SEXUAL SELECTION IN THE BARN SWALLOW (HIRUNDO RUSTICA). IV. PATTERNS OF FLUCTUATING ASYMMETRY AND SELECTION AGAINST ASYMMETRY

The patterns of variation in fluctuating asymmetry were studied in four morphological characters of the barn swallow Hirundo rustica. The level of absolute and relative asymmetry was larger in the secondary sexual character “outer tail length” than in three nonsexual morphological traits (wing, central tail, and tarsus length). The extent of individual asymmetry in outer tail length was negatively correlated with tail-ornament size, whereas the relationship between asymmetry of all other morphological characters and their size was flat or U-shaped. Asymmetry in outer tail length was unrelated to asymmetry in other morphological characters, whereas asymmetries in the length of wing, central tail, and tarsus were positively correlated. Male bam swallows exhibited larger asymmetry in outer tail length than females. Asymmetry of most morphological traits exhibited intermediate repeatabilities between years, with the exception of male and female outer tail length, which were highly repeatable. Tail asymmetry of offspring weakly, though significantly, resembled that of their parents. Asymmetry in wing and outer tail length was also significantly related to several fitness components. Male barn swallows that acquired a mate were less asymmetric in wing and outer tail length than unmated males. Females with more asymmetrical tails laid eggs significantly later. Annual reproductive success was unrelated to fluctuating asymmetry. Male barn swallows that survived were less asymmetric in wing and outer tail length than nonsurvivors, whereas female survivors were less asymmetric in outer tail length than nonsurvivors. These results suggest that levels of fluctuating asymmetry in barn swallows are associated with differences in fitness.     

Multiple elements of the black-billed magpie's tail correlate with variable honest information on quality in different age/sex classes

Animals may be able to assess the quality of other individuals by using information contained in single or multiple traits. We investigated the honesty of the information potentially conveyed through different elements of the tail of the black-billed magpie, Pica pica, which develops in a similar way in both sexes while still in the nest. Variability and age and sex dimorphism were all higher for tail features than for other morphological characters. Tail length of first-year males and females was negatively correlated with the number of feathers with fault bars, which may reflect stressful environmental conditions during feather growth in the nest. Adult males may indicate current body condition through overall tail expression, while the current condition of adult females may be signalled by indicators of stressful environmental conditions during feather growth in the previous year, although both these correlations were weak. Tail length correlated negatively with testes volume of first years, and tails tended to be less damaged in both first-year and adult males with large testes. First-year males that had reached sexual maturity had shorter tails than those that had not. There was no association between parasitism and tail expression, but tail length was positively related to spleen size in first-year males. This study provides evidence that multiple information about individual quality may be conveyed by different features of the tail in ways that vary in different age and sex classes.     

Sexual size dimorphism and assortative mating for morphological traits in Passer domesticus

In this study, assortative mating for different morphological traits was studied in a captive population of house sparrows (Passer domesticus). Males were larger than females. Assortative mating was found for tail length, wing length and general body size. Males with larger badge size mated with females with longer tails. The strongest assortative mating occurred for tail length (r=0.77), and this assortative mating remained significant after controlling for wing length, mass and tarsus length, suggesting that it was not an artefact of assortative mating for body size. The possibility of sexual selection for tail length in the house sparrow is discussed.     

Female Mate Choice, Calling Song and Genetic Variance in the Cricket, Gryllodes sigillatus

Female preferences for song patterns of males of Gryllodes sigillatus and genetic variance of morphological traits correlated with them were analyzed. Females preferred short pulses associated with large males. The males’ thorax width, wing length and femur III length showed stronger relationship with the song pulse duration, whereas the relationship between pulse duration and wing width was not significant. Interestingly, this last trait was the only one that showed significant levels of genetic variance. Perhaps these results could be explained by the evolutionary response to sexual selection. Sexual selection could deplete the genetic variance in the male traits related to male‐mating success.     

