Stem Tetrapods Research Articles

Giant stem tetrapod was apex predator in Gondwanan late Palaeozoic ice age.

Current hypotheses of early tetrapod evolution posit close ecological and biogeographic ties to the extensive coal-producing wetlands of the Carboniferous palaeoequator with rapid replacement of archaic tetrapod groups by relatives of modern amniotes and lissamphibians in the late Carboniferous (about 307 million years ago). These hypotheses draw on a tetrapod fossil record that is almost entirely restricted to palaeoequatorial Pangea (Laurussia)1,2. Here we describe a new giant stem tetrapod, Gaiasia jennyae, from high-palaeolatitude (about 55° S) early Permian-aged (about 280 million years ago) deposits in Namibia that challenges this scenario. Gaiasia is represented by several large, semi-articulated skeletons characterized by a weakly ossified skull with a loosely articulated palate dominated by a broad diamond-shaped parasphenoid, a posteriorly projecting occiput, and enlarged, interlocking dentary and coronoid fangs. Phylogenetic analysis resolves Gaiasia within the tetrapod stem group as the sister taxon of the Carboniferous Colosteidae from Euramerica. Gaiasia is larger than all previously described digited stem tetrapods and provides evidence that continental tetrapods were well established in the cold-temperate latitudes of Gondwana during the final phases of the Carboniferous-Permian deglaciation. This points to a more global distribution of continental tetrapods during the Carboniferous-Permian transition and indicates that previous hypotheses of global tetrapod faunal turnover and dispersal at this time2,3 must be reconsidered.

Evolutionary origin of Hoxc13-dependent skin appendages in amphibians

Cornified skin appendages, such as hair and nails, are major evolutionary innovations of terrestrial vertebrates. Human hair and nails consist largely of special intermediate filament proteins, known as hair keratins, which are expressed under the control of the transcription factor Hoxc13. Here, we show that the cornified claws of Xenopus frogs contain homologs of hair keratins and the genes encoding these keratins are flanked by promoters in which binding sites of Hoxc13 are conserved. Furthermore, these keratins and Hoxc13 are co-expressed in the claw-forming epithelium of frog toe tips. Upon deletion of hoxc13, the expression of hair keratin homologs is abolished and the development of cornified claws is abrogated in X. tropicalis. These results indicate that Hoxc13-dependent expression of hair keratin homologs evolved already in stem tetrapods, presumably as a mechanism for protecting toe tips, and that this ancestral genetic program was coopted to the growth of hair in mammals.

New information on the neurocranium of Archeria crassidisca and the relationships of the Embolomeri

Abstract Embolomeres have been considered candidate stem-amniotes by many authors, but recent work has shown that anatomical and phylogenetic support for this hypothesis may be ambiguous. Newer phylogenetic approaches focus on the anatomy of the endochondral bones surrounding the brain (neurocranium) and associated dermal investing bones, so accurate description of the neurocranium of tetrapods, and particularly embolomeres, is critical to assess the phylogenetic context of amniote origins. I here amend prior descriptions of the neurocranium of the abundant Early Permian embolomere, Archeria crassidisca, with a description of a previously unreported partial braincase. This specimen reveals features broadly associated with stem-tetrapods of the fin-to-limb transition and recently described in aïstopod stem-tetrapods, including a cranial notochordal remnant, a possible buccohypophyseal canal, and anteroventrally projecting saccular fossae that extend deep into the basioccipital. I present an amended description and reconstruction of the neurocranium and a phylogenetic analysis integrating these codings, which recovers embolomeres more robustly within the tetrapod stem. I review characters previously used to assign embolomeres to the amniote stem and find this evidence is ambiguous or incorrect. I then outline strong character support for a stem-tetrapod position of embolomeres, and implications for the ancestral body plan of crown-group tetrapods.

Fossil bone histology reveals ancient origins for rapid juvenile growth in tetrapods

Patterns of growth throughout the lifetime of an animal reflect critical life history traits such as reproductive timing, physiology, and ecological interactions. The ancestral growth pattern for tetrapods has traditionally been described as slow-to-moderately paced, akin to modern amphibians, with fast growth and high metabolic rates considered a specialized physiological trait of amniotes. Here, we present bone histology from an ontogenetic series of the Early Carboniferous stem tetrapod Whatcheeria deltae, and document evidence of fibrolamellar bone—primary bone tissue associated with fast growth. Our data indicate that Whatcheeria juveniles grew rapidly and reached skeletal maturity quickly, allowing them to occupy a large-bodied predator niche in their paleoenvironment. This life history strategy contrasts with those described for other stem tetrapods and indicates that a diversity of growth patterns existed at the origins of tetrapod diversification. Importantly, Whatcheeria marks an unexpectedly early occurrence of fibrolamellar bone in Tetrapoda, both temporally and phylogenetically. These findings reveal that elevated juvenile growth is not limited to amniotes, but has a deep history in the tetrapod clade and may have played a previously unrecognized role in the tetrapod invasion of land.

