Abstract: Author Summary Bird songs range in form from the simple notes of a Chipping Sparrow to the complex repertoire of the nightingale. Recent studies suggest that bird songs may contain non-adjacent dependencies where the choice of what to sing next depends on the history of what has already been produced. However, the complexity of these rules has not been examined statistically for the most elaborate avian singers. Here we show that one complex singer—the domesticated canary—produces a song that is strongly influenced by long-range rules. The choice of how long to repeat a given note or which note to choose next depends on the history of the song, and these dependencies span intervals of time much longer than previously assumed for birdsong. Like most forms of human music, the songs of canaries contain patterns expressed over long timescales, governed by rules that apply to multiple levels of a temporal hierarchy. This vocal complexity provides a valuable model to examine how ordered behaviors are assembled from more elementary neural components in a relatively simple neural circuit.
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