High fidelity on islands: a comparative study of extrapair paternity in passerine birds

It is commonly assumed that the intensity of sexual selectionis lower in island populations. Extrapair paternity (EPP) iswidespread within passerine birds and is indicative of sexualselection. A conservative analysis of the levels of EPP inisland and equivalent mainland populations of passerines revealsthat insular populations are indeed characterized by low levelsof EPP. This supports the idea that the intensity of sexualselection is lower on islands. This relationship has previouslybeen predicted, based on the assumption of low levels of geneticvariation for fitness in such populations. The evidence fromthis analysis suggests that this is just one of several nonmutuallyexclusive hypotheses that may explain the high fidelity of island-living females.     

杂色山雀的婚外父权与配偶间的遗传相容性无关

婚配制度作为一种进化稳定对策是动物对某一环境包括种群内部环境适应的结果。据统计,约92%的鸟类为社会性单配制,但单配制鸟类中很多都存在婚外父权现象,表现出社会性婚配制度与遗传性婚配制度的不一致性。杂色山雀(Parus varius)是一种社会性单配制的小型森林洞巢鸟,其是否存在婚外父权现象至今尚未见报道。通过对杂色山雀进行亲权鉴定以确定其有无婚外父权现象及婚外父权的比例,结果显示:45.45%(20/44)的巢存在婚外父权,14.39%(38/264)的后代为婚外子代,说明杂色山雀具有较高的婚外父权水平。进一步探究其婚外父权的发生原因,结果如下:(1)有、无婚外父权巢的社会性亲本之间的遗传相似性无显著差异(P=0.504);(2)有婚外父权巢中婚内子代和无婚外父权巢中子代的杂合度(P=0.118)以及有婚外父权巢中婚外子代与婚内子代的杂合度(P=0.206)均无显著差异。(3)有婚外父权巢中的婚内子代与婚外子代间8项体征指标比较,差异均不显著(Ps0.05)。综上,社会性单配制杂色山雀婚外父权的发生与配偶间的遗传相容性无关,还有待从其他角度进行探究。     

Extra-pair paternity does not result in differential sexual selection in the mutually ornamented black swan (Cygnus atratus)

We studied patterns of parentage in 85 broods (332 cygnets) of black swans during three breeding seasons, using a set of eight polymorphic microsatellite markers. We detected both intraspecific brood parasitism (IBP; < 5% of cygnets per year) and extra-pair paternity (EPP). In these years, 10-17% (mean = 15.1%) of cygnets resulted from EPP, and 27-40% (mean 37.6%) of broods contained at least one extra-pair cygnet. Compared with levels of EPP in closely related species with similar life histories, these values are unexpectedly high. EPP in black swans appears unrelated to ecological factors (breeding density and synchrony) or genetic factors (genetic similarity between pair members or genetic quality of the offspring). We found no evidence that a mutual sexual feather ornament known to play a role in social mate choice in black swans (curled wing feathers) is involved in extra-pair mate choice. EPP does not lead to greater variance in reproductive success in males, relative to females in this species. We therefore suggest that EPP does not result in differential sexual selection on males and females, explaining why they are ornamented to the same degree.     

The Magnetic Field as a Reference System for Genetically Encoded Migratory Direction in Pied Flycatchers (Ficedula hypoleuca Pallas)

To see whether the migratory orientation of pied flycatchers (Ficedula hypoleuca Pallas) is genetically encoded with respect to the earth magnetic field a group of young birds was hand-raised. They were thus prevented from ever experiencing the sky. The birds were tested in autumn 1980 and 1981 in the local geomagnetic field (Fig. 1) and in three artificial fields (Fig. 2a-c). The results show that their magnetic compass matures independent of any experience with the sky and contains sufficient information for the birds to orient toward their migratory direction.     

