Variation In Extrapair Paternity Research Articles

Combined roles for breeding synchrony, habitat and scale as predictors of extrapair paternity

Despite extensive research on passerine reproduction, it remains difficult to predict which traits of individuals, nesting populations and landscapes drive extrapair paternity (EPP). Two broad categories of drivers, habitat structure and population level factors, have been considered separately or at a single spatial or temporal scale. We used a 6-year nesting data set of scissor-tailed flycatcher, Tyrannus forficatus , which have high rates of cuckoldry (73% of nests, 59% of nestlings), to explore how nest density, breeding synchrony, habitat structure and clutch initiation date affect EPP rates. We further considered synchrony at two temporal and habitat structure at two spatial scales. We predicted that visual occlusion from vertical habitat structuring on breeding territories would allow extrapair males to go unnoticed and provide cover for extrapair copulations. Predictions for effects of nest density and fertility synchrony were double-edged: EPP may either increase as extrapair mate availability and ease of comparison with social mates increases or decrease as social mates increase their efforts to assure paternity. Using Bayesian techniques, we found a combination of population level factors and habitat structure, including interactions among the latter, at different scales best accounted for variation in EPP. EPP declined with increasing population synchrony. Variation in EPP was also explained by fine-scale habitat measures, decreasing with increasing nest tree diameter at breast height and woody vegetation cover but increasing with tree density. Notably, EPP increased with a coarse-scale habitat measure, linear extent of fence or powerline, suggesting a role for human alteration of habitat. Fences are used as communal perches by neighbours and floater males, potentially increasing interactions between asynchronous individuals. Our study demonstrates that breeding synchrony influences the probability of EPP and that habitat structure on individual territories is a strong predictor of paternity that acts independently of breeding density or synchrony at our scale of measurement. • We explore factors influencing cuckoldry in scissor-tailed flycatchers across 6 years. • Cuckoldry increased with populationwide fertility synchrony. • Cuckoldry was not related to spatial distribution of nests. • Cuckoldry increased with tree number and powerline/fence length and low shrub cover. • These correlates may impact cuckoldry via their effects on male mate guarding.

Experimental extra-pair copulations provide proof of concept for fertility insurance in a socially monogamous bird.

Extra-pair paternity is common among socially monogamous birds, but whether females benefit from having extra-pair copulations remains unresolved. In this study, I staged single extra-pair copulations in captive pairs of socially monogamous Japanese quail (Coturnix japonica) and used extra-pair males of a different colour to identify extra-pair fertilizations. This eliminated among-female variation in extra-pair copulations as a source of variation in extra-pair fertilizations and tested whether a single extra-pair copulation would insure against infertility of the social male. Overall, the probability of extra-pair fertilization was 0.46 for the first egg that was fertilized after extra-pair copulation, but this rapidly declined over consecutive eggs in the laying sequence. However, a single extra-pair copulation was effective in ensuring fertilization of the majority of a typical clutch in the few cases where the social male was completely infertile. Hence, my results show that variation in extra-pair paternity can be independent from variation in extra-pair copulation behaviour and that a single, strategically timed, extra-pair copulation can largely insure against social male infertility. Among-female variation in extra-pair fertilizations, and their declining probability over the laying sequence, as typical for many bird species, can thus, in principle, be parsimoniously explained by a uniform female fertility insurance strategy.

Analysis of within-individual variation in extrapair paternity in blue tits (Cyanistes caeruleus) shows low repeatability and little effect of changes in neighborhood.