Computed tomography and three-dimensional reconstruction of the skull of the stem tetrapod Crassigyrinus scoticus Watson, 1929

ABSTRACT The early tetrapod Crassigyrinus scoticus was a large aquatic predator known from the lower- to mid-Carboniferous (upper Tournasian to upper Visean/lower Serpukovian, approximately 350–330 Ma) of Scotland and Canada. Crassigyrinus is enigmatic in terms of its phylogenetic position due to its unusual morphology, which features a mixture of primitive and derived characters. Previous reconstructions, based on five incomplete and deformed specimens, have suggested a dorsoventrally tall skull with a short and broad snout, large orbits and external nares, and an extended postorbital region. In this study, we scanned four specimens using computed tomography and segmented imaging data to separate bone from matrix and individual bones from each other. Based on these data, we present a revised description of the upper and lower jaws, including sutural morphology and abundant new anatomical information. Damage was repaired and the skull retrodeformed to create a hypothetical three-dimensional reconstruction of the skull of Crassigyrinus that is dorsoventrally flatter than earlier reconstructions, yet still morphologically unique amongst early tetrapods. Overall skull shape, the size and distribution of the teeth, sutural morphology, and the specialized anatomy of the jaw joint and mandibular symphysis all suggest that Crassigyrinus was a powerful aquatic predator capable of hunting and subduing large prey.

Control of Breathing in Ectothermic Vertebrates.

The ectothermic vertebrates are a diverse group that includes the Fishes (Agnatha, Chondrichthyes, and Osteichthyes), and the stem Tetrapods (Amphibians and Reptiles). From an evolutionary perspective, it is within this group that we see the origin of air-breathing and the transition from the use of water to air as a respiratory medium. This is accompanied by a switch from gills to lungs as the major respiratory organ and from oxygen to carbon dioxide as the primary respiratory stimulant. This transition first required the evolution of bimodal breathing (gas exchange with both water and air), the differential regulation of O2 and CO2 at multiple sites, periodic or intermittent ventilation, and unsteady states with wide oscillations in arterial blood gases. It also required changes in respiratory pump muscles (from buccopharyngeal muscles innervated by cranial nerves to axial muscles innervated by spinal nerves). The question of the extent to which common mechanisms of respiratory control accompany this progression is an intriguing one. While the ventilatory control systems seen in all extant vertebrates have been derived from common ancestors, the trends seen in respiratory control in the living members of each vertebrate class reflect both shared-derived features (ancestral traits) as well as unique specializations. In this overview article, we provide a comprehensive survey of the diversity that is seen in the afferent inputs (chemo and mechanoreceptor), the central respiratory rhythm generators, and the efferent outputs (drive to the respiratory pumps and valves) in this group. © 2022 American Physiological Society. Compr Physiol 12: 1-120, 2022.

Kinematic comparisons between mudskipper fins and salamander limbs during terrestrial locomotion.

Terrestrial and aquatic environments present drastically different challenges, yet amphibious behaviors evolved multiple times in vertebrates. Terrestrial salamanders are often used to model the locomotion of crownward stem tetrapods, but amphibious fishes may model earlier evolutionary stages as vertebrates became terrestrial. For instance, some early tetrapods may have moved on land with a mudskipper-like gait. Previously published kinetic data found that the ground reaction forces produced by the pectoral fins of mudskippers (Periophthalmus barbarus) were more medial than the limbs of tiger salamanders (Ambystoma tigrinum), which might elevate bending stresses in the fins. However, kinematic data are needed to explain these kinetic differences. Therefore, we quantified the three-dimensional kinematics of mudskipper pectoral fins and compared these to published data on tiger salamander limbs. We found that mudskipper pectoral fins generally remained more retracted, extended, and adducted compared to salamander limbs. Kinematic patterns in mudskipper pectoral fins were aligned with published kinetic data and shared a restricted range of motion found in early tetrapods. Our findings demonstrate that mudskipper pectoral fins provide weight support and propulsion but have lower mobility in the proximal versus distal elements, for which greater flexibility in the latter might compensate. Broadly, these data provide new insights into the biomechanics of using fins versus limbs for moving over land and factors that may favor the evolution of different terrestrial gaits.

