primates in our own case,so that it is clear that vocal learning evolved
quite separately in the two cases.Indeed it also appears to have evolved
at least twice elsewhere among birds (parrots,hummingbirds) and three
times among mammals (seals,whales and dolphins,bats;Janik and Slater
1997).Although vocal learning is a key feature in the evolution of sound
communication,and of complex repertoires in particular,I will not
discuss it further here.It is reviewed extensively elsewhere (Catchpole
and Slater 1995;Slater 1989;Whaling,this volume).
Repertoires and Their Use
Repertoires have a wide range of sizes even among close relatives.
Thrushes,for example,range from the European redwing (Turdus iliacus),
which only has a single type,to the song thrush (T.philomelos) with its
repertoire of over 200 (Ince and Slater 1985).Here we are talking about
distinct songs,with little if any sharing of elements between them.
However,some birds build up what sounds like a formidable repertoire
from just a few elements assorted in all sorts of different ways.Catchpole
(1976) suggested that a male sedge warbler,in the course of his life,may
never repeat exactly the same sequence of elements twice.This is because
the song is long,and the few dozen elements follow each other in highly
varied orders.At one level (that of the song) his repertoire size is enor-
mous,but at another (that of the element) it is not very large at all.
Of course,if a bird does not often sing the same song type twice it may
be either because it has a huge repertoire of types or because it is impro-
vising so that its repertoire is continuously changing.The latter may
occur in some species,as suggested for the Sardinian warbler (Sylvia
melanocephala;Luschi 1993),but it is certainly rare.In some other
species,songs change more slowly from one part of the season to another
or from one year to the next (e.g.,thrush nightingale;Sorjonen 1987).
When they appear to generate new songs much more rapidly than this,
so that the same one is seldom repeated,it may be because a limited
repertoire of elements is reassorted between songs.Jumbled and ram-
bling though many birdsongs may seem to be,examination of sonograms
reveals that they are in reality usually far from that;exactly the same
elements or song phrases occur again and again,albeit perhaps in very
different orders.
When a repertoire is large,it may be quite difficult to measure by
looking through the bird’s output and searching for repetitions
(Kroodsma 1982).The brown thrasher (Toxostoma rufum) from North
America is the best example here,and it is the current record holder for
song repertoire size.As each bird has over 1,000 song types,arriving at
52 Peter J.B.Slater
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