmental studies reveal a key difference in the embryonic origin of cranial vault bones between representatives of two amniote lineages, mammals and birds, thereby challenging this view. In the mouse, the frontal is derived from cranial neural crest (CNC) but the parietal is derived from mesoderm, placing the CNC–mesoderm boundary at the suture between these bones. In the chicken, this boundary is located within the frontal. This difference and related data have led several recent authors to suggest that bones of the avian cranial vault are misidentified and should be renamed. To elucidate this apparent conflict, we fate-mapped CNC and mesoderm in axolotl to reveal the contributions of these two embryonic cell populations to the cranial vault in a urodele amphibian. The CNC–mesoderm boundary in axolotl is located between the frontal and parietal bones, as in the mouse but unlike the chicken. If, however, the avian frontal is regarded instead as a fused frontal and parietal (i.e. frontoparietal) and the parietal as a postparietal, then the cranial vault of birds becomes developmentally and topologically congruent with those of urodeles and mammals. This alternative hypothesis of cranial vault homology is also phylogenetically consistent with data from the tetrapod fossil record, where frontal, parietal and postparietal bones are present in stem lineages of all extant taxa, including birds. It further implies that a postparietal may be present in most non-avian archosaurs, but fused to the parietal or supraoccipital as in many extant mammals.

High levels of morphological homoplasy have hindered progress in understanding morphological evolution within gymnophione lissamphibians. Stemming from the hypothesis that the braincase has the potential to yield phylogenetic information, the braincases of 27 species (23 genera) of gymnophione amphibians were examined using high-resolution micro-computed tomography and histologically prepared specimens. Morphology of the brain and its relationship to features of the braincase is described, and it is shown that eight different patterns exist in the distribution of foramina in the antotic region. The distribution of variants is congruent with molecule-based phylogeny. Additionally, all variants are shown to correspond directly to stages along developmental continua, suggesting that the evolutionary truncation of development in the antotic region at various stages has driven the evolution of morphology in this region. Attempts to correlate the observed morphology with proxies of putative heterochronic events (including those attributable to burrowing, life history, and size) fail to explain the distribution of morphology if each proxy is considered separately. Thus, it is concluded that either currently unrecognized causes of heterochrony or combinations thereof have influenced morphology in different lineages independently. These data identify clades whose morphology can now be reconsidered in light of previously unrecognized heterochronic events, thereby providing a foundation for future analyses of the evolution of morphology within Gymnophiona as a whole. Most significantly, these data confirm, for the first time in a lissamphibian group, that the braincase can preserve important phylogenetic information that is otherwise obscured in regions of the skull that experience strong influences from functional constraints.