Extra-pair paternity in tree swallows: why do females mate with more than one male?

Recent studies of monogamous tree swallows (Tachycineta bicolor) suggest that females may receive some type of genetic benefit from extra-pair fertilizations. In this study we attempted to determine what type of genetic benefits might be gained by females. We compared numerous morphological and behavioral traits (Table 1) of every male nesting on one grid of nest-boxes (n = 23) to determine what male traits were correlated with male success at gaining extra-pair fertilizations. DNA fingerprinting revealed an increase in the level of extra-pair paternity from the previous year (50% of broods contained extra-pair young in 1990 vs. 87% of broods in 1991), but no significant correlates of paternity. We found six extra-pair fathers at seven nests (20 nests had extra-pair young). The traits of these extra-pair males did not differ from those of the males they cuckolded. We discuss several reasons for this lack of difference, but argue that our results are not inconsistent with females choosing extra-pair males to enhance the genotypic quality of their offspring. Despite a complete search of the nest-box grid for extra-pair fathers, we were able to explain the paternity of just 21% (13/63) of all extra-pair young. This suggests that extra-pair fathers were either residents off our study grid or non-territorial floaters. Tree swallows are quite mobile and spend only part of the day at their nest prior to laying. In addition, we rarely see swallows visiting other grids of nest-boxes. Therefore, we suggest that most extra-pair copulations occur at some unknown location, possibly at a feeding or roosting area where females may be able to choose from many more potential extra-pair fathers than at their nest-site.     

Extra-pair paternity in eastern bluebirds: effects of manipulated density and natural patterns of breeding synchrony

The causes of variation in rates of extra-pair paternity among avian populations remain unclear, but could include environmental factors such as breeding density and synchrony. By experimentally manipulating nest site availability, we tested the effects of breeding density on the frequency of extra-pair paternity in eastern bluebirds (Sialia sialis). We also examined the role of breeding synchrony on extra-pair paternity using natural timing of nests. Microsatellite analysis revealed 34 of 305 nestlings (11.2%) were the result of extra-pair fertilizations; and 21 of 79 broods (26.6%) had at least one extra-pair nestling. Several measures of breeding density had independent effects on extra-pair paternity. First, experimental plot type affected extra-pair paternity, with 28 of 34 (82.4%) extra-pair young from nests in high density areas, and only six (17.6%) from nests in low density areas. Independently of plot type, the number of breeding neighbors within a 320-m radius was a significant predictor of the likelihood of extra-pair paternity at the nest. Extra-pair paternity was associated with temporal factors such as absolute timing of breeding and natural levels of local breeding synchrony, but only in bivariate comparisons. We found a positive interaction between density of neighbors within a 320-m radius and local breeding synchrony; this term reduced the main effects of synchrony and number of neighbors, but not experimental treatment. Our results demonstrate the importance of utilizing multiple aspects of proximity in breeding density analyses and testing for interactions between ecological factors that can influence the behavioral events leading to extra-pair fertilizations.     

Colonial and single breeding in fieldfares,Turdus pilaris L.: a comparison of nesting success in early and late broods

Summary Pairs of fieldfares breed singly or in colonies. Their breeding season was subdivided into an early part, when the trees were still leafless, and a late part, when the trees had leaves. Early colonial pairs has a significantly higher nesting success than single pairs. However, amongst late broods the nesting success of both categories of pairs was the same. Colonies of only two pairs show a weak trend suggesting that it could be advantageous for early broods to have a neighbour nearby, while the nesting success of late broods seems to be highest if the neighbour is farther away. The number of neighbours is more important than the distance to the nearest neighbour when colonies of different sizes are examined. The nesting success of colonial broods is also influenced by the time of breeding. Success increases with colony size, in early colonies, but the opposite applies for the late breeding season.A little owl was placed near single and colonial nests to show how communal defence contributed to the greater success of early colonial nests. Predators entering a colony risk being contaminated by significantly more faeces due to the higher rate of attacks by fieldfares using aimed defecation near colonial nests and these could affect the predators' flying ability.The possibility of fieldfare colonies serving as information centres for food finding is also discussed.     

