性选择、配偶外父亲身份确认程度、遗传变异性和保护

岛屿动物中的性选择强度不高,其原因可能是由于岛屿种群的遗传变异性水平较低。本文作者检验了鸟类岛屿种群是否具有较低的遗传变异性、性选择强度大的种群是否具有较高的突变输入率（rate of mutational input),在鸟类岛屿种群中是否具有较低的性选择强度（可以根据配偶外父亲身份的频率来估计）。小卫星共有谱带系数（minisatellite band sharing coefficient)可确定无亲源关系个体之间的遗传变异性,对与遗传变异性有关的雄性个体的父亲（paternity)进行了成对比较以检验如下假说：在具有较多遗传变异的种群中,雌性个体更经常地进行配偶外交配。在小卫星谱带系数较低的鸟类种群中,配偶外父亲的频次较高。对岛屿和大陆鸟类进行的第二个比较分析表明：岛屿种群中的配偶外父亲频次较低,遗传变异性也较低,其部分原因在于突变输入（mutational input)减少。上述发现表明：（1）父亲确认程度（parternity)随遗传变异性的数量而增加;（2）在遗传变异性较大的种群中,突变率较高,性选择的程度更激烈;（3）岛屿种群中性选择的强度一般比大陆种群弱。这对于理解遗传变异性的空间变异、理解岛屿种群和其它隔离种群的保护问题有重要启示。     

Extra-pair paternity in the socially monogamous mountain bluebird Sialia currucoides and its effect on the potential for sexual selection

Sexual selection theory posits that ornamental traits can evolve if they provide individuals with an advantage in securing multiple mates. That male ornamentation occurs in many bird species in which males pair with a single female is therefore puzzling. It has been proposed that extra-pair mating can substantially increase the variance in reproductive success among males in monogamous species, thus increasing the potential for sexual selection. We documented the frequency of extra-pair paternity and examined its effect on variation in male reproductive success in the mountain bluebird Sialia currucoides , a socially monogamous songbird in which males possess brilliant plumage ornamentation. Extra-pair paternity was common in our Wyoming study population, with 72% of broods containing at least one extra-pair offspring. The standardized variance in actual male reproductive success (i.e., the total number of within-pair and extra-pair offspring sired) was more than seven times higher than the variation in apparent success (i.e., success assuming that no extra-pair mating occurred). Success at siring within-pair and extra-pair offspring both contributed to the variation in overall male reproductive success. Within-pair success, however, did not predict a male's level of extra-pair success, suggesting that males do not sacrifice within-pair paternity to gain extra-pair paternity. Calculation of the sexual selection (Bateman) gradient showed that males sire approximately two additional offspring for each extra-pair mate that we identified. Thus, in this sexually dichromatic species, extra-pair mating increases the variance in male reproductive success and provides the potential for sexual selection to act.     

Contrasting levels of extra-pair paternity in mainland and island populations of the house sparrow (Passer domesticus): is there an 'island effect'?

Despite the many studies that have investigated the genetic mating system of socially monogamous birds, very little is known about the underlying causes of extra-pair paternity and few studies have attempted to test those hypotheses which have been suggested. This study describes die analysis of die genetic mating system of two populations of the house sparrow [Passer domesticus) , and uses the results from four other populations to test existing hypodieses using an intra-specific comparative approach. The parentage analysis was conducted using a combination of published and newly presented microsatellite loci isolated from the house sparrow. One population in Kentucky, U.S.A. was found to contain what may be considered to be a typical level of extra-pair paternity for mis species (10.5%, 19/185 offspring). The second, a population on the island of Lundy, UK, exhibited a very low level (1.3%, 4/305 offspring), significandy lower dian that in all the other populations studied so far. The finding of such diverse rates of extra-pair paternity, along with the existing estimates from ofher populations, has allowed us to test the effects of breeding density and genetic variation on die level of extra-pair paternity. We found no effect of either factor on the frequency of extra-pair paternity in the house sparrow, leaving the cause of this variation open to fresh ideas.     

