Interval singing links to phenotypic quality in a songbird.

Darwin was fascinated by melodic performances of insects, fish, birds, mammals, and men. He considered the ability to produce musical notes without direct use the most mysterious endowment of mankind. Bird song is attributed to sexual selection, but it remains unknown how the expected relationship between melodic performance and phenotypic quality arises. Melodies consist of sequences of notes, and both Pythagoras and music theorists in the Middle Ages found that their tonal frequencies form simple ratios that correspond to small-integer proportions derived from the harmonic series. Harmonics are acoustically predictable, and thus form the basis of the natural, just tuning system in music. Here I analyze the songs of the great tit (), a bird with a stereotyped song of typically two notes, and test the prediction that the deviations of the intervals from small-integer frequency ratios based on the harmonic series are related to the quality of the singer. I show that the birds with the smallest deviations from small-integer ratios possess the largest melanin-based black ventral tie, a signal that has been demonstrated to indicate social status and dominance, past exposure to parasites, and reproductive potential. The singing of notes with exact frequency relationships requires high levels of motor control and auditory sensory feedback. The finding provides a missing link between melodic precision and phenotypic quality of individuals, which is key for understanding the evolution of vocal melodic expression in animals, and elucidates pathways for the evolution of melodic expression in music.