The importance of extrapair paternity (EPP) as an aspect of mixed reproductive strategies is currently the focus of many studies. Since females have at least some control over the occurrence of inseminations, they are expected to engage in extrapair copulations only if they benefit, for instance through gaining highquality or compatible genes for their offspring, or insurance against infertility. To distinguish between these benefits, we studied characteristics related to intermale variation in fertilization success as well as differences in fitness between half-siblings in the reed bunting, Emberiza schoeniclus, a socially monogamous passerine with high levels of EPP (50% of young). We found that older males were more successful at siring offspring in other broods and a nonsignificant tendency for them to be less cuckolded in their own broods. The success of older males does not support the fertility insurance hypothesis: a male's sperm storage capacity did not predict his fertilization success, the occurrence of infertile eggs was unrelated to the occurrence of EPP in a nest and older males had more infertile eggs in nests with no EPP. Extrapair young had longer tarsi at fledging than their maternal half-siblings, but we found no other differences between maternal or paternal half-siblings in several presumed quality traits. Owing to the absence of long-term fitness data, we are unable to determine whether the difference in tarsus length is related to a difference in fitness benefits. At this time, we cannot exclude the possibility that females do gain genetic benefits through EPP.

The tragedy of the commons predicts social collapse when public goods are jointly exploited by individuals attempting to maximize their fitness at the expense of other social group members. However, animal societies have evolved many times despite this vulnerability to exploitation by selfish individuals. Kin selection offers a solution to this social dilemma, but in large social groups mean relatedness is often low. Sociable weavers (Philetairus socius) live in large colonies that share the benefits of a massive communal nest, which requires individual investment for construction and maintenance. Here, we show that despite low mean kinship within colonies, relatives are spatially and socially clustered and that nest-building males have higher local relatedness to other colony members than do non-building males. Alternative hypotheses received little support, so we conclude that the benefits of the public good are shared with kin and that cooperative investment is, despite the large size and low relatedness of these communities, kin directed.

Do the two parents at a nest make simultaneous decisions whether to care for their offspring or to desert? If a single parent is sufficient for rearing young, one parent (typically, the male) may desert and reproduce with a new mate within the same breeding season, leaving the other parent with the brunt of care. As each parent is expected to maximize its own reproductive success, the interests of the two parents do not necessarily coincide, and a sexual conflict over care may emerge. Here we investigate the process of clutch desertion in a small passerine bird, the Penduline Tit Remiz pendulinus. Among birds, this species has a remarkably variable breeding system, because a single parent (either the male or the female) may provide the full care of the young, whereas about 30% of clutches are abandoned by both parents. First, we show that biparental desertion occurs within a single day in 73.7% of the clutches (n = 14), whereas desertion decisions are sequential in 26.3% of the clutches (n = 5) (male first: 10.5% (n = 2); female first: 15.8% (n = 3); n = 19 clutches in total). Secondly, we observed the behaviour of both parents before desertion, and investigated whether desertion can be predicted from their behaviour. However, neither singing nor nest‐building behaviour predicted whether the male or the female would desert. We therefore suggest that biparental desertion may be simultaneous by male and female in our population of Penduline Tits. Furthermore, the parents do not appear to signal their intention to desert their mate. We argue that the parents’ interest may be actually to disguise their intention to desert.

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Eurasian penduline tits (Remiz pendulinus) have an unusually diverse breeding system consisting of frequent male and female polygamy, and uniparental care by the male or the female. Intriguingly, 30 to 40 per cent of all nests are deserted by both parents. To understand the evolution of this diverse breeding system and frequent clutch desertion, we use 6 years of field data to derive fitness expectations for males and females depending on whether or not they care for their offspring. The resulting payoff matrix corresponds to an asymmetric Snowdrift Game with two alternative evolutionarily stable strategies (ESSs): female-only and male-only care. This, however, does not explain the polymorphism in care strategies and frequent biparental desertion, because theory predicts that one of the two ESSs should have spread to fixation. Using a bootstrapping approach, we demonstrate that taking account of individual variation in payoffs explains the patterns of care better than a model based on the average population payoff matrix. In particular, a model incorporating differences in male attractiveness closely predicts the observed frequencies of male and female desertion. Our work highlights the need for a new generation of individual-based evolutionary game-theoretic models.