Many studies investigated variation in the frequency of extrapair paternity (EPP) among individuals. However, our understanding of within-individual variation in EPP remains limited. Here, we comprehensively investigate variation in EPP at the within-individual level in a population of blue tits (Cyanistes caeruleus). Our study is based on parentage data comprising >10 000 genotyped offspring across 11 breeding seasons. First, we examined the repeatability of the occurrence of EPP, the number of extrapair offspring, the number of extrapair partners, and the occurrence of paternity loss using data from males and females that bred in multiple years. Second, we tested whether within-individual changes in EPP between breeding seasons relate to between-year changes in the local social environment. Repeatabilities were generally low but significant for the occurrence and number of extrapair young in females and for whether a male sired extrapair young or not. We found no evidence that the presence of the former social partner or changes in the proportion of familiar individuals or in phenotypic traits of the neighbors influenced changes in levels of EPP in females. However, in adult males, a decrease in the average body size of male neighbors was associated with higher extrapair siring success. If confirmed, this result suggests that the competitive ability of a male relative to its neighbors influences his extrapair mating success. We suggest that alternative hypotheses, including the idea that within-individual changes in EPP are due to “chance events” rather than changes in an individual’s social breeding environment, deserve more consideration.

Female need for paternal care shapes variation in extra-pair paternity in a cooperative breeder

AbstractSocially monogamous females regularly mate with males outside the pair bond. The prevailing explanation for this behavior is that females gain genetic benefits resulting from increased fitness of extra-pair offspring. Furthermore, because of the risk of reduced paternal care in response to cuckoldry, females are expected to seek extra-pair copulations when they can rear offspring with little help from their social partner (“constrained female” hypothesis). We tested these hypotheses and analyzed variation in paternal care in the Afrotropical, facultative cooperative breeding placid greenbul (Phyllastrephus placidus). Overall, approximately 50% of the offspring resulted from extra-pair (and extra-group) mating. Identified extra-pair males were in most cases neighboring dominant males, yet never within-group subordinates. As predicted by the constrained female hypothesis, the occurrence of extra-pair paternity (EPP) increased with the number of cooperative helpers (and not with total group size). However, dominant males did not adjust their food provisioning rates in response to EPP. Although extra-pair males were more strongly related to the dominant female and less heterozygous than the latter’s social mate, this did not result in more inbred extra-pair offspring, likely because identified extra-pair males were not representative of the extra-pair male population. While earlier studies on EPP mainly focused on male genetic quality, results from this study provide evidence that female’s social context may affect extra-pair strategies too.

Extra‐pair paternity in birds

Since the first molecular study providing evidence for mating outside the pair bond in birds over 30 years ago, >500 studies have reported rates of extra‐pair paternity (EPP) in >300 bird species. Here, we give a detailed overview of the current literature reporting EPP in birds and highlight the sampling biases and patterns in the data set with respect to taxonomy, avian phylogeny and global regions, knowledge of which will be crucial for correct interpretation of results in future comparative studies. Subsequently, we use this comprehensive dataset to simultaneously test the role of several ecological and life history variables. We do not find clear evidence that variation in EPP across socially monogamous species can be explained by latitude, density (coloniality), migration, generation length, genetic structuring (dispersal distance), or climatic variability, after accounting for phylogeny. These results contrast previous studies, most likely due to the large heterogeneity within species in both EPP and the predictor of interest, indicating that using species averages might be unreliable. Despite the absence of broadscale ecological drivers in explaining interspecific variation in EPP, we suggest that certain behaviours and ecological variables might facilitate or constrain EPP, as indicated by our finding that EPP was negatively associated with latitude within noncolonial species, suggesting a role of breeding synchrony. Thus, rather than focussing on general explanations for variation in EPP across all species, a future focus should be on how various aspects of ecology or life history might have driven variation in EPP among groups of species or populations of the same species. Hence, we argue that variation in EPP can be partly explained when taking the right perspective. This comprehensive overview, and particularly the dataset provided herein will create a foundation for further studies.

