Extra-pair Success Research Articles

Female differential allocation in response to extrapair offspring and social mate attractiveness.

Renewed debate over what benefits females might gain from producing extra‐pair offspring emphasizes the possibility that apparent differences in quality between within‐pair and extra‐pair offspring are confounded by greater maternal investment in extra‐pair offspring. Moreover, the attractiveness of a female's social mate can also influence contributions of both partners to a reproductive attempt. Here, we explore the complexities involved in parental investment decisions in response to extra‐pair offspring and mate attractiveness with a focus on the female point of view. Adult zebra finches paired and reproduced in a colony setting. A male's early‐life diet quality and his extra‐pair reproductive success were used as metrics of his mating attractiveness. Females paired with males that achieved extra‐pair success laid heavier eggs than other females and spent less time attending their nests than their mates or other females. Extra‐pair nestlings were fed more protein‐rich hen's egg than within‐pair nestlings. Females producing extra‐pair offspring had more surviving sons than females producing only within‐pair offspring. Collectively, results show that females differentially allocate resources in response to offspring extra‐pair status and their social mate's attractiveness. Females may also obtain fitness benefits through the production of extra‐pair offspring.

Extrapair mating and the strength of sexual selection: insights from a polymorphic species.

Extrapair mating could drive sexual selection in socially monogamous species, but support for this hypothesis remains equivocal. We used lifetime fitness data and a unique model species, the dimorphic white-throated sparrow (Zonotrichia albicollis), to examine how extrapair mating affects the potential for sexual selection. In this species, the morphs employ distinct reproductive strategies, with white males pursuing extrapair mating at higher rates than tan counterparts. Social and extrapair mating is disassortative by morph, with paternity exchange occurring primarily between pairs composed of white males and tan females. Bateman gradients and Jones indexes indicated stronger sexual selection via mate numbers in white males than in females and tan males, and generally did not differ between females as compared with tan males. Extrapair mating contributed more to the Bateman gradient for white than tan males, and white males also had higher variance in annual reproductive success. However, variance in lifetime reproductive success did not differ between morphs or sexes. Moreover, extrapair mating did not increase variance in male reproductive success relative to apparent patterns, and within-pair success accounted for much more variance than extrapair success. Thus, extrapair mating by white males increases Bateman gradients and the potential for sexual selection via mate numbers. However, our latter results support previous research suggesting that extrapair mating may play a limited role in driving the overall potential for sexual selection.

Male ornamentation and within-pair paternity are not associated with male provisioning rates in scarlet rosefinches Carpodacus erythrinus

As proposed by the ‘good parent model’ for evolution of secondary male ornamentation, secondary ornaments may signal male provisioning rates and, therefore, direct benefit to females. On the other hand, male parental care intensity can potentially be affected by the occurrence of extra-pair offspring in its nest. According to ‘parental investment theory’, males that lose paternity in their nests should reduce their parental care. In this study, we analyse potential relationships between intensity of parental care, male ornamentation, the occurrence of extra-pair paternity and male extra-pair fertilisation success in the scarlet rosefinch Carpodacus erythrinus. Our results based on 50 observed nests indicate no effect of paternity loss on the rate of food provisioning to nestlings in scarlet rosefinches. Simultaneously, we found no evidence for an association between male ornamentation and male provisioning rates. The only male trait associated with provisioning was the ability to sire extra-pair offspring. Our data indicate that direct selection against female promiscuity is weak or absent in rosefinches.

Multimodal sexual selection in a warbler: plumage and song are related to different fitness components

Many species possess multiple sexual signals that are produced in different modalities. Signals from different modalities may convey different or similar (redundant) information, or be intended for different receivers. We studied sexual selection on two such signals (plumage and song) in the common yellowthroat, Geothlypis trichas. We measured the coloration and size of both a carotenoid-based and a melanin-based plumage ornament and defined singing consistency as the lack of variability between repeated renditions of the same song. Overall, we found that plumage ornaments predicted within-pair mating success, while song consistency predicted extrapair success. Only plumage was related to male condition but, among experienced males, consistent singers had higher survival rates. Singing consistency was more variable than plumage, as performance varied across observations of the same male in relation to social context and date. Our results suggest that visual and acoustic signals in this species are most important in different sexual selection contexts.

