Evolution of the respiratory system in nonavian theropods: evidence from rib and vertebral morphology

Recent reports of region-specific vertebral pneumaticity in nonavian theropod dinosaurs have brought attention to the hypothesis that these animals possessed an avian-style respiratory system with flow-through ventilation. This study explores the thoracic rib and vertebral anatomy of Sinraptor, Allosaurus, Tyrannosaurus, and Deinonychus; four nonavian theropods that all show well-preserved thoracic vertebrae and ribs. Comparisons to the osteology and soft tissue anatomy of extant saurians provide new evidence supporting the hypothesis of flow-through ventilation in nonavian theropods. Analyses of diapophyseal and parapophyseal position and thoracic rib morphology suggest that most nonavian theropods possessed lungs that were deeply incised by the adjacent bicapitate thoracic ribs. This functionally constrains the lungs as rigid nonexpansive organs that were likely ventilated by accessory nonvascularized air sacs. The axial anatomy of this group also reveals that a crocodilian-like hepatic-piston lung would be functionally and biomechanically untenable. Taken together with the evidence that avian-like air sacs were present in basal theropods, these data lead us to conclude that an avian-style pulmonary system was likely a universal theropod trait.