Birds' tails as signaling devices: markings, shape, length, and feather quality

While exaggerated length and ornamental shapes are confirmed sexually selected tail traits in birds, the signal function of tail markings has received less study. Signal roles for tail markings as amplifiers of length, shape, and feather quality are discussed, and the role of tail markings as feather-quality handicaps is proposed: absence of melanin increases damage and abrasion. Predicted correlations of tail markings with other tail traits are derived for these signal roles. A comparative study of the relationships between these tail traits in an entire avifauna, the western Palearctic, tested the predictions. Tail displays were present in nearly 80% of species, associated with greater long-tailedness, but not all displayed tails had markings or ornamental shape. The incidence of marks across tail shapes and the combinations of marks indicate that tail markings act as handicaps or amplifying handicaps of tail feather quality. Tail elongation and ornamental shapes could act as additional handicaps of feather quality-that is, they could be multipurpose signals. Incorporation of revealing indicators such as feather damage and associated handicap and/or amplifying traits may allow a reduction in the cost of signaling while maintaining signal reliability and, hence, influence sexual selection in complex signaling systems.     

青海沙蜥的两性异型和雌性繁殖

作者研究了青海沙蜥(Phrynocephalus vlangalii)形态特征的两性异形和雌体繁殖特征。蜥蜴于2005年5月初捕自西宁以西约150km的倒淌河,被检形态特征包括体色、体长、腹长、尾长、头长和头宽,新排卵雌体维持在实验室梯度热环境中直至产仔。成体两性异形显著,而性未成熟个体缺乏两性异形。最大的成年雄体和雌体分别为70.2mmSVL(snout-vent length)和82.8mmSVL。雄性成体具有相对较大的头长、头宽和尾长,雌性成体SVL大于雄体且具有相对较大的腹长。对4个形态特征进行主成分分析(特征值≥0.5)区分出2个主成分,共解释83.9%的两性相关形态特征的变异。去除SVL差异的影响后,尾长、头长和头宽在第一主成分有较高的正负载系数(解释57.8%的变异),腹长在第二主成分有较高的负负载系数(解释26.1%的变异)。实验室梯度热环境下的雌体于6月下旬至7月中旬产单窝、2-6个后代。窝仔数和窝仔重与母体SVL呈正相关,幼仔重与母体SVL无关。未在青海沙蜥中检测到后代数量与大小之间的权衡。     

Sexual selection in the barn swallow (Hirundo rustica). V. Geographic variation in ornament size

Models of sexual selection in a cline predict the patterns of clinal variation in female mate preference and male secondary sexual characters. These predictions were tested for the nominate subspecies of the barn swallow Hirundo rustica which demonstrates clinal variation in morphology, with several characters in both sexes showing increasing size at higher latitudes. Sexual size dimorphism in the length of the tail ornament and the short, central tail feathers increase with increasing latitude while size dimorphism in other morphological characters is independent of latitude. The main reason for the two divergent patterns of sexual size dimorphism appears to be the higher foraging cost of having a long tail ornamental at low latitudes. The control of development decreases with increasing latitude as demonstrated by an increasing latitudinal cline in fluctuating asymmetry of tail length. Phenotypic variance in tail length increases with latitude in males, but not in females, as shown by the coefficients of variation. Clinal variation in morphology is not due to natural selection associated with a latitudinal increase in the distance between breeding and wintering areas. The geographic patterns of morphological variation suggest that the tail character has diverged geographically as a result of a sexual process of reliable signalling.     

