Consilient evidence affirms expansive stabilizing ligaments in the tyrannosaurid foot

Tyrannosaurid dinosaurs were ecologically unique vertebrates as the sole clade of large terrestrial carnivores (adults >400 kg) in their continent-spanning habitats. Expanded ligaments between metatarsals, inferred by gross morphology of attachment correlates, have been hypothesized to have strengthened their specialized arctometatarsus. We tested the hypothesis of ligament presence with scanning electron microscopy and histological thin sections of putative attachment sites in a third metatarsal of the tyrannosaurid Gorgosaurus libratus, compared with a thin section from the unspecialized metatarsals of the early theropod Coelophysis bauri. In the Gorgosaurus metatarsal, Sharpey's fibers and rough, pitted surface textures consistent with ligament coalescence occur at expansive distal regions and localized rugosities are ideally located for resisting torsional loading on the foot. Sparser Sharpey’s fibers occur at expected locations in other arctometatarsus-bearing coelurosaurs. In contrast, the Coelophysis metatarsal lacked Sharpey’s fibers or rugosity at the sectioned location, acting as a definitive negative control for identifying these features in tyrannosaurids. With soft-tissue correlates confirmed as ligament entheses, we conclude that tyrannosaurids possessed distinctive and specific ligament attachments to the third metatarsal lacking in other large carnivorous dinosaurs. Histological evidence for extensive distal intermetatarsal ligaments is consistent with greater inferred agility in derived tyrannosaurid dinosaurs than in earlier lineages of large theropods.