The braincase of Euparkeria, and the evolutionary relationships of birds and crocodilians

Abstract

The Triassic reptile Euparkeria has been frequently given a pivotal position in interpretations of the evolution of archosaurs. Most recently, Welman (1995) has argued from braincase data that Euparkeria is more closely related to birds than are either theropod dinosaurs or crocodilians – a conclusion clearly at odds with the current orthodoxy. The braincase of a single specimen of Euparkeria is described in detail and compared with previous descriptions and with the braincases of other diapsids. Variations among the known specimens are documented. The homology of various braincase structures are reassessed in light of the study by Welman (1995). We argue that the braincase of Euparkeria has an undivided metotic fissure, an incompletely ossified medial wall of the otic capsule, a well-defined ‘semilunar depression’, and posteroventrally positioned foramina in the parabasisphenoid for the entrance of the cerebral branches of the internal carotid arteries. It lacks enclosure of the Eustachian system in bone, well-developed tympanic sinuses, or a well-defined recess for the lagena. A review of braincase morphology in extinct and extant diapsids suggests that braincase features of Euparkeria are largely plesiomorphic for Archosauria. The evolutionary relationships between Euparkeria and extant archosaurs (birds and crocodilians) are considered by reviewing braincase morphology in extant and extinct diapsids. No shared derived characters could be found that support the resolutions (crocodilians (Euparkeria+birds)) or (birds (Euparkeria+crocodilians)). Three derived characters shared by extant archosaurs support the resolution (Euparkeria (crocodilians+birds)), but only the presence of laterally positioned foramina in the parabasisphenoid for the entrance of the cerebral branches of the internal carotid arteries appears to represent strong evidence. The other two features are a degree of ossification (of the medial wall of the otic capsule) that exhibits some homoplasy among archosaurs, and an absence (of the ‘semilunar depression’), and therefore do not represent particularly robust hypotheses of homology. Our interpretation of the braincase of Euparkeria is fully congruent with the consensus among recent explicit phylogenetic analyses that this taxon is close to, but not a member of, the archosaur crown group. Birds and crocodilians share a number of other derived similarities (subdivided metotic fissure, elongated and tubular cochlear recess, enclosed Eustachian system, extensive tympanic sinuses, quadrate-prootic articulation) that are probably not homologous because of their absence in a number of non-avian dinosaurs and crocodilian-line crown-group archosaurs.