Behavioural observations and DNA fingerprinting were used to determine
the relationship between male and female behaviours and levels of ext
ra-pair paternity in the wheatear, Oenanthe oenanthe. Behavioural obse
rvations were consistent with the hypothesis that males attempted to e
nsure paternity by mate guarding, while pursuing extra-pair copulation
s (EPCs) primarily outside the fertile period of their pair female. Th
e intensity of guarding varied with time of season and was greater at
late nests. However, although males on territories with late nests als
o experienced high intrusion rates, the intensity of guarding was infl
uenced more by the operational sex ratio (which was female skewed at e
arly nests) than by intrusion rates per se. We suggest that early bree
ding males adopted a strategy of territorial defence to ensure paterni
ty, as opposed to guarding their female directly (which late breeding
males did), to capitalize on the increased opportunities to pursue EPC
s in neighbouring territories. Females were less conspicuous than male
s in their pursuit of EPCs, were never seen off territory or observed
to solicit extra-pair males directly, and rejected the majority of EPC
s. The frequency of extra-pair paternity was 11% of 73 offspring, in 2
9% of 17 broods, and was not correlated with the intensity of guarding
. Female cooperation appeared to be important for successful copulatio
n, and extra-pair paternity is therefore likely to be a consequence of
solicited, or at least accepted, EPCs. We discuss why females might h
ave participated in EPCs. (C) 1998 The Association for the Study of An
imal Behaviour.