The ecological–evolutionary interplay: density‐dependent sexual selection in a migratory songbird

Little is understood about how environmental heterogeneity influences the spatial dynamics of sexual selection. Within human-dominated systems, habitat modification creates environmental heterogeneity that could influence the adaptive value of individual phenotypes. Here, we used the gray catbird to examine if the ecological conditions experienced in the suburban matrix (SM) and embedded suburban parks (SP) influence reproductive strategies and the strength of sexual selection. Our results show that these habitats varied in a key ecological factor, breeding density. Moreover, this ecological factor was closely tied to reproductive strategies such that local breeding density predicted the probability that a nest would contain extra-pair offspring. Partitioning reproductive variance showed that while within-pair success was more important in both habitats, extra-pair success increased the opportunity for sexual selection by 39% at higher breeding densities. Body size was a strong predictor of relative reproductive success and was under directional selection in both habitats. Importantly, our results show that the strength of sexual selection did not differ among habitats at the landscape scale but rather that fine-scale variation in an ecological factor, breeding density, influenced sexual selection on male phenotypes. Here, we document density-dependent sexual selection in a migratory bird and hypothesize that coarse-scale environmental heterogeneity, in this case generated by anthropogenic habitat modification, changed the fine-scale ecological conditions that drove the spatial dynamics of sexual selection.

Superiority of extra-pair offspring: maternal but not genetic effects as revealed by a mixed cross-fostering design

Extra-pair copulations (EPC) are the rule rather than an exception in socially monogamous birds, but despite widespread occurrences, the benefits of female infidelity remain elusive. Most attention has been paid to the possibility that females gain genetic benefits from EPC, and fitness comparisons between maternal half-siblings are considered to be a defining test of this hypothesis. Recently, it was shown that these comparisons may be confounded by within-brood maternal effects where one such effect may be the distribution of half-siblings in the laying order. However, this possibility is difficult to study as it would be necessary to detect the egg from which each chick hatched. In this study, we used a new approach for egg-chick assignment and cross-fostered eggs on an individual basis among a set of nests of the collared flycatcher Ficedula albicollis. After hatching, chicks were ascribed to mothers and therefore to individual eggs by molecular genetic methods. Extra-pair young predominated early in the laying order. Under natural conditions, this should give them a competitive advantage over their half-siblings, mediated by hatching asynchrony. However, we experimentally synchronized hatching, and after this treatment, extra-pair young did not outperform within-pair young in any studied trait including survival up to recruitment and several indicators of reproductive success and attractiveness. We obtained only modest sample sizes for the last two traits and did not test for extra-pair success of male offspring. Thus, we cannot exclude the possibility of advantages of extra-pair young during the adult phase of life. However, our data tentatively suggest that the more likely reason for females' EPCs is the insurance against the infertility of a social mate.

Polygyny and extra-pair paternity enhance the opportunity for sexual selection in blue tits

Polygyny and extra-pair paternity are generally thought to enhance sexual selection. However, the extent to which these phenomena increase variance in male reproductive success will depend on the covariance between success at these two strategies. We analysed these patterns over four breeding seasons in facultatively polygynous blue tits Cyanistes caeruleus. We found that both polygyny and extra-pair paternity increased variance in male reproductive success and that standardised variance in annual number of genetic fledglings was 2.6 times higher than standardised variance in apparent success when assuming strict monogamy. Nevertheless, male success at securing within-pair paternity was unrelated to success at gaining extra-pair paternity and, when considering the positive effect of age on extra-pair success and attracting a second female, polygynous males were no more likely to sire extra-pair fledglings. Overall, polygynous males fledged more genetic offspring than monogamous males, but first-year polygynous males lost a greater share of within-pair paternity. A literature review suggests that this adverse effect of polygyny on within-pair paternity is frequent among birds, inconsistent with the prediction that females engage in extra-pair copulation with successful males to obtain good genes. Furthermore, a male's share of paternity was repeatable between years, and among females of polygynous males within years, such that a compatibility function of extra-pair copulations was likewise unsupported. Instead, we suggest that the observed patterns are most consistent with a fertility insurance role for extra-pair copulations, which does not exclude the greater opportunity for sexual selection through differential ability of males to gain paternity.