Terrestrial force production by the limbs of a semi-aquatic salamander provides insight into the evolution of terrestrial locomotor mechanics.

Amphibious fishes and salamanders are valuable functional analogs for vertebrates that spanned the water-land transition. However, investigations of walking mechanics have focused on terrestrial salamanders and, thus, may better reflect the capabilities of stem tetrapods that were already terrestrial. The earliest tetrapods were likely aquatic, so salamanders that are not primarily terrestrial may yield more appropriate data for modeling the incipient stages of terrestrial locomotion. In the present study, locomotor biomechanics were quantified from semi-aquatic Pleurodeles waltl, a salamander that spends most of its adult life in water, and then compared with those of a primarily terrestrial salamander (Ambystoma tigrinum) and a semi-aquatic fish (Periophthalmus barbarus) to evaluate whether terrestrial locomotion was more comparable between species with ecological versus phylogenetic similarities. Ground reaction forces (GRFs) from individual limbs or fins indicated that the pectoral appendages of each taxon had distinct patterns of force production, but GRFs from the hindlimbs were comparable between the salamander species. The rate at which force is produced can affect musculoskeletal function, so we also calculated 'yank' (first time derivative of force) to quantify the dynamics of GRF production. Yank was sometimes slower in P. waltl but there were some similarities between the three species. Finally, the semi-aquatic taxa (P. waltl and P. barbarus) had a more medial inclination of the GRF compared to terrestrial salamanders, potentially elevating bone stresses among more aquatic taxa and limiting their excursions onto land.

Locomotor energetics in the Indonesian blue-tongued skink (Tiliqua gigas) with implications for the cost of belly-dragging in early tetrapods.

During the last decade, biomechanical and kinematic studies have suggested that a belly-dragging gait may have represented a critical locomotor stage during tetrapod evolution. This form of locomotion is hypothesized to facilitate animals to move on land with relatively weaker pectoral muscles. The Indonesian blue-tongued skink (Tiliqua gigas) is known for its belly-dragging locomotion and is thought to employ many of the same spatiotemporal gait characteristics of stem tetrapods. Conversely, the savannah monitor (Varanus exanthematicus) employs a raised quadrupedal gait. Thus, differences in the energetic efficiency of locomotion between these taxa may elucidate the role of energetic optimization in driving gait shifts in early tetrapods. Five Tiliqua and four Varanus were custom-fitted for 3D printed helmets that, combined with a Field Metabolic System, were used to collect open-flow respirometry data including O2 consumption, CO2 production, water vapor pressure, barometric pressure, room temperature, and airflow rates. Energetic data were collected for each species at rest, and when walking at three different speeds. Energetic consumption in each taxon increased at greater speeds. On a per-stride basis, energetic costs appear similar between taxa. However, significant differences were observed interspecifically in terms of net cost of transport. Overall, energy expenditure was ~20% higher in Tiliqua at equivalent speeds, suggesting that belly-dragging does impart a tangible energetic cost during quadrupedal locomotion. This cost, coupled with the other practical constraints of belly-dragging (e.g., restricting top-end speed and reducing maneuverability in complex terrains) may have contributed to the adoption of upright quadrupedal walking throughout tetrapod locomotor evolution.

Evolution, Homology, and Development of Tetrapod Limb Muscles

Since the early 1900s, researchers have attempted to unravel the origin and evolution of tetrapod limb muscles using a combination of comparative anatomy, phylogeny, and development. The methods for reconstructing soft tissues in extinct animals have been refined over time as our ability to determine muscle homology and phylogenetic relationships between tetrapods has improved. Since many muscles do not leave osteological correlates, muscle reconstruction in extinct animals is largely based on anatomy and development in extant animals. While muscle anatomy in extant tetrapods is quite conservative, the homologies of certain muscles between taxonomic groups are still uncertain. Comparative developmental studies can help to resolve these controversies, as well as revealing general patterns of muscle morphogenesis across tetrapod groups. We review the methods, results, and controversies in the muscle reconstructions of early members of the amniote, mammalian, and lissamphibian lineages, including recent attempts to reconstruct limb muscles in members of the tetrapod stem group. We also review the contribution of recent comparative developmental studies toward understanding the evolution of tetrapod limb muscles, including morphogenic gradients, the origin of paired fins, and the evolution of morphological complexity. Finally, we discuss the role of broad, comparative myological studies as part of an integrative research program on vertebrate evolutionary biology.

