Endocranial development in non-avian dinosaurs reveals an ontogenetic brain trajectory distinct from extant archosaurs.

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form.

Ontogenetic endocranial shape change in alligators and ostriches and implications for the development of the non-avian dinosaur endocranium.

Birds and crocodiles show radically different patterns of brain development, and it is of interest to compare these to determine the pattern of brain growth expected in dinosaurs. Here we provide atlases of 3D brain (endocast) reconstructions for Alligator mississippiensis (alligator) and Struthio camelus (ostrich) through ontogeny, prepared as digital restorations from CT scans of stained head and dry skull specimens. Our morphometric analysis confirms that ostrich brains do not change significantly in shape during postnatal growth, whereas alligator brains unfold from a cramped bird-like shape in the hatchling to an elongate, straight structure in the adult. We confirm that birds exhibit paedomorphic dinosaur endocranial traits such as retaining an enlarged and compact brain shape in the adult, whereas crocodiles show peramorphic traits where the brain elongates with growth as the skull elongates. These atlases of ontogenetic stages of modern bird and crocodilian endocrania provide a basis for comparison of non-avian dinosaur endocasts and consideration of the divergence of the "avian" and "crocodilian" modes of brain development and heterochronic change on phylogenies.

Ontogeny of the Middle-Ear Air-Sinus System in Alligator mississippiensis (Archosauria: Crocodylia).

Modern crocodylians, including Alligator mississippiensis, have a greatly elaborated system of pneumatic sinuses invading the cranium. These sinuses invade nearly all the bones of the chondrocranium and several bony elements of the splanchnocranium, but patterns of postnatal paratympanic sinus development are poorly understood and documented. Much of crocodylomorph--indeed archosaurian--evolution is characterized by the evolution of various paratympanic air sinuses, the homologies of which are poorly understood due in large part to the fact that individual sinuses tend to become confluent in adults, obscuring underlying patterns. This study seeks to explore the ontogeny of these sinuses primarily to clarify the anatomical relations of the individual sinuses before they become confluent and thus to provide the foundation for later studies testing hypotheses of homology across extant and extinct Archosauria. Ontogeny was assessed using computed tomography in a sample of 13 specimens covering an almost 19-fold increase in head size. The paratympanic sinus system comprises two major inflations of evaginated pharyngeal epithelium: the pharyngotympanic sinus, which communicates with the pharynx via the lateral (true) Eustachian tubes and forms the cavum tympanicum proprium, and the median pharyngeal sinus, which communicates with the pharynx via the median pharyngeal tube. Each of these primary inflations gives rise to a number of secondary inflations that further invade the bones of the skull. The primary sinuses and secondary diverticula are well developed in perinatal individuals of Alligator, but during ontogeny the number and relative volumes of the secondary diverticula are reduced. In addition to describing the morphological ontogeny of this sinus system, we provide some preliminary exploratory analyses of sinus function and allometry, rejecting the hypothesis that changes in the volume of the paratympanic sinuses are simply an allometric function of braincase volume, but instead support the hypothesis that these changes may be a function of the acoustic properties of the middle ear.