Genetic similarity, breeding distribution range and sexual selection

Large populations with extensive breeding distributions may sustain greater genetic variability, thus producing a positive relationship between genetic variation and population size. Levels of genetic variability may also be affected by sexual selection, which could either reduce levels because a small fraction of males contribute to the following generation, or augment them by generating genetic variability through elevated rates of mutations. We investigated to what extent genetic variability, as estimated from band sharing coefficients for minisatellite markers, could be predicted by breeding distribution range, population size and intensity of sexual selection (as reflected by degree of polygyny and extra-pair paternity). Across a sample of 62 species of birds in the Western Palearctic, we found extensive interspecific variation in band sharing coefficients. High band sharing coefficients (implying low local genetic variability among individuals) were associated with restricted breeding distributions, a conclusion confirmed by analysis of statistically independent linear contrasts. Independently, species with large population sizes had small band sharing coefficients. Furthermore, bird species with a high richness of subspecies for their breeding distribution range had higher band sharing coefficients. Finally, bird species with high levels of polygyny and extra-pair paternity had small band sharing coefficients. These results suggest that breeding distribution range, population size and intensity of sexual selection are important predictors of levels of genetic variability in extant populations.     

Yearling male great tits, Parus major, suffer more strongly from cuckoldry than older males

In the majority of socially monogamous bird species, females have offspring sired by males other than their social mate as the result of extra-pair copulations. While it is widely recognised that there is considerable variation in the frequency of extra-pair paternity between species, between populations of a species and between individuals of a population, determinants of this variation are surprisingly difficult to establish. With respect to individual variation within a population, it is an important step to test for male and female correlates of cuckoldry to better understand the patterns as well as the adaptive significance of extra-pair mating behaviour. Here, we analysed patterns of extra-pair paternity in relation to male age, female age and their interaction in the great tit Parus major, a socially monogamous passerine with a moderate frequency of extra-pair paternity. Based on a large sample of 316 genotyped first broods from five successive years, we failed to demonstrate interaction effects of male and female age on both the proportion of extra-pair offspring and the likelihood that at least one extra-pair offspring is present within a brood. However, both the proportion of extra-pair offspring and the likelihood of paternity loss were higher for yearling as compared to older males, while this was not the case for yearling vs. older females. Furthermore, the proportion of extra-pair offspring within a brood decreased with increasing age of the attending male in within-individual analyses. We found a comparable effect also for attending females in within-individual analyses, but only when excluding two individuals with 100% extra-pair paternity. A female (extra-pair) mating preference for older males and/or a limited ability of yearling males to prevent cuckoldry in their broods could explain these age-related patterns of paternity loss. Effect sizes, however, were not particularly large and substantial residual variation within age categories suggests the importance of further yet unidentified determinants of variation in paternity loss in the study population.     

Immune defence,extra-pair paternity,and sexual selection in birds

Secondary sexual characters have been suggested to reliably reflect the ability of individuals to resist debilitating parasites, and females may gain direct or indirect fitness benefits from preferring the most extravagantly ornamented males. Extra-pair paternity provides an estimate of an important component of sexual selection in birds. Species with a high frequency of extra-pair paternity have a variance in realized reproductive success that is greater than the variance in apparent reproductive success, and extra-pair copulations and hence extra-pair paternity by females are often directly associated with the expression of male secondary sexual characters. If sexually dichromatic species have experienced a long period of antagonistic coevolution with their parasites, such species should have evolved larger immune defence organs than sexually monochromatic species. Bird species with sexual dichromatism had larger spleens for their body size than monochromatic species in a comparative analysis. Furthermore, species with a high frequency of extra-pair paternity were sexually dichromatic and had large spleens for their body size. These results are consistent with the hypothesis that females of dichromatic bird species seek extra-pair copulations to obtain indirect fitness benefits in terms of superior resistance of their offspring to virulent parasites.     

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.     

Male tail streamer length does not predict apparent or genetic reproductive success in North American barn swallows Hirundo rustica erythrogaster

In the socially monogamous barn swallow, previous studies of individuals in the European subspecies Hirundo r. rustica have shown that a male's tail streamer length is under strong sexual selection and is positively associated with several measures of reproductive success, including a low probability of being cuckolded by other males. The prominence of these results has led to subsequent experimental and correlational investigations of individuals in the phenotypically divergent subspecies H. r. erythrogaster in North America, where it has been shown that male tail streamer length is not as strongly associated with reproductive success as in European populations. We examined relationships between male tail streamer length and patterns of: (1) social mate selection and reproductive success, and (2) extra-pair paternity in 265 progeny of 53 social fathers within a New York barn swallow population. Although tail streamers in this population were sexually dimorphic, male tail streamer length did not predict patterns of mate selection, seasonal reproductive success, or extra-pair paternity. Moreover, in contrast to the strong positive relationships between paternity and male streamer length in European populations, summarized in this paper, we found no positive relationship between a male's paternity of young in the nest he is attending and his tail streamer length in our study population in New York. Our results further corroborate recent suggestions that the function of sexual signals varies geographically in this species, although we await additional experimental analyses on streamer lengths to understand the maintenance of sexual dimorphism in this trait.     

