The singing of adult birds shows a clear hierarchical organization and, due
to its development through vocal imitation of sound patterns. makes an exc
ellent biological model to examine the variables that influence the imitati
on of patterns on different hierarchy levels, e.g., songs and elements comp
osing the songs. We studied such variables in the nightingale (Luscinia meg
arhynchos), i.e., a species that in the wild uses a large repertoire of son
gs. Subjects were raised in the lab and tutored by presenting them with a s
erial learning task, here a sequence of differently patterned songs. Analys
es of singing, which the trained subjects performed at the end of their voc
al ontogeny, allowed us to uncover hierarchy-related accomplishments and al
so specific constraints in learning by imitation. Our results showed that a
n imitation of elements and element sequences reflected a kind of "gestalt"
learning, which finally became visible in the form of specific song patter
ns. An imitation of song sequences, on the other hand, reflected a kind of
"list" learning, which finally emerged into the formation of specific subre
pertoires of song types. The structure of such subrepertoires mirrored the
serial order of imitated songs but allowed, nevertheless, a serially flexib
le retrieval of single types of songs. These findings contribute to a bette
r understanding of relationships between signal imitation and signal use du
ring vocal interactions among songbirds.