Summary1. Most studies on extra-pair paternity (EPP) focus either on a specific male's extra-pair gains or his extra-pair losses. For an individual bird however, mate choice or mate availability may underlie strong spatial restrictions. Disregarding this spatial aspect may underestimate or mask effects of parameters influencing observed EPP patterns. 2. Here, we propose a spatially explicit model for investigating the probability of having extra-pair offspring (EPO) within local networks of breeding pairs. The data set includes all realized and unrealized potential extra-pair matings. This method is biologically meaningful because it allows (a) considering both members of an extra-pair mating and their social mates, and (b) direct modelling of the spatial context in which extra-pair behaviour occurs. The method has the advantage that it can provide inference about the relative contribution of spatial and non-spatial parameters, and about the relative importance of male and female neighbourhoods. 3. We apply this method to parentage data from 1025 broods collected over 12 breeding seasons in two independent study populations of blue tits (Cyanistes caeruleus). We investigate a set of predictions based on the EPP literature, namely that EPP depends on male age and body size, breeding density and breeding synchrony. In all analyses, we control for breeding distance, a parameter that is expected to influence EPP even under random mating. 4. The results show that older and larger males were more likely to sire EPO, but both effects decreased with increasing breeding distance. Local breeding density but not synchrony predicted whether a particular male-female combination had EPO, at least in one of the study areas. Apart from breeding distance, male age had the strongest effect on EPP, followed by a measure of breeding density. The method thus allows a comprehensive assessment of the relative importance of different types of spatial and non-spatial parameters to explain variation in the occurrence of EPP, while controlling for the fact that individuals that breed further apart are less likely to have EPO. 5. The proposed approach is not limited to investigate EPP, but can be applied to other behavioural interactions between two individuals, such as dominance, competition and (social) mating.

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In several bird species, the period around dawn seems important for extrapair behavior. For example, a study on great tits (Parus major) showed that females that emerged earlier from their roosting place during the peak of their fertile period were more likely to have extrapair young in their brood. We investigated the potential effect of female emergence times on extrapair behavior in the blue tit (Cyanistes caeruleus). First, we tested the relationship between natural female emergence times from the nest-box and the presence or frequency of extrapair offspring in the brood, using 4 years of data. Females progressively emerged earlier from the nest-box as egg laying approached, with the earliest emergence 2 days before the start of laying. However, we found no relationship between female emergence time and the occurrence of extrapair young in the brood. Secondly, in 2 breeding seasons, we experimentally advanced female emergence times by supplying the roosting females with additional light in the early morning. Although the experiment had inconsistent effects on the occurrence of extrapair young in the brood, we found no evidence that female emergence time during peak fertility is directly linked to extrapair paternity. Interestingly, females exposed to artificial light were more likely to return to breed in the next year.

Thiessen polygons are often used to model territory characteristics. However, information about the quality of Thiessen polygon-based estimates is currently lacking. We used published data to investigate the match between Thiessen polygons and mapped bird territories regarding territory size, shape and neighbourhood. Although territory sizes and the number of neighbours were strongly correlated between these two methods, both parameters were overestimated by the Thiessen polygons. Therefore, caution is required when Thiessen polygons are used as a model for absolute values and when the assumptions of Thiessen polygons, such as formation of discrete territories and a contiguous study area, are not met.