Postcranial Skeletal Pneumaticity in Cuculidae.

Postcranial skeletal pneumaticity (i.e., epithelial-lined, air-filled bones) is a condition unique to birds among extant tetrapods. Previous research reveals extensive variation in the expression of this trait in different bird species, from taxa that pneumatize nearly the entire skeleton to others that do not pneumatize a single bone. These studies, however, have primarily focused on aquatic/semi-aquatic birds, specifically Anseriformes (screamers, ducks, geese, swans) and Aequorlitornithes (loons, gulls, penguins, storks, etc.). This is the first clade-centric study of pneumaticity in an exclusively terrestrial clade (i.e., a group without any proclivities for water), Cuculidae. Given the variation in body size and ecology exhibited by cuckoos, they represent an ideal group for evaluating previously established trends in pneumaticity patterns. Similar to previous studies, our results indicate that cuckoos do exhibit extensive postcranial skeletal pneumaticity but with much more limited variation in expression. Of the surveyed species, 30 of 41 display an identical expression pattern, pneumatizing all postaxial vertebrae, the humerus, sternum, and pelvic girdle. The remaining species (11/41) deviate from this pattern by no more than two elements (i.e., the femur or the scapula/coracoid). All variable species expand upon the basic cuckoo pattern, with five species pneumatizing the femur and the remaining six taxa pneumatizing both the scapula and coracoid. Furthermore, most variation occurs in early diverging clades, with distinct subclades associated with specific anatomical expansions in pneumaticity (e.g., pneumatic femora in Neomorphinae and pneumatic scapulae/coracoids in select members of Couinae and Centropodinae). Limited variation noted in Cuculidae may be the result of the relatively high base level of pneumaticity when compared with previously sampled groups of water-oriented birds. Additional analyses indicate a positive relationship between body mass and pneumaticity, with possible (i.e., non-quantifiable) relationships noted between the limited expansions from the basic cuckoo pattern and specific locomotor behaviors (e.g., pneumatic femora present in species with enhanced cursorial behavior). These basic trends have also been observed in other densely sampled neognath clades. Taken together, the data presented herein supports the hypothesis that changes in pneumaticity expression may be correlated with shifts in biomechanical loading regimes rather than solely as a weight saving (i.e., density-altering) mechanism.