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The acquisition of terrestrial, limb-based locomotion during tetrapod evolution has remained a subject of debate for more than a century1,2. Our current understanding of the locomotor transition from water to land is largely based on a few exemplar fossils such as Tiktaalik3, Acanthostega4, Ichthyostega5 and Pederpes6. However, isolated bony elements may reveal hidden functional diversity, providing a more comprehensive evolutionary perspective7. Here we analyse 40 three-dimensionally preserved humeri from extinct tetrapodomorphs that span the fin-to-limb transition and use functionally informed ecological adaptive landscapes8–10 to reconstruct the evolution of terrestrial locomotion. We show that evolutionary changes in the shape of the humerus are driven by ecology and phylogeny and are associated with functional trade-offs related to locomotor performance. Two divergent adaptive landscapes are recovered for aquatic fishes and terrestrial crown tetrapods, each of which is defined by a different combination of functional specializations. Humeri of stem tetrapods share a unique suite of functional adaptations, but do not conform to their own predicted adaptive peak. Instead, humeri of stem tetrapods fall at the base of the crown tetrapod landscape, indicating that the capacity for terrestrial locomotion occurred with the origin of limbs. Our results suggest that stem tetrapods may have used transitional gaits5,11 during the initial stages of land exploration, stabilized by the opposing selective pressures of their amphibious habits. Effective limb-based locomotion did not arise until loss of the ancestral ‘L-shaped’ humerus in the crown group, setting the stage for the diversification of terrestrial tetrapods and the establishment of modern ecological niches12,13. Analysis of humeri from fossils that span the fin-to-limb transition reveal that the change in the humerus shape is driven by both ecology and phylogeny, and is associated with functional trade-offs related to locomotor performance.

The well-known late Mississippian/early Carboniferous locality of East Kirkton in Scotland has the earliest described fauna of terrestrial tetrapods. Seven species are now known, represented by articulated skeletons of moderate-sized animals with snout-vent length of up to 200 mm, and each is unique to East Kirkton. Here we describe the skull bones of a much larger tetrapod that closely resembles those of embolomeres from the Pennsylvanian. Although the new material is too incomplete to be named as a new species, it enhances the taxonomic diversity of the East Kirkton tetrapod fauna, predates the embolomeres from other sites in Scotland and extends the range of the group earlier into the Mississippian.

The late Viséan anthracosauroid Eldeceeon rolfei from the East Kirkton Limestone of Scotland is re-described. Information from two originally described and two newly identified specimens broadens our knowledge of this tetrapod. A detailed account of individual skull bones and a revision of key axial and appendicular features are provided, alongside the first complete reconstructions of the skull and lower jaw and a revised reconstruction of the postcranial skeleton. In comparison to Silvanerpeton, the only other anthracosauroid from East Kirkton, Eldeceeon is characterised by a proportionally wider semi-elliptical skull, comparatively smaller nostrils set farther apart, smaller and more rounded orbits, a shorter skull table with gently convex lateral margins, and a deeper suspensorium with a straight posterior margin and a small dorsal embayment. The remarkably large hind feet and elongate toes of Eldeceeon presumably represent an adaptation for attaining high locomotory speed through increased stride length and reduced stride frequency. This would necessitate great muscle force but few muscle contractions. At the beginning of a new stride cycle, repositioning the pes anteriorly and lifting the toes off the ground would require a strong and large muscle to pull the femur upward and rotate it inward and forward. It is hypothesised that such muscle might correspond to the puboischiofemoralis internus 2, which would extend along the posterior half of the vertebral column, consistent with the occurrence of long, curved ribs in the anterior half of the trunk. Using maximum parsimony and Bayesian inference, cladistic analyses of all major groups of stem amniotes retrieve a sister group relationship between Eldeceeon and Silvanerpeton, either as the most plesiomorphic stem amniote clade or as a clade immediately crownward of anthracosauroids.

