Courtship vocalizations in male ducks: spectral composition and resonance of the syringeal bulla.

Ducks display a unique and dramatic sexual dimorphism in their vocal organ, the syrinx. Males have a left-sided bulla that is not present in females and that has been long hypothesized to play a role in courtship vocalizations, though this connection has never been tested. The large, hollow morphology of the bulla and its proximity to the sound-producing vocal folds introduce the possibility that it may work as a Helmholtz resonator, which makes it possible to predict the resonance frequencies enhanced by this structure. We found that during early ontogeny, the distribution of energy across the harmonic spectrum of contact calls is not different between males and females. We then used microcomputed tomography (µCT) scans of duck syringes to estimate resonance frequencies of the bullae and compared these with spectral features of their vocalizations. This comparison overall supports the idea that the bulla resonance may specifically enhance aspects of courtship vocalizations, especially in species that have a tonal courtship whistle. This was further supported when we tested the frequencies produced when air was blown through 3D printed bullae. We also saw potential influence of the bulla in non-courtship vocalizations, which could be explored further with a greater understanding of the input of other vocal tract features that influence vocalization. We observed that, in general, excepting the common eider, bulla size shows a weak positive correlation with male bird body mass. This study provides support for the long-held hypothesis that the adult male duck bulla influences resonance frequencies, in particular in courtship vocalizations.