Romer's Gap Plugged: 5 new Tetrapods From Tournaisian Mississippian Carboniferous Scotland

Phylogenetic and environmental context of a Tournaisian tetrapod fauna

Authors:

Clack et al

Abstract:

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.

pop sci write up.

Consider Romer's Gap Plugged! New Floodplain Siltstone Deposit From Scotland Preserves Tetrapods From Tournaisian Carboniferous

Early Mississippian sandy siltstones preserve rare vertebrate fossils in seasonal flooding episodes

Authors:

Bennett et al

Abstract:

Flood-generated sandy siltstones are under-recognised deposits that preserve key vertebrate (actinopterygians, rhizodonts, and rarer lungfish, chondrichthyans and tetrapods), invertebrate and plant fossils. Recorded for the first time from the Lower Mississippian Ballagan Formation of Scotland, more than 140 beds occur throughout a 490 m thick core succession characterised by fluvial sandstones, palaeosols, siltstones, dolostone ‘cementstones’ and gypsum from a coastal–alluvial plain setting. Sandy siltstones are described as a unique taphofacies of the Ballagan Formation. They are matrix-supported siltstones with millimetre-sized siltstone and very fine sandstone lithic clasts. Common bioclasts include plants and megaspores, fish, ostracods, eurypterids and bivalves. Fossils have a high degree of articulation compared with those found in other fossil-bearing deposits such as conglomerate lags at the base of fluvial channel sandstones. Bed thickness and distribution varies throughout the formation, with no stratigraphic trend. The matrix sediment and clasts are sourced from the reworking of floodplain sediments including desiccated surfaces and palaeosols. Secondary pedogenic modification affects 30% of the sandy siltstone beds and most (71%) overlie palaeosols or desiccation cracks. Sandy siltstones are interpreted as cohesive debris flow deposits that originated by the overbank flooding of rivers and due to localised floodplain sediment transport at times of high rainfall; their association with palaeosols and desiccation cracks indicates seasonally wet to dry cycles throughout the Tournaisian. Tetrapod and fish fossils derived from floodplain lakes and land surfaces are concentrated by local erosion and reworking and are preserved by deposition into temporary lakes on the floodplain; their distribution indicates a local origin, with sediment distributed across the floodplain in seasonal rainfall episodes. These deposits are significant new sites that can be explored for the preservation of rare non-marine fossil material and provide unique insights into the evolution of early terrestrial ecosystems.

Lungfish Remained Morphologically Diverse Through Romer's Gap

Lungfish diversity in Romer's Gap: reaction to the end-Devonian extinction

Authors:

Smithson et al

Abstract:

Romer's Gap, the interval following the end-Devonian extinction event, has been described as a post-extinction trough for vertebrates. It is a time roughly equivalent to the Tournaisian stage of the early Carboniferous and has been characterized by a lull in diversity of survivors. Lungfish typified this description. One species was known from one locality. Recently, a diverse collection of lungfish tooth plates, representing seven new forms, was recovered from new Tournaisian vertebrate localities in northern Britain. They display a range of previously unknown morphologies, with tooth shape and wear patterns not seen in other post-Devonian forms. A comparison of tooth ridge number and tooth ridge angle in lungfishes from the Famennian, Tournaisian and Visean reveals marked differences between late Devonian and early Carboniferous taxa. The most common tooth plate shape in the Famennian is absent from our sample of Tournaisian taxa. Two completely new shapes have evolved, one with a relatively low tooth ridge angle, no greater than 40°, in which most of the tooth ridges are essentially parallel, and the other with a much higher tooth ridge angle of up to 180° where the tooth ridges are highly divergent. This high level of morphological diversity over a narrow time period suggests that, following the end-Devonian extinction, gaps in ecospace left by the extinction of major groups of fishes were exploited by a previously unrecorded radiation of lungfishes. Whilst taxonomic diversity of lungfishes declined following the end-Devonian extinction, recovery and diversification among tooth-plated forms was rapid, and morphological disparity among these forms subsequently increased. Contrary to previous assumptions, morphological disparity among lungfish did not decline until much later in the Carboniferous.