Multiple ornamentation, female breeding synchrony, and extra-pair mating success of golden whistlers (Pachycephala pectoralis)

Considerable variation exists in rates of extra-pair paternity between species, and across and within populations of the same species. Explanations for this variation include ecological (e.g. breeding synchrony), morphological (e.g. ornamentation), and genetic (e.g. relatedness) factors, but it is rare for studies to simultaneously explore these factors within a single population. This is especially true for highly ornamented species, where mate choice based on ornamentation may be more complex than in less-adorned species. We conducted such a study in a migratory population of the highly ornamented golden whistler (Pachycephala pectoralis). We quantified male genetic reproductive success and related it to a range of factors putatively involved in determining extra-pair mating success. We found no effects of genetic factors (male heterozygosity and relatedness) on extra-pair success, nor of territory size, male age, or incubation effort. Instead, males possessing yellower breast plumage and large song repertoires enjoyed higher reproductive success. Additionally, we found a negative relationship between local breeding synchrony and male extra-pair mating success. This may be a consequence of mate guarding during the female fertile period and an inability of males to simultaneously mate-guard and pursue extra-pair fertilisations. In this species, the opportunity for extra-pair matings appears to vary temporally with an ecological variable (local breeding synchrony), while fine-scale, inter-male differences in mating success may be influenced by individual attributes (male ornamentation). The migratory nature of the study population and its lack of natal philopatry may mean that relatedness and inbreeding avoidance are less important considerations in mate choice.

Dawn song in superb fairy-wrens: a bird that seeks extrapair copulations during the dawn chorus

Functional explanations of the dawn chorus in birds remain elusive. One hypothesis suggests that this acoustic display may play a role in female choice of extrapair males. Most young in cooperatively breeding superb fairy-wrens, Malurus cyaneus, are sired by extra-group males. Females initiate extra-group copulations exclusively through predawn forays to males singing in the dawn chorus. We measured variation between males in dawn singing at three levels of song production: song components, structured into songs, which make up recitals. We related this variation to independent measures of male quality and social status. Males sing two distinct categories of songs during the dawn chorus: a complex and variable chatter song and a more repeatable trill song. Dominant males with male subordinate helpers produced chatter songs at a greater rate than either dominant males without helpers or subordinates, suggesting a role in the competition between male group members. However, the trill song is implicated in female choice because older males sing songs with a longer trill component and have greater extrapair success, and trade-offs between phrases within the trill component imply constraints on the length of the trill component in entirety that could enforce honesty. The dawn chorus of the superb fairy-wren may thus have a duel role, involving enforcement of dominance among male group members (male–male competition), and signalling attractiveness to mates (female choice).

Fertilization success and UV ornamentation in blue tits Cyanistes caeruleus: correlational and experimental evidence

Cases where less ornamented males are favored through sexual selection are rare among birds. Here we show, based on data from 3 consecutive breeding seasons, that male blue tits with less ultraviolet (UV)-ornamented crown feathers sire more offspring. This pattern was mainly driven by the higher success of older, less UV-ornamented males at siring extrapair offspring. The reason behind this relationship is unclear although we hypothesize that being less UV-ornamented may enable adult males to intrude into nearby territories by mimicking juveniles. To test causality, we experimentally manipulated male crown coloration creating 2 groups, one with higher (UV(+) treatment) and one with lower UV reflectance (UV(−) treatment). Contrary to our expectations, UV(−) males were less likely to sire extrapair offspring than UV(+) males. The treatment had no effect on the likelihood of losing paternity in a male's own nest. Because the experimental evidence does not support the observational data, a direct effect of male crown color on extrapair success cannot be confirmed. However, potential pitfalls of this and other such color manipulation experiments, like fading of treatment with time and mismatches between behavior and coloration, call for new improved manipulation techniques and detailed behavioral observations to conclusively test for the effect of blue tit crown coloration on male extrapair success.

Extrapair paternity in chestnut-sided warblers is correlated with consistent vocal performance

The elaborateness of many bird songs is commonly presumed to have evolved under the influence of sexual selection by female mate choice. Thus, aspects of acoustic diversity, such as song repertoire size, are seen as likely targets of female choice. In many songbird species with song repertoires, however, the repertoires are small. In such species, female choice might be based on song features other than, or in addition to, song diversity. To investigate this conjecture, I assessed singing and paternity in a population of chestnut-sided warblers (Dendroica pensylvanica), a species in which song repertoires are of modest size. Twenty-two song traits were evaluated to determine which ones best predicted male extrapair reproductive success. The candidate traits encompassed measures of song diversity (e.g., song repertoire size), gross-scale song performance (e.g., singing rate), and fine-scale song performance (e.g., variability among songs in a bout). Regression analysis revealed that the best predictor of extrapair success was singing with little variability. In particular, the most successful males sang with consistent pitch and timing, as well as high pitch. The greater extrapair success of males with more consistent vocal performance may be due to female preference for such performance, which could be an indicator of male quality. Copyright 2007.