Temperate-Tropical Variation in Breeding Synchrony and Extra-Pair Paternity Among New World Tachycineta Swallows

Extra-pair paternity rates vary markedly across avian taxa, but patterns of variation in this trait have been obscured by a paucity of data on closely related species, especially those spanning broad environmental gradients. Here we compare variation in extra-pair paternity rates among five species in the widespread swallow genus Tachycineta. Rates of extra-pair paternity vary widely in this group, ranging from 13 to 87% of nests having extra-pair young. The inter-specific variation in extra-pair paternity within this small group of closely related swallows has a range equivalent to that found among all Hirundinidae and is close to the range of variation across all birds. Despite theory that predicts extra-pair paternity rates to be explained by latitudinal variation in breeding synchrony our results show that extra-pair paternity rates in this genus do not closely track a latitudinal gradient, as predicted by studies of other life-history traits, and are not explained by differences in breeding synchrony as previously suggested. The genetic mating systems of birds, described by the rates of extra-pair paternity, are connected to all other life-history traits through a complex network of trade-offs with organismal (phylogenetic) and ecological (environmental) factors. Disentangling each of these interactions to understand latitudinal patterns in any given life-history trait remains a daunting task.

Small, Variable Sperm in a Barn Swallow Population with Low Extra-Pair Paternity

Sperm competition can theoretically affect sperm morphology; however, it remains unclear whether and how sperm morphology tracks the intensity of sperm competition in each population. The barn swallow Hirundo rustica is a model species used in the study of sexual selection, and exhibits considerable variation in extra-pair paternity (percentage extra-pair young, ca. 3-30%) among populations. In the Joetsu population of the barn swallow, extra-pair paternity is virtually absent (< 3%), providing a rare opportunity to study sperm morphology under limited sperm competition, and to compare it with those reported in populations with frequent extra-pair paternity (>15%). We found that head, midpiece, and total sperm length were significantly shorter in the Joetsu population than in populations with frequent extra-pair paternity. Moreover, the variability in total sperm length, measured as the coefficient of variation in the Joetsu population, was twice as high as that of populations with frequent extra-pair paternity. These results are consistent with a positive, directional, and stabilizing effect of sperm competition on sperm morphology. Together with previous studies in populations with frequent extra-pair paternity, the current study provides one of few sets of evidence to show a link between the intensity of sperm competition and the mean and variance of sperm morphology within a wild bird species.

Linking the fine-scale social environment to mating decisions: a future direction for the study of extra-pair paternity.

Variation in extra-pair paternity (EPP) among individuals of the same population could result from stochastic demography or from individual differences in mating strategies. Although the adaptive value of EPP has been widely studied, much less is known about the characteristics of the social environment that drive the observed patterns of EPP. Here, we demonstrate how concepts and well-developed tools for the study of social behaviour (such as social network analysis) can enhance the study of extra-pair mating decisions (focussing in particular on avian mating systems). We present several hypotheses that describe how characteristics of the social environment in which individuals are embedded might influence the levels of EPP in a socially monogamous population. We use a multi-level social approach (Hinde, 1976) to achieve a detailed description of the social structure and social dynamics of individuals in a group. We propose that the pair-bond, the direct (local) social environment and the indirect (extended) social environment, can contribute in different ways to the variation observed in the patterns of EPP, at both the individual and the population level. A strength of this approach is that it integrates into the analysis (indirect) interactions with all potential mates in a population, thus extending the current framework to study extra-pair mating behaviour. We also encourage the application of social network methods such as temporal dynamic analysis to depict temporal changes in the patterns of interactions among individuals in a group, and to study how this affects mating behaviour. We argue that this new framework will contribute to a better understanding of the proximate mechanisms that drive variation in EPP within populations in socially monogamous species, and might ultimately provide insights into the evolution and maintenance of mating systems.

Multiple hypotheses explain variation in extra-pair paternity at different levels in a single bird family.