Ward Takes a hit, Romer's Gap not Real: Diverse Tournaisian Mississipian Carboniferous Tetrapods From Nova Scotia Include Several Devonian Taxa

A Diverse Tetrapod Fauna at the Base of 'Romer's Gap'

Authors:

Anderson et al

Abstract:

The lack of fossil tetrapod bearing deposits in the earliest Carboniferous (‘Romer’s Gap’) has provoked some recent discussions regarding the proximal cause, with three explanations being offered: environmental, taphonomic, and collection failure. One of the few, and earliest, windows into this time is the locality of Blue Beach exposed in the Tournaisian deposits at Horton Bluff lying along the Avon River near Hantsport, Nova Scotia, Canada. This locality has long been known but, because the fossils were deposited in high energy settings they are almost always disarticulated, so the fauna has not been described in detail. Recent intensive collection has revealed a diverse assemblage of material, including for the first time associated elements, which permits an evaluation of the faunal constituents at the locality. Although not diagnosable to a fine taxonomic level, sufficient apomorphies are present to identify representatives from numerous clades known from more complete specimens elsewhere. The evidence suggests a diverse fauna was present, including whatcheeriids and embolomeres. A single humerus previously had been attributed to a colosteid, but there is some uncertainty with this identification. Additional elements suggest the presence of taxa otherwise only known from the late Devonian. Depositional biases at the locality favor tetrapod fossils from larger individuals, but indirect evidence from trackways and tantalizing isolated bones evidences the presence of small taxa that remain to be discovered. The fossils from Blue Beach demonstrate that when windows into the fauna of ‘Romer’s Gap’ are found a rich diversity of tetrapods will be shown to be present, contra arguments that suggested this hiatus in the fossil record was due to extrinsic factors such as atmospheric oxygen levels. They also show that the early tetrapod fauna is not easily divisible into Devonian and Carboniferous faunas, suggesting that some tetrapods passed through the end Devonian extinction event unaffected.

Studying Modern Belly Dragging Lizards to Understand Early Carboniferous Tetrapod Locomotion

Bridging “Romer’s Gap”: Limb Mechanics of an Extant Belly-Dragging Lizard Inform Debate on Tetrapod Locomotion During the Early Carboniferous

Authors:

Nyakatura et al

Abstract:

Devonian stem tetrapods are thought to have used ‘crutching’ on land, a belly-dragging form of synchronous forelimb action-powered locomotion. During the Early Carboniferous, early tetrapods underwent rapid radiation, and the terrestrial locomotion of crown-group node tetrapods is believed to have been hindlimb-powered and ‘raised’, involving symmetrical gaits similar to those used by modern salamanders. The fossil record over this period of evolutionary transition is remarkably poor (Romer’s Gap), but we hypothesize a phase of belly-dragging sprawling locomotion combined with symmetrical gaits. Since belly-dragging sprawling locomotion has differing functional demands from ‘raised’ sprawling locomotion, we studied the limb mechanics of the extant belly-dragging blue-tongued skink. We used X-ray reconstruction of moving morphology to quantify the three-dimensional kinematic components, and simultaneously recorded single limb substrate reaction forces (SRF) in order to calculate SRF moment arms and the external moments acting on the proximal limb joints. In the hindlimbs, stylopodal long-axis rotation is more emphasized than in the forelimbs, and much greater vertical and propulsive forces are exerted. The SRF moment arm acting on the shoulder is at a local minimum at the instant of peak force. The hindlimbs display patterns that more closely resemble ‘raised’ sprawling species. External moment at the shoulder of the skink is smaller than in ‘raised’ sprawlers. We propose an evolutionary scenario in which the locomotor mechanics of belly-dragging early tetrapods were gradually modified towards hindlimb-powered, raised terrestrial locomotion with symmetrical gait. In accordance with the view that limb evolution was an exaptation for terrestrial locomotion, the kinematic pattern of the limbs for the generation of propulsion preceded, in our scenario, the evolution of permanent body weight support.