Person:

Jenkins, Farish

The scant fossil record of caecilians has obscured the origin and evolution of this lissamphibian group. Eocaecilia micropodia from the Lower Jurassic of North America remains the only stem-group caecilian with an almost complete skull preserved. However, this taxon has been controversial, engendering re-evaluation of traits considered to be plesiomorphic for extant caecilians. Both the validity of the placement of E. micropodia as a stem caecilian and estimates of the plesiomorphic condition of extant caecilians have been questioned. In order to address these issues, the braincase of E. micropodia was examined via micro-computed tomography. The braincase is considered to be a more reliable phylogenetic indicator than peripheral regions of the skull. These data reveal significant new information, including the possession of an ossified nasal septum, ossified anterior wall of the sphenethmoid, long anterolateral processes on the sphenethmoid, and paired olfactory nerve foramina, which are known only to occur in extant caecilians; the latter are possibly related to the evolution of the tentacle, a caecilian autapomorphy. A phylogenetic analysis that included 64 non-amniote taxa and 308 characters represents the first extensive test of the phylogenetic affinities of E. micropodia. The results place E. micropodia securely on the stem of extant caecilians, representing a clade within Temnospondyli that is the sister taxon to batrachians plus Gerobatrachus. Ancestral character state reconstruction confirms the braincase of E. micropodia to be largely representative of the plesiomorphic condition of extant caecilians. Additionally, the results refine the context within which the evolution of the caecilian form can be evaluated. The robust construction and pattern of the dermal skull of E. micropodia is interpreted as symplesiomorphic with advanced dissorophoid temnospondyls, rather than being autapomorphic in its robust construction. Together these data increase confidence in incorporating E. micropodia into discussions of caecilian evolution.

Three-dimensional skeletal movement is often impossible to accurately quantify from external markers. X-ray imaging more directly visualizes moving bones, but extracting 3-D kinematic data is notoriously difficult from a single perspective. Stereophotogrammetry is extremely powerful if biplanar fluoroscopy is available, yet implantation of three radio-opaque markers in each segment of interest may be impractical. Herein we introduce scientific rotoscoping (SR), a new method of motion analysis that uses articulated bone models to simultaneously animate and quantify moving skeletons without markers. The three-step process is described using examples from our work on pigeon flight and alligator walking. First, the experimental scene is reconstructed in 3-D using commercial animation software so that frames of undistorted fluoroscopic and standard video can be viewed in their correct spatial context through calibrated virtual cameras. Second, polygonal models of relevant bones are created from CT or laser scans and rearticulated into a hierarchical marionette controlled by virtual joints. Third, the marionette is registered to video images by adjusting each of its degrees of freedom over a sequence of frames. SR outputs high-resolution 3-D kinematic data for multiple, unmarked bones and anatomically accurate animations that can be rendered from any perspective. Rather than generating moving stick figures abstracted from the coordinates of independent surface points, SR is a morphology-based method of motion analysis deeply rooted in osteological and arthrological data.

The skull morphology of Kayentachelys aprix Gaffney et al., 1987, a turtle from the Early Jurassic Kayenta Fm of northern Arizona, demonstrates the presence of cryptodiran synapomorphies in agreement with Gaffney et al. (1987, 1991, 2007), and contrary to the conclusions of Sterli and Joyce (2007), Joyce (2007), Sterli (2008), and Anquetin et al. (2008). Specific characters found in Kayentachelys and diagnostic of cryptodires include the processus trochlearis oticum, the curved processus pterygoideus externus with a vertical plate, and the prefrontal–vomer contact, which are confirmed as absent in the outgroups, specifically the Late Triassic Proganochelys. The Joyce (2007) analysis suffers from the reduction of the signal from skull characters, with a consequently greater reliance on shell characters, resulting in pleurodires being resolved at various positions within the cryptodires. Kayentachelys reveals what a primitive cryptodire would be expected to look like: a combination of primitive and derived characters, with the fewer derived characters providing the best test of its relationships to other turtles. Although incompletely known, the Mid-Late Jurassic Condorchelys, Heckeremys, and Eileanchelys may be early cryptodires close to Kayentachelys. We confirm the Late Triassic Proterochersis as a pleurodire, dating the pleurodire–cryptodire split as Late Triassic or earlier.

