Tonality over a broad frequency range is linked to vocal learning in birds.

Many birds emit tonal song syllables even though the sound sources generate sound with rich upper harmonic energy content. This tonality is thought to arise in part from dynamically adjusted filtering of harmonic content. Here, we compare tonality of song syllables between vocal learners and non-learners to assess whether this characteristic is linked to the increased neural substrate that evolved with vocal learning. We hypothesize that vocal learning ability is correlated with enhanced ability for generating tonal sounds, because vocal production learners might also have an enhanced ability to articulate their vocal tracts and sound source for producing tonality. To test this hypothesis, we compared vocal learners and non-learners from two groups (186 passerines and 42 hummingbirds) by assessing tonality of song syllables. The data suggest that vocal learners in both clades have evolved to sing songs with higher tonality than the related, non-vocal learning clades, which is consistent with stronger roles for broadband dynamic filtering and adjustments to the sound source. In addition, oscine songs display higher tonality than those of hummingbirds. A complex interplay of vocal tract biomechanics, anatomical differences of the sound source as well as increased motor control through vocal learning facilitates generation of broad tonality.