The relationship of limbed vertebrates (tetrapods) to lobe-finned fish (sarcopterygians) is well established, but the origin of major tetrapod features has remained obscure for lack of fossils that document the sequence of evolutionary changes. Here we report the discovery of a well-preserved species of fossil sarcopterygian fish from the Late Devonian of Arctic Canada that represents an intermediate between fish with fins and tetrapods with limbs, and provides unique insights into how and in what order important tetrapod characters arose. Although the body scales, fin rays, lower jaw and palate are comparable to those in more primitive sarcopterygians, the new species also has a shortened skull roof, a modified ear region, a mobile neck, a functional wrist joint, and other features that presage tetrapod conditions. The morphological features and geological setting of this new animal are suggestive of life in shallow-water, marginal and subaerial habitats.The evolution of tetrapods from sarcopterygian fish is one of the major transformations in the history of life and involved numerous structural and functional innovations, including new modes of locomotion, respiration and hearing. Fish and tetrapod fossils across this transition can reveal how these innovations were assembled. During the origin of tetrapods in the Late Devonian (385-359 million years ago), the proportions of the skull were remodelled, the series of bones connecting the head and shoulder was lost, and the region that was to become the middle ear was modified. At the same time, robust limbs with digits evolved, the shoulder girdle and pelvis were altered, the ribs expanded, and bony connections between vertebrae developed. Few of these features, however, are seen in the closest relatives of tetrapods-the elpistostegalian fishes-which are incompletely known. Elpistostege, for example, is represented only by two partial dermal skull roofs and a segment of the axial skeleton from the early Frasnian Escuminac Formation in Quebec 1-3 . The best-known elpistostegalian, Panderichthys, consists of complete specimens of Middle to Late Devonian age (late Givetian and early Frasnian stages) mostly from the Lode quarry in Latvia 4-10 . Panderichthys possesses relatively few tetrapod synapomorphies, and provides only partial insight into the origin of major features of the skull, limbs and axial skeleton of early tetrapods. In view of the morphological gap between elpistostegalian fish and tetrapods, the phylogenetic framework for the immediate sister group of tetrapods has been incomplete and our understanding of major anatomical transformations at the fish-tetrapod transition has remained limited.The discovery of a new elpistostegalian sarcopterygian from the Fram Formation in Nunavut Territory, Canada (Fig. 1) significantly enhances our knowledge of the fish-tetrapod transition. Many articulated specimens from a single site are used to describe a taxon that is a remarkable intermediate between Panderichthys and early tetrapods. The material provides oppor...

Wrists, ankles and digits distinguish tetrapod limbs from fins, but direct evidence on the origin of these features has been unavailable. Here we describe the pectoral appendage of a member of the sister group of tetrapods, Tiktaalik roseae, which is morphologically and functionally transitional between a fin and a limb. The expanded array of distal endochondral bones and synovial joints in the fin of Tiktaalik is similar to the distal limb pattern of basal tetrapods. The fin of Tiktaalik was capable of a range of postures, including a limb-like substrate-supported stance in which the shoulder and elbow were flexed and the distal skeleton extended. The origin of limbs probably involved the elaboration and proliferation of features already present in the fins of fish such as Tiktaalik.

A tetrapod humerus from the Late Devonian of Pennsylvania has a novel mix of primitive and derived characters. A comparative analysis of this fossil and other relevant humeri from the Devonian shows that the role of the limb in propping the body arose first in fish fins, not tetrapod limbs. The functional diversity of the earliest known limbs includes several different kinds of appendage design. This functional diversity was achieved with a humeral architecture that was remarkably conserved during the Devonian.

A major challenge in understanding the origin of terrestrial vertebrates has been knowledge of the pelvis and hind appendage of their closest fish relatives. The pelvic girdle and appendage of tetrapods is dramatically larger and more robust than that of fish and contains a number of structures that provide greater musculoskeletal support for posture and locomotion. The discovery of pelvic material of the finned elpistostegalian, Tiktaalik roseae, bridges some of these differences. Multiple isolated pelves have been recovered, each of which has been prepared in three dimensions. Likewise, a complete pelvis and partial pelvic fin have been recovered in association with the type specimen. The pelves of Tiktaalik are paired and have broad iliac processes, flat and elongate pubes, and acetabulae that form a deep socket rimmed by a robust lip of bone. The pelvis is greatly enlarged relative to other finned tetrapodomorphs. Despite the enlargement and robusticity of the pelvis of Tiktaalik, it retains primitive features such as the lack of both an attachment for the sacral rib and an ischium. The pelvic fin of Tiktaalik (NUFV 108) is represented by fin rays and three endochondral elements: other elements are not preserved. The mosaic of primitive and derived features in Tiktaalik reveals that the enhancement of the pelvic appendage of tetrapods and, indeed, a trend toward hind limb-based propulsion have antecedents in the fins of their closest relatives.A t first glance, the origin of tetrapods (limbed vertebrates) from finned precursors seems an almost insurmountable transition between life in water and life on land. If the basis of comparison were living taxa alone, then the anatomical and behavioral differences among finned and limbed vertebrates could appear vast: for example, fin structure and function differ dramatically from those of limbs. Fossil evidence, in particular vertebrates from the middle and late part of the Devonian period (393-359 Mya), offers intermediate conditions that bridge this gap (1). The fossils that provide the most informative anatomical intermediates are from the tetrapodomorph lineage (also known as stem tetrapods) and have been recovered from a variety of nonmarine and marginal marine deposits from around the globe (2-4). The creatures closest to the node containing the most basal limbed vertebrates-elpistostegalids, such as Panderichthys, Tiktaalik, and Elpistostege-are most enlightening in understanding the primitive conditions from which tetrapods arose. Although most work has focused on revealing homologies and function of the pectoral appendage of these forms (4-7), relatively little is known of the pelvic appendage beyond limited material of Panderichthys (8). Consequently, analyses of the pelvic fin have been given only sporadic attention over the past decades (4, 8-11) largely because they are often poorly preserved or not preserved at all. In most cases, it is thought that this poor preservation of the pelvic appendage is due to its putative small size and fragility (10).Pelv...

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.

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An early tetrapod fossil from the Upper Devonian of Pennsylvania (Catskill Formation) extends the temporal range of tetrapods in North America and suggests that they attained a virtually global equatorial distribution by the end of the Devonian. Derived features of the shoulder girdle indicate that appendicular mechanisms of support and propulsion were well developed even in the earliest phases of tetrapod history. The specialized morphology of the pectoral skeleton implies that the diversity of early tetrapods was great and is suggestive of innovative locomotor patterns in the first tetrapods.

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