Social and ecological factors affecting paternity allocation in American robins with overlapping broods

Factors that affect extra-pair mating in birds are likely to vary across the breeding season. Changing densities of active nests may alter the opportunities for extra-pair mating, and parental duties may alter a male’s opportunity to guard his mate from extra-pair mating. The latter affects species with multiple broods, where males care for fledglings from first nests while females initiate second nests. We studied a population of multi-brooded American robins (Turdus migratorius) to assess how seasonal changes in nesting density and changes in mate-guarding opportunity influenced paternity patterns over successive breeding attempts. Extra-pair paternity (EPP) occurred in 71.9% of broods and accounted for 48.1% of young. High nesting densities in the study population may explain the high overall rate of EPP, but seasonal variation in breeding density did not explain patterns of EPP among nests. Contrary to the predictions of the mate-guarding hypothesis, EPP did not increase in the second nests that followed successful first nests, and the percentage of extra-pair young in second nests did not decline as the overlap between successive nests increased. The fact that EPP was actually lower when the interval between clutches was shorter suggests that the sooner the males can assume sole care of first broods and allow their mates to renest (indicative of superior paternal quality), the more paternity they realize in the next nest. These results suggest that mate-guarding opportunity does not influence paternity in this population of American robins and that female robins may allocate paternity based on their assessment of male parental performance at first nests.     

Extra-pair paternity and tail ornamentation in the barn swallow Hirundo rustica

Females of socially monogamous species may copulate with attractive non-mates to obtain access to the genes of such males, and a preference for attractive copulation partners may result in sexual selection. Extra-pair copulations are common in the socially monogamous barn swallow Hirundorustica, and a 2-year study of paternity using multi-locus DNA fingerprinting demonstrated that 33% of 63 broods and 28% of 261 offspring were sired by extra-pair males. The frequency of extra-pair offspring within broods was highly skewed with the majority of all broods having either no extra-pair offspring or only extra-pair offspring. Individual pairs were consistent in their frequency of extra-pair paternity among broods, and the repeatability of extra-pair paternity of multiple broods of the same female was statistically significant. The proportion of extra-pair offspring was negatively related to the tail length of the male attending the nest. Behavioural observations showed that extra-pair fertilizations were more likely in broods raised by females that had been observed to engage in extra-pair copulations. The frequency of extra-pair offspring was unrelated to the intensity of two male paternity guards, mate guarding and the rate of intra-pair copulations. In an analysis of extra-pair paternity and male parental care in different broods of the same male, male barn swallows fed their offspring relatively less frequently if the brood contained more extra-pair offspring. Therefore, female barn swallows pursue extra-pair copulations with attractive males, which may result in sexual selection, even though extra-pair paternity is costly for females due to the reduction of paternal care by their social mates. Received: 24 January 1997 / Accepted after revision: 2 August 1997     

The high frequency of extra-pair paternity in tree swallows is not an artifact of nestboxes

A common criticism of nestbox studies is one of creating artificial nesting conditions and breeding behavior different from what would be seen under natural conditions. We assessed the frequency of extra-pair paternity (percentage of broods with at least one extra-pair young) in 25 families of tree swallows (Tachycineta bicolor) nesting in natural cavities and compared it to that in a nestbox population. We found that 84% of females nesting in natural cavities obtained fertilizations from extra-pair males. These extra-pair males fathered 69% of all nestlings. Studies of tree swallows breeding in nestboxes have shown that 50–87% of broods contained extra-pair young, with extra-pair males fathering 38–53% of all the young. In broods with extra-pair paternity, natural cavities contained a significantly greater proportion of extra-pair young than did nestboxes. Despite differences in nesting habitat and female age structure, the frequency of extra-pair paternity did not differ significantly between the natural-cavity and nestbox populations. Therefore, the presence of extra-pair paternity in tree swallows is not an artifact of nestboxes or of artificial nesting conditions. Received: 2 May 1995/Accepted after revision: 14 January 1996     

Breeding synchrony and paternity in the barn swallow (Hirundo rustica)