The Carboniferous lungfish genusSagenodusis reviewed from all available British specimens and described in detail for the first time. We identify two species exclusive to the UK:Sagenodusinaequalis, the type species, deriving from the late Carboniferous (=Pennsylvanian); andSagenodus quinquecostatusderived from the early Carboniferous (=Mississippian). The genus is probably the most widespread of the known Carboniferous lungfish genera, but the British species have not been formally described since their discovery in the mid–late 19th Century. This work will provide data to help resolve existing questions about the position ofSagenodusin the phylogeny of Palaeozoic lungfishes, and provide a template for the recognition of isolated elements in museum collections and the finds from recent and future field work. The early Carboniferous species,S. quinquecostatus, shows a so far unique functional mechanism in which the lower tooth plates appear to rotate relative to the upper plates during jaw closure, implying a kinetic function at the symphysis or jaw joint.

The taxonomically diverse terrestrial tetrapod fauna from the late Mississippian East Kirkton Limestone includes the earliest known members of stem Amphibia and stem Amniota. Here we name and describe a new East Kirkton tetrapod with an unusual hindlimb morphology reminiscent of that of several stem- and primitive crown amniotes. It displays a unique ilium with two slender and elongate processes and a 5-digit pes with a long, stout metatarsal IV and a greatly elongate digit IV. The new taxon broadens our knowledge of East Kirkton tetrapods, adding to the remarkable diversity of their hindlimb constructions, functional specializations, locomotory modes, and adaptations to a wide variety of substrates. An unweighted character parsimony analysis places the new taxon in a polytomy alongside some other Carboniferous groups. Conversely, weighted parsimony and Bayesian analyses retrieve it among the earliest diverging stem amniotes, either as the basalmost anthracosaur or within a clade that includes also Eldeceeon and Silvanerpeton, crownward of an array of chroniosaurs plus anthracosaurs.

Until recently the immediate aftermath of the Hangenberg event of the Famennian Stage (Upper Devonian) was considered to have decimated sarcopterygian groups, including lungfish, with only two taxa...

The Ballagan Formation of northern Britain provides an exceptional record of Early Mississippian ecosystems that developed as tetrapods emerged onto land. In this paper, we study two 500-metre sections of the formation near Berwick-upon-Tweed, which are characterised by abundant ferroan dolostone beds. Five lithofacies are identified: cemented siltstone and sandstone, homogeneous dolomicrite, mixed dolomite and siltstone, mixed calcite and dolomite, and dolomite with evaporite minerals. Cemented sediments have non-planar to planar subhedral dolomite crystals, up to 40 μm in size, whereas other facies predominantly comprise dolomicrite or planar euhedral dolomite rhombs 15 μm in size, with patches of larger rhombs indicating partial recrystallisation. The macro- and microfossil content of the dolostones is dominated by sarcopterygian (rhizodont) and actinopterygian fish, bivalves, Serpula, ostracods and Chondrites trace fossils; with rarer Spirorbis, chondrichthyans (Ageleodus, hybodonts and ?ctenacanths, xenacanths), non-gyracanth acanthodians, gastropods, eurypterids, brachiopods, plant debris, wood, lycopsid roots, charcoal, megaspores, phycosiphoniform burrows, Zoophycos? and Rhizocorallium. The oxygen and carbon isotope composition of dolomites range from –3.6‰ to –1.7‰ (for δ18O) and –2.6‰ to +1.6‰ (for δ13C) respectively indicating dolomite growth in mixed salinity waters. Frequent marine storm-surge events transported marine waters and animals into floodplain lakes, where evaporation, interstitial sulphate-reducing bacteria, iron reduction and methanogenesis allowed dolomite growth in the shallow sub-surface. Secondary pedogenic modification (by roots, brecciation, desiccation, and soil forming processes) is common and represents lake evaporation with, in some cases, saline marsh development. The dolostone facies are part of a complex environmental mosaic of sub-aerial dry floodplain, wet marshy floodplains, rivers, and lakes ranging in salinity from freshwater to hypersaline. Marine influence is strongest at the base of the formation and decreases over time, as the floodplain became drier, and forested areas became more established. Coastal lakes were an important habitat for animals recovering from the end-Devonian Hangenberg Crisis and may have acted as a pathway for euryhaline fishes, molluscs and arthropods to access freshwater environments.