Variation in Articular Cartilage Thickness Among Extant Salamanders and Implications for Limb Function in Stem Tetrapods

The size and shape of articular cartilage in the limbs of extant vertebrates are highly variable, yet they are critical for understanding joint and limb function in an evolutionary context. For example, inferences about unpreserved articular cartilage in early tetrapods have implications for how limb length, joint range of motion, and muscle leverage changed over the tetrapod water-land transition. Extant salamanders, which are often used as functional models for early limbed vertebrates, have much thicker articular cartilage than most vertebrate groups, but the exact proportion of cartilage and how it varies across salamander species is unknown. I aimed to quantify this variation in a sample of 13 salamanders representing a broad range of sizes, modes of life, and genera. Using contrast-enhanced micro-CT, cartilage dimensions and bone length were measured non-destructively in the humerus, radius, ulna, femur, tibia, and fibula of each specimen. Cartilage correction factors were calculated as the combined thickness of the proximal and distal cartilages divided by the length of the bony shaft. Articular cartilage added about 30% to the length of the long bones on average. Cartilage was significantly thicker in aquatic salamanders (42 ± 14% in the humerus and 35 ± 8 in the femur) than in terrestrial salamanders (21 ± 7% in both humerus and femur). There was no consistent relationship between relative cartilage thickness and body size or phylogenetic relatedness. In addition to contributing to limb length, cartilage caps increased the width and breadth of the epiphyses by amounts that varied widely across taxa. To predict the effect of salamander-like cartilage correction factors on muscle leverage, a simplified model of the hindlimb of the Devonian stem tetrapodAcanthostegawas built. In this model, the lever arms of muscles that cross the hip at an oblique angle to the femur was increased by up to six centimeters. Future reconstructions of osteological range of motion and muscle leverage in stem tetrapods and stem amphibians can be made more rigorous by explicitly considering the possible effects of unpreserved cartilage and justifying assumptions based on available data from extant taxa, including aquatic and terrestrial salamanders.

Microcomputed tomography of the holotype of the early tetrapodIchthyerpeton bradleyae(Huxley in Wright and Huxley, 1866) from the Pennsylvanian of Ireland

AbstractIchthyerpeton bradleyae(Huxley in Wright and Huxley, 1866) is one of the seven tetrapods originally described by Huxley from the Jarrow Assemblage (Pennsylvanian, Langsettian Regional Substage equated with the Bashkirian International Stage) in south-eastern Ireland. The holotype, one of only two specimens considered to represent the taxon, consists of the postcranial skeleton, which has been highly compressed and has undergone extensive replacement of bone by carbonaceous material. The holotype is studied using microcomputed tomography, which reveals that the vertebral column has at least 25 diplospondylous vertebrae with cylindrical centra. Neural arches and a haemal arch are described for the first time. Neural arches in the caudal region are paired and neural spines only contact one another dorsally. The hemal arch is fused and wraps around the ventral margin of the centrum. A stout femur and tibia are described. The morphology of the femur is unique for early tetrapods, with fibular and tibial condyles of similar length and lacking an adductor crest. The morphology of the femur, and its length relative to the tibia, suggests that the holotype ofI. bradleyaepreserves an immature individual. The tibia is a flat bone characteristic of stem tetrapods. Phalanges from the right and left pes are present. Because the phalanges are disarticulated, a phalangeal count cannot be determined. Despite the new anatomical information, the systematic position ofI. bradleyaeis still difficult to establish; however, it does not belong within the colosteids, temnospondyls, or embolomeres, to which it has previously been assigned.

Functional adaptive landscapes predict terrestrial capacity at the origin of limbs.

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 tetrapodlandscape, 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.

