Functional morphology of the forelimb musculature reflects flight and foraging styles in aquatic birds

Aquatic birds show a great diversity of locomotion styles and wing morphologies, from penguins that are fully specialized for an aquatic life to species of aerial flyers that also use their wings for underwater propulsion (e.g. auks and shearwaters). Moving between the air–water interface exerts conflicting pressures on the body and wing anatomy of diving birds. In this work, we investigated the functional morphology of the forelimb musculature of 18 species of aquatic birds that display a variety of flight and foraging styles. Muscle architecture was related to function, with special emphasis on muscle mass. Dissections of one of the forelimbs of 20 specimens of waterbirds were performed to obtain numerical data of muscle architecture. Total wing muscle mass scaled isometrically to body mass1.0, whereas fascicle length scaled to muscle mass0.284, which is consistent with previous results of scaling in wings of raptors. A principal component analysis (PCA) of normalised muscle masses resulted in a biplot where three main morphological groups can be distinguished. Anatids (ducks and geese) occupy a space represented by muscles that are activated during downstroke. Auks and penguins clustered together in a region dominated by muscles that assist in wing elevation and showed a degree of hypertrophy. The rest of the species grouped together in the lower limits of both PCs where muscles that facilitate wing flexion–extension and stabilisation are loaded. The distribution of mass in the wing muscles of the aquatic birds seemed to be related to flight and foraging style and showed non-significant influence of shared phylogenetic history (Kmult: 0.71, p value: 0.083, 10,000 permutations).