The end-Devonian to mid-Mississippian time interval has long been known for its depauperate palaeontological record, especially for tetrapods. This interval encapsulates the time of increasing terrestriality among tetrapods, but only two Tournaisian localities previously produced tetrapod fossils. Here we describe five new Tournaisian tetrapods (Perittodus apsconditus, Koilops herma, Ossirarus kierani, Diploradus austiumensis and Aytonerpeton microps) from two localities in their environmental context. A phylogenetic analysis retrieved three taxa as stem tetrapods, interspersed among Devonian and Carboniferous forms, and two as stem amphibians, suggesting a deep split among crown tetrapods. We also illustrate new tetrapod specimens from these and additional localities in the Scottish Borders region. The new taxa and specimens suggest that tetrapod diversification was well established by the Tournaisian. Sedimentary evidence indicates that the tetrapod fossils are usually associated with sandy siltstones overlying wetland palaeosols. Tetrapods were probably living on vegetated surfaces that were subsequently flooded. We show that atmospheric oxygen levels were stable across the Devonian/Carboniferous boundary, and did not inhibit the evolution of terrestriality. This wealth of tetrapods from Tournaisian localities highlights the potential for discoveries elsewhere.

Here we describe the postcranial skeleton and present the first full-body reconstruction of the early tetrapod Whatcheeria deltae from the Viséan of Iowa. The skeletal proportions, including an elongate neck and large limbs, are unlike those of other Devonian and Mississippian tetrapods. The robust limbs of Whatcheeria appear adapted for a walking gait, but the lateral lines of the cranium are fundamentally unsuited for sustained subaerial exposure. Thus, although Whatcheeria bears a general resemblance to certain terrestrially adapted Permian and Triassic members of crown tetrapod lineages, its unusual form signals a broader range of early amphibious morphologies and habits than previously considered. From the exceptionally rich collection it is evident that most Whatcheeria specimens represent immature individuals. Rare specimens suggest an adult body size of at least 2 m, over twice that of the holotype. Further comparison suggests that the Pederpes holotype might also be a juvenile and reveals a combination of hindlimb characters unique to Whatcheeria and Pederpes. These new data contribute to a revised diagnosis of the family Whatcheeriidae and a re-evaluation of fragmentary Devonian–Carboniferous fossils reported as ‘whatcheeriid’ but sharing no synapomorphies with the more precisely defined clade.

The end‐Devonian mass extinction has been framed as a turning point in vertebrate evolution, enabling the radiation of tetrapods, chondrichthyans and actinopterygians. Until very recently ‘Romer's Gap’ rendered the Early Carboniferous a black box standing between the Devonian and the later Carboniferous, but now new Tournaisian localities are filling this interval. Recent work has recovered unexpected tetrapod and lungfish diversity. However, the composition of Tournaisian faunas remains poorly understood. Here we report on a Tournaisian vertebrate fauna from a well‐characterized, narrow stratigraphic interval from the Ballagan Formation exposed at Burnmouth, Scotland. Microfossils suggest brackish conditions and the sedimentology indicates a low‐energy debris flow on a vegetated floodplain. A range of vertebrate bone sizes are preserved. Rhizodonts are represented by the most material, which can be assigned to two taxa. Lungfish are represented by several species, almost all of which are currently endemic to the Ballagan Formation. There are two named tetrapods, Aytonerpeton and Diploradus, with at least two others also represented. Gyracanths, holocephalans, and actinopterygian fishes are represented by rarer fossils. This material compares well with vertebrate fossils from other Ballagan deposits. Faunal similarity analysis using an updated dataset of Devonian–Carboniferous (Givetian–Serpukhovian) sites corroborates a persistent Devonian/Carboniferous split. Separation of the data into marine and non‐marine partitions indicates more Devonian–Carboniferous faunal continuity in non‐marine settings compared to marine settings. These results agree with the latest fossil discoveries and suggest that the Devonian–Carboniferous transition proceeded differently in different environments and among different taxonomic groups.