No Evidence of Current Sexual Selection on Sexually Dimorphic Traits in a Bird with High Variance in Mating Success

Sexual dimorphism, particularly in ornamental traits, is likely to have arisen by sexual selection. Most empirical and theoretical studies of sexual dimorphism assume that ongoing sexual selection also maintains the dimorphism. Over four seasons, I measured the sexual selection acting on three sexually dimorphic attributes (epaulet size, body size, and the blackness of the body plumage) of male red-winged blackbirds and found no consistent directional or stabilizing selection on any of them. Correlational selection was also negligible. I used path analysis to explore potential relationships in more detail but found no direct or indirect effects of male traits on either within- or extrapair success. Males who were resident on the marsh for more years had higher within-pair success, primarily because they spent more of the season on their territory. Experimental manipulations of epaulet size and color and the extent of nonblack feathers in the black body plumage had no detectable effect on the number of within-pair mates, paternity, or the number of extrapair offspring sired in nearby territories. These results combine with data from other studies of red-winged blackbirds to suggest that, despite high variation in male mating success and hence a strong opportunity for sexual selection, several morphological attributes that differ between the sexes and vary among males are not under current sexual selection. The possible explanations for why add complexity to our understanding of how sexual selection operates.

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.

Do female great tits (Parus major) assess males by eavesdropping? A field study using interactive song playback

Male singing behaviour correlates with extra-pair success in several passerine birds. Singing interactions during territorial contests provide relative information on the males involved. Such information may be important in female extra-pair behaviour and eavesdropping on singing interactions among males may allow females to make such relative assessments. We used interactive playback to instigate singing contests with male great tits during the peak fertile period of their mate in an attempt to alter females' assessment of mates' quality relative to neighbours (potential extra-pair partners). We escalated a contest to one male (by overlapping his songs) and then subsequently de-escalated a contest (by alternating) to a neighbour. Intrusions onto neighbouring territories by females mated to either treatment male were then monitored. Females mated to escalation treatment males were more likely to intrude following playbacks than females mated to de-escalation treatment males. Although the absolute song output of males did not differ between treatments, males produced more song relative to playback in de-escalation treatments and relative song output was positively correlated with female intrusions. Therefore, female great tits eavesdrop on singing interactions and change their visitation rates to neighbouring territories according to their mate's singing performance relative to neighbours.

Reproductive synchrony and extra-pair mating strategy in a socially monogamous bird,Dendroica petechia

Depending on the circumstances under which extra-pair mating occurs, theory makes opposing predictions about how reproductive synchrony should influence extra-pair paternity. This study investigated which sex initiated extra-pair mating in the yellow warbler, whether extra-pair behaviour and extra-pair paternity were related to reproductive synchrony, and whether synchrony affected the mating success of all males equally. Observations and captures of individuals making territorial intrusions indicated that males initiated extra-pair mating. Spatial patterns of male territorial intrusions and of extra-pair paternity were similar. Males’ extra-pair activity was reduced when their social partner was fertile. More offspring were sired by the extra-pair male when the female nested asynchronously with that sire's social mate. Neither population-wide synchrony, nor synchrony with neighbours, however, seemed to predict the incidence of extra-pair parentage or the identity of the sire, indicating factors other than synchrony were also important. Males with more breast streaking (a plumage ornament) were more successful as extra-pair sires, and were least affected by the constraint of synchrony. Larger males were less often cuckolded, and achieved extra-pair success mainly when their partner was not fertile. Thus, male yellow warblers apparently use different mating tactics depending on their plumage and size. More generally, the results suggest how mating strategies are affected by which sex initiates extra-pair mating and by the relative contributions of within-pair and extra-pair paternity to total reproductive success.

Extra–pair mating, male plumage coloration and sexual selection in yellow warblers (Dendroica petechia)

Extra-pair mating has been proposed as a source of sexual selection responsible for secondary sexual traits that are common among socially monogamous birds, although supporting evidence is scant. In the socially monogamous yellow warbler, males are larger than females, and unlike females, have extensive reddish streaking on their breasts. Using DNA fingerprinting we show that within-pair parentage was positively related to male size, and that extra-pair mating success was positively related to the amount of streaking on the breast. To our knowledge, this is the first intraspecific evidence of an association between a male plumage ornament and gains of extra-pair paternity that is apparently independent of age. This study confirms that extra-pair mating can be an important mechanism of sexual selection even when the most successful sires are commonly cuckolded, and refutes a previous hypothesis that the variation in plumage and behaviour among male yellow warblers is an example of alternative, equally successful, evolutionarily stable strategies (ESS). More generally, the demonstrated independence of within-pair and extra-pair success and their associated traits indicates that where animals have multiple secondary sexual traits, different traits may be selected by different mechanisms that contribute to total reproductive success.

