Phylogenetic analysis of early tetrapod evolution has resulted in a consensus across diverse data sets in which the tetrapod stem group is a relatively homogenous collection of medium- to large-sized animals showing a progressive loss of 'fish' characters as they become increasingly terrestrial, whereas the crown group demonstrates marked morphological diversity and disparity. The oldest fossil attributed to the tetrapod crown group is the highly specialized aïstopod Lethiscus stocki, which shows a small size, extreme axial elongation, loss of limbs, spool-shaped vertebral centra, and a skull with reduced centres of ossification, in common with an otherwise disparate group of small animals known as lepospondyls. Here we use micro-computed tomography of the only known specimen of Lethiscus to provide new information that strongly challenges this consensus. Digital dissection reveals extremely primitive cranial morphology, including a spiracular notch, a large remnant of the notochord within the braincase, an open ventral cranial fissure, an anteriorly restricted parasphenoid element, and Meckelian ossifications. The braincase is elongate and lies atop a dorsally projecting septum of the parasphenoid bone, similar to stem tetrapods such as embolomeres. This morphology is consistent in a second aïstopod, Coloraderpeton, although the details differ. Phylogenetic analysis, including critical new braincase data, places aïstopods deep on the tetrapod stem, whereas another major lepospondyl lineage is displaced into the amniotes. These results show that stem group tetrapods were much more diverse in their body plans than previously thought. Our study requires a change in commonly used calibration dates for molecular analyses, and emphasizes the importance of character sampling for early tetrapod evolutionary relationships.

The Early Permian recumbirostran lepospondyl Rhynchonkos stovalli has been identified as a possible close relative of caecilians due to general similarities in skull shape as well as similar robustness of the braincase, a hypothesis that implies the polyphyly of extant lissamphibians. In order to better assess this phylogenetic hypothesis, we studied the morphology of the holotype and three specimens previously attributed to R. stovalli. With the use of micro-computed x-ray tomography (μCT) we are able to completely describe the external and internal cranial morphology of these specimens, dramatically revising our knowledge of R. stovalli and recognizing two new taxa, Aletrimyti gaskillae gen et sp. n. and Dvellacanus carrolli gen et sp. n. The braincases of R. stovalli, A. gaskillae, and D. carrolli are described in detail, demonstrating detailed braincase morphology and new information on the recumbirostran supraoccipital bone. All three taxa show fossorial adaptations in the braincase, sutural articulations of skull roof bones, and in the lower jaw, but variation in cranial morphology between these three taxa may reflect different modes of head-first burrowing behaviors and capabilities. We revisit the homology of the supraoccipital, median anterior bone, and temporal bone of recumbirostrans, and discuss implications of alternate interpretations of the homology of these elements. Finally, we evaluate the characteristics previously used to unite Rhynchonkos stovalli with caecilians in light of these new data. These proposed similarities are more ambiguous than previous descriptions suggest, and result from the composite nature of previous descriptions, ambiguities in external morphology, and functional convergence between recumbirostrans and caecilians for head-first burrowing.

Recumbirostran ‘microsaurs,’ a group of early tetrapods from the Late Carboniferous and Early Permian, are the earliest known example of adaptation to head-first burrowing in the tetrapod fossil record. However, understanding of the diversity of fossorial adaptation within the Recumbirostra has been hindered by poor anatomical knowledge of the more divergent forms within the group. Here we report the results of μCT study of Quasicaecilia texana, a poorly-known recumbirostran with a unique, broad, shovel-like snout. The organization of the skull roof and braincase of Quasicaecilia is found to be more in line with that of other recumbirostrans than previously described, despite differences in overall shape. The braincase is found to be broadly comparable to Carrolla craddocki, with a large presphenoid that encompasses much of the interorbital septum and the columella ethmoidalis, and a single compound ossification encompassing the sphenoid, otic, and occipital regions. The recumbirostran braincase conserves general structure and topology of braincase regions and cranial nerve foramina, but it is highly variable in the number of ossifications and their extent, likely associated with the reliance on braincase ossifications to resist compression during sediment compaction and mechanical manipulation by epaxial and hypaxial musculature. Expansion of the deep ventral neck musculature in Quasicaecilia, autapomorphic among recumbirostrans, may reflect unique biomechanical function, and underscores the importance of future attention to the role of the cervical musculature in contextualizing the origin and evolution of fossoriality in recumbirostrans.

Morphological data sets are misleading in salamander (Caudata) phylogeny due to the relative homoplasy of the dermal skull observed in paedomorphic forms, leading to the trend of excluding morphology when exploring questions of salamander phylogeny. Investigations in caecilians (Gymnophiona) have demonstrated that the inclusion of braincase morphology can rescue morphological phylogenetic analyses and produce topologies congruent with molecular data sets. We scanned 28 species (25 genera) of salamander, representing all 10 families, with high‐resolution micro‐computed tomography to investigate braincase variation. We describe the morphology of the braincase for all 10 families and distinguish between paedomorphic and metamorphic morphologies. Our results demonstrate a general uniformity amongst metamorphic species with variation largely restricted to the occipito‐otic region. A greater range of variation is observed within paedomorphic forms than would be expected when considering the homoplasy of the dermal skull. Obligate paedomorphic forms demonstrate considerably more variation in the anterior braincase than do facultative paedomorphs, which we suggest is evidence of a greater complexity in the evolution and development of these forms than neoteny alone would produce. This raises the question of character independence within morphological data sets and warrants further investigation into the correlation of other characters before morphological data are omitted.

Christensen et al. recently published a study of hearing in neotenic and experimentally metamorphosed axolotls (Ambystoma mexicanum) and a larval and adult tiger salamander (A. tigrinum), which contributes greatly to our understanding of salamander sound perception in water and air [1]. They demonstrate that premetamorphic, atympanic aquatic salamanders are capable of perceiving high frequency airborne sounds similar to terrestrial adults, demonstrating an interesting capacity for an atympanic ear to function for sound perception in a variety of media. This important research, in combination with other recent studies on the sound perception capabilities of lungfish [2], is helping to better clarify issues related to the water-to-land transition. However, the implication of their experimental design demands a consideration of the evolutionary history of their selected exemplar taxa so these valuable data can be interpreted in a broader context. Our criticisms focus on two major points: the use of salamanders as a proxy for hearing ability intermediate between aquatic and terrestrial vertebrates, and the use of microsaurs as exemplars of the early tetrapod condition.The currently accepted hypothesis for the origin of a tympanic ear is as a derivation from the spiracle of sarcopterygian fish [3]. In 'fish' the spiracle is associated with gill breathing: water passes through the spiracular notch in the rear of the skull, past the braincase and the bracing hyomandibula, to the oropharynx and gill arches [4]. With the loss of gills, the persisting spiracular lumen is co-opted into a middle ear chamber, and the cranial support role of the hyomandibula (now termed stapes) is eliminated, leaving this bone to extend freely into the middle ear chamber (figure 1). In derived tetrapods, the spiracular opening is expanded and covered by a membrane (the tympanum) that collects sound. Vibrations are transmitted to the inner ear via the stapes, which becomes more delicate and better able to propagate fine vibrations [7][8][9][10].