Extra-pair Partners Research Articles

Extra‐pair paternity confirmed in wild white‐handed gibbons

Knowledge of the genetic mating system of animal species is essential for our understanding of the evolution of social systems and individual reproductive strategies. In recent years, genetic methods have uncovered an unexpected diversity of paternal genetic contributions across diverse animal social mating systems, but particularly in pair-living species. In most pair-living birds, for example, genetic and behavioral observations have confirmed a previously unknown significance of extra-pair copulations (EPCs) and extra-pair paternity. Among mammals, white-handed gibbons (Hylobates lar) are also known to live in pairs and are traditionally believed to be single-male single-female breeders. However, at Khao Yai National Park, Thailand, behavioral observations have confirmed the occurrence of both EPCs and functional multi-male grouping, but knowledge about the genetic mating system is still unavailable. In this study, we genotyped 89 white-handed gibbons of the Khao Yai population based on fecal samplings and were able to determine paternity for 41 offspring through short tandem repeat analysis. We found that females' stable social partners sired the majority (90.5%) of offspring (N = 38), while only a few (7.1%) offspring (n = 2 confirmed cases; n = 1 inferred case) were conceived with extra-pair partners. The paternity of one offspring remained inconclusive (2.4%), because the offspring's genotype did not mismatch with the genotypes of two potential sires. Like other predominantly pair-living species, gibbons appear to follow a mixed-reproductive strategy. The genetic mating system of wild white-handed gibbons is best described as flexible, primarily monogamous and opportunistically promiscuous. Inc.

Experimentally Simulating Paternity Uncertainty: Immediate and Long-Term Responses of Male and Female Reed Warblers Acrocephalus scirpaceus

In many socially monogamous species, both sexes seek copulation outside the pair bond in order to increase their reproductive success. In response, males adopt counter-strategies to combat the risk of losing paternity. However, no study so far has tried to experimentally prove the function of behaviour for paternity assurance. Introducing a potential extra-pair partner during the female fertile period provides a standardised method to examine how pair members respond immediately (e.g. increase mate guarding or copulation frequency) or long term (e.g. later parental investment and paternity uncertainty). In this study on a socially monogamous passerine species, we experimentally confronted pairs of reed warblers with a conspecific male (caged male simulating an intruder) during egg-laying. Our results revealed that occurrence of an intruder during that period triggered aggression against the intruder, depending on the presence of the female. The male territory owner also attacked the female partner to drive her away from the intruder. Thus territory defence in reed warblers also serves to protect paternity. The increase in paternity uncertainty did not affect later paternal investment. Paternal investment was also independent of the actual paternity losses. In females, the experiment elicited both, immediate and long-term responses. E.g. female copulation solicitations during the intruder experiment were only observed for females which later turned out to have extra-pair chicks in their nest. In relation to long term response females faced with an intruder invested later less in offspring feeding, and had less extra-pair chicks in their nests. Extra-pair paternity also seems to be affected by female quality (body size). In conclusion female reed warblers seem to seek extra-pair fertilizations but we could demonstrate that males adopt paternity assurance tactics which seems to efficiently help them to reduce paternity uncertainty.

Behavioural roles in booby mate switching

The behaviour that mediates divorce and partner change in socially monogamous species is largely unstudied and unknown, although roles and adaptive functions in some birds have been inferred from breeding outcomes, partial behavioural records and captive studies. Here, roles and functions of natural within-season mate switching of a colonial bird were characterized by describing interactions over many days during the whole daylight period. Switching occurred in 5.9% of bluefooted booby pairs and was initiated by desertion of males or, less commonly, females. Three male desertions were consistent with either the Errors of Mate Choice or Incompatibility hypotheses but were better explained by our new Unfaithful Mate hypothesis because infidelity of the mate preceded desertion. Another two male desertions were more consistent with the Better Options hypothesis because the males switched to their ongoing extra-pair (EP) partners. One female desertion was consistent with the Errors of Mate choice or Incompatibility hypotheses, and another with the Better Options hypothesis. All five deserted females switched promptly to their EP partners. Thus, most switches consisted of (1) males replacing a possibly unsatisfactory partner with an ew (already identif ied or to be identif ied) partner, or (2) deserted females pairing with their EP partners.