GENETIC ESTIMATES OF ANNUAL AND LIFETIME REPRODUCTIVE SUCCESS IN MALE RED-WINGED BLACKBIRDS

DNA-based methods of determining parentage in birds have revealed that apparent reproductive success can be very different from true reproductive success. Our main goal in this study was to provide estimates of true annual and lifetime reproductive success of male Red-winged Blackbirds (Agelaius phoeniceus) to determine the importance of extra-pair mating to reproductive success. Males that were more successful at extra-pair mating, both annually and over their lifetimes, were also more successful on their own territories, so extra-pair success was not achieved at the expense of within-pair success. Relative to within-pair success, extra-pair success accounted for only one-fifth as much variance in total success (annual and lifetime). For some individuals, however, extra-pair success was a substantial part of their total success. Standardized variance in true annual success was 16% higher overall than the variance in apparent success, although the effect was inconsistent from year to year. Standardized variance in lifetime success increased 39% due to extra-pair paternity. Thus, extra-pair mating modestly increased the opportunity for sexual selection. Males with the highest annual success on their own territories mated with more females on their territories, had fewer of those females mating with other males, and lost fewer nests to predation. Annual extra-pair success was highly correlated with the number of extra-pair mates. The number of years a male bred and the number of females with which he mated contributed significantly to the variance in lifetime success, both within pairs and extra-pair. Reproductive success in the first breeding season was significantly positively correlated with lifetime success. Positive correlations between true and apparent reproductive success and between true mating success and harem size suggest that conventional (i.e., nongenetic) estimates of success provide reasonable estimates of population patterns, despite being unreliable for estimating individual success. Many of the ecological factors that affected lifetime success (e.g., territory acquisition, longevity) of males and females appeared similar between our Canadian population and a population in Washington state, USA. However, variance in male success appears to be much higher in Washington, suggesting that exploiting this geographical variation might be a productive way to study sexual selection in this species.

MEASURING THE EFFECTS OF PAIRING SUCCESS, EXTRA-PAIR COPULATIONS AND MATE QUALITY ON THE OPPORTUNITY FOR SEXUAL SELECTION.

Sexual selection can act through variation in the number of social mates obtained, variation in mate quality, or variation in success at obtaining extra-pair fertilizations. Because within-pair fertilizations (WPF) and extra-pair fertilizations (EPF) are alternate routes of reproduction, they are additive, rather than multiplicative, components of fitness. We present a method for partitioning total variance in reproductive success (a measure of the opportunity for selection) when fitness components are both additive and multiplicative and use it to partition the variance into components that correspond to each mechanism of sexual selection. Computer simulations show that extra-pair fertilizations can either increase or decrease total variance, depending on the covariance between within-pair and extra-pair success. Simulations also suggest that for socially monogamous species, extra-pair fertilizations have a greater effect than variation in mate quality or pairing status on the opportunity for selection. Application of our model to data gathered for a population of red-winged blackbirds (Agelaius phoeniceus) indicates that most of the variance in male reproductive success was attributable to within-pair sources of variance. Nevertheless, extra-pair copulations increased the opportunity for selection because males varied both in the proportion of their social young that they sired and in the number of extra-pair mates that they obtained. Furthermore, large and positive covariances existed between the number of extra-pair mates a male obtained and both social pairing success and within-pair paternity, indicating that, in this population, males preferred as social mates also were preferred as extra-pair mates.

Single-locus DNA fingerprinting reveals that male reproductive success increases with age through extra-pair paternity in the house sparrow (Passer domesticus)

Single-locus profiles produced with four cloned minisatellites were used to investigate the paternity of 44 house sparrow nestlings sired through extra-pair copulations. Comparison of the paternal alleles with a profile database of 144 candidate males resulted in 33 paternity assignments to 13 males. Both age and proximity were related to success in cuckoldry. Males achieving extra-pair paternity were usually breeding near to the cuckolded pair. Those at least two years old sired significantly more young through extra-pair matings, resulting in an average reproductive output 36% higher than that of yearlings. Whereas older males gained extra-pair fertilizations during their own breeding attempts, yearlings only achieved extra-pair success outside their own breeding cycle and did not significantly increase their own reproductive success. In contrast to other species in which extra-pair males are apparently chosen for their high quality, male sparrows siring extra-pair young were cuckolded at the same rate as other males.