Evolution of variation in male secondary sexual characteristics. A test of eight hypotheses applied to pied flycatchers

The male pied flycatcher Ficedula hypoleuca exhibits sexual dimorphism in its plumage colour, varying from a female-like brown to jet black. The evoltution of this variation in male plumage colour can be explained by at least eight hypotheses viz., (I) neutral mutation; (II) individual recognition; (III) three forms of inter-sexual selection; (IIIa) mate selection for phenotypes, (IIIb) Fisherian selection, (IIIc) handicap selection; (IV) intrasexual selection; (V) delayed maturation; and (VI) female mimicry. The assumptions and predictions derived from all these hypotheses were tested by analysing the observed variation in male plumage colour in relation to age, body size, physical condition, survival rate, aggressivity in territorial defence, territorial quality, female choice of mate, sex ratio, and reproductive success. We found that: males became blacker with age; black males were larger than browner ones; however, browner males survived better between breeding seasons than did blacker ones; black males were more aggressive against black than brown intruders; blacker males occupied better territories; blacker males were paired earlier than browner males; and finally blacker males produced heavier offspring than did brown males. A reasonable conclusion from the results of these analyses is that four of the hypotheses considered, viz. ‘mate selection for phenotypes’ (IIIa), Fisherian selection (IIIb), ‘handicap selection’ (IIIc), and intra-sexual selection (IV) could not be rejected. We therefore suggest that the evolution of the observed variation in the secondary sexual characteristics of the male pied flycatcher can be explained by a combination of these hypotheses.