When mothers make sons sexy: maternal effects contribute to the increased sexual attractiveness of extra-pair offspring

Quality differences between offspring sired by the social and by an extra-pair partner are usually assumed to have a genetic basis, reflecting genetic benefits of female extra-pair mate choice. In the zebra finch (Taeniopygia guttata), we identified a colour ornament that is under sexual selection and appears to have a heritable basis. Hence, by engaging in extra-pair copulations with highly ornamented males, females could, in theory, obtain genes for increased offspring attractiveness. Indeed, sons sired by extra-pair partners had larger ornaments, seemingly supporting the genetic benefit hypothesis. Yet, when comparing ornament size of the social and extra-pair partners, there was no difference. Hence, the observed differences most likely had an environmental basis, mediated, for example, via differential maternal investment of resources into the eggs fertilized by extra-pair and social partners. Such maternal effects may (at least partly) be mediated by egg size, which we found to be associated with mean ornament expression in sons. Our results are consistent with the idea that maternal effects can shape sexual selection by altering the genotype-phenotype relationship for ornamentation. They also caution against automatically attributing greater offspring attractiveness or viability to an extra-pair mate's superior genetic quality, as without controlling for differential maternal investment we may significantly overestimate the role of genetic benefits in the evolution of extra-pair mating behaviour.

Natural extrapair paternity matches receptivity patterns in unguarded females: evidence for importance of female choice

While observational and experimental field studies have demonstrated that extrapair paternity is associated with a variety of male traits, there are still few published studies associating measures of paternity with behavioural preferences of females for extrapair males. In this sense, the majority of studies of extrapair paternity have failed to separate male competition from female choice. In this study we compared patterns of extrapair paternity in western bluebirds, Sialia mexicana, to previously published data on extrapair mating preferences from the same population. Results from previous experiments, where females’ mates were prevented from guarding or otherwise interfering with extrapair mating, accurately predicted patterns of paternity at 256 unmanipulated nests examined over a 6-year period. Specifically, as predicted based on which males were accepted as extrapair copulation partners, genetic extrapair sires were older but not larger than a female’s social mate. At the same time, relatively older extrapair males were not proportionally more successful, in terms of proportion of offspring sired, than younger males. Together these findings indicate that female choice for older males is expressed, but male competition, sperm competition and intersexual conflict also influence the magnitude of paternity. We suggest that experimental approaches, combined with measures of paternity under unmanipulated conditions, provide important opportunities to test the potential role of male versus female behaviours as selective factors in extrapair fertilizations.

Inclusive fitness of ‘kissing cousins’: new evidence of a role for kin selection in the evolution of extra-pair mating in birds

Social monogamy is nearly ubiquitous across avian taxa,but evidence from a proliferation of studies utilizing molecular paternity analysis suggests that sexual monogamy is the rare exception rather than the rule (Griffith et al. 2002). Efforts to explain the prevalence of extra-pair paternity (EPP) have largely focused on the potential fitness benefits for offspring genetic quality, as females are less likely to benefit directly from seeking extra-pair mates. In particular, there has been considerable interest in the degree to which EPP may represent an adaptive female strategy to avoid inbreeding (or outbreeding)depression when paired with a highly related (or unrelated)social mate (Kempenaers 2007). Others have argued that, because relatives share many genes identical by descent,females might increase their own inclusive fitness by providing additional breeding opportunities to genetically related males (Waser et al. 1986; Kokko & Ots 2006). Thus, in the absence of significant inbreeding depression, pursuing EPP with relatives should be favoured by kin selection, although there exist few unambiguous empirical examples of such preferences in the literature. In this issue of Molecular Ecology, Wang &Lu (2011) present an analysis of mating patterns with respect to genetic relatedness of social and extra-pair partners in the ground tit (Parus humilis), a facultative cooperative breeder in which socially monogamous pairs occasionally form cooperative groups with unpaired helper males (Fig. 1). Consistent with the predictions of the kin-selection hypothesis, females in both bi-parental and cooperative groups preferentially engaged in extra-pair matings with relatives, irrespective of relatedness to their social mates, and while suffering no apparent costs of inbreeding depression in their progeny. These finding shave several exciting implications for our understanding of avian mating system diversity and the evolution of cooperative breeding.

Female ground tits prefer relatives as extra-pair partners: driven by kin-selection?