Variations in morphological and life-history traits under extreme temperatures in <Emphasis Type="Italic">Drosophila ananassae</Emphasis>

Using half-sib analysis, we analysed the consequences of extreme rearing temperatures on genetic and phenotypic variations in the morphological and life-history traits of Drosophila ananassae. Paternal half-sib covariance contains a relatively small proportion of the epistatic variance and lacks the dominance variance and variance due to maternal effect, which provides more reliable estimates of additive genetic variance. Experiments were performed on a mass culture population of D. ananassae collected from Kanniyakumari (India). Two extremely stressful temperatures (18°C and 32°C) and one standard temperature (25°C) were used to examine the effect of stressful and non-stressful environments on the morphological and life-history traits in males and females. Mean values of various morphological traits differed significantly among different temperature regimens in both males and females. Rearing at 18°C and 32°C resulted in decreased thorax length, wing-to-thorax (w/t) ratio, sternopleural bristle number, ovariole number, sex comb-tooth number and testis length. Phenotypic variances increased under stressful temperatures in comparison with non-stressful temperatures. Heritability and evolvability based on among-sires (males), among-dams (females), and the sum of the two components (sire + dam) showed higher values at both the stressful temperatures than at the non-stressful temperature. These differences reflect changes in additive genetic variance. Viability was greater at the high than the low extreme temperature. As viability is an indicator of stress, we can assume that stress was greater at 18°C than at 32°C in D. ananassae. The genetic variations for all the quantitative and life-history traits were higher at low temperature. Variation in sexual traits was more pronounced as compared with other morphometric traits, which shows that sexual traits are more prone to thermal stress. Our results agree with the hypothesis that genetic variation is increased in stressful environments.     

大山雀身体大小的性二态及其季节变化

动物中普遍存在雌雄个体身体大小的性二态现象。了解近缘种之间身体大小性二态现象的差异，可为深入探讨身体大小性二态现象的潜在驱动机制提供证据。国外对欧亚大山雀（Parus major）的研究发现，其喙长、跗跖长、翅长等 6 项身体大小指标存在着明显的性二态，且喙长的性二态存在季节间差异。大山雀（P. cinereus）曾被作为欧亚大山雀的一个亚种，其形态和行为与欧亚大山雀存在着诸多相似之处。为探讨大山雀是否也存在身体大小性二态及季节性差异，本研究分析了 2018 至 2020 年间在河南董寨国家级自然保护区捕捉的 226 只（雌性 96 只和雄性 130 只）大山雀的喙长、头喙长、跗跖长、翅长、尾长和体长这 6 项体征指标的两性差异及其季节变化。结果显示，大山雀上述 6 项身体大小指标均存在不同程度的性二态现象，且雄性个体仅喙长与雌性的差异不显著，其余 5 项指标均显著大于雌性。此外，身体大小指标的两性差异不随季节显著变化，但两性的跗跖长在秋季均显著短于冬季和繁殖季，尾长在繁殖季均显著长于秋季和冬季。上述结果表明，大山雀身体大小的性二态及其季节性差异与欧亚大山雀并不完全相似。无论其身体大小存在性二态和季节变化的原因，还是其与欧亚大山雀在身体大小性二态模式上的差别，都有待今后进一步的研究。     

Condition and brightness of structural blue‐green: motmot tail‐racket brightness is related to speed of feather growth in males,but not in females

Coloration plays an important role in sexual and social communication, and in many avian species both males and females maintain elaborate colours. Recent research has provided strong support for the hypothesis that elaborate female traits can be maintained by sexual or social selection; however, most research on female ornamentation has focused on pigment‐based colours, and less is known about how structural colours are maintained. Both sexes of the turquoise‐browed motmot (Eumomota superciliosa) have a blue‐green racket‐tipped tail, and it remains unknown if tail coloration serves as a sexual or social signal in one or both sexes. Here, we describe sexual dichromatism in the blue‐green portion of the tail racket, and we test for a relationship between coloration and condition, as indicated by growth bars. Tail colour of both sexes has a similar spectral shape, and there is significant, although moderate, sexual dichromatism: males are brighter than females, and males have marginally greater blue‐green saturation than females. The length of feather grown per day is positively related to overall feather brightness, but this relationship is only present in males. The relationship between male coloration and condition suggests that tail colour has the potential to convey information about individual quality during mate choice or contest competition. The lack of a similar relationship in females suggests that female tail colour does not convey the same condition‐dependent information that we suggest may be reflected by male colour. Female tail colour may therefore reflect other aspects of condition, be involved in other (non‐condition‐dependent) forms of communication, or be expressed as a non‐functional byproduct of genetic correlation between the sexes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 673–681.     