Adaptive coloration in pied flycatchers (Ficedula hypoleuca)—The devil is in the detail

Understanding the origin and persistence of phenotypic variation within and among populations is a major goal in evolutionary biology. However, the eagerness to find unadulterated explanatory models in combination with difficulties in publishing replicated studies may lead to severe underestimations of the complexity of selection patterns acting in nature. One striking example is variation in plumage coloration in birds, where the default adaptive explanation often is that brightly colored individuals signal superior quality across environmental conditions and therefore always should be favored by directional mate choice. Here, we review studies on the proximate determination and adaptive function of coloration traits in male pied flycatchers (Ficedula hypoleuca). From numerous studies, we can conclude that the dark male color phenotype is adapted to a typical northern climate and functions as a dominance signal in male–male competition over nesting sites, and that the browner phenotypes are favored by relaxed intraspecific competition with more dominant male collared flycatchers (Ficedula albicollis) in areas where the two species co‐occur. However, the role of avoidance of hybridization in driving character displacement in plumage between these two species may not be as important as initially thought. The direction of female choice on male coloration in pied flycatchers is not simply as opposite in direction in sympatry and allopatry as traditionally expected, but varies also in relation to additional contexts such as climate variation. While some of the heterogeneity in the observed relationships between coloration and fitness probably indicate type 1 errors, we strongly argue that environmental heterogeneity and context‐dependent selection play important roles in explaining plumage color variation in this species, which probably also is the case in many other species studied in less detail.     

Parentage assignment and extra-group paternity in a cooperative breeder: the Seychelles warbler (Acrocephalus sechellensis)

We describe the development and initial application of a semiautomated parentage testing system in the Seychelles warbler (Acrocephalus sechellensis). This system used fluorescently labelled primers for 14 polymorphic microsatellite loci in two multiplex loading groups to genotype efficiently over 96% of the warbler population on Cousin island. When used in conjunction with the program CERVUS, this system provided sufficient power to assign maternity and paternity within the Seychelles warbler, despite the complications associated with its cooperative breeding system and a relatively low level of genetic variation. Parentage analyses showed that subordinate 'helper' females as well as the dominant 'primary' females laid eggs in communal nests, indicating that the Seychelles warbler has an intermediate level of female reproductive skew, in between the alternative extremes of helper-at-the-nest and joint nesting systems. Forty-four per cent of helpers bred successfully, accounting for 15% of all offspring. Forty per cent of young resulted from extra-group paternity.     

High multiple paternity and low last-male sperm precedence in a hermaphroditic planarian flatworm: consequences for reciprocity patterns

It is difficult to predict a priori how mating success translates into fertilization success in simultaneous hermaphrodites with internal fertilization. Whereas insemination decisions will be determined by male interests, fertilization will depend on female interests, possibly leading to discrepancies between insemination and fertilization patterns. The planarian flatworm Schmidtea polychroa, a simultaneous hermaphrodite in which mating partners trade sperm was studied. Sperm can be stored for months yet individuals mate frequently. Using microsatellites, maternity and paternity data were obtained from 748 offspring produced in six groups of 10 individuals during four weeks. Adults produced young from four mates on average. Reciprocal fertilization between two mates was found in only 41 out of 110 registered mate combinations, which is clearly less than what is predicted from insemination patterns. Multiple paternity was high: > 80% of all cocoons had two to five fathers for only three to five offspring per cocoon. Because animals were collected from a natural population, 28% of all hatchlings were sired by unknown sperm donors in the field, despite a 10-day period of acclimatization and within-group mating. This percentage decreased only moderately throughout the experiment, showing that sperm can be stored and used for at least a month, despite frequent mating and sperm digestion. The immediate paternity a sperm donor could expect to obtain was only about 25%. Male reproductive success increased linearly with the number of female partners, providing support for Bateman's principle in hermaphrodites. Our results suggest that hermaphrodites do not trade fertilizations when trading sperm during insemination, lending support to the view that such conditional sperm exchange is driven by exchange of resources.     