Socially monogamous female birds routinely mate with males outside the pair bond. Three alternative hypotheses consider genetic benefits as the major driver behind the female strategy. The inbreeding avoidance hypothesis predicts that females paired with closely related males should seek copulations with distantly related extra-pair partners to avoid fitness loss from inbreeding depression; the outbreeding avoidance hypothesis predicts the opposite; the kin-selection hypothesis suggests that regardless of social mate relatedness, females should give related males extra-pair fertilization opportunities to gain inclusive fitness if the costs from inbreeding are minor. We test these hypotheses with a facultative cooperative breeder, the ground tit (Parus humilis). Social pairs of ground tits formed randomly with respect to genetic relatedness. In both bi-parental and cooperative groups, a female's engaging in extra-pair mating was independent of relatedness to her social mate; however, females preferred extra-pair sires to which they were more related than to their social mates. Moreover, females had higher relatedness with either their extra-group extra-pair sires in both bi-parental and cooperative groups, or within-group helper sires in cooperative groups, than expected by chance. When more than one potential extra-pair partner was available around a female's nest, she tended to select a relative. There was no indication of fitness reduction from extra-pair mating, which occurred at an intermediate level of inbreeding. These data support the kin-selection hypothesis, although there might be alternative nongenetic reasons associated with the extra-pair mating preference. Our finding offers a new explanation for why female birds pursue extra-pair mating. It also may broaden our understanding of the role of kin-selection in the evolution of cooperative society.

Genetic monogamy across variable demographic landscapes in cooperatively breeding Florida scrub-jays

Variation in ecological and demographic characteristics may alter the value of extrapair paternity (EPP) for socially monogamous species, thereby leading to variation in mating strategies among conspecific populations. Environmental factors influencing the need for parental care, and demographic factors influencing relatedness of social pairs or availability of unrelated extrapair partners, are both predicted to influence the direct and indirect benefits of EPP in cooperatively breeding birds. We examined genetic mating strategies in 3 long-term study populations of cooperatively breeding Florida scrub-jays (FSJs; Aphelocoma coerulescens) in which the value of EPP—or opportunities for it—was likely to vary: a fragmented site with a high frequency of inbreeding (potentially elevating the value of EPP as a means of increasing offspring heterozygosity); a suburban population with high rates of brood reduction (potentially elevating the value of shared parental investment); and a wildland site with a high frequency of unrelated breeders and opposite-sex auxiliaries (potentially elevating the opportunity for shared within-group parentage). Despite these differences, genetic monogamy dominated at all sites: 100% of the offspring sampled from the suburban site (144 offspring) and fragmented site (258 offspring), and 99.5% of offspring from the wildland site (367 of 369 offspring) were produced monogamously. Rare exceptions in our study populations demonstrate that, even in the FSJ, genetic monogamy is a plastic trait. The near ubiquity of genetic monogamy across 3 ecologically different study sites, however, suggests that this tendency toward monogamy is impervious to the population-level environmental and social variation that we documented. Key words: Aphelocoma coerulescens, extrapair paternity, Florida scrub-jay, genetic monogamy. [Behav Ecol]

Variation in social organization influences the opportunity for sexual selection in a social lizard

Social monogamy has traditionally been suggested to be maintained because of weak sexual selection on male partner acquisition. However, the ubiquitous incidence of extra-pair paternity suggests that sexual selection can be strong in monogamous systems, although studies partitioning variance in male reproductive success have come to mixed conclusions. Here, we use detailed field data to examine variance in male reproductive success and its implications for the maintenance of sociality in a population of the socially monogamous lizard Egernia whitii. We show that both within-pair and, to a lesser extent, extra-pair partner acquisition contribute to the variance in male reproductive success, resulting in considerable opportunity for sexual selection. Despite this, levels of multiple mating are lower in Egernia compared to other reptiles, suggesting that male partner acquisition is constrained. We suggest that this constraint may be a result of strong territoriality combined with sexual conflict over multiple mating generated by costs of extra-pair paternity to females as a result of facultative male care. This has the potential to limit sexual selection by reducing variance in male reproductive success and therefore contribute to the maintenance of complex social organization.