Extra-pair paternity (EPP), where offspring are sired by a male other than the social male, varies enormously both within and among species. Trying to explain this variation has proved difficult because the majority of the interspecific variation is phylogenetically based. Ideally, variation in EPP should be investigated in closely related species, but clades with sufficient variation are rare. We present a comprehensive multifactorial test to explain variation in EPP among individuals in 20 populations of nine species over 89years from a single bird family (Maluridae). Females had higher EPP in the presence of more helpers, more neighbours or if paired incestuously. Furthermore, higher EPP occurred in years with many incestuous pairs, populations with many helpers and species with high male density or in which males provide less care. Altogether, these variables accounted for 48% of the total and 89% of the interspecific and interpopulation variation in EPP. These findings indicate why consistent patterns in EPP have been so challenging to detect and suggest that a single predictor is unlikely to account for the enormous variation in EPP across levels of analysis. Nevertheless, it also shows that existing hypotheses can explain the variation in EPP well and that the density of males in particular is a good predictor to explain variation in EPP among species when a large part of the confounding effect of phylogeny is excluded.

Genetic diversity does not explain variation in extra-pair paternity in multiple populations of a songbird.

Many songbirds are socially monogamous but genetically polyandrous, mating with individuals outside their pair bonds. Extra-pair paternity (EPP) varies within and across species, but reasons for this variation remain unclear. One possible source of variation is population genetic diversity, which has been shown in interspecific meta-analyses to correlate with EPP but which has limited support from intraspecific tests. Using eight populations of the genetically polyandrous red-winged blackbird (Agelaius phoeniceus), including an island population, we investigated whether population-level differences in genetic diversity led to differences in EPP. We first measured genetic diversity over 10 microsatellite loci and found, as predicted, low genetic diversity in the island population. Additional structure analyses with multilocus genotypes and mtDNA showed the island population to be distinct from the continental populations. However, the island population's EPP rate fell in the middle of the continental populations' distribution, whereas the continental populations themselves showed significant variation in EPP. This result suggests that genetic diversity by itself is not a predictor of EPP rate. We discuss reasons for the departure from previous results, including hypotheses for EPP that do not solely implicate female-driven behaviour.

Habitat structure and colony structure constrain extrapair paternity in a colonial bird

Individual variation in sexual fidelity and extrapair paternity (EPP) is widely attributed to environmental heterogeneity, but the only variables known to be influential are food abundance and density of conspecific breeders (potential extrapair partners). Habitat structure is thought to impact EPP but is rarely measured and, when considered, is usually confounded with food abundance and predation pressure. To sidestep these confounds, we tested whether EPP is associated with habitat structure variables and with local conspecific density in a species whose nesting habitat is not used for feeding and lacks predators. In a blue-footed booby, Sula nebouxii, colony, the probability of EPP in a female's nest was highest in parts of the study plot where there were few obstacles to locomotion, and was quadratically related to local density of sexually active males, even though local males did not sire the EP chicks. The probability of a male breeder siring EP (extrapair) chicks elsewhere was quadratically related to local density of sexually active males around his nest. From these patterns we infer that both sexes may foray for EP interactions, that males and females nesting at intermediate density are most likely to be accessed by forayers, and that obstacles in the vicinity of a female's nest constrain access of foraying males. To our knowledge, this is the first demonstration that individual variation in EPP is associated with habitat structure in the absence of confounding variation in food availability, predation pressure or breeder quality, and the first evidence that EPP opportunities of female and male breeders are reduced by high

Habitat geometry does not affect levels of extrapair paternity in an extremely unfaithful fairy-wren