Carotenoid‐based bill coloration functions as a social,not sexual,signal in songbirds (Aves: Passeriformes)

Many animals use coloration to communicate with other individuals. Although the signalling role of avian plumage colour is relatively well studied, there has been much less research on coloration in avian bare parts. However, bare parts could be highly informative signals as they can show rapid changes in coloration. We measured bill colour (a ubiquitous bare part) in over 1600 passerine species and tested whether interspecific variation in carotenoid‐based coloration is consistent with signalling to potential mates or signalling to potential rivals in a competitive context. Our results suggest that carotenoid bill coloration primarily evolved as a signal of dominance, as this type of coloration is more common in species that live in social groups in the nonbreeding season, and species that nest in colonies; two socio‐ecological conditions that promote frequent agonistic interactions with numerous and/or unfamiliar individuals. Additionally, our study suggests that carotenoid bill coloration is independent of the intensity of past sexual selection, as it is not related to either sexual dichromatism or sexual size dimorphism. These results pose a significant challenge to the conventional view that carotenoid‐based avian coloration has evolved as a developmentally costly, condition‐dependent sexual signal. We also suggest that bare part ornamentation may often signal different information than plumage ornaments.     

A tale of two siblings: multiple paternity in big brown bats (Eptesicus fuscus) demonstrated using microsatellite markers

In many bat species, the opportunity for sperm competition or other mechanisms of post-copulatory paternity biasing is thought to be great, due to the long delay between copulation and fertilization, demonstrated sperm storage capabilities, and observed promiscuity. We present the results of the first study to assess whether litters of big brown bats (Eptesicus fuscus) containing dizygotic twins share the same father. We sampled 26 mother-offspring triads from three colonies in Indiana and Illinois, as well as 299 additional adults (237 females and 62 males) from these colonies and six other maternity colonies in the same area in 1997-1998. All individuals were genotyped at nine highly variable autosomal microsatellite loci and one X-linked locus. We assessed multiple paternity using autosomal and X-linked locus exclusions, and using maximum-likelihood methods. All methods confirmed multiple paternity within litters, and the maximum-likelihood analyses indicated that almost half of the sampled litters were composed of maternal half-siblings rather than full-siblings. Our results highlight the potential importance of post-copulatory mechanisms of paternity determination in the mating system of big brown bats, and have important implications for gene flow and population structuring in this species.     

Individual color variation and male quality in pied flycatchers (Ficedula hypoleuca): a role of ultraviolet reflectance

Bright coloration of males in many animal species has inspiredresearchers for more than a century. In this field study, weinvestigated whether color variation between individuals isrelated to individual quality in pied flycatcher (Ficedulahypoleuca) males in terms of arrival time at the breeding sites.In addition to traditional visual color scoring, plumage color was measured using spectroradiometric measurements between 320and 700 nm. This range includes the near-ultraviolet wavebandfrom 320 to 400 nm. Males that arrived earlier at breedingsites had higher proportional UV reflectance in the crown andmantle. The proportional UV reflectance in the crown and mantlewas not related to traditionally scored general brownness inmales. However, adult males had a higher proportion of ultravioletin the plumage than yearling males or females. These resultssuggest that in pied flycatcher males, the UV reflectance ofplumage may be positively correlated with individual quality.     

Amplified fragment length polymorphism (AFLP) markers reveal extra-pair parentage in a bird species: the bluethroat (Luscinia svecica)

We tested the use of amplified fragment length polymorphism (AFLP) to assess the frequency of extra-pair parentage in a bluethroat (Luscinia svecica namnetum) population. Thirty-six families totalling 162 nestlings were analysed. Using a combination of three primer pairs, we reached an exclusion probability of 93% for the population. This probability can reach 99% considering families independently. We revealed that extra-pair fertilizations are very common: 63.8% of all broods contain at least one extra-pair young, totalling 41.9% of all young analysed. However, with the technique and the three primer pairs used it was not possible to attribute the parentage exclusions to extra-pair paternity, maternity or both. As brood parasitism has never been reported in this species, it seems likely that the exclusions are due to extra-pair males. This study shows that dominant AFLP markers can be useful for studying the mating system of taxa for which no microsatellite primers are available. This technique allows the approximate estimation of parentage exclusions despite the fact that it is not possible to know which parent has to be excluded.     