Variation in Apical Hook Length Reflects the Intensity of Sperm Competition in Murine Rodents

Background

Post-copulatory sexual selection has been shown to shape morphology of male gametes. Both directional and stabilizing selection on sperm phenotype have been documented in vertebrates in response to sexual promiscuity.

Methodology

Here we investigated the degree of variance in apical hook length and tail length in six taxa of murine rodents.

Conclusions

Tail sperm length and apical hook length were positively associated with relative testis mass, our proxy for levels of sperm competition, thus indicating directional post-copulatory selection on sperm phenotypes. Moreover, our study shows that increased levels of sperm competition lead to the reduction of variance in the hook length, indicating stabilizing selection. Hence, the higher risk of sperm competition affects increasing hook length together with decreasing variance in the hook length. Species-specific post-copulatory sexual selection likely optimizes sperm morphology.     

Variation in the peacock’s train shows a genetic component

Female peafowl (Pavo cristatus) show a strong mating preference for males with elaborate trains. This, however, poses something of a paradox because intense directional selection should erode genetic variation in the males’ trains, so that females will no longer benefit by discriminating among males on the basis of these traits. This situation is known as the ‘lek paradox’, and leads to the theoretical expectation of low heritability in the peacock’s train. We used two independent breeding experiments, involving a total of 42 sires and 86 of their male offspring, to estimate the narrow sense heritabilities of male ornaments and other morphometric traits. Contrary to expectation, we found significant levels of heritability in a trait known to be used by females during mate choice (train length), while no significant heritabilities were evident for other, non-fitness related morphological traits (tarsus length, body weight or spur length). This study adds to the building body of evidence that high levels of additive genetic variance can exist in secondary sexual traits under directional selection, but further emphasizes the main problem of what maintains this variation.     

EVOLUTION,PHYLOGENY, SEXUAL DIMORPHISM AND MATING SYSTEM IN THE GRACKLES (QUISCALUS SPP.: ICTERINAE)

According to theory, two consequences of sexual selection are sexual dimorphism in size and secondary sexual characteristics, due to either intra- or intersexual selection. In this paper I suggest three criteria for the test of an evolutionary hypothesis involving quantitative morphological characters. First, the postulated change must be shown to have occurred in evolutionary time. Second, this change must be positively correlated with a change in the proposed selective agent. Third, given two taxa with different degrees of sexual size dimorphism and different mating system, the possible influence of drift must be rejected. If the hypothesis is not rejected by these three criteria, then we still have no proof of causality, but we can at least be more confident about its plausibility. This is applied to the particular hypothesis that sexual dimorphism in the Boat-tailed and Great-tailed grackles (Quiscalus spp; Icterinae; Aves) is caused by the highly polygynous mating system in these species. In relation to an outgroup, both species have increased disproportionately in male tarsus and tail size, creating an increased sexual dimorphism. This has cooccurred with the evolution of their particular mating system. However, the variance among species in male tarsus size can be accounted for by drift, and need not be a result of selection for increased size. In contrast, the variance among species in male tail size was much larger than expected under a null model of drift, indicating directional selection for long tails. The variance in female tail size was not larger than expected by drift, whereas the variance in female tarsus size was in fact lower than expected by drift, indicating stabilizing selection. The data are consistent with the hypothesis with regard to tail size, but not with regard to body size.     