The barn swallow (Hirundo rustica) is a socially monogamous passerine which usually breeds in colonies where extra-pair copulations are frequent. Males intensively guard their mates during the female fertile period. Since males are more likely to be available for extra-pair copulations when their mate is not fertile, synchrony in timing of breeding may affect paternity of individual males. In this study, we analysed the change in mate-guarding rate by males in relation to the fertility condition of the female, and the relationships between breeding synchrony and density with paternity in first broods of 52 male barn swallows. Paternity (proportion of nestlings fathered in own brood) was assessed by typing of three highly polymorphic microsatellite loci. Mate guarding by males peaked during the fertile period of their mates. Paternity increased as breeding synchrony in the colony increased. Paternity of barn swallows is positively associated with the degree of exaggeration of male tail ornaments. The relationship between male ornamentation and paternity was partly mediated by an effect of ornament size on breeding synchrony. We suggest that females might delay breeding with low-quality males to enhance their opportunities for being fertilised by high-quality extra-pair males. Received: 16 January 1998 / Accepted after revision: 25 October 1998     

Genetic mating system and timing of extra-pair fertilizations in the Kentish plover

It is still unclear why females in many bird species pursue extra-pair copulations. Current hypotheses focus mainly on indirect benefits such as obtaining particular good genes for their offspring or maximizing genetic compatibility between themselves and the father of their offspring. Supporting the latter, a recent study of shorebirds suggests that extra-pair matings may function to avoid the negative effects of genetic similarity between mates. Here, we further investigate genetic parentage in the Kentish plover, Charadrius alexandrinus, a shorebird with a highly variable social mating system. DNA fingerprinting revealed that most pairs were genetically monogamous: 7.9% of the broods (7/89) contained extra-pair young, comprising 3.9% of all chicks (9/229). These cases represented, however, three alternative reproductive behaviors: extra-pair paternity, quasi-parasitism (extra-pair maternity) and intraspecific brood parasitism. This is the first study showing the occurrence of all three behaviors in one shorebird species. We also found that extra-pair fertilizations (extra-pair paternity and quasi-parasitism) were more frequent later in the breeding season. There was no consistent relationship between genetic similarity of mates and laying date; the pattern, as well as the degree of genetic similarity, differed among breeding sites within the study population.Communicated by M. Webster     

Is female promiscuity constrained by the presence of her social mate? An experiment with bluethroats <Emphasis Type="Italic">Luscinia svecica</Emphasis>

Extra-pair paternity is common in birds and much research has focussed on the selective advantage of extra-pair matings for both sexes. In contrast, little attention has been given to the fact that in most species the majority of offspring are sired by the social male. We investigated whether extra-pair matings of female bluethroats (Luscinia svecica) are constrained by the presence of the pair male, by detaining males in cages on their territories for one morning during the peak of female fertility. The proportion of offspring sired by extra-pair males was higher in broods where males had been detained (35%) than in control broods (16%), while the proportion of broods that had at least one extra-pair offspring did not differ significantly between experimental (65%) and control broods (44%). Within the experimental group, levels of extra-pair paternity were not related to the day of experiment in relation to start of egg laying, but males caught early in the morning lost more paternity than males caught later on. Our results show that pair males exert constraints on the frequency of extra-pair paternity by being present during the period of peak fertility, which could be a direct effect of their mate guarding effort and/or due to an advantage in sperm competition for pair males.     

Multiple paternity and offspring quality in tree swallows

There is mounting evidence in a variety of taxa that females increase offspring quality by mating with multiple males, often resulting in multiple paternity. In birds, however, few studies have explicitly examined the benefits of mating with several different males; instead, the focus has been on whether or not extra-pair mating occurs, and its adaptive significance remains controversial. We examined the hypothesis that offspring quality, particularly immune response (phytohaemagglutinin assay) and growth, increases with the number of sires in broods of socially monogamous tree swallows (Tachycineta bicolor). We found one of the highest known levels of multiple paternity in birds (84% of nests with two or more extra-pair young had at least two extra-pair sires). Among nests with extra-pair young, the number and diversity of sires continued to increase linearly with the number of extra-pair young, so there was no evidence that some males monopolized paternity at high levels of extra-pair fertilization. Indeed, the number of sires was actually greater than expected in large broods, suggesting that some females might be seeking more mates. We found no effect of the number of sires on nestling immune response or growth. In mixed paternity broods, the immune response of extra-pair young did not differ from that of their within-pair half-siblings. However, among all broods, nestlings had a stronger immune response in nests with at least one extra-pair nestling than in nests with all within-pair nestlings. These results are not consistent with a good genes benefit of extra-pair mating, but they do suggest that there are environmental effects associated with extra-pair mating that increase nestling immune response. These environmental effects could produce indirect genetic effects on sexual selection if they are heritable. The extraordinarily high number of sires in this species highlights a relatively unexplored source of sexual selection in birds.     

