Recent studies of lekking animals suggest that a suite of characters m
ay be favoured by sexual selection. Examples of such traits are high s
urvival, increased androgen levels, territory features and morphologic
al characters including exaggerated morphological ornaments. Here we a
rgue that such selection is often mediated by behavioural differences
and we use results from a field study of lekking black grouse, Tetrao
tetrix, as an example of our argument. In the absence of females, male
s display a range of stereotyped behaviours including vocalizations, v
isual displays and fighting. When females attend the lek, the behaviou
ral repertoire of the males becomes more limited. Time budget analyses
reveal that in the vast majority of cases, males in the presence of f
emales perform only three types of behaviour: a vocal display called r
ookooing; a courtship behaviour called circling; and fighting. This su
ggests that female preference could be based on male courtship and fig
hting behaviour while females visit individual male territories and th
at displays not used in the presence of females could be ruled out as
important for female preference. Sexual selection in black grouse, how
ever, is a complex interplay between male-male competition and female
choice and therefore female preference is not the only determinant of
male success. Successful males were more often involved in fights in t
he absence of females on the lek and defended territories that were la
rger than expected. Therefore, we hypothesize that differences in male
fighting ability result in some males occupying relatively large terr
itories at the centre of the leks. Relatively large territories seem t
o be needed for successful courtship. Therefore behavioural difference
s mediate differences in male copulation success both through female p
reference and male-male competition. (C) 1997 The Association for the
Study of Animal Behaviour.