Promiscuity, paternity and personality in the great tit

Understanding causes of variation in promiscuity within populations remain a major challenge. While most studies have focused on quantifying fitness costs and benefits of promiscuous behaviour, an alternative possibility--that variation in promiscuity within populations is maintained because of linkage with other traits-has received little attention. Here, we examine whether promiscuity in male and female great tits (Parus major)--quantified as extra-pair paternity (EPP) within and between nests--is associated with variation in a well-documented personality trait: exploration behaviour in a novel environment. Exploration behaviour has been shown to correlate with activity levels, risk-taking and boldness, and these are behaviours that may plausibly influence EPP. Exploration behaviour correlated positively with paternity gained outside the social pair among males in our population, but there was also a negative correlation with paternity in the social nest. Hence, while variation in male personality predicted the relative importance of paternity gain within and outside the pair bond, total paternity gained was unrelated to exploration behaviour. We found evidence that males paired with bold females were more likely to sire extra-pair young. Our data thus demonstrate a link between personality and promiscuity, with no net effects on reproductive success, suggesting personality-dependent mating tactics, in contrast with traditional adaptive explanations for promiscuity.     

Extra-pair paternity and song characteristics in the willow warbler Phylloscopus trochilus

Complexity in bird song is often argued to be advantageous in processes of sexual selection, and numerous studies show that characteristics of song are associated with increased success in territory defence or mate attraction. Less evidence exists on the relationship between bird song characteristics and patterns of extra-pair paternity. We tested whether males that suffered from extra-pair paternity differed in song characteristics from males with no extra pair paternity in the willow warbler Phylloscopus trochilus . In the Scottish population that we studied, we found that 23.5% of young were not related to the social father, and that 47% of nests contained at least one extra pair young. Older males were less likely to suffer paternity loss. While song repertoire size was not related to loss of paternity, males with short songs suffered higher paternity loss than males with long songs. Although arrival date is a good correlate of social mate choice in this population, it was not related to extra-pair paternity. These results suggest that females use song length, or a trait correlated with this song characteristic, as a cue for choosing extra-pair partners in this species, or alternatively that variance in the success of mate guarding or female coercion is related to this song variable.     

社会性单配制鸟类婚外父权的发生和影响因素

一雌一雄单配制鸟类中,雌性个体与配偶外雄性发生交配的行为称为婚外交配,继而导致了婚外受精产生婚外子代的现象称为产生了婚外父权。婚外父权广泛存在于鸟类中,针对其发生和影响因素已经成为了鸟类行为生态学研究的热点。本文收集了近十年社会性单配制鸟类婚外父权方面的研究文献,从婚外父权的发生及其影响因素两个方面综述了单配制鸟类婚外父权的研究进展。婚外父权发生原因的探讨主要包括：1、从两性的角度探讨雌雄两性在婚外行为中不同的进化繁殖策略。雄性策略旨在增加自身的繁殖输出;有关雌性策略则提出了确保受精假说、食物供给假说、遗传利益假说等,但目前尚存争议;2、在遗传利益假说中较常见的又分为3个假说：“优秀基因”假说、“遗传相容性”假说和“遗传多样性”假说,该三种假说是针对雌性从遗传方面获得的利益而提出的,不断有报道指出雌性配偶选择会被潜在的雄性遗传特性所影响;3、非遗传利益——母系效应影响婚外父权的进化。一些研究指出遗传质量参数,如体重、身体大小、存活率和免疫应答等方面可能会存在母系效应。婚外父权发生的影响因素这里主要指环境因素,包括繁殖同步性、繁殖密度、栖息地环境、产卵及孵化时机等。由于物种不同,受到环境压力不同,导致婚外父权发生率千差万别。最后本文针对未来的研究方向做出了展望。尽管近十年的研究进一步解释了鸟类婚外父权现象,但是该领域仍然存在并且产生了许多新的未解决的问题,而相关实验操作和理论的完善是深入探讨这些问题的关键。     

Causes of extra-pair paternity and its inter-specific variation in socially monogamous birds