Multiple paternity in the cooperatively breeding fish <Emphasis Type="Italic">Neolamprologus pulcher</Emphasis>

In cooperative breeders, mature males may compete for fertilizations. In this study, we measured the degree of multiple paternity in a natural population of a cooperatively breeding fish. Neolamprologus pulcher (Perciformes: Cichlidae) is a highly social cichlid endemic to Lake Tanganyika. We used highly variable microsatellite loci to survey 12 groups with an average number of 10.6 brood care helpers per group and a total of 43 offspring (mean 3.6 per brood). In 11 of 12 groups, all young were assigned to the dominant female. The dominant male sired all offspring in three groups, part of the offspring in four groups, and in five groups, he had no paternity at all. In total, 44.2% of young were not fathered by the current male territory owner. Multiple paternity was found in 5 of 12 broods (41.7 %), with 8 of 35 young (22.9 %) being sired by males other than the respective territory owners. This is an exceptionally high rate of extra-pair paternity among cooperatively breeding vertebrates. Neither helpers present in these territories during collection nor neighbouring males were unequivocally assigned to have sired these extra-pair young. However, behavioural observations suggest that male helpers may have produced these young before being expelled from the territory in response to this reproductive parasitism. We discuss these results in the light of reproductive skew theory, cooperative breeding in vertebrates and alternative reproductive tactics in fish.     

Extra-pair paternity, offspring mortality and offspring sex ratio in the socially monogamous coal tit (Parus ater)

Females of many socially monogamous bird species commonly engage in extra-pair copulations. Assuming that extra-pair males are more attractive than the females’ social partners and that attractiveness has a heritable component, sex allocation theory predicts facultative overproduction of sons among extra-pair offspring (EPO) as sons benefit more than daughters from inheriting their father’s attractiveness traits. Here, we present a large-scale, three-year study on sex ratio variation in a passerine bird, the coal tit (Parus ater). Molecular sexing in combination with paternity analysis revealed no evidence for a male-bias in EPO sex ratios compared to their within-pair maternal half-siblings. Our main conclusion, therefore, is that facultative sex allocation to EPO is absent in the coal tit, in accordance with findings in several other species. Either there is no net selection for a deviation from random sex ratio variation (e.g. because extra-pair mating may serve goals different from striving for ‘attractiveness genes’) or evolutionary constraints preclude the evolution of precise maternal sex ratio adjustment. It is interesting to note that, however, we found broods without EPO as well as broods without mortality to be relatively female-biased compared to broods with EPO and mortality, respectively. We were unable to identify any environmental or parental variable to co-vary with brood sex ratios. There was no significant repeatability of sex ratios in consecutive broods of individual females that would hint at some idiosyncratic maternal sex ratio adjustment. Further research is needed to resolve the biological significance of the correlation between brood sex ratios and extra-pair paternity and mortality incidence, respectively.     

Positive association between social and extra-pair mating in a polygynous songbird, the dickcissel (Spiza americana)

In polygynous species, males appear to gain additional offspring by pairing with multiple females simultaneously. However, this may not be true if some females copulate outside of the social pair bond. Polygynous males could experience lower paternity because of trade-offs among gaining multiple social mates, guarding fertility with these mates, and pursuing extra-pair matings. Alternatively, polygynous males could simultaneously gain extra social mates and have high paternity, either because of female preferences or because of male competitive attributes. We tested four predictions stemming from these hypotheses in a facultatively polygynous songbird, the dickcissel (Spiza americana). Unlike most previous studies, we found that males with higher social mating success (harem size) also tended to have higher within-pair paternity and that the number of extra-pair young a male sired increased significantly with his social mating success. Females that paired with mated males were not more likely to produce extra-pair young. In contrast, extra-pair paternity was significantly lower in the nests of females whose nesting activity overlapped that of another female on the same territory. This pattern of mating was robust to differences in breeding density. Indeed, breeding density had no effect on either extra-pair mating or on the association between polygyny and paternity. Finally, nest survival increased with harem size. This result, combined with the positive association between polygyny and paternity, contributed to significantly higher realized reproductive success by polygynous male dickcissels.     

