Air Space Proportion in Pterosaur Limb Bones Using Computed Tomography and Its Implications for Previous Estimates of Pneumaticity

Air Space Proportion (ASP) is a measure of how much air is present within a bone, which allows for a quantifiable comparison of pneumaticity between specimens and species. Measured from zero to one, higher ASP means more air and less bone. Conventionally, it is estimated from measurements of the internal and external bone diameter, or by analyzing cross-sections. To date, the only pterosaur ASP study has been carried out by visual inspection of sectioned bones within matrix. Here, computed tomography (CT) scans are used to calculate ASP in a small sample of pterosaur wing bones (mainly phalanges) and to assess how the values change throughout the bone. These results show higher ASPs than previous pterosaur pneumaticity studies, and more significantly, higher ASP values in the heads of wing bones than the shaft. This suggests that pneumaticity has been underestimated previously in pterosaurs, birds, and other archosaurs when shaft cross-sections are used to estimate ASP. Furthermore, ASP in pterosaurs is higher than those found in birds and most sauropod dinosaurs, giving them among the highest ASP values of animals studied so far, supporting the view that pterosaurs were some of the most pneumatized animals to have lived. The high degree of pneumaticity found in pterosaurs is proposed to be a response to the wing bone bending stiffness requirements of flight rather than a means to reduce mass, as is often suggested. Mass reduction may be a secondary result of pneumaticity that subsequently aids flight.