Based on molecular technology, researchers find that extra-pair paternity (EPP) prevails among socially monogamous bird species. This phenomenon challenges traditional views of sexual selection and mating system, and has become one of the hot-spots in the avian behavior ecology. This review focuses on the evolutionary causes leading to EPP and the potential factors affecting it. Early studies of EPP in birds used a wide variety of tools, including plumage color polymorphism, polymorphic enzymes, and sex-differences in estimation of the heritability of morphological traits. Although each of these methods can be used to estimate the likelihood that EPP are present or absent in a population, none of them provide enough accurate estimation to allow meaningful cross-species comparison. Currently, studies of EPP mainly use “DNA-methods”, namely multi-locus mini-satellite fingerprints, single-locus mini-satellite fingerprints, and micro-satellite genotyping, because they can provide accurate outcome of paternity identification and their results are reproducible. From the point of the female, the driving forces for EPP are that females may gain direct benefits or indirect genetic benefits from EPP. Hypotheses explaining the benefits include fertility assurance, good genes, genetic compatibility, and genetic diversity. Despite large numbers of theoretically plausible explanations for EPP, there have been few direct empirical tests that can provide unambiguous support for only one type of explanation. The most straightforward test of the genetic benefit hypothesis of extra-pair copulation is a comparison of the performance of maternal half-siblings from multiply sired broods. Some highly-cited landmark studies spectacularly support the genetic benefit hypothesis, while other studies failed to reveal any systematic differences in maternal half-sibling performance, even in the same species or in taxonomically closely-related species. Briefly, the point that purpose of extra-pair copulation is to gain genetic benefits is still facing great challenges. Therefore, some researchers suggest that more attention should be focused on the interactions between parties involved in the extra-pair paternity phenomenon. Others, however, believe that such interactions can also be of a cooperative nature, and involve exchange of direct benefits. In brief, an approach focusing on interactions (involving either conflict or cooperation) seems to be the most promising direction to improve our understanding of the phenomenon of extra-pair paternity. As for the factors leading to intra- and inter-specific variation in the level of EPP, current researches mainly focus on the breeding density, breeding synchrony, the complexity of the habitat, paternal care, adult mortality, food availability, and genetic diversity. Explaining intra- and inter-specific variation in the extent of EPP has been difficult, but an appreciation of the problems of small sample sizes, and an ever-increasing comparative database have led to several recent advances. It now seems probable that differences between species in the rate of EPP are due to a combination of differences in life history, pattern of parental care, and local opportunities for promiscuity. In a word, although there have been a lot of theoretical and empirical researches about EPP of birds, no consensus on the basic questions has been received in this area. Thus, more scientific statistical analysis method and more empirical experiments are still badly needed to improve the theoretical system.     

Paternity analysis reveals opposing selection pressures on crown coloration in the blue tit (Parus caeruleus)

In socially monogamous species, extra-pair paternity can increase the variance in reproductive success and thereby the potential for sexual selection on male ornaments. We studied whether male secondary sexual ornaments are selected through within- and/or extra-pair reproductive success in the blue tit (Parus caeruleus). Male blue tits display a bright blue crown plumage, which reflects substantially in the ultraviolet (UV) and previously has been indicated to be an important sexual signal. We show that males with a more UV-shifted crown hue were less cuckolded, which probably resulted from female preference for more ornamented mates. By contrast, however, older males and males with a less UV-shifted hue sired more extra-pair young. This probably did not reflect direct female preference, since cuckolders were not less UV-ornamented than the males they cuckolded. Alternatively, a trade-off between UV ornamentation and other traits that enhance extra-pair success could explain this pattern. Our results might reflect two alternative male mating tactics, where more UV-ornamented males maximize within-pair success and less UV-ornamented males maximize extra-pair success. Since crown colour was selected in opposite directions by within-pair and extra-pair paternity, directional selection through extra-pair matings seemed weak, at least in this population and breeding season. Reduced intensity of sexual selection due to alternative mating tactics constitutes a potential mechanism maintaining additive genetic variance of male ornaments.     

Extra-pair paternity in the Skylark Alauda arvensis

We present the first quantitative data on the genetic breeding system of a lark (Alaudidae), the Skylark Alauda arvensis . Using a set of eight microsatellite loci isolated in a variety of passerine species, we genotyped 171 offspring from 52 broods of Skylark and detected 35 extra-pair offspring (20%), in 14 different broods (27%). All offspring matched their putative mother, so there was no evidence of intraspecific brood parasitism. Previous non-genetic studies had suggested that the species was predominantly socially monogamous, with only rare occurrences of social polygyny and polyandry, although some behaviours, such as mate guarding, did suggest the possibility of extra-pair copulations. The relatively high level of extra-pair paternity in this species is likely to affect the variation in male reproductive success because extra-pair paternity was non-randomly distributed amongst males, with those with shorter wings more likely to be cuckolded.     

