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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 genus Sagenodus is 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); and Sagenodus quinquecostatus derived 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 of Sagenodus in 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.
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 tetrapods Crassigyrinus scoticus and Eoherpeton watsoni from 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.
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 splendens and Ctenodus 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 of Ctenodus tooth plates and are here renamed Clackodus angustulus. A further taxon, Conchopoma sp., 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.
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 fish Remigolepis is 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. The Schopfites species that occurs through most of the succession is Schopfites delicatus rather than Schopfites 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), contains Retispora lepidophyta and 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.
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.
A new early tetrapod, Mesanerpeton woodi gen. et sp. nov., collected by Stan Wood from the Ballagan Formation, Tournaisian CM palynozone, at Willie's Hole, Scottish Borders, is described. It includes vertebrae like those of Crassigyrinus, with poorly developed neural arches, a well ossified ulna with a large olecranon, and a humerus that is structurally intermediate between the pleisiomorphic condition of Devonian taxa and that of all later forms. A comparative analysis of this new material and other tetrapodomorph humeri revealed how an increase in humeral torsion transformed the course of the brachial artery and median nerve through the bone, from an entirely ventral path to one in which the blood vessel and nerve passed through the entepicondyle from the dorsal to the ventral surface. Increasing humeral torsion is suggested to improve walking in early tetrapods by potentially contributing to an increase in stride length, and is one of a number of changes to limb morphology during the Early Carboniferous that led to the development of terrestrial locomotion.
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.
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.
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