Declining extra-pair paternity with laying order associated with initial incubation behavior,but independent of final clutch size in the blue tit

Although functional explanations for female engagement in extra-pair copulation have been studied extensively in birds, little is known about how extra-pair paternity is linked to other fundamental aspects of avian reproduction. However, recent studies indicate that the occurrence of extra-pair offspring may generally decline with laying order, possibly because stimulation by eggs induces incubation, which may suppress female motivation to acquire extra-pair paternity. Here we tested whether experimental inhibition of incubation during the laying phase, induced by the temporary removal of eggs, resulted in increased extra-pair paternity, in concert with a later cessation of laying, in blue tits (Cyanistes caeruleus). As expected, experimental females showed a more gradual increase in nocturnal incubation duration over the laying phase and produced larger clutches than controls. Moreover, incubation duration on the night after the first egg was laid predicted how extra-pair paternity declined with laying order, with less incubation being associated with more extra-pair offspring among the earliest eggs in the clutch. However, incubation duration on this first night was unrelated to our experimental treatment and independent of final clutch size. Consequently, the observed decline in extra-pair paternity with laying order was unaffected by our manipulation and larger clutches included proportionally fewer extra-pair offspring. We suggest that female physiological state prior to laying, associated with incubation at the onset of laying, determines motivation to acquire extra-pair paternity independent of final clutch size. This decline in proportion of extra-pair offspring with clutch size may be a general pattern within bird species.     

Sperm storage reflects within- and extra-pair mating opportunities in a cooperatively breeding bird

In passerine birds, storage and maturation of sperm takes place in the cloacal protuberance (CP), an external swelling of the reproductive organ. The considerable variation in CP size among species is presumed to be a consequence of varying levels of sperm-competition, but whether individual variation in CP size within a species also reflects sperm competition is not well established. Here, we study temporal variation in male CP size in relation to within-pair and extra-pair mating opportunities and cuckoldry risk in purple-crowned fairy-wrens Malurus coronatus. This is a socially monogamous cooperatively breeding passerine that can breed year-round and has low levels of extra-pair paternity (in 6?% of broods). We show that male CP size sharply increased a few weeks before, and rapidly regressed after his partner laid eggs, consistent with a cost of its maintenance and/or sperm production. Surprisingly, despite low levels of extra-pair paternity, CP size of non-breeding and pre-breeding males was positively correlated with the number of breeding females in the population, suggesting that CP size is sensitive to extra-pair mating opportunities. However, CP sizes do not seem to reflect cuckoldry risk: CP size of dominant males was unaffected by the presence of a subordinate that was unrelated to the dominant female, although those subordinates occasionally sire offspring, and had a larger CP than subordinates living with their mother. Our results suggest that, even in a species with low levels of extra-pair paternity, individual investment in sperm storage reflects both within-pair and, albeit to lesser extent, extra-pair mating opportunities.     

Sperm competition and sexual selection: a meta-analysis of paternity studies of birds

Sperm competition (the competition among the sperm of different males for fertilization of the eggs of a female) has been suggested to be an important component of sexual selection, but no general assessment has been made of this proposition. We used a meta-analytic approach to assess the extensive literature on paternity (the proportion of offspring in a focal nest sired by an attending male) in birds based on allozyme and molecular techniques. The relative variance in male mating success was on average increased by a factor of 4.6 over the variance in apparent male success. Males with more extravagant secondary sexual characters had higher paternity in their own nests than less adorned males. There was a weak effect of male age being positively associated with paternity in own nests. Male body size measured as the length of wing and tarsus was weakly positively associated with paternity in own nests. Male survival prospect was positively associated with paternity in own nests. Polygynous males generally had decreased paternity of their broods compared to monogamous males. Paternity of the resident male decreased with increasing population density and breeding asynchrony. The intensity of paternity guards such as within-pair copulation rate and mate guarding were not significantly related to extra-pair paternity. Sperm competition was thus an important component of sexual selection by increasing the variance in male mating success, and by being associated with the expression of secondary sexual characters, in particular in dense and asynchronously breeding populations of birds. Received: 12 February 1998 / Accepted after revision: 31 May 1998     