Limb-Bone Development of Seymouriamorphs: Implications for the Evolution of Growth Strategy in Stem Amniotes

Tetrapod life on land was the result of a lengthy process, the final steps of which resulted in full independence of amniotic tetrapods from the aquatic environment. Developmental strategies, including growth rate and the attainment of sexual maturity, played a major role in this transition. Early amniotes, such as Ophiacodon, tended to reach sexual maturity in a year while most non-amniotic Paleozoic tetrapods (including Devonian tetrapods and temnospondyls) became adult after 3 to 11 years. This ontogenetic transition is accompanied by a drastic change in growth rate and bone microstructure suggesting faster growth dynamics in early amniotes than in Devonian tetrapods and temnospondyls. Was the acquisition of a faster development (earlier sexual maturity and faster growth rate) a drastic evolutionary event or an extended process over geological time? To answer this question, the limb bone histology of two Early Permian (i.e. 270-290 million-year-old) stem-amniote seymouriamorphs, Seymouria sanjuanensis and Discosauriscus austriacus, were investigated. We used three-dimensional bone paleohistology based on propagation phase-contrast synchrotron microtomography. Both seymouriamorphs display relatively fast bone growth and dynamics (even though cyclic in the humerus of D. austriacus). This significantly contrasts with the slow primary bone deposition encountered in the stylopods of temnospondyls and Devonian (i.e. 360 million-year-old) stem tetrapods of similar sizes. On the basis of skeletochronological data, the seymouriamorph D. austriacus retained a long pre-reproductive period as observed in Devonian tetrapods and most temnospondyls. The combination of characteristics (faster growth rate but long pre-reproductive period) suggests that the shift towards an amniotic developmental strategy was an extended process in the evolutionary history of amniotes.

Fin ray patterns at the fin-to-limb transition

The fin-to-limb transition was marked by the origin of digits and the loss of dermal fin rays. Paleontological research into this transformation has focused on the evolution of the endoskeleton, with little attention paid to fin ray structure and function. To address this knowledge gap, we study the dermal rays of the pectoral fins of 3 key tetrapodomorph taxa-Sauripterus taylori (Rhizodontida), Eusthenopteron foordi (Tristichopteridae), and Tiktaalik roseae (Elpistostegalia)-using computed tomography. These data show several trends in the lineage leading to digited forms, including the consolidation of fin rays (e.g., reduced segmentation and branching), reduction of the fin web, and unexpectedly, the evolution of asymmetry between dorsal and ventral hemitrichia. In Eusthenopteron, dorsal rays cover the preaxial endoskeleton slightly more than ventral rays. In Tiktaalik, dorsal rays fully cover the third and fourth mesomeres, while ventral rays are restricted distal to these elements, suggesting the presence of ventralized musculature at the fin tip analogous to a fleshy "palm." Asymmetry is also observed in cross-sectional areas of dorsal and ventral rays. Eusthenopteron dorsal rays are slightly larger than ventral rays; by contrast, Tiktaalik dorsal rays can be several times larger than ventral rays, and degree of asymmetry appears to be greater at larger sizes. Analysis of extant osteichthyans suggests that cross-sectional asymmetry in the dermal rays of paired fins is plesiomorphic to crown group osteichthyans. The evolution of dermal rays in crownward stem tetrapods reflects adaptation for a fin-supported elevated posture and resistance to substrate-based loading prior to the origin of digits.

Carboniferous-Permian climate change constrained early land vertebrate radiations.

The Carboniferous-Permian transition (CPT) was Earth's last pre-Quaternary icehouse-greenhouse transition, recording major shifts in late Palaeozoic climate regimes and increased continental seasonality over approximately 40 Myr. Its parallels to Quaternary climate change have inspired recent investigations into the impacts of purported rainforest collapse on palaeotropical vertebrate diversity, but little is known about how the protracted spatial dynamics of this transition impacted the emergence of modern tetrapod lineages. Here, we apply ecological ordinance analyses on a dataset of 286 CPT fossil vertebrate localities binned across four physiographic regions forming a palaeoequatorial transect. Our results clarify the spatiotemporal expansion of land-living vertebrates, demonstrating that the reduction of tropical wetlands accommodated emerging dryland-adapted amniote faunas from a western Pangaean epicentre. We call this west-east lag the 'Vaughn-Olson model': CPT climatic transitions were regionally diachronous with delayed proliferation of amniote-dominated dryland assemblages in the east. By combining our ecological analyses with a phylogenetic approach, we demonstrate that this pattern also applies to some co-occurring total-group amphibians, suggesting that there was pervasive selection for such dryland adaptations across the crown tetrapod tree, in contrast with stem tetrapods and 'fishes'.

