Functional variation of neck muscles and their relation to feeding style in Tyrannosauridae and other large theropod dinosaurs.

Reconstructed neck muscles of large theropod dinosaurs suggest influences on feeding style that paralleled variation in skull mechanics. In all examined theropods, the head dorsiflexor m. transversospinalis capitis probably filled in the posterior dorsal concavity of the neck, for a more crocodilian- than avian-like profile in this region. The tyrannosaurine tyrannosaurids Daspletosaurus and Tyrannosaurus had relatively larger moment arms for latero-flexion by m. longissimus capitis superficialis and m. complexus than albertosaurine tyrannosaurids, and longer dorsiflexive moment arms for m. complexus. Areas of dorsiflexor origination are significantly larger relative to neck length in adult Tyrannosaurus rex than in other tyrannosaurids, suggesting relatively large muscle cross-sections and forces. Tyrannosaurids were not particularly specialized for neck ventro-flexion. In contrast, the hypothesis that Allosaurus co-opted m. longissimus capitis superficialis for ventro-flexion is strongly corroborated. Ceratosaurus had robust insertions for the ventro-flexors m. longissimus capitis profundus and m. rectus capitis ventralis. Neck muscle morphology is consistent with puncture-and-pull and powerful shake feeding in tyrannosaurids, relatively rapid strikes in Allosaurus and Ceratosaurus, and ventroflexive augmentation of weaker jaw muscle forces in the non tyrannosaurids.