The Devonian tetrapodAcanthostega gunnariJarvik: postcranial anatomy, basal tetrapod interrelationships and patterns of skeletal evolution

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Abstract
The postcranial skeleton of Acanthostega gunnari from the Famennian of East Greenland displays a unique, transitional, mixture of features conventionally associated with fishand tetrapod-like morphologies. The rhachitomous vertebral column has a primitive, barely differentiated atlas-axis complex, encloses an unconstricted notochordal canal, and the weakly ossified neural arches have poorly developed zygapophyses. More derived axial skeletal features include caudal vertebral proliferation and, transiently, neural radials supporting unbranched and unsegmented lepidotrichia. Sacral and post-sacral ribs reiterate uncinate cervical and anterior thoracic rib morphologies: a simple distal flange supplies a broad surface for iliac attachment. The octodactylous forelimb and hindlimb each articulate with an unsutured, foraminate endoskeletal girdle. A broad-bladed femoral shaft with extreme anterior torsion and associated flattened epipodials indicates a paddle-like hindlimb function. Phylogenetic analysis places Acanthostega as the sister-group of Ichthyostega plus all more advanced tetrapods. Tulerpeton appears to be a basal stemamniote plesion, tying the amphibian-amniote split to the uppermost Devonian. Caerorhachis may represent a more derived stem-amniote plesion. Postcranial evolutionary trends spanning the taxa traditionally associated with the fish-tetrapod transition are discussed in detail. Comparison between axial skeletons of primitive tetrapods suggests that plesiomorphic fish-like morphologies were re-patterned in a cranio-caudal direction with the emergence of tetrapod vertebral regionalisation. The evolution of digited limbs lags behind the initial enlargement of endoskeletal girdles, whereas digit evolution precedes the elaboration of complex carpal and tarsal articulations. Pentadactylous limbs appear to have stabilised independently in amniote and amphibian lineages; the colosteid Greererpeton has a pentadactylous manus, indicating that basal amphibian forelimbs may not be restricted to patterns of four digits or less.