Unique pelvic fin in a tetrapod-like fossil fish, and the evolution of limb patterning

All living tetrapods have a one-to-two branching pattern in the embryonic proximal limb skeleton, with a single element at the base of the limb (the humerus or femur) that articulates distally with two parallel radials (the ulna and radius or the tibia and fibula). This pattern is also seen in the fossilized remains of stem-tetrapods, including the fishlike members of the group, in which despite the absence of digits, the proximal parts of the fin skeleton clearly resemble those of later tetrapods. However, little is known about the developmental mechanisms that establish and canalize this highly conserved pattern. We describe the well-preserved pelvic fin skeleton of Rhizodus hibberti, a Carboniferous sarcopterygian (lobe-finned) fish, and member of the tetrapod stem group. In this specimen, three parallel radials, each robust with a distinct morphology, articulate with the femur. We review this unexpected morphology in a phylogenetic and developmental context. It implies that the developmental patterning mechanisms seen in living tetrapods, now highly constrained, evolved from mechanisms flexible enough to accommodate variation in the zeugopod (even between pectoral and pelvic fins), while also allowing each element to have a unique morphology.

Long-bone development and life-history traits of the Devonian tristichopterid Hyneria lindae

ABSTRACTHyneria lindae is one of the largest Devonian sarcopterygians. It was found in the Catskill Formation (late Famennian) of Pennsylvania, USA. The current study focuses on the palaeohistology of the humerus of this tristichopterid and supports a low ossification rate and a late ossification onset in the appendicular skeleton. In addition to anatomical features, the large size of the cell lacunae in the cortical bone of the humerus mid-shaft may suggest a large genome size and associated neotenic condition for this species, which could, in turn, be a partial explanation for the large size of H. lindae. The low metabolism of H. lindae revealed here by bone histology supports the hypothesis of an ambush predatory behaviour. Finally, the lines-of-arrested-growth pattern and late ossification of specimen ANSP 21483 suggest that H. lindae probably had a long juvenile stage before reaching sexual maturity. Although very few studies address the life-history traits of stem tetrapods, they all propose a slow limb development for the studied taxa despite different ecological conditions and presumably distinct behaviours. The bone histology of H. lindae would favour the hypothesis that a slow long-bone development could be a general character for stem tetrapods.

New insights into the origins and radiation of the mid-Palaeozoic Gondwanan stem tetrapods

ABSTRACTThe earliest tetrapodomorph fishes appear in Chinese deposits of Early Devonian age, and by the Middle Devonian they were widespread globally. Evidence for the earliest digitated tetrapods comes from largely uncontested Middle Devonian trackways and Late Devonian body fossils. The East Gondwana Provence (Australasia, Antarctica) fills vital gaps in the phylogenetic and biogeographic history of the tetrapods, with the Gondwanan clade Canowindididae exhibiting a high degree of endemism within the early part of the stem tetrapod radiation. New anatomical details of Koharalepis, from the Middle Devonian Aztec Siltstone of Antarctica, are elucidated from synchrotron scan data. These include the position of the orbit, the condition of the hyomandibular, the shape of the palate and arrangement of the vomerine fangs. Biogeographical and phylogenetic models of stem tetrapod origins and radiations are discussed.

An enigmatic braincase from Five Points, Ohio (Westphalian D) further supports a stem tetrapod position for aïstopods

ABSTRACTWe describe a new specimen of the aïstopodOestocephalusfrom Five Points, Ohio, which preserves much of the posterior braincase. The specimen, the largest aïstopod skull described, preserves the postorbital region to the occiput. The posterior braincase has coossified the basioccipital, exoccipitals, and opisthotic. The parasphenoid is rostrally restricted, toothless, and highly vaulted along the cultriform process. The lateral walls of the cultriform process are further reinforced by large longitudinally running, ventral flanges from the parietals. Two large endochondral ventral projections from the basioccipital, previously interpreted as basal tuberosities for hypaxial muscle insertion, are here instead interpreted as articulations for the branchial skeleton. This interpretation is further supported by traces of vasculature that is consistent with what is seen in gill-bearing species. A model for the reorganisation of the basicranial region on the transition from hyomandibula to stapes is proposed, which suggests that gills, or gill-support skeletal elements, might be further distributed along the tetrapod stem than previously thought. These data further support the placement of aïstopods in the tetrapod stem group and require a reconsideration of our understanding of early tetrapod evolution.