Neural and endocranial anatomy of Triassic phytosaurian reptiles and convergence with fossil and modern crocodylians

Phytosaurs are a clade of large, carnivorous pseudosuchian archosaurs from the Late Triassic with a near cosmopolitan distribution. Their superficial resemblance to longirostrine (long-snouted) crocodylians, such as gharials, has often been used in the past to infer ecological and behavioural convergence between the two groups. Although more than thirty species of phytosaur are currently recognised, little is known about the endocranial anatomy of this clade. Here, we describe the endocranial anatomy (including the brain, inner ear, neurovascular structures and sinus systems) of the two non-mystriosuchine phytosaurs Parasuchus angustifrons (="Paleorhinus angustifrons") and Ebrachosuchus neukami from the Late Triassic of Germany based on digital reconstructions. Results show that the endocasts of both taxa are very similar to each other in their rostrocaudally elongate morphology, with long olfactory tracts, weakly demarcated cerebral regions and dorsoventrally short endosseous labyrinths. In addition, several sinuses, including large antorbital sinuses and prominent dural venous sinuses, were reconstructed. Comparisons with the endocranial anatomy of derived phytosaurs indicate that Phytosauria is united by the presence of elongate olfactory tracts and longitudinally arranged brain architecture-characters which are also shared with Crocodyliformes. However, a substantial morphological variability is observed in the cephalic and pontine flexure and the presence of a pineal organ across the different phytosaur species. These results suggest that the endocranial anatomy in Phytosauria generally follows a plesiomorphic pattern, with moderate variation within the clade likely resulting from divergent sensory and behavioural adaptations.