Vocal constraints on song amplitude in star finches .

Given the two main functions of birdsong, mate attraction and territory defence, song amplitude is crucial for communication because it determines the communication range and it can also carry information. To understand the evolution of signals, it is helpful to consider the constraints on signal production, as physical constraints set the limits for signal plasticity and how signals can respond to selection. Previous work on the constraints of song amplitude was restricted to species that use loud vocalisations for long-distance communication. However, the low song amplitudes of some non-territorial species may hint at unknown limitations considering that females may prefer loud song. The star finch () is one such species. In this study, we recorded the songs of male star finches in the laboratory and investigated the relationship between vocal amplitude and other acoustic parameters of their song syllables. We found that vocal amplitude was linked to the phonetic structure of the syllables. More complex sounds (measured as higher syllable bandwidth and higher Wiener entropy) were produced at lower amplitudes than less complex sounds. To the best of our knowledge, this is the first report of a trade-off between song complexity and vocal amplitude. More complex song syllables possibly require a more intricate control of the syrinx and vocal tract, which might only be possible at lower amplitudes. We speculate that if female star finches prefer complex songs, this would cause males to sing quietly, as they cannot produce complex syllables with high amplitudes. We also provided a phonetogram (vocal range profile) for the star finch, indicating a positive correlation between amplitude and peak frequency. This finding corroborates earlier studies on species that use loud vocalisations for long-range communication, which, like the star finches in our study, were also unable to produce low frequencies at high amplitudes. This suggests that the frequency-amplitude correlation is a more general phenomenon in bird vocalisations, independent of the overall source amplitude of a species. While the evolution of birdsong is often viewed as constrained by neural costs and body size, or selective pressures from predators and social aggression, our results emphasise the importance of understanding phonetic features as well. At the same time, our study fills an important gap on non-territorial species that produce soft songs. The absence of the ecological demands for long-distance signalling has probably led birds to use soft yet complex songs that function within the pair bond, as we report here for star finches.