Long tails matter in sugarbirds—positively for extrapair but negatively for within-pair fertilization success

Extrapair mating is known to occur in many animals and potentially has a significant influence on reproductive success. Female extrapair mate choice may explain the occurrence of exaggerated ornaments in socially monogamous species, but the influence of ornamentation on extrapair mating success has rarely been investigated experimentally. Cape sugarbirds (Promerops cafer) are socially monogamous, almost always lay 2 egg clutches and have long-term pair bonds often lasting several seasons. However, they also display sexually dimorphic extravagant ornamentation in the form of a long, graduated tail and have one of the highest rates (65% of young) of extrapair paternity recorded in birds. We provide a test of the hypothesis that the ornamented tail is used in mate choice for extrapair partners by conducting an experiment in which tail length was manipulated after social mating but before copulation. This experiment therefore allowed females to respond to the manipulation of male tail length when making a choice of copulation partner but not social mate. We show that the tail length of male sugarbirds has a significant effect on the success of males in gaining extrapair paternity, with long-tailed males obtaining significantly more extrapair young than shorttailed males. However, males with short tails sire a significantly greater proportion of the nestlings in their own nests than longtailed individuals. This result suggests that males may adopt alternative strategies dependent on their ornamentation. Key words: extrapair mating, mate choice, Pomerops cafer, sexual selection, sugarbird, tail ornamentation. [Behav Ecol]

Signalling for food and sex? Begging by reproductive female white-throated magpie-jays

Food begging is common in nutritionally dependent young of many animals, but structurally homologous calls recur in adult signal repertoires of many species. We propose eight functional hypotheses for begging in adults; these stem from observations in birds but apply broadly to other taxa in which begging occurs. Adult cooperatively-breeding white-throated magpie-jays (Calocitta formosa) use loud begging vocalizations, particularly near the nest site during reproduction. We analysed the social context and behavioural phenology of loud calling and allofeeding in this species and compared these with predictions from each functional hypothesis. We found that reproductive females are the primary producers of beg calls, and their begging peaks during the fertile period when reproductive conflict among males and females was highest. Loud begging rates correlated positively with provisioning rates, but females called more in the pre-incubation fertile period than after they initiated incubation. Based on the context, phenology and active space of the signal, we conclude that female loud begging vocalizations function to signal nutritional need to group members, but also have been evolutionarily co-opted to advertise fertility to potential extra-pair partners. The location of calling is likely a consequence of nest guarding by breeding females to prevent intraspecific brood parasitism.

Sex differences in the motivation and mitigation of jealousy-induced interrogations

Given the differential costs sexual and emotional infidelity posed for each sex, we hypothesized that humans have sex-differentiated damage assessment strategies to investigate the nature of a partner’s infidelity. Study 1 tested this hypothesis using a forced-choice dilemma in which participants ( N = 172) indicated whether they would be more likely to inquire about the sexual or emotional nature of a partner’s extra-pair relationship. Results confirmed our predictions that (a) men more so than women inquire about the sexual nature of the extra-pair relationship and (b) women more so than men inquire about the emotional nature of the extra-pair relationship. We further hypothesized that humans have sex-differentiated damage control strategies to mitigate the costs of being caught committing infidelity. Study 2 tested this hypothesis using a forced-choice dilemma in which participants ( N = 117) indicated whether they would be more likely to deny the sexual or emotional nature of their involvement with an extra-pair partner. Results confirmed our predictions that if their partner discovered that they were involved with someone else, (a) men more so than women deny any emotional involvement with the extra-pair partner, whereas (b) women more so than men deny any sexual involvement with the extra-pair partner.

Evidence that pairing with genetically similar mates is maladaptive in a monogamous bird

BackgroundEvidence of multiple genetic criteria of mate choice is accumulating in numerous taxa. In many species, females have been shown to pair with genetically dissimilar mates or with extra-pair partners that are more genetically compatible than their social mates, thereby increasing their offsprings' heterozygosity which often correlates with offspring fitness. While most studies have focused on genetically promiscuous species, few studies have addressed genetically monogamous species, in which mate choice tends to be mutual.ResultsHere, we used microsatellite markers to assess individual global heterozygosity and genetic similarity of pairs in a socially and genetically monogamous seabird, the black-legged kittiwake Rissa tridactyla. We found that pairs were more genetically dissimilar than expected by chance. We also identified fitness costs of breeding with genetically similar partners: (i) genetic similarity of pairs was negatively correlated with the number of chicks hatched, and (ii) offspring heterozygosity was positively correlated with growth rate and survival.ConclusionThese findings provide evidence that breeders in a genetically monogamous species may avoid the fitness costs of reproducing with a genetically similar mate. In such species that lack the opportunity to obtain extra-pair fertilizations, mate choice may therefore be under high selective pressure.