Polyandrous females provide sons with more competitive sperm: Support for the sexy‐sperm hypothesis in the rattlebox moth (Utetheisa ornatrix)

Given the costs of multiple mating, why has female polyandry evolved? Utetheisa ornatrix moths are well suited for studying multiple mating in females because females are highly polyandrous over their life span, with each male mate transferring a substantial spermatophore with both genetic and nongenetic material. The accumulation of resources might explain the prevalence of polyandry in this species, but another, not mutually exclusive, possibility is that females mate multiply to increase the probability that their sons will inherit more‐competitive sperm. This latter “sexy‐sperm” hypothesis posits that female multiple mating and male sperm competitiveness coevolve via a Fisherian runaway process. We tested the sexy‐sperm hypothesis by using competitive double matings to compare the sperm competition success of sons of polyandrous versus monandrous females. In accordance with sexy‐sperm theory, we found that in 511 offspring across 17 families, the male whose polyandrous mother mated once with each of three different males sired significantly more of all total offspring (81%) than did the male whose monandrous mother was mated thrice to a single male. Interestingly, sons of polyandrous mothers had a significantly biased sex ratio of their brood toward sons, also in support of the hypothesis.     

Evolutionary dynamics of a sexual ornament in the house sparrow (Passer domesticus): the role of indirect selection within and between sexes

The relative contribution of sexual and natural selection to evolution of sexual ornaments has rarely been quantified under natural conditions. In this study we used a long-term dataset of house sparrows in which parents and offspring were matched genetically to estimate the within- and across-sex genetic basis for variation and covariation among morphological traits. By applying two-sex multivariate "animal models" to estimate genetic parameters, we estimated evolutionary changes in a male sexual ornament, badge size, from the contribution of direct and indirect selection on correlated traits within males and females, after accounting for overlapping generations and age-structure. Indirect natural selection on genetically correlated traits in males and females was the major force causing evolutionary change in the male ornament. Thus, natural selection on female morphology may cause indirect evolutionary changes in male ornaments. We observed however no directional phenotypic change in the ornament size of one-year-old males during the study period. On the other hand, changes were recorded in other morphological characters of both sexes. Our analyses of evolutionary dynamics in sexual characters require application of appropriate two-sex models to account for how selection on correlated traits in both sexes affects the evolutionary outcome of sexual selection.     

It is not only group size: Soldiers also modulate the tolerance to insecticide in termites (Blattodea: Isoptera)

The survivorship of social insects is known to increase with group size, even in situations of stress, such as starvation and exposure to insecticides. However, in termites, studies have been undertaken only with workers, disregarding the possible effect of soldiers. The role of soldiers in the termite colonies goes beyond defense, mainly in Nasutitermes species. It is already known that soldiers initiate the foraging as well as improve the decision-making of food resources. Here, we evaluated the effect of group size and the presence of soldiers on exposure to sublethal doses of the insecticide imidacloprid in Nasutitermes corniger (Termitidae: Nasutitermitinae). To do so, toxicity bioassays were undertaken initially to determine the dose of the insecticide required to kill 50% of the N. corniger population (LD₅₀) to be used in the main experiments. Survival bioassays were then carried out with termite groups, with and without soldiers, in different sizes (6, 10, 14, 22, 26, 30 and 60), exposed and nonexposed to insecticide. In general, the mean time to death of termites increases linearly with group size. However, the mean time to death in groups with soldiers was significantly longer only in groups exposed to the insecticide. Our results indicate that soldiers can help to increase the tolerance of nasute termite groups to insecticides, in addition to the group size, as already shown in the previous study. The size of the group and social context could, therefore, modulate behavioral and/or physiological responses that enhance the ability to survive under stressful situations.     

Horn length is not correlated with calling efforts in the horn‐headed cricket Loxoblemmus doenitzi (Orthoptera: Gryllidae)

Field cricket species are ideal model organisms for the study of sexual selection because cricket calling songs, used to attract mating partners, are pronouncedly sexually dimorphic. However, few studies have focused on other sexually dimorphic traits of field crickets. The horn‐headed cricket, Loxoblemmus doenitzi, exhibits exaggerated sexual dimorphism in head shape: males have flat heads with triangular horns, while females lack horns. This study examines the relationship between horn length, male calling efforts and diet quality. Horn length was not found to be significantly correlated with calling efforts. When diet was manipulated for late‐stage nymphs, calling efforts in the group with poor‐quality diet treatment was significantly lower than that of crickets in the group with high‐quality diet treatment. However, horn length was not affected by diet quality. The implication of these results in the context of the evolution of multiple signals and sexual dimorphism is discussed.     

