seem to be a random outpouring is in fact highly organized (Eens,
Pinxten,and Verheyen 1989,1991b).A bout of singing lasts an average
of twenty-five seconds and consists of a sequence of phrases or song
types with very short intervals (mostly less than 0.1 second) between
them.Each phrase tends to be repeated several times before the bird
moves on to the next.Repertoire size varies:among twenty-seven males
it ranged from twenty-one to sixty-seven song types (Eens,Pinxten,and
Verheyen 1991b).
The order of song types within a bout of singing is relatively fixed.
Types fall into four broad categories,which also tend to occur at partic-
ular points in the sequence:
1.The bout normally starts with a number of whistles,each male having
a repertoire of seven to twelve (Hausberger and Guyomarc’h 1981).
2.The second section consists of a series of variable and complex
phrases including cases of mimicry;each male has fifteen to twenty dif-
ferent imitations in his repertoire (Hindmarsh 1984).
3.The third part of the bout consists of rattle song types.These phrases
include a rapid succession of clicks sounding like a rattle,and each male
has a repertoire of two to fourteen of them.
4.Most song bouts that are not interrupted earlier end with some loud,
high-frequency song types;a male may have up to six of these.
Again,as with the nightingale,starling song may seem endlessly varied
to the ear,but closer analysis reveals that each male has a limited reper-
toire of types and that these are ordered according to quite well-
specified rules.
Marsh Warbler
Most birds learn only the song of their own species.The fact that they
are reared by,and normally imprint upon and develop social relations
with,members of that species is one reason for this.But some species,of
which the starling is one,usually include imitations within their song.
Given the rarity of improvisation among birds,the best guess is that this
is a way in which males can enhance the variety,and hence the attrac-
tiveness,of their songs.
One of the most remarkable cases of mimicry,which forms a good case
study,is that of the European marsh warbler (Acrocephalus palustris)
studied by Dowsett-Lemaire (1979).This species breeds in Europe and
migrates to East Africa.Young birds are thought to learn their song
entirely in the first few months of life.They cannot base it on other
members of their own species,as adult males cease to sing before their
chicks hatch.Instead,Dowsett-Lemaire estimated that each young male
56 Peter J.B.Slater
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