Among the morphological changes that occurred during the ‘fish-to-tetrapod’ transition was a marked reorganization of the cranial endoskeleton. Details of this transition, including the sequence of character acquisition, have not been evident from the fossil record. Here we describe the braincase, palatoquadrate and branchial skeleton of Tiktaalik roseae, the Late Devonian sarcopterygian fish most closely related to tetrapods. Although retaining a primitive configuration in many respects, the cranial endoskeleton of T. roseae shares derived features with tetrapods such as a large basal articulation and a flat, horizontally oriented entopterygoid. Other features in T. roseae, like the short, straight hyomandibula, show morphology intermediate between the condition observed in more primitive fish and that observed in tetrapods. The combination of characters in T. roseae helps to resolve the relative timing of modifications in the cranial endoskeleton. The sequence of modifications suggests changes in head mobility and intracranial kinesis that have ramifications for the origin of vertebrate terrestriality.

Vertebrate claws are used in a variety of important behaviours and are typically composed of a keratinous sheath overlying the terminal phalanx of a digit. Keratinous claws, however, are rare in living amphibians; their microstructure and other features indicate that they probably originated independently from those in amniotes. Here we show that certain African frogs have a different type of claw, used in defence, that is unique in design among living vertebrates and lacks a keratinous covering. These frogs have sectorial terminal phalanges on their hind feet that become functional by cutting through the skin. In the resting state, the phalanx is subdermal and attached to a distal bony nodule, a neomorphic skeletal element, via collagen-rich connective tissue. When erected, the claw breaks free from the nodule and pierces the ventral skin. The nodule, suspended by a sheath attached to the terminal phalanx and supported by collagenous connections to the dermis, remains fixed in place. While superficially resembling the shape of claws in other tetrapods, these are the only vertebrate claws known to pierce their way to functionality.

The plagiosaurine Gerrothorax pulcherrimus, a ubiquitous faunal component of the Fleming Fjord Formation, is recognized by tubercular ornamentation, contact between the postfrontal and supratemporal, at least two tooth rows on the posterior coronoid, and a posttemporal fenestra that is small or absent. Gerrothorax pulcherrimus also possesses a derived pectoral morphology that includes an interlocking claviculocleithral complex and an interclavicle with paired posterolateral projections and a transversely truncated posterior margin. Gerrothorax pustuloglomeratus is a junior synonym of G. pulcherrimus; other Gerrothorax species cannot be differentiated from G. pulcherrimus. Flat-headed temnospondyls have been interpreted as having achieved a large gape by raising the skull rather than by lowering the jaw. No detailed, corroborating anatomical evidence from the atlanto-occipital joint, however, has ever been adduced. In G. pulcherrimus the widths of the atlantal and condylar facets are comparable, but dorsoventrally the condylar facets are 45% longer than the comparable dimension of the atlantal facets. Elevation of the skull occurred by atlanto-occipital rotation, and was facilitated by a radius of curvature of the dorsal part of the condylar facets that is shorter than that of the inferior part. From a resting, closed mouth position, G. pulcherrimus was capable of elevating the skull through an excursion of approximately 50°, a movement that rotated the quadrate forward and protruded the lower jaw. An elongate, broadly open neural canal in the atlanto-occipital region, and dorsal displacement of the occipital condyles, are related to relieving the spinal medulla of the sharp angular deformation that head lifting might entail.

The concavoconvex glenoid and bulbous humeral head of modern birds represent both structurally and functionally a hemisellar (half saddle) joint modified from a pattern common among post-Paleozoic tetrapods. Birds share with crocodilians, their nearest living relatives, similarities in joint architecture, including scapulohumeral and coracohumeral ligaments. The glenoid underwent a major reorientation during the evolution of the avian shoulder from the primitive condition of being posteroventrally directed as in Deinonychus antirrhopus and coelurosaurs generally to being dorsolaterally directed as in modern forms. The laterally facing glenoid of Archaeopteryx lithographica was intermediate in orientation and provided for a substantial degree of wing elevation but not the fully abducted, sagittal positioning that modern birds employ at the upstroke-downstroke transition.