Harmonic Hopping, and Both Punctuated and Gradual Evolution of Acoustic Characters in Selasphorus Hummingbird Tail-Feathers

Models of character evolution often assume a single mode of evolutionary change, such as continuous, or discrete. Here I provide an example in which a character exhibits both types of change. Hummingbirds in the genus Selasphorus produce sound with fluttering tail-feathers during courtship. The ancestral character state within Selasphorus is production of sound with an inner tail-feather, R2, in which the sound usually evolves gradually. Calliope and Allen's Hummingbirds have evolved autapomorphic acoustic mechanisms that involve feather-feather interactions. I develop a source-filter model of these interactions. The 'source' comprises feather(s) that are both necessary and sufficient for sound production, and are aerodynamically coupled to neighboring feathers, which act as filters. Filters are unnecessary or insufficient for sound production, but may evolve to become sources. Allen's Hummingbird has evolved to produce sound with two sources, one with feather R3, another frequency-modulated sound with R4, and their interaction frequencies. Allen's R2 retains the ancestral character state, a ∼1 kHz "ghost" fundamental frequency masked by R3, which is revealed when R3 is experimentally removed. In the ancestor to Allen's Hummingbird, the dominant frequency has 'hopped' to the second harmonic without passing through intermediate frequencies. This demonstrates that although the fundamental frequency of a communication sound may usually evolve gradually, occasional jumps from one character state to another can occur in a discrete fashion. Accordingly, mapping acoustic characters on a phylogeny may produce misleading results if the physical mechanism of production is not known.