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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.
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.
The early tetrapod Crassigyrinus scoticus was a large aquatic predator from the mid Carboniferous (late Viséan or early Serpukovian) of Scotland, around 330 My in age. There are five main specimens with cranial remains: an articulated skeleton; two incomplete skulls; and two lower jaws. Crassigyrinus retains several apparently primitive features of the palatal dentition and lower jaw, and its phylogenetic position is disputed. A partial lower jaw resembling that of Crassigyrinus was discovered at Burnmouth in the Borders region of Scotland. The horizon in which it was found is dated as late Tournaisian, CM palynozone, around 350 My in age. Though it lacks dentition, the jaw preserves much of the postsplenial, angular and surangular, whose appearance externally and internally is almost identical to that of C. scoticus. Internally, the jaw shows a similarly limited extent of the suturing between the splenial series and the prearticular, a primitive condition. Externally, the type and distribution of dermal ornamentation closely matches that of C. scoticus, as does the deeply excavated and marginally positioned lateral line groove. As well as external and internal features, all specimens of C. scoticus are of similar skull size, though the Burnmouth jaw is somewhat smaller. If correctly attributable to Crassigyrinus, this specimen extends the existence of the genus by approximately 20 million years towards the base of the Carboniferous.
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.
Previously described and new specimens of the anthracosaur Silvanerpeton miripedes from the Scottish Viséan of East Kirkton yield important new data which allow us to provide a more complete reconstruction of the skull roof, palate, braincase and lower jaw. A stout sacral rib and an incompletely ossified tarsus with a subquadrangular intermedium are also recorded for the first time. A remarkably well preserved humerus in extensor view shows similarities with humeri of immature specimens of the embolomere Proterogyrinus. A new cladistic analysis, built from combining characters used in two recent matrices, places Silvanerpeton in a basal position relative to embolomeres and more derived stem amniotes. Data from Silvanerpeton inform character polarity near the base of the amniote total group. We discuss some morphofunctional implications of character changes at the root of total group amniotes, acquisition of terrestrial habits, and patterns of early disparity in this clade.
The lower jaw of the Devonian tetrapod Acanthostega is described for the first time. Redescriptions are provided for the lower jaws of the elpistostegid Panderichthys, the Devonian tetrapods Elginerpeton, Obruchevichthys, Metaxygnathus, Ventastega and Ichthyostega, and the Carboniferous tetrapods Crassigyrinus, Megalocephalus and Gephyrostegus. The character distri- butions thus revealed differ considerably from previous accounts, particularly in the wide distribution of certain primitive characters. Meckelian ossification in the middle part of the jaw is widespread among Devonian tetrapods, being demonstrably absent only in Acanthostega. Among Carboniferous tetrapods, a tooth-bearing parasymphysial plate is shown to be present in Crassigyrinus and Megalocephalus (having already been demonstrated by other authors in Whatcheeria and Greererpeton). A phylogenetic analysis of 26 early tetrapods including all the aforementioned genera, scored for 51 lower jaw characters, produces at least 2,500 equally parsimonious trees. However, the lack of resolution lies largely in a big top end polychotomy containing anthracosaurs, temnospondyls, seymouriamorphs, microsaurs and a nectridean-amniote clade. Below this polycho- tomy, which may correspond approximately to the tetrapod crown group, there is a well-resolved stem-group containing, in descending order, Megalocephalus, Greererpeton, Crassigyrinus, (jaws associated with) Tulerpeton, Whatcheeria, Acanthostega, Metaxygnathus, Ichthyostega, Ventastega and Metaxygnathus (unresolved), an Elginerpeton-Obruchevichthys clade, and Panderichthys. This conflicts with recently published phylogenies by Coates and Lebedev & Coates, which place Tulerpeton and all post-Devonian tetrapods in the amphibian or amniote branches of the tetrapod crown group.
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