Sperm competition and sexually size dimorphic brains in birds

Natural selection may favour sexually similar brain size owing to similar selection pressures in males and females, while sexual selection may lead to sexually dimorphic brains. For example, sperm competition involves clear-cut sex differences in behaviour, as males display, mate guard and copulate with females, while females choose among males, and solicit or reject copulations. These behaviours may require fundamentally different neural government in the two sexes leading to sex-dependent brain evolution. Using two phylogenetic approaches in a comparative study, we tested for roles of both natural and sexual-selection pressures on brain size evolution of birds. In accordance with the natural-selection theory, relative brain size of males coevolved with that of females, which may be the result of adaptation to similar environmental constraints such as feeding innovation. However, the mode of brain size evolution differed between the sexes, and factors associated with sperm competition as reflected by extra-pair paternity may give rise to sexually size dimorphic brains. Specifically, species in which females have larger brains than males were found to have a higher degree of extra-pair paternity independently of potentially confounding factors, whereas species in which males have relatively larger brains than females appeared to have lower rates of extra-pair paternity. Hence, the evolution of sperm competition may select for complex behaviours together with the associated neural substrates in the sex that has a higher potential to control extra-pair copulations at the observed levels. Brain function may thus be affected differently in males and females by sexual selection.     

Sexual selection favours male parental care, when females can choose

Explaining the evolution of male care has proved difficult. Recent theory predicts that female promiscuity and sexual selection on males inherently disfavour male care. In sharp contrast to these expectations, male-only care is often found in species with high extra-pair paternity and striking variation in mating success, where current theory predicts female-only care. Using a model that examines the coevolution of male care, female care and female choice; I show that inter-sexual selection can drive the evolution of male care when females are able to bias mating or paternity towards parental males. Surprisingly, female choice for parental males allows male care to evolve despite low relatedness between the male and the offspring in his care. These results imply that predicting how sexual selection affects parental care evolution will require further understanding of why females, in many species, either do not prefer or cannot favour males that provide care.     

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.     

Extra pair paternity in birds: a review of interspecific variation and adaptive function

The application of molecular genetic techniques has revolutionized our view of avian mating systems. Contrary to prior expectations, birds are only very rarely sexually monogamous, with 'extra-pair offspring' found in approximately 90% of species. Even among socially monogamous species, over 11% of offspring are, on average, the result of extra-pair paternity (EPP). Based on over 150 molecular genetic studies of EPP in birds, we review two topical areas: (i) ecological explanations for interspecific variation in the rate of EPP; and (ii) evidence bearing on the adaptive function of EPP. We highlight the remaining challenges of understanding the relative roles of genes and ecology in determining variation between taxa in the rate of extra paternity, and testing for differences between extra-pair offspring and those sired within-pair.     

Do male paternity guards ensure female fidelity in a duetting fairy-wren?

Most socially monogamous bird species engage in extra-pair mating,and consequently males may adopt various behavioral strategiesto guard paternity. However, the relationship between mate guardingand extra-pair paternity is unclear: low levels of extra-pairpaternity can be associated either with no mate guarding orwith intense mate guarding. We investigate paternity guardsin the purple-crowned fairy-wren (Malurus coronatus), a duettingspecies where extra-pair paternity is rare. This species isunusual in a genus known for extremely high levels of extra-pairmating. We examine the behavioral interactions between the sexesunderlying these low rates of extra-pair paternity and showthat male purple-crowned fairy-wrens do not use frequent copulationor courtship feeding to assure paternity or guard females acousticallyby duetting. Males maintain close proximity to females bothwhen they are fertile and when they are not breeding and donot invest in courtship displays to extra-pair females. Consistentwith predictions of theoretical models, low extra-pair paternityin this species may be related to female fidelity rather thanmale paternity assurance strategies, but short-term removalof males would be necessary to test this experimentally.     

Low incidence of extra-pair paternity in the colonially nesting common swift Apus apus

The frequency of extra-pair paternity in a wild colony of swifts Apus apus was determined by multilocus DNA fingerprinting in two successive breeding seasons. The data were used to examine the expectation that extra-pair paternity is frequent in colonial-nesting species, either for proximal reasons such as the increased opportunity for extra-pair matings, or because extra-pair matings are important in the evolution and maintenance of coloniality. Forty-two broods containing 88 chicks were analysed. The genetic analysis revealed four cases of extra-pair paternity (4.5% of chicks) from four (9.5%) nests. Rapid mate-switching was considered unlikely to be the cause of extra-pair paternity since three of the cases were in the nests of previously established breeding pairs. Extra-pair copulations were not observed, but were assumed to be the cause of extra-pair paternity. The data show that high levels of extra-pair paternity are not an inevitable feature of high-density nesting.