Ramsay Heatley Traquair, the eminent Victorian Scottish palaeoichthyologist and museum curator, procured an extensive collection of Palaeozoic fishes from across Scotland with the help of local miners and quarrymen. One very productive locality near Edinburgh was Loanhead. Traquair described numerous fossil fish from this Serpukhovian site, including four lungfish taxa:Ctenodus interruptus,Sagenodus quinquecostatus,Uronemus splendensandCtenodus angustulus. The first three are now quite well known, but the fourth was only briefly described and never figured. It is based entirely on tooth plates, which are unusual both in their very small size and the arrangement of the tooth ridges. They lack the diagnostic characters ofCtenodustooth plates and are here renamedClackodus angustulus. A further taxon,Conchopomasp., has recently been identified. Represented by a spade-shaped parasphenoid and denticulated jaw elements, it is the earliest known occurrence of the genus, extending its range into the Mississippian. A sixth taxon may be represented by an isolated parasphenoid bearing an anterior process, previously only seen in Devonian lungfish. The presence of up to six lungfish taxa at a single locality is unprecedented in the Carboniferous and indicates that the high level of lungfish diversity encountered in the Tournaisian of the Scottish Borders continued throughout the Mississippian, adding to the growing evidence that post-Devonian lungfish evolution was not as limited as previously proposed. This may have been due to changes in tooth plate growth, enabling greater variation in dentition and diet. In most Devonian taxa, tooth plate growth can be explained by comparison with that in extant forms, but analysis of Carboniferous tooth plates suggest growth was different in many taxa, possibly based on more than one pioneer tooth, allowing for novel patterns of tooth ridges and different types of teeth to develop on the same plate.

Stan Wood had a gift for finding exceptional Early Carboniferous fossils. Among them are 32 type specimens. His discoveries significantly changed our understanding of the history of life on Earth. Many of the fossils he collected are on display in museums across the UK and the localities he discovered continue to yield important new material. Here we briefly review some of Stan Wood's key achievements and describe the legacy he left.

Stan Wood was an exceptional fossil collector who, over a collecting career of more than 40 years, provided British palaeontology with an abundance and variety of new Carboniferous fossils, the like of which had not been collected since Victorian times. So, what made him a great collector? Here, with the help of Stan's family, his friends and colleagues, we try to provide the answer. There is no single factor that stands out, but a complex mixture of innate and learned behaviours that together produced a unique talent. Although he acquired an Open University degree in geology, Stan was largely self-taught as a collector and, in doing so, became an accomplished and confident field geologist. He was naturally curious, persistent and very observant, with a photographic memory. He was tough, very strong and enjoyed hard physical work. He was congenial, unorthodox and a calculated risk taker. He asked questions, tested ideas and had a healthy disregard for authority. He was systematic, kept detailed records and shared his discoveries. He not only loved collecting fossils but, in the process, discovered in himself the essential qualities of a true scientist.

Bone healing is an important survival mechanism, allowing vertebrates to recover from injury and disease. Here we describe newly recognized paleopathologies in the hindlimbs of the early tetrapodsCrassigyrinus scoticusandEoherpeton watsonifrom the early Carboniferous of Cowdenbeath, Scotland. These pathologies are among the oldest known instances of bone healing in tetrapod limb bones in the fossil record (about 325 Ma). X-ray microtomographic imaging of the internal bone structure of these lesions shows that they are characterized by a mass of trabecular bone separated from the shaft's trabeculae by a layer of cortical bone. We frame these paleopathologies in an evolutionary context, including additional data on bone healing and its pathways across extinct and extant sarcopterygians. These data allowed us to synthesize information on cell-mediated repair of bone and other mineralized tissues in all vertebrates, to reconstruct the evolutionary history of skeletal tissue repair mechanisms. We conclude that bone healing is ancestral for sarcopterygians. Furthermore, other mineralized tissues (aspidin and dentine) were also capable of healing and remodeling early in vertebrate evolution, suggesting that these repair mechanisms are synapomorphies of vertebrate mineralized tissues. The evidence for remodeling and healing in all of these tissues appears concurrently, so in addition to healing, these early vertebrates had the capacity to restore structure and strength by remodeling their skeletons. Healing appears to be an inherent property of these mineralized tissues, and its linkage to their remodeling capacity has previously been underappreciated.

