Basal tetrapods display a wide spectrum of vertebral centrum morphologies that can be used to distinguish different tetrapod groups. The vertebral types range from multipartite centra in stem-tetrapods, temnospondyls, and seymouriamorphs up to monospondylous centra in lepospondyls and have been drawn upon for reconstructing major evolutionary trends in tetrapods that are now considered textbook knowledge. Two modes of vertebral formation have been postulated: the multipartite vertebrae formed first as cartilaginous elements with subsequent ossification. The monospondylous centrum, in contrast, was formed by direct ossification without a cartilaginous precursor. This study describes centrum morphogenesis in basal tetrapods for the first time, based on bone histology. Our results show that the intercentra of the investigated stem-tetrapods consist of a small band of periosteal bone and a dense network of endochondral bone. In stereospondyl temnospondyls, high amounts of calcified cartilage are preserved in the endochondral trabeculae. Notably, the periosteal region is thickened and highly vascularized in the plagiosaurid stereospondyls. Among “microsaur” lepospondyls, the thickened periosteal region is composed of compact bone and the notochordal canal is surrounded by large cell lacunae. In nectridean lepospondyls, the periosteal region has a spongy structure with large intertrabecular spaces, whereas the endochondral region has a highly cancellous structure. Our observations indicate that regardless of whether multipartite or monospondylous, the centra of basal tetrapods display first endochondral and subsequently periosteal ossification. A high interspecific variability is observed in growth rate, organization, and initiation of periosteal ossification. Moreover, vertebral development and structure reflect different lifestyles. The bottom-dwelling Plagiosauridae increase their skeletal mass by hyperplasy of the periosteal region. In nectrideans, the skeletal mass decreases, as the microstructure is spongy and lightly built. Additionally, we observed that vertebral structure is influenced by miniaturization in some groups. The phylogenetic information that can be drawn from vertebral development, however, is limited.

The histology of the vertebral centrum of the morphologically diverse Temnospondyli is poorly known. In this study, the variability of the histological framework of various taxa from several Permian and Triassic localities was investigated for the first time. Twelve intercentra, forming the anterior part of the diplospondylous centra of temnospondyls, were examined histologically. The intercentra of all studied taxa share a highly vascularized cortex on the ventro-lateral side and primary and secondary trabecular, endochondral bone on the dorsal side. A high variability is present, among others, within the arrangement of the vascular cavities, the density of the trabeculae and the distribution of calcified cartilage. The Stereospondyli possess a high amount of calcified cartilage between the trabeculae, in all other taxa the calcified cartilage covers only the dorsal surface of the intercentrum. Among the plagiosaurids, despite morphological similarities, the intercentra show a different development. In Gerrothorax and Plagiosaurus, periosteal bone is also present on the dorsal side around the neural canal, tentatively indicating a fusion of pleurocentrum and intercentrum. The different histological framework of the investigated intercentra may indicate the phylogenetic value of intercentra microstructure, however further studies are necessary. The preservation of calcified cartilage between the trabeculae seems to be a paedomorphic character typical for all Stereospondyli.

A variety of vertebral centrum morphologies have evolved within early tetrapods which range from multipartite centra consisting of intercentra and pleurocentra in stem-tetrapods, temnospondyls, seymouriamorphs, and anthracosaurs up to monospondylous centra in lepospondyls. With the present study, we aim to determine the formation of both intercentrum and pleurocentrum and asked whether these can be homologized based on their bone histology. Both intercentra and pleurocentra ossified endochondrally and periosteal bone was subsequently deposited on the outer surface of the centra. Our observations indicate low histological variation between intercentrum and pleurocentrum in microstructural organization and growth which inhibits the determination of homologies. However, intercentrum and pleurocentrum development differs during ontogeny. As previously assumed, the intercentrum arises from ventrally located and initially paired ossification centers that fuse ventromedially to form the typical, crescentic, rhachitomous intercentrum. In contrast, presacral pleurocentra may be ancestrally represented by four ossification centers: a ventral and a dorsal pair. Subsequently, two divergent developmental patterns are observed: In stem-tetrapods and temnospondyls, the pleurocentrum evolves from the two dorsally located ossification centers which may occasionally fuse to form a dorsal crescent. In some dvinosaurian temnospondyls, the pleurocentrum may even ossify to full rings. In comparison, the pleurocentrum of stem-amniotes (anthracosaurs, chroniosuchids, seymouriamorphs, and lepospondyls) arises from the two ventrally located ossification centers whereby the ossification pattern is almost identical to that of temnospondyls but mirror-inverted. Thus, the ring-shaped pleurocentrum of Discosauriscus ossifies from ventral to dorsal. We also propose that the ossified portions of the intercentrum and pleurocentrum continued as cartilaginous rings or discs that surrounded the notochord in the living animals.

The origin of lissamphibians remains unresolved and different origins within Paleozoic early tetrapods have been proposed. Both the vertebral morphology as well as the ossification sequence of the vertebrae are potentially important character traits to test these different hypotheses. Here, we aim to determine if vertebral formation and ossification sequence in lissamphibians and early tetrapods are valuable indicatives of phylogenetic relationships. To answer this question, we examined the vertebral development in growth series of different salamander taxa and compared the results with literature data on vertebral development in frogs, caecilians and early tetrapods. In salamanders, caecilians and early tetrapods, the vertebral centrum develops by chordacentral and perichordal centrum formation, whereas in frogs, no chordacentral centrum formation has been observed so far. In the salamander taxa studied here, the following chondrification and ossification sequences are observed: chondrification first of the neural arches and then of the centra followed by ossification first of the centra and then of the neural arches. In frogs, in contrast, the neural arches ossify prior to the centra and it can be assumed that this developmental sequence represents the ancestral condition for tetrapods, as it characterizes some stem‐tetrapods and all temnospondyls from which growth series are known. However, the ossification sequence of the vertebral column is only incompletely known in stem‐amniotes which makes a comparison with extant lissamphibians difficult. This indicates that the mode of centrum formation and the ossification sequence are highly variable and cannot be used to determine the origin of lissamphibians within early tetrapods.

The basal tetrapod Solenodonsaurus janenschi Broili, 1924, from Nýřany (Westphalian D, Late Carboniferous), Czech Republic, is redescribed and its phylogenetic position reevaluated. A distinct groove at the base of the maxillary teeth is regarded as an autapomorphic character, which is present in both the large and small specimens. Other characteristic features, which are not unique to S. janenschi, are: an extension of the lacrimal that forms the anteroventral margin of the orbit; a long posterior extension of the jugal; spool-shaped vertebrae, and small, wedge-like intercentra. A phylogenetic analysis based on the data matrix of Ruta, Coates and Quicke suggests that S. janenschi is the sister taxon of the Lepospondyli. Shared characters include the shape of the vertebrae, non-swollen neural arches, and absence of an intertemporal. Although nested within the amniote stem, S. janenschi is not as closely related to basal amniotes as previously suggested. A rather long, slender humerus argues for a predominantly terrestrial mode of life, and the curved, slender ribs, as well as the comparatively small skull, suggest costal ventilation of the lungs similar to that in amniotes, rather than buccal pumping. The morphology of the shallow squamosal embayment in which an ossified dorsal margin is absent, renders the presence of a tympanum unlikely.