Soft tissues influence nasal airflow in diapsids: Implications for dinosaurs.

The nasal passage performs multiple functions in amniotes, including olfaction and thermoregulation. These functions would have been present in extinct animals as well. However, fossils preserve only low-resolution versions of the nasal passage due to loss of soft-tissue structures after death. To test the effects of these lower resolution models on interpretations of nasal physiology, we performed a broadly comparative analysis of the nasal passages in extant diapsid representatives, e.g., alligator, turkey, ostrich, iguana, and a monitor lizard. Using computational fluid dynamics, we simulated airflow through 3D reconstructed models of the different nasal passages and compared these soft-tissue-bounded results to similar analyses of the same airways under the lower-resolution limits imposed by fossilization. Airflow patterns in these bony-bounded airways were more homogeneous and slower flowing than those of their soft-tissue counterparts. These data indicate that bony-bounded airway reconstructions of extinct animal nasal passages are far too conservative and place overly restrictive physiological limitations on extinct species. In spite of the diverse array of nasal passage shapes, distinct similarities in airflow were observed, including consistent areas of nasal passage constriction such as the junction of the olfactory region and main airway. These nasal constrictions can reasonably be inferred to have been present in extinct taxa such as dinosaurs.

Septal deviation in the nose of the longest faced crocodylian: A description of nasal anatomy and airflow in the Indian gharial (Gavialis gangeticus) with comments on acoustics.

The remarkably thin rostrum in the Indian gharial (Gavialis gangeticus) imparts challenges to nasal physiology. Competition for space in the slim jaws necessitates a thin nasal septum, leaving this taxon susceptible to nasal passage abnormalities such as septal deviation. Here we describe the nasal anatomy of gharials based on multiple individuals including one that showcases an extreme instance of nasal septum deviation. We found that gharials have both confluent nostrils and choanae, which may be important for their unique nasal acoustics. The deviated nasal septum in the female showed distinct waviness that affected the nasal passages by alternately compressing them. We performed a computational fluid dynamic analysis on the nasal passages to visualize the effects of septal deviation on airflow. Our analysis found the deviated septum increased nasal resistance and wall shear stress during respiration, resulting in unequal distribution of the air field between both sides of the nasal passage. Our findings indicate that gharials-and potentially other longirostrine crocodylians-may be particularly susceptible to septal deviations. Lastly, we observed pterygoid bullae to be present in both sexes, though their morphology differed. Airflow in the male pterygoid bullae produced a Bernoulli effect which may be responsible for the unique "pop" sounds recorded in this species.

Convoluted nasal passages function as efficient heat exchangers in ankylosaurs (Dinosauria: Ornithischia: Thyreophora).

Convoluted nasal passages are an enigmatic hallmark of Ankylosauria. Previous research suggested that these convoluted nasal passages functioned as heat exchangers analogous to the respiratory turbinates of mammals and birds. We tested this hypothesis by performing a computational fluid dynamic analysis on the nasal passages of two ankylosaurs: Panoplosaurus mirus and Euoplocephalus tutus. Our models predicted that Panoplosaurus and Euoplocephalus would have required 833 and 1568 thermal calories, respectively, to warm a single breath of air by 20°C. Heat recovery during exhalation resulted in energy savings of 65% for Panoplosaurus and 84% for Euoplocephalus. Our results fell well within the range of values for heat and water savings observed in extant terrestrial amniotes. We further tested alternate airway reconstructions that removed nasal passage convolutions or reduced nasal vestibule length. Our results revealed that the extensive elaboration observed in the nasal vestibules of ankylosaurs was a viable alternative to respiratory turbinates with regards to air conditioning. Of the two dinosaurs tested, Euoplocephalus repeatedly exhibited a more efficient nasal passage than Panoplosaurus. We suggest that the higher heat loads associated with the larger body mass of Euoplocephalus necessitated these more efficient nasal passages. Our findings further indicate that the evolution of complicated airways in dinosaurs may have been driven by the thermal requirements of maintaining cerebral thermal homeostasis.

Nasal conchae function as aerodynamic baffles: Experimental computational fluid dynamic analysis in a turkey nose (Aves: Galliformes).

We tested the aerodynamic function of nasal conchae in birds using CT data from an adult male wild turkey (Meleagris gallopavo) to construct 3D models of its nasal passage. A series of digital "turbinectomies" were performed on these models and computational fluid dynamic analyses were performed to simulate resting inspiration. Models with turbinates removed were compared to the original, unmodified control airway. Results revealed that the four conchae found in turkeys, along with the crista nasalis, alter the flow of inspired air in ways that can be considered baffle-like. However, these baffle-like functions were remarkably limited in their areal extent, indicating that avian conchae are more functionally independent than originally hypothesized. Our analysis revealed that the conchae of birds are efficient baffles that-along with potential heat and moisture transfer-serve to efficiently move air to specific regions of the nasal passage. This alternate function of conchae has implications for their evolution in birds and other amniotes.

Breathing life into dinosaurs: tackling challenges of soft-tissue restoration and nasal airflow in extinct species.

The nasal region plays a key role in sensory, thermal, and respiratory physiology, but exploring its evolution is hampered by a lack of preservation of soft-tissue structures in extinct vertebrates. As a test case, we investigated members of the "bony-headed" ornithischian dinosaur clade Pachycephalosauridae (particularly Stegoceras validum) because of their small body size (which mitigated allometric concerns) and their tendency to preserve nasal soft tissues within their hypermineralized skulls. Hypermineralization directly preserved portions of the olfactory turbinates along with an internal nasal ridge that we regard as potentially an osteological correlate for respiratory conchae. Fossil specimens were CT-scanned, and nasal cavities were segmented and restored. Soft-tissue reconstruction of the nasal capsule was functionally tested in a virtual environment using computational fluid dynamics by running air through multiple models differing in nasal soft-tissue conformation: a bony-bounded model (i.e., skull without soft tissue) and then models with soft tissues added, such as a paranasal septum, a scrolled concha, a branched concha, and a model combining the paranasal septum with a concha. Deviations in fluid flow in comparison to a phylogenetically constrained sample of extant diapsids were used as indicators of missing soft tissue. Models that restored aspects of airflow found in extant diapsids, such as appreciable airflow in the olfactory chamber, were judged as more likely. The model with a branched concha produced airflow patterns closest to those of extant diapsids. These results from both paleontological observation and airflow modeling indicate that S. validum and other pachycephalosaurids could have had both olfactory and respiratory conchae. Although respiratory conchae have been linked to endothermy, such conclusions require caution in that our re-evaluation of the reptilian nasal apparatus indicates that respiratory conchae may be more widespread than originally thought, and other functions, such as selective brain temperature regulation, could be important.