Social Interactions among San Joaquin Kit Foxes Before, During, and After the Mating Season

Direct observations of small foxes are rare because of their secretive nature and nocturnal activity patterns. We observed 10 adult (� 1-year-old) San Joaquin kit foxes (Vulpes macrotis mutica) at night over a 7-month period that included the mating season to quantify the types, rates, and durations of social interactions. Of 191 interactions between members of this seasonally breeding, socially monogamous species, 52 were with adult foxes in the same social group, 43 with adult foxes from other known social groups, 48 with juveniles (,1 year old), and 48 with unknown foxes. Most interactions (n ¼ 173), especially those between pair-mates, between females, and with juveniles, were affiliative. Agonistic encounters were rare (n ¼ 18) and occurred mostly between males of different social groups and with unknown foxes. Interaction rates with social group members, most of which were pair-mates (88%), peaked in the months before the annual mating season, whereas those with non‐social group foxes, most of which represented potential extrapair copulation partners (79%), peaked later and more sharply. The duration of interactions with adult social and non‐social group foxes was highest during the mating season. Increased social interactions at this time may help foxes strengthen bonds with pair-mates and become familiar with potential extrapair partners.

Parental investment with a superior alien in the brood

When a parent's parentage differs across breeding attempts, established theory predicts that the parent should invest more in a brood when perceived parentage is high. We present a model of parental investment in which offspring unrelated to the parent have a competitive advantage over the parent's own offspring and take a larger share of investment. We show that this can weaken or, if the competitive advantage is great, reverse the predicted relationship between perceived parentage and parental investment. A moderate competitive advantage of extra-pair young over within-pair young could partly explain the lack of any clear relationship between paternal care and paternity in many studies, and could easily arise if females choose extra-pair partners for good genes. Our results are also relevant to interspecific avian brood parasitism. As parasites reared together with host offspring are often superior competitors, their hosts could benefit from increasing investment in response to suspected parasitism.

MHC-associated mating strategies and the importance of overall genetic diversity in an obligate pair-living primate

Mate choice is one of the most important evolutionary mechanisms. Females can improve their fitness by selectively mating with certain males. We studied possible genetic benefits in the obligate pair-living fat-tailed dwarf lemur (Cheirogaleus medius) which maintains life-long pair bonds but has an extremely high rate of extra-pair paternity. Possible mechanisms of female mate choice were investigated by analyzing overall genetic variability (neutral microsatellite marker) as well as a marker of adaptive significance (major histocompatibility complex, MHC-DRB exon 2). As in human medical studies, MHC-alleles were grouped to MHC-supertypes based on similarities in their functional important antigen binding sites. The study indicated that females preferred males both as social and as genetic fathers for their offspring having a higher number of MHC-alleles and MHC-supertypes, a lower overlap with female’s MHC-supertypes as well as a higher genome wide heterozygosity than randomly assigned males. Mutual relatedness had no influence on mate choice. Females engaged in extra-pair mating shared a significant higher number of MHC-supertypes with their social partner than faithful females. As no genetic differences between extra-pair young (EPY) and intra-pair young (IPY) were found, females might engage in extra-pair mating to ‘correct’ for genetic incompatibility. Thus, we found evidence that mate choice is predicted in the first place by the ‘good-genes-as-heterozygosity hypothesis’ whereas the occurrence of extra-pair matings supports the ‘dissassortative mating hypothesis’. To the best of our knowledge this study represents the first investigation of the potential roles of MHC-genes and overall genetic diversity in mate choice and extra-pair partner selection in a natural, free-living population of non-human primates.

Behavioural compatibility, extrapair copulation and mate switching in a socially monogamous parrot

Extrapair sexual behaviour may facilitate mate switching in monogamous species by allowing individuals to appraise and acquire a different mate. An individual benefits from mate switching if the new mate enhances the switcher's reproductive success more than the current mate. Several factors, including mate compatibility, may influence whether another individual represents a better option than the current mate. In a large, communally housed, captive colony of cockatiels, Nymphicus hollandicus, we tested three related hypotheses. (1) Members of pairs that show low mate behavioural compatibility should be more likely to engage in extrapair sexual behaviour than members of pairs with high mate behavioural compatibility. (2) Individuals that pursue extrapair sexual behaviour should be members of pairs with a high likelihood of reproductive failure. (3) Individuals that engage in extrapair sexual behaviour should later pair with their extrapair partner. We operationally defined pairmate relationships as more compatible when they involved lower within-pair aggression, less distance between mates and greater within-pair allopreening responsiveness, behavioural synchrony and copulation frequency than the opposite pattern of behaviour. Pairs in which neither pair member engaged in extrapair sexual behaviour had significantly higher behavioural compatibility scores than pairs in which at least one member did so. Every individual that engaged in extrapair copulation subsequently re-paired with their extrapair partner. This extrapair copulation and subsequent mate switching effectively dissolved relationships that had poor reproductive prospects and established new, reproductively successful relationships. This result suggests that extrapair sexual behaviour facilitates mate switching in cockatiels, and that behavioural compatibility is important in mate retention and divorce.

