J. Omithol. 139, 287-296 (1998)
© Deutsche Ornithologen-Gesellschaft/BlackwellWissensehafts-Verlag, Berlin
ISSN 0021-8375
Song structure in the Hoopoe (Upupa epops) -
Strophe length reflects male condition
Manuel Martin-Vivaldi, Jos6 Javier Palomino and Manuel Soler
Departamento de Biologfa Animal y Ecologfa, Facultad de Ciencias, Universidad de Granada,
E-18071 Granada, Spain (e-mall: MVIVALDI@GOLIAT.UGR.ES)
Summary
The structure of Hoopoe (Upupa epops) song is analysed in a colour ringed population in southern
Spain. The song of males in this species is very simple, with a repertoire size of one. The strophes
of a male only differ in the number of elements that they include (strophe length), and strophe length
is the main song feature differing between males. Daring the prelaying period each individual used
mainly strophes of only two lengths, between 2 and 6 elements per strophe, and mean strophe length
of males during this period was highly repeatable. However, some males changed the range of strophe
types produced and decreased their mean strophe length after unsuccessful breeding or spending long
periods of time singing (unpaired males). These changes show that strophe length is phenotypically
plastic, and suggest that singing long strophes is energetically more costly than singing short ones.
The significantrelationship between strophe length and body condition, and the fact that long strophes
were associated with longer previous pauses than short strophes, also suggest that increasing strophe
length is costly. All these findings are in accordance with the hypothesis that strophe length reflects
male condition in the Hoopoe, although it is not clear what the actual cost of singing long strophes
is.
Key words: Sexual selection, honest signalling, non-passerines
Zusammenfassung
Struktur des Gesangs des Wiedehopfs (Upupa epops) -
Strophenl~inge reflektiert M[innchen-Qualit~it
Die Stmkmr des Gesangs des Wiedehopfs wurde in einer farbberingten stidspanischen Population
untersucht. Der Gesang des Mfinnchens ist sehr einfach und umfaBt nur ein Repertoire. Die Strophen
eines Mfinnchens differieren nur in der Anzahl Elemente (Strophenlange), und die Mannchen unter-
scheiden sich vor allem in der Strophenl~nge voneinander. In der Vorbmtphase verwendeten die
Mfinnchen meist nur zwei verschieden lange Strophen, bestehend aus 2 und 6 Elementen je Strophe,
und die durchschnittliche Strophenl/inge war in dieser Phase sehr konstant. Nach Bmtverlust oder bei
kontinuierlich singenden, ledigen MS_nnchen kam es zu einer Verkiirzung der mittleren Strophenl/m-
ge. Die Strophenl~ge ist ph~notypisch plastisch, und lange Strophen zu singen scheint energetisch
anfwendiger als kurze Strophen. Der enge Zusammenhang zwischen Strophenl/inge und m~nnlicher
Konstitution einerseits und die Beobachtung, dab lange Strophen mit langen Pausen korrelieren
andererseits, zeigen ebenfalls, dab lhngere Strophen energetisch aufwendiger sind. Die wirklichen
Kosten daftir sind jedoch noch nicht bekannt.
Introduction
In most bird species several song features have
been shown to function both in intersexual and
intrasexual communication ("dual function"
hypothesis, Albrecht & Oring 1995, Hassel-
quist 1990, McDonald 1989, Searcy 1988), in-
dicating that they provide information useful
for both sexes. For instance, male quality, in
particular male condition, is worth knowing for
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