Density of potential mates has often been proposed to explain the enormous variation in extrapair paternity. However, density is often confounded by other ecological factors that might affect extrapair paternity in their own way. Furthermore, extrapair mating shows strong phylogenetic inertia, making both meaningful intra- and interspecific comparisons difficult. An extreme way to change density is through habitat fragmentation that reduces connectivity between territories. Recently, habitat connectivity was hypothesized to explain the surprising discovery of a virtually monogamous species among the world’s most unfaithful bird genus. The monogamous Malurus coronatus lives in narrow riparian strips that limit contact with neighbors to both extreme ends of territories, whereas Malurus species with high levels of extragroup paternity typically live in high-connected habitat in which they are surrounded by neighbors. Here, we test the habitat geometry hypothesis by comparing levels of extragroup paternity of Malurus elegans living in fragmented low-connected habitat and in high-connected habitat. We found that M. elegans does not have lower levels of extragroup paternity in low-connected habitat (68%) than in high-connected habitat (56% of offspring), indicating that connectivity does not limit opportunities for extragroup paternity. Furthermore, there was no evidence that females in low-connected habitat gained extragroup paternity further away or from less sires or that they were more likely to be closely related to their social mate. We conclude that behavioral plasticity in response to density-dependent cost and benefits of mating behavior does not explain intrageneric variation in extragroup paternity in Malurus. Furthermore, habitat fragmentation may not strongly affect inbreeding risk in this species.

Extra-pair paternity in relation to regional and local climate in an Arctic-breeding passerine

Reproductive processes are affected by local and regional climate variation. Birds breeding in the Arctic may experience strong energetic constraints, which will affect their reproductive output. Recent research has emphasized the importance of extra-pair copulation as a means of improving reproductive output. In this paper, we explore ecological and climatic determinants that may explain variation in extra-pair paternity (EPP) in an arctic-breeding passerine, the snow bunting Plectrophenax nivalis. EPP occurred in 10.8 % of the young and 20.9 % of the broods sampled from 1999 to 2003. We found that the proportion of extra-pair young in a nest was positively related to the body size and age of the social male and weakly negatively related to the local average minimum temperature prior to the onset of egg laying. These results suggest that older and larger males lost a larger share of paternity than smaller and younger males, and that the relative loss of paternity decreased with cold weather during the female’s fertile period. Large and old males spend less time mate guarding compared to small and young males and may allocate more time towards extra-pair forays, and thus lose more paternity in their own nest. Climatic conditions most likely constrain the total energy budget with less energy available for extra-pair activity in cold weather.

Breeding synchronization facilitates extrapair mating for inbreeding avoidance

Extrapair (EP) mating can enable females to reduce the negative effects of inbreeding. However, opportunities for EP mating are often ecologically or demographically constrained, and it is unclear whether and how females can overcome these constraints. Here, we show that fitness costs from inbreeding in purple-crowned fairy-wrens (Malurus coronatus) are severe (≥30% reduction in hatchability of eggs in incestuous pairs). Although EP paternity was low for unrelated pairs (3% of broods), and negatively affected by low breeding density, 46% of broods produced by incestuously paired females contained EP young that were almost always sired by a direct neighbor. Moreover, females in incestuous pairs initiated egg laying more synchronously with direct neighbors than females in an unrelated pair, and more synchronous broods were more likely to contain EP young. Because male sperm stores are highest around their own partner’s fertile period, our results imply that incestuously paired females achieve high levels of EP paternity by adjusting the timing of egg laying to match peak fertility of neighboring males. This example of a dramatic increase in EP paternity due to subtle behavioral adjustments illustrates that negative effects of inbreeding exert strong selection for female strategies to facilitate EP mating, and this may contribute to unexplained variation in EP paternity.

