Two, not mutually exclusive, hypotheses for how female choice of males
could explain male territory settlement patterns on leks have recentl
y been proposed. First, the 'temporal spillover hypothesis assumes tha
t females return to previously popular mating sites. If the previous o
wner of such a popular site disappears it will pay other males to defe
nd such areas of the lek. Second, the spatial spillover hypothesis ass
umes that unsuccessful males gather around the most successful males t
o achieve copulations from females with which top males for some reaso
n fail to mate. In the black grouse, Tetrao tetrix, there were no indi
cations of temporal spillover between successive years. While individu
al males retained their popularity to some degree, there was no consis
tency in the popularity df the same locations per se. Instead, some sp
atial spillover is suggested by the fact that new territories in each
season were established closer to the copulation centre than to the le
k centre of the previous year. Furthermore, males close to the preferr
ed males seemed to achieve copulations because of their proximity to s
uch males. Successful males did not change their territorial position,
but less preferred males tended to move closer to the lek centre fill
ing the vacancies arising from any yearly mortality of males. While sp
atial spillover explains only a minor proportion of matings, most fema
les choosing males independent of their site, it may be the only chanc
e for poor quality males to obtain matings.