The enigmatic tetrapod Acherontiscus caledoniae from the Pendleian stage of the Early Carboniferous shows heterodontous and durophagous teeth, representing the earliest known examples of significant adaptations in tetrapod dental morphology. Tetrapods of the Late Devonian and Early Carboniferous (Mississippian), now known in some depth, are generally conservative in their dentition and body morphologies. Their teeth are simple and uniform, being cone-like and sometimes recurved at the tip. Modifications such as keels occur for the first time in Early Carboniferous Tournaisian tetrapods. Acherontiscus , dated as from the Pendleian stage, is notable for being very small with a skull length of about 15 mm, having an elongate vertebral column and being limbless. Cladistic analysis places it close to the Early Carboniferous adelospondyls, aïstopods and colosteids and supports the hypothesis of ‘lepospondyl’ polyphyly. Heterodonty is associated with a varied diet in tetrapods, while durophagy suggests a diet that includes hard tissue such as chitin or shells. The mid-Carboniferous saw a significant increase in morphological innovation among tetrapods, with an expanded diversity of body forms, skull shapes and dentitions appearing for the first time.

In Scotland, the base of the Ballagan Formation has traditionally been placed at the first grey mudstone within a contiguous Late Devonian to Carboniferous succession. This convention places the Devonian–Carboniferous boundary within the Old Red Sandstone (ORS) Kinnesswood Formation. The consequences of this placement are that tetrapods from the Ballagan Formation were dated as late Tournaisian in age and that the ranges of typically Devonian fish found in the Kinnesswood Formation continued into the Carboniferous. The Pease Bay specimen of the fishRemigolepisis from the Kinnesswood Formation. Comparisons with its range in Greenland, calibrated against spores, show it was Famennian in age. Detailed palynological sampling at Burnmouth from the base of the Ballagan Formation proves that the early Tournaisian spore zones (VI and HD plus Cl 1) are present. TheSchopfitesspecies that occurs through most of the succession isSchopfites delicatusrather thanSchopfites claviger. The latter species defines the late Tournaisian CM spore zone. The first spore assemblage that has been found in Upper ‘ORS' strata underlying the Ballagan Formation (Preston, Whiteadder Water), containsRetispora lepidophytaand is from the early latest Famennian LL spore zone. The spore samples are interbedded with volcaniclastic debris, which shows that the Kelso Volcanic Formation is, in part, early latest Famennian in age. These findings demonstrate that the Ballagan Formation includes most of the Tournaisian with the Devonian–Carboniferous boundary positioned close to the top of the Kinnesswood Formation. The Stage 6 calcrete at Pease Bay can be correlated to the equivalent section at Carham, showing that it represents a time gap equivalent to the latest Famennian glaciation(s). Importantly, some of the recently described Ballagan Formation tetrapods are older than previously dated and now fill the key early part of Romer's Gap.

A conglomerate bed from the Tournaisian Ballagan Formation of Scotland preserves a rich array of vertebrate and other nonmarine fossils providing an insight into the wider ecosystem and paleoenvironment that existed during this pivotal stage of Earth history. It challenges hypotheses of a long-lasting post-extinction trough following the end-Devonian extinction event. The fauna recovered includes a wide size range of tetrapods, rhizodonts, and dipnoans, from tiny juveniles or small-bodied taxa up to large adults, and more than one taxon of each group is likely. Some fauna, such as actinopterygians and chondrichthyans, are rare as macrofauna but are better represented in the microfossil assemblage. The fauna provides evidence of the largest Carboniferous lungfish ever found. The specimens are preserved in a localized, poorly-sorted conglomerate which was deposited in the deepest part of a river channel, the youngest of a group of channels. In addition to the fossils (micro- and macro-), the conglomerate includes locally-derived clasts of paleosols and other distinctive elements of the surrounding floodplains. Charcoal fragments represent small woody axes and possible larger trunk tissue from arborescent pteridosperms. Preservation of the fossils indicates some aerial exposure prior to transport, with abrasion from rolling. The findings presented here contrast with other published trends in vertebrate size that are used to interpret a reduction in maximum sizes during the Tournaisian. The richness of the fauna runs counter to the assumption of a depauperate nonmarine fauna following the end-Devonian Hangenberg event, and charcoal content highlights the occurrence of fire, with the requisite levels of atmospheric oxygen during that stage.