New support for an old hypothesis: density affects extra‐pair paternity

Density has been suggested to affect variation in extra-pair paternity (EPP) in avian mating systems, because increasing density promotes encounter rates and thus mating opportunities. However, the significance of density affecting EPP variation in intra- and interspecific comparisons has remained controversial, with more support from intraspecific comparisons. Neither experimental nor empirical studies have consistently provided support for the density hypothesis. Testing the density hypothesis is challenging because density measures may not necessarily reflect extra-pair mating opportunities, mate guarding efforts may covary with density, populations studied may differ in migratory behavior and/or climatic conditions, and variation in density may be insufficient. Accounting for these potentially confounding factors, we tested whether EPP rates within and among subpopulations of the reed bunting (Emberiza schoeniclus) were related to density. Our analyses were based on data from 13 subpopulations studied over 4 years. Overall, 56.4% of totally 181 broods contained at least one extra-pair young (EPY) and 37.1% of totally 669 young were of extra-pair origin. Roughly 90% of the extra-pair fathers were from the adjacent territory or from the territory after the next one. Within subpopulations, the proportion of EPY in broods was positively related to local breeding density. Similarly, among subpopulations, proportion of EPY was positively associated with population density. EPP was absent in subpopulations consisting of single breeding pairs, that is, without extra-pair mating opportunities. Our study confirms that density is an important biological factor, which significantly influences the amount of EPP within and among subpopulations, but also suggests that other mechanisms influence EPP beyond the variation explained by density.

Sperm Length Variation as a Predictor of Extrapair Paternity in Passerine Birds

BackgroundThe rate of extrapair paternity is a commonly used index for the risk of sperm competition in birds, but paternity data exist for only a few percent of the approximately 10400 extant species. As paternity analyses require extensive field sampling and costly lab work, species coverage in this field will probably not improve much in the foreseeable future. Recent findings from passerine birds, which constitute the largest avian order (∼5 900 species), suggest that sperm phenotypes carry a signature of sperm competition. Here we examine how well standardized measures of sperm length variation can predict the rate of extrapair paternity in passerine birds.Methodology/Principal FindingsWe collected sperm samples from 55 passerine species in Canada and Europe for which extrapair paternity rates were already available from either the same (n = 24) or a different (n = 31) study population. We measured the total length of individual spermatozoa and found that both the coefficient of between-male variation (CVbm) and within-male variation (CVwm) in sperm length were strong predictors of the rate of extrapair paternity, explaining as much as 65% and 58%, respectively, of the variation in extrapair paternity among species. However, only the CVbm predictor was independent of phylogeny, which implies that it can readily be converted into a currency of extrapair paternity without the need for phylogenetic correction.Conclusion/SignificanceWe propose the CVbm index as an alternative measure to extrapair paternity for passerine birds. Given the ease of sperm extraction from male birds in breeding condition, and a modest number of sampled males required for a robust estimate, this new index holds a great potential for mapping the risk of sperm competition across a wide range of passerine birds.

Synchrony does not explain extrapair paternity rate variation in northern or southern house wrens

Rates of extrapair paternity (EPP) vary widely among and within bird species, and 2 hypotheses suggest that this variation is driven by variation in breeding synchrony. These hypotheses make contradictory predictions, and each has some support from field studies, but the general relationship between EPP and synchrony remains unclear. We investigated EPP in relation to population-wide and local breeding synchrony in 2 populations of house wrens (Troglodytes aedon)—a migratory northern temperate (New York, USA) and a sedentary southern temperate (Buenos Aires, Argentina) population—that differ in numerous life-history traits. The northern population had significantly higher EPP rates and modestly but significantly higher local breeding synchrony. Population-wide breeding synchrony did not differ between populations. The proportion of extrapair young within a nest was not related to the nest’s population-wide or local synchrony index in either population. These results suggest that across divergent life histories in this species, breeding synchrony does not account for within-population variation in EPP.