Lifetime reproductive success,selection on lifespan,and multiple sexual ornaments in male European barn swallows

Natural and sexual selection arise when individual fitness varies according to focal traits. Extra‐pair paternities (EPPs) can affect the intensity of selection by influencing variance in fitness among individuals. Studies of selection require that individual fitness is estimated using proxies of lifetime reproductive success (LRS). However, estimating LRS is difficult in large, open populations where EPPs cause reallocation of biological paternity. Here, we used extensive field sampling to estimate LRS in a population of barn swallows (Hirundo rustica) to estimate selection on lifespan and ornamental traits of males. We found selection on lifespan mediated both by within‐ and extra‐pair fertilization success and selection on tail length mediated by within‐ but not extra‐pair fertilization success. In addition, we found selection on tail white spots via extra‐pair fertilization success after controlling for selection on other traits. These results were not confounded by factors that hamper studies of LRS, including nonexhaustive sampling of offspring and biased sampling of males. Hence, natural and sexual selection mediated by LRS operates on lifespan, tail length, and size of the tail white spots in barn swallows.     

Genetic and environmental effects on morphology and fluctuating asymmetry in nestling barn swallows

A barn swallow Hirundo rustica partial cross‐fostering experiment with simultaneous brood size manipulation was conducted in two years with contrasting weather conditions, to estimate heritable variation in tarsus, tail and wing size and fluctuating asymmetry. Environmental stress had contrasting effects depending on trait type. Significant heritabilities for tarsus, tail and wing size were found only in enlarged broods irrespective of year effects, while tarsus asymmetry was significantly heritable in the year with benign weather conditions irrespective of brood size manipulation effects. Tail, wing and composite (multicharacter) asymmetry were never significantly heritable. The environment with the higher heritability generally had higher additive genetic variance and lower environmental variance, irrespective of trait type. Heritability was larger for trait size than for trait asymmetry. Patterns of genetic variation in nestlings do not necessarily translate to the juvenile or adult stage, as indicated by lack of correlation between nestling and fledgling traits.     

Phenotypic variation and fluctuating asymmetry in sexually dimorphic feather ornaments in relation to sex and mating system

Secondary sexual characters have been hypothesized to demonstrate increased phenotypic variation between and within individuals as compared to ordinary morphological traits. We tested whether this was the case by studying phenotypic variation, expressed as the coefficient of variation (CV), and developmental instability, measured as fluctuating asymmetry (FA), in ornamental and non-ornamental traits of 70 bird species with feather ornamentation while controlling for similarity among species due to common descent. Secondary sexual characters differed from ordinary morphological traits by showing large phenotypic CV and FA. This difference can be explained by the different mode of selection operating on each kind of trait: a history of intense directional (ornaments) and stabilizing selection (non-ornaments). Phenotypic variation is reduced in the sex with more intense sexual selection (males), but does not differ among species with different mating systems. The strength of stabilizing selection arising from natural selection is associated with decreased CV (wing CV is smaller than tarsus or tail CVs). We found evidence of FA being reduced in ornamental feathers strongly affected by aerodynamics (tail feathers) compared to other ornaments, but only in females. In conclusion, CV and FA were not related, suggesting mat phenotypic plasticity and developmental instability are independent components of phenotypic variation.     

Parasites differentially increase the degree of fluctuating asymmetry in secondary sexual characters

The degree of fluctuating asymmetry has been demonstrated to reflect the ability of individuals to cope with different kinds of environmental stress (Parsons 1990). Parasites and diseases are one kind of environmental stress which most individuals encounter during their lifetime. Parasites have also been suggested to play an important role in sexual selection and the development of ornaments, since the full expression of ornaments may reflect the ability of hosts to cope with the debilitating effects of parasites. Here I report for the first time that a parasite, the haematophagous tropical fowl mite Ornithonyssus bursa (Macronyssidae, Gamasida), directly affects the degree of fluctuating asymmetry in a secondary sexual character of its host, the elongated tail of the swallow Hirundo rustica (Aves: Hirundinidae). I experimentally manipulated the mite load of swallow nests during one season by either increasing or reducing the number of mites, or keeping nests as controls. The degree of fluctuating asymmetry was measured in the subsequent year after the swallows had grown new tail ornaments under the altered parasite regime. The degree of fluctuating asymmetry was larger at increasing levels of parasites for male tail length, but not for the length of the shortest tail feather or wing length or for tail and wing length in females. These results suggest that the degree of fluctuating asymmetry in tail ornaments, but not in other feather traits, reliably reveals the level of parasite infestation. This has important implications for the ability of conspecifics to use the size and the expression of ornaments in assessment of phenotypic quality and thus in sexual selection.     