Breeding synchrony and extra-pair mating in red-winged blackbirds

Using data from a 6-year paternity study of red-winged blackbirds, I tested the hypotheses that increased nesting synchrony should either promote extra-pair mating by increasing the advantage of extra-pair mating to females, or decrease extra-pair mating by constraining males from seeking extra-pair copulations. Contrary to these hypotheses, the occurrence of extra-pair paternity did not vary with nesting synchrony over the breeding season, or vary with the number of synchronous nests within territories or within marshes, or with nesting order on territories. However, for nearly all nests with extra-pair young, there were fewer females synchronous with that nest on the cuckolder's territory than on the territory of the cuckolded male. This “advantage” of a synchrony difference was less pronounced for older males that cuckolded younger males, particularly when the two males were not neighbors. Collectively, these results suggest that breeding synchrony affects extra-pair mating by affecting mate guarding, but that breeding synchrony alone can not be used to predict which females are more likely to engage in extra-pair mating, nor with which extra-pair males they will mate. Understanding why extra-pair mating by older males is less affected by breeding synchrony may explain much about both the proximate and ultimate causes of extra-pair mating in red-winged blackbirds. Received: 7 June 1996 / Accepted after revision: 25 November 1996     

Male tail streamer length predicts fertilization success in the North American barn swallow (Hirundo rustica erythrogaster)

In socially monogamous species, extra-pair paternity has the potential to increase the variance in male reproductive success, thereby affecting the opportunity for sexual selection on male extravagant ornamentation. In the European barn swallow (Hirundo rustica rustica), the tail streamer length is a sexually selected male ornament and an honest indicator of viability. The North American barn swallow (Hirundo rustica erythrogaster) also shows sexual dimorphism in tail streamer length, but whether this trait holds the same signalling function in this subspecies is a controversial issue, and the available literature is presently scarce. Here, we present data on paternity in the North American barn swallow, including a complete sampling of extra-pair sires in four colonies. We analysed how extra-pair paternity affected the variance in male fertilization success and examined whether male tail streamer (i.e. the outermost tail feather) length correlated with fertilization success (n=86 males). Extra-pair paternity constituted 31% of all offspring and significantly increased the variance in male fertilization success. The number of offspring sired by extra-pair males accounted for almost half of the total variance in male fertilization success. Males with naturally long tail streamers had a higher fertilization success than males with shorter tail streamers, and this pattern was mainly caused by a higher extra-pair success for males with long tail streamers. Males with long tail streamers also paired with early breeding females in prime body condition. These results are consistent with the idea that there is directional sexual selection on male tail streamer length, possibly mediated through male extra-pair mating success or the timing of breeding onset.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

Extra-pair paternity and the reproductive role of male floaters in the tree swallow (Tachycineta bicolor)

In many avian species, a part of the population is present at the breeding grounds but does not breed. Current theories generally assume that floaters are younger or lower-quality individuals, and empirical data confirm this. However, floating could also arise as an alternative strategy to breeding, if floaters are able to reproduce via extra-pair copulations. Until the present study, there has been no evidence that floaters father offspring. We studied a population of tree swallows (Tachycineta bicolor), a species with one of the highest levels of extra-pair paternity known in birds. Using microsatellite markers, we determined the biological fathers of 65% of the extra-pair young. Of a total of 53 extra-pair young (52% of all offspring), 47% were fathered by local residents, 6% by residents breeding elsewhere (up to 2 km from the focal grid), and 13% by floaters. Residents seemed to be more successful and they were also more likely to return as territory holders in the next breeding season compared to floaters. Extra-pair males were on average in better condition than the within-pair males they cuckolded. Interestingly, resident males that disappeared (possibly to float) during the fertile period were heavier than males that stayed, and floaters were heavier than residents, but not different in any other characteristic. Although alternative interpretations of the data are possible, we propose that floating might be a conditional strategy in tree swallows whereby males in good condition gain more paternity via extra-pair copulations, whereas males in worse condition are more successful by providing parental care.