What makes the diverse flight of birds possible? Phylogenetic comparative analysis of avian alula morphology

Studies on the evolution of avian flight have failed to clarify why wing shape is weakly correlated with flight styles and phylogenetically constrained. Birds seem to have achieved their diverse flight styles owing to changes in the shapes of localized parts of the wing, despite a highly conserved wing outline. The alula, which stems from the first digit of the wing, is a part of the wing that might have enabled various flight styles to develop. To test this hypothesis, we measured alula length and width using museum specimens of 187 species from 21 orders of birds and used a phylogenetic comparative method to investigate the relationship between alula morphology and flight style, body mass, habitat and migratory distance. We found that: (1) phylogenetic signals were weak for the alula width and aspect ratio but moderate for length; (2) alula morphology exhibited an allometric relationship with body mass; and (3) fewer flapping birds had large alulae. Alula morphology might have evolved in response to changes in body mass and flight styles. Variations in the shape of localized wing parts, such as the alula, which modifies airflow around a wing, might be key to the evolution of the diverse flight styles of birds.