Radical loss of an extreme extra-pair mating system

BackgroundMating outside the pair-bond is surprisingly common in socially monogamous birds, but rates of extra-pair paternity (EPP) vary widely between species. Although differences in life-history and contemporary ecological factors may explain some interspecific variation, evolutionary forces driving extra-pair (EP) mating remain largely obscure. Also, since there is a large phylogenetic component to the frequency of EPP, evolutionary inertia may contribute substantially to observed EP mating patterns. However, the relative importance of plasticity and phylogenetic constraints on the incidence of EP mating remains largely unknown.ResultsWe here demonstrate very low levels of EPP (4.4% of offspring) in the purple-crowned fairy-wren Malurus coronatus, a member of the genus with the highest known levels of EPP in birds. In addition, we show absence of the suite of distinctive behavioral and morphological adaptations associated with EP mating that characterize other fairy-wrens. Phylogenetic parsimony implies that these characteristics were lost in one speciation event. Nonetheless, many life-history and breeding parameters that are hypothesized to drive interspecific variation in EPP are not different in the purple-crowned fairy-wren compared to its promiscuous congeners.ConclusionSuch radical loss of an extreme EP mating system with all associated adaptations from a lineage of biologically very similar species indicates that evolutionary inertia does not necessarily constrain interspecific variation in EPP. Moreover, if apparently minor interspecific differences regularly cause large differences in EPP, this may be one reason why the evolution of EP mating is still poorly understood.

No evidence of genetic benefits from extra-pair fertilisations in female sand martins (Riparia riparia)

Genetic parentage studies of socially monogamous birds reveal a widespread prevalence of extra-pair paternity. Variation in extra-pair paternity among individuals may depend on how different individuals benefit from extra-pair fertilisations and on the opportunity to pursue extra-pair copulations. A long-term study of sand martins (Riparia riparia) in Hungary allowed us to examine patterns of extra-pair fertilisations in a large colony of over 3,000 breeding pairs with many known age individuals. We used multi-locus DNA fingerprinting to determine whether extra-pair fertilisations occur when females are paired to (1) presumably low quality mates, or (2) genetically similar or dissimilar mates, and whether extra-pair fertilisations result in offspring of higher quality. Extra-paternal young were found in 38% of 47 broods and comprised 19% of 190 offspring. Males that lost paternity did not differ significantly from others in age or body condition. Social mates of broods containing extra-pair offspring did not differ in genetic similarity from pairs without extra-pair offspring. Furthermore, there was no significant difference in body condition between extra-pair young and their maternal half-siblings. We were unable to assign paternity and therefore cannot exclude the possibility that extra-pair males differed from the within-pair males they cuckolded, in age, body condition or genetic similarity with the female. We found a positive relationship between paternity losses and breeding density, suggesting that low breeding density may constrain opportunities for seeking extra-pair copulations.

Pair identity — An important factor concerning variation in extra-pair paternity in the coal tit (Parus ater)

AbstractIn the majority of socially monogamous bird species, extra-pair paternity (EPP) has been shown to be common, while its frequency varies considerably among as well as within species. Our knowledge of factors affecting this variation, however, still remains comparatively poor. A thorough analysis of EPP patterns on an individual level may lead to a better understanding of the general diversity in EPP. Here, we analyse intra- and inter-annual as well as individual variation in the occurrence of EPP in first and second broods of coal tits (Parus ater) in three successive years. Based on a sample of 483 broods with 3559 offspring genotyped, we show that EPP was extraordinarily frequent in the study population, surpassing the values observed in other species of the genus Parus by far. On the population level, the rates of EPP were found to be remarkably similar between years, while there was pronounced intra-annual variation such that EPP rates increased significantly from first to second broods. Considering consecutive broods of individual females and males, the general occurrence of EPP was 'inconsistent' (i.e. EPP often affected only one of two broods), but the proportion of extra-pair young (EPY) per brood showed significant repeatability for both sexes in case of mate retention. When mate change occurred, repeatability collapsed, indicating that pair identity was more important in determining the repeatability of EPP than female or male identity alone. This was further supported by the fact that, besides female and male age, also their interaction (reflecting pair age combination) significantly predicted the proportion of EPY, though not in all breeding periods surveyed. Hence, the identity of the breeding pair (reflecting possible interactions of male and female characteristics) should be explicitly considered and accounted for in future studies investigating patterns of EPP on an individual level. Finally, we show that, depending on the boundary conditions, in principle, all coal tits experience EPP: When analysing three or more broods of the same individual, there were no completely 'faithful' females and also nearly no males, which were not cuckolded at least once.