Variability of a consistent trait: The size of the white wing patch in European Stonechats (Saxicola rubicola rubicola)

Numerous bird species exhibit striking white patches on their plumages that have been reported as signals of individual quality in the context of sexual selection. Whereas differences in white plumage traits between individuals have been well studied, phenotypic variation and the factors affecting their consistency within the individual have received less attention. Here, we studied the consistency in the size of the white wing patch and its components of variation in a Spanish population of European Stonechats Saxicola rubicola rubicola monitored over 6 years (2007–2012). The wing patch was larger in males than in females for all age-classes, and first-year individuals exhibited smaller wing patches than older birds, particularly males. This trait also varied within the plumage-year at both population and individual level, with slight changes from the moulting period to the breeding season and a sharp decrease afterwards. In addition, patch size varied both between and within individuals across years. The daily mean temperature experienced in the month immediately prior to the moult (i.e. May) had a positive effect on male wing patch size. In females, however, no variable was found to explain wing patch variation satisfactorily. Despite being a dynamic trait throughout the moults, baseline and adjusted repeatabilities indicated a moderately high consistency for white wing patch size in both sexes. Therefore, the white wing patch in Stonechats is a dynamic trait that changes throughout individuals' lives, but is also a consistent trait at the individual level. These results indicate that this depigmented patch could convey information about both the long-term quality and the current state of individuals, harmonizing the existence of intra-individual changes in the size of sexual traits with its potential role as a genetic quality indicator.     

Heightened condition dependence is not a general feature of male eyespan in stalk-eyed flies (Diptera: Diopsidae)

Stalk-eyed flies are exemplars of sexual selection leading to the evolution of exaggerated male ornaments (eyespan). In Sphyracephala beccarri, there is no evidence for female mate choice for exaggerated male eyespan and only minor sex differences in eyespan. We used S. beccarri to test whether heightened condition dependence only evolves when male eyespan becomes sexually exaggerated. Male eyespan showed heightened condition dependence under food stress compared with a control trait (wing length). However, female eyespan displayed a similar pattern and there was no sex difference in the degree of increased eyespan sensitivity. The finding that eyespan is a sensitive indicator of food stress, even in an unexaggerated state, suggests that this may have acted as a pre-adaptation to the role of eyespan in sexual signalling in other Diopsid species. These results are consistent with handicap theory and Fisher's view of how sexual selection is initiated.     

雄性川金丝猴睾酮分泌与其社群环境变化的关系

采用化学免疫发光法,跟踪测定了上海野生动物园一个半放养川金丝猴群中3只先后参与交配繁殖活动的成年雄猴一年中尿中睾酮含量,并结合行为学记录对其性激素和性行为间的相互影响关系进行了初步研究。发现成年雄猴睾酮分泌峰的出现更多的是由其社会因素的变化引起的,而且,睾酮分泌峰出现的时期并不与雌性受孕高峰期相重叠,提示雄性繁殖具非季节性特点。睾酮分泌峰的出现,一方面说明此时社群的社会关系处于不稳定状态,另一方面发现睾酮水平的提高对雄猴攻击能力的增强起促进作用。由此,本研究认为川金丝猴雌雄性个体在繁殖中采用了不尽相同的繁殖策略[动物学报49(3)：325—33l,2003]。     

Social selection and the evolution of a female weapon in queens of the ant Messor pergandei (Hymenoptera: Formicidae)

The evolution of weaponry occurs less frequently in females than in males and is most often important for protecting ecological resources or offspring rather than winning mates. The purpose of female weapons is often confounded by the presence of similar weapons in males, so cases where only females need weapons provide important tests of our understanding of how and why weapons evolve. In some populations of the ant, Messor pergandei (Mayr), newly mated queens initiate new nests in social groups that subsequently break down when queens engage in battles for control. The incipient social environment differs geographically, so that lethal fighting occurs in some populations but not others. Consistent with the hypothesis that queens in populations where lethal fighting occurs should show selection for weaponry (broad heads and strong mandibles), we found that heads of queens from sites where lethal fighting occurs were broader than those at sites with non‐fighting queens and a site with solitary queens. Evolution of weaponry is specific to queens, because regression results from workers often did not follow this pattern. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1011–1020.