Chondrichthyan teeth from a new locality in the Scottish Borders supply additional evidence of Early Carboniferous chondrichthyans in the UK. The interbedded dolostones and siltstones of the Ballagan Formation exposed along Whitrope Burn are interpreted as representing a restricted lagoonal environment that received significant amounts of land-derived sediment. This site is palynologically dated to the latest Tournaisian–early Viséan. The diverse dental fauna documented here is dominated by large crushing holocephalan toothplates, with very few, small non-crushing chondrichthyan teeth. Two new taxa are named and described. Our samples are consistent with worldwide evidence that chondrichthyan crushing faunas are common following the Hangenberg extinction event. The lagoonal habitat represented by Whitrope Burn may represent a temporary refugium that was host to a near-relict fauna dominated by large holocephalan chondrichthyans with crushing dentitions. Many of these had already become scarce in other localities by the Viséan and become extinct later in the Carboniferous. This fauna provides evidence of early endemism or niche separation within European chondrichthyan faunas at this time. This evidence points to a complex picture in which the diversity of durophagous chondrichthyans is controlled by narrow spatial shifts in niche availability over time.

Tetrapod postcranial bones are described from Scotland: from the Limestone Coal Group (Early Carboniferous, Serpukhovian) at the Dora open cast site, Fife, and from beds equivalent to the Burdiehouse Limestone (Early Carboniferous, Visean) on the island of Inchkeith, Firth of Forth. The elements from Dora are derived relative to Devonian and Tournaisian tetrapods in having a diamond-shaped interclavicle with no parasternal process, a humerus with a triangular-shaped entepicondyle, a rod-like ilium lacking a post-iliac process and a gracile femur with a prominent internal trochanter but no adductor blade. These bones share characters with their homologues in colosteids and temnospondyls and may be attributable to Doragnathus woodi . The femur from Inchkeith most closely resembles that of the embolomere Proterogyrinus scheelei .

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 ($\textit{Perittodus apsconditus}$, $\textit{Koilops herma}$, $\textit{Ossirarus kierani}$, $\textit{Diploradus austiumensis}$ and $\textit{Aytonerpeton microps}$) 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.

Devonian tetrapods (limbed vertebrates), known from an increasingly large number of localities, have been shown to be mainly aquatic with many primitive features. In contrast, the post-Devonian record is marked by an Early Mississippian temporal gap ranging from the earliest Carboniferous (Tournaisian and early Viséan) to the mid-Viséan. By the mid-Viséan, tetrapods had become effectively terrestrial as attested by the presence of stem amniotes, developed an essentially modern aspect, and given rise to the crown group. Up to now, only two localities have yielded tetrapod specimens from the Tournaisian stage: one in Scotland with a single articulated skeleton and one in Nova Scotia with isolated bones, many of uncertain identity. We announce a series of discoveries of Tournaisian-age localities in Scotland that have yielded a wealth of new tetrapod and arthropod fossils. These include both terrestrial and aquatic forms and new taxa. We conclude that the gap in the fossil record has been an artifact of collection failure.

Synopsis Formal diagnosis of Westlothiana lizziae, a new genus and species is given, to differentiate this taxon and to comply with the International Code of Zoological Nomenclature. This is the most primitive known reptile and establishes the presence in the Lower Carboniferous of a reptile fauna sufficiently ancient to have been ancestral to all later amniotes.