The epipterygoid of crocodyliforms and its significance for the evolution of the orbitotemporal region of eusuchians

New fossils of the giant African crocodyliform Sarcosuchus imperator clarify its skeletal anatomy, growth patterns, size, longevity, and phylogenetic position. The skull has an expansive narial bulla and elongate jaws studded with stout, smooth crowns that do not interlock. The jaw form suggests a generalized diet of large vertebrates, including fish and dinosaurs. S. imperator is estimated to have grown to a maximum body length of at least 11 to 12 meters and body weight of about 8 metric tons over a life-span of 50 to 60 years. Unlike its closest relatives, which lived as specialized piscivores in marginal marine habitats, S. imperator thrived in fluvial environments.

The homologies of jaw muscles among archosaurs and other sauropsids have been unclear, confounding interpretation of adductor chamber morphology and evolution. Relevant topological patterns of muscles, nerves, and blood vessels were compared across a large sample of extant archosaurs (birds and crocodylians) and outgroups (e.g., lepidosaurs and turtles) to test the utility of positional criteria, such as the relative position of the trigeminal divisions, as predictors of jaw muscle homology. Anatomical structures were visualized using dissection, sectioning, computed tomography (CT), and vascular injection. Data gathered provide a new and robust view of jaw muscle homology and introduce the first synthesized nomenclature of sauropsid musculature using multiple lines of evidence. Despite the great divergences in cephalic morphology among birds, crocodylians, and outgroups, several key sensory nerves (e.g., n. anguli oris, n. supraorbitalis, n. caudalis) and arteries proved useful for muscle identification, and vice versa. Extant crocodylians exhibit an apomorphic neuromuscular pattern counter to the trigeminal topological paradigm: the maxillary nerve runs medial, rather than lateral to M. pseudotemporalis superficialis. Alternative hypotheses of homology necessitate less parsimonious interpretations of changes in topology. Sensory branches to the rictus, external acoustic meatus, supraorbital region, and other cephalic regions suggest conservative dermatomes among reptiles. Different avian clades exhibit shifts in some muscle positions, but maintain the plesiomorphic, diapsid soft-tissue topological pattern. Positional data suggest M. intramandibularis is merely the distal portion of M. pseudotemporalis separated by an intramuscular fibrocartilaginous sesamoid. These adductor chamber patterns indicate multiple topological criteria are necessary for interpretations of soft-tissue homology and warrant further investigation into character congruence and developmental connectivity. Copyright (c) 2007 Wiley-Liss, Inc.