Ecological, social, and genetic contingency of extrapair behavior in a socially monogamous bird

Extrapair mating strategies are common among socially monogamous birds, but vary widely across ecological and social contexts in which breeding occurs. This variation is thought to reflect a compromise between the direct costs of mates’ extrapair behavior and indirect benefits of extrapair fertilizations (EPF) to offspring fitness. However, in most free‐living populations, the complete spatial and temporal distribution of mating attempts, genetic characteristics of available mates, and their relative contribution to EPF strategies are difficult to assess. Here we examined prevalence of EPF in relation to breeding density, synchrony, and genetic variability of available mates in a wild population of house finches Carpodacus mexicanus where all breeding attempts are known and all offspring are genotyped. We found that 15% of 59 nests contained extra‐pair offspring and 9% of 212 offspring were sired by extra‐pair males. We show experimentally that paired males and females avoided EPF displays in the presence of their social partners, revealing direct selection against EPF behavior. However, at the population level, the occurrence of EPF did not vary with nests dispersion, initiation date, synchrony, or with distance between the nests of extrapair partners. Instead, the occurrence of EPF closely covaried with genetic relatedness of a pool of available mates and offspring of genetically dissimilar mating tended to be resistant to a novel pathogen. These results corroborate findings that, in this population, strong fitness benefits of EPF are specific to each individual, thus highlighting the ecological, social, and genetic contingency of costs and benefits of an individual's extrapair behaviors.

A spatial genetic structure and effects of relatedness on mate choice in a wild bird population

Inbreeding depression, as commonly found in natural populations, should favour the evolution of inbreeding avoidance mechanisms. If natal dispersal, the first and probably most effective mechanism, does not lead to a complete separation of males and females from a common origin, a small-scale genetic population structure may result and other mechanisms to avoid inbreeding may exist. We studied the genetic population structure and individual mating patterns in blue tits (Parus caeruleus). The population showed a local genetic structure in two out of four years: genetic relatedness between individuals (estimated from microsatellite markers) decreased with distance. This pattern was mainly caused by immigrants to the study area; these, if paired with fellow immigrants, were more related than expected by chance. Since blue tits did not avoid inbreeding with their social partner, we examined if individuals preferred less related partners at later stages of the mate choice process. We found no evidence that females or males avoided inbreeding through extra-pair copulations or through mate desertion and postbreeding dispersal. Although the small-scale genetic population structure suggests that blue tits could use a simple rule of thumb to select less related mates, females did not generally prefer more distantly breeding extra-pair partners. However, the proportion of young fathered by an extra-pair male in mixed paternity broods depended on the genetic relatedness with the female. This suggests that there is a fertilization bias towards less related copulation partners and that blue tits are able to reduce the costs of inbreeding through a postcopulatory process.

Early birds are sexy: male age, dawn song and extrapair paternity in blue tits, Cyanistes (formerly Parus) caeruleus

Sexual selection theory predicts that signals reflecting the relative quality of individuals should be used in mate choice. Females could base their choice of copulation partners on male secondary sexual traits that honestly signal male age, as predicted by the age-based indicator mechanism. Studies have shown that female blue tits prefer older males and that aspects of dawn song reflect male quality, but it remains unknown whether dawn song characteristics correlate with male age. We compared dawn song characteristics of second-year (SY) and older (ASY) male blue tits (cross-sectional analysis), and tested for age-related changes within individuals (longitudinal analysis) and differential overwinter survival of SY males. We further investigated the relation between dawn song and paternity gain and loss. We found that ASY male blue tits began to sing earlier relative to sunrise than did SY males. This difference in the onset of dawn singing was due to age-related changes in individual performance rather than differential survival of individuals with varying expression of the trait. Males that began to sing earlier at dawn had more mating partners, and were more likely to gain extrapair paternity. Our findings suggest that the onset of dawn song can provide a simple mechanism for females to assess the relative quality of their mate and of neighbouring males. We propose that females use the onset of singing as a cue for their choice of extrapair partners.