SEX CHROMOSOME LINKED GENETIC VARIANCE AND THE EVOLUTION OF SEXUAL DIMORPHISM OF QUANTITATIVE TRAITS

Theory predicts that sex chromsome linkage should reduce intersexual genetic correlations thereby allowing the evolution of sexual dimorphism. Empirical evidence for sex linkage has come largely from crosses and few studies have examined how sexual dimorphism and sex linkage are related within outbred populations. Here, we use data on an array of different traits measured on over 10,000 individuals from two pedigreed populations of birds (collared flycatcher and zebra finch) to estimate the amount of sex‐linked genetic variance (h²_z). Of 17 traits examined, eight showed a nonzero h²_Z estimate but only four were significantly different from zero (wing patch size and tarsus length in collared flycatchers, wing length and beak color in zebra finches). We further tested how sexual dimorphism and the mode of selection operating on the trait relate to the proportion of sex‐linked genetic variance. Sexually selected traits did not show higher h²_Z than morphological traits and there was only a weak positive relationship between h²_Z and sexual dimorphism. However, given the relative scarcity of empirical studies, it is premature to make conclusions about the role of sex chromosome linkage in the evolution of sexual dimorphism.     

Morphological flexibility across an environmental gradient in the epiphytic orchid,Tolumnia variegata: complicating patterns of fitness

Deceit‐pollinated orchid species show substantial variation in floral traits, which may be maintained by genetic drift or various forms of selection, or may reflect phenotypic plasticity. We explored how much plasticity occurs in both vegetative and floral traits of Tolumnia variegata (Oncidiinae, Orchidaceae) across two different light environments in Puerto Rico using data from a reciprocal transplant experiment. We also examined how fruit set, a measure of reproductive success and a surrogate for fitness, is associated with this morphological variation, and whether it changes over time. Tolumnia variegata responded to environmental variables in multiple ways. Vegetative characters were more plastic than those associated with sexual reproduction. Transplant effects accounted for significant variation in flower length, lip length, number of inflorescences, peduncle length, leaf length and the total number of ramets, but responses were not always consistent among years. Phenotypic selection on morphological characters was dependent on plant location. The trends detected were complex, and often inconsistent across years, probably as a result of wetter and drier years than average. Overall fruit set was quite variable among plants, averaging 15%, with no significant differences among sun and shade plants. Although reproductive success was similar among sites, habitat heterogeneity and annual variation had an effect on morphological expression, which sometimes modified the trajectories of phenotypic selection. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 431–446.     

Low inbreeding depression in a sexual trait in the stalk-eyed fly <Emphasis Type="Italic">Teleopsis dalmanni</Emphasis>

The genic capture hypothesis implies that the expression of sexual ornaments largely depends on genes affecting resource acquisition and use. The ornaments should thus show high degree of directional dominance typical of life-history traits, and consequently, they should be severely affected by inbreeding. Here we investigated the effect of inbreeding on a sexual ornament (male eyespan) in stalk-eyed fly, Teleopsis dalmanni. For comparison, we also measured inbreeding depression in non-sexual morphological traits: female eyespan as well as wing and thorax lengths in both sexes. Both eyespan, and other morphological traits we measured, showed significant inbreeding depression. In accord with predictions of genic capture hypothesis, male eyespan did decrease under inbreeding significantly more than female eyespan. However, the decline in male eyespan was fully explained by overall decline in body length. Moreover, the magnitude of inbreeding depression in male eyespan was considerably lower than that typically observed for life-histories; in fact, it fitted within the range typically characterizing morphological traits. We therefore conclude that our results provide weak support for genic capture hypothesis.