SEXUAL SELECTION IN THE MONOGAMOUS BARN SWALLOW (HIRUNDO RUSTICA). I. DETERMINANTS OF TAIL ORNAMENT SIZE

The proportion of phenotypic variance in the length of the sexually selected tail of the monogamous barn swallow Hirundo rustica that is attributable to genetic variance was studied in the field in Denmark during a seven-year period. Tail length was on average 20% greater in males than in females. Tail length correlated with wing length, but not with other morphological traits. Tail length increased with the first molt, but remained constant during subsequent years. Changes in tail length between years, owing to molt were significantly affected by sex and by degree of infection with an haematophagous mite (Ornithonyssus bursa). There were significant differences in sexual size dimorphism between years, apparently as a result of environmental conditions in the African winter quarters during molt. Tail length was a highly repeatable morphological trait, and standardization of tail length for age effects only marginally increased repeatability. Heritability of tail length as estimated from regression of values for sons on those of their fathers was 0.59. This suggests that secondary sexual traits affected by strong directional selection still may show a statistically significant heritability.     

Better‐surviving barn swallow mothers produce more and better‐surviving sons

Sex allocation theory predicts that parents are selected to bias their progeny sex ratio (SR) toward the sex that will benefit the most from parental quality. Because parental quality may differentially affect survival of sons and daughters, a pivotal test of the adaptive value of SR adjustment is whether parents overproduce offspring of the sex that accrues larger fitness advantages from high parental quality. However, this crucial test of the long‐term fitness consequences of sex allocation decisions has seldom been performed. In this study of the barn swallow (Hirundo rustica), we showed a positive correlation between the proportion of sons and maternal annual survival. We then experimentally demonstrated that this association did not depend on the differential costs of rearing offspring of either sex. Finally, we showed that maternal lifespan positively predicted lifespan of sons but not of daughters. Because in barn swallows lifespan is a strong determinant of lifetime reproductive success, the results suggest that mothers overproduce offspring of the sex that benefits the most from maternal quality. Hence, irrespective of mechanisms causing the SR bias and mother–son covariation in lifespan, we provide strong evidence that sex allocation decisions of mothers can highly impact on their lifetime fitness.     

Selection on multiple sexual signals in two Central and Eastern European populations of the barn swallow

Variation in intensity and targets of sexual selection on multiple traits has been suggested to play a major role in promoting phenotypic differentiation between populations, although the divergence in selection may depend on year, local conditions or age. In this study, we quantified sexual selection for two putative sexual signals across two Central and East European barn swallow (Hirundo rustica rustica) populations from Czech Republic and Romania over multiple years. We then related these differences in selection to variation in sexual characters among barn swallow populations. Our results show that tail length and ventral coloration vary between populations, sexes, and age classes (first‐time breeders vs. experienced birds). We found that selection on tail length was stronger in first‐time breeders than in experienced birds and in males than in females in the Romanian population, while these differences between age groups and sexes were weak in Czech birds. We suggest that the populational difference in selection on tail length might be related to the differences in breeding conditions. Our results show that ventral coloration is darker (i.e., has lower brightness) in the Romanian than in the Czech population, and in experienced birds and males compared with first‐time breeders and females, respectively. The sexual difference in ventral coloration may suggest sexual selection on this trait, which is supported by the significant directional selection of ventral coloration in first‐time breeding males on laying date. However, after controlling for the confounding effect of wing length and tarsus length, the partial directional selection gradient on this trait turned nonsignificant, suggesting that the advantage of dark ventral coloration in early breeding birds is determined by the correlated traits of body size. These findings show that ventral coloration may be advantageous over the breeding season, but the underlying mechanism of this relationship is not clarified.     

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.     

Ectoparasites cause increased bilateral asymmetry of naturally selected traits in a colonial bird

Abstract Parasitism has been shown to correlate with levels of bilateral symmetry in some organisms, with more asymmetric individuals often having more parasites. However, few studies have shown experimentally that parasitism directly causes increased asymmetry. By fumigating some cliff swallow (Petrochelidon pyrrhonota) colonies and leaving others untreated, we investigated experimentally whether ectoparasitism by the cimicid swallow bug led to higher levels of asymmetry in length of wings, outer tail feathers, and tarsus among juvenile and adult birds. Juveniles from fumigated colonies measured soon after fledging had significantly less asymmetry in wing and outer tail length than juveniles from nonfumigated colonies; asymmetry in tarsus length was unaffected by parasitism. Adults that had undergone one or more post‐juvenal molts on the wintering grounds showed no differences in asymmetry between those reared in fumigated vs. nonfumigated colonies. These results show that ectoparasitism directly leads to increased feather asymmetry in cliff swallows, probably through parasite‐induced nutritional stress. Because wing and tail asymmetry impair flight performance and reduce foraging efficiency, the increased asymmetry caused by parasites represents a fitness cost to cliff swallows. This is among the few experimental studies to show an effect of parasites on asymmetry of naturally selected characters.     

Performance of potential European biological control agents of Vincetoxicum spp. with notes on their distribution

Exploratory surveys were conducted in Europe to locate potential biological control agents of Vincetoxicum nigrum and Vincetoxicum rossicum and to collect known specialist herbivores of Vincetoxicum hirundinaria (Abrostola asclepiadis, Chrysolina a. asclepiadis, Eumolpus asclepiadeus and Euphranta connexa). Development of the most abundant herbivores found attacking leaves, roots, and developing seeds was evaluated in the laboratory on three Vincetoxicum spp. Field surveys revealed that the highest diversity of herbivores was associated with the abundant and widespread V. hirundinaria. No new herbivores were reported from V. hirundinaria or V. nigrum, but the leaf‐feeding noctuid Hypena opulenta was recorded for the first time attacking V. rossicum and V. scandens in Ukraine. Based upon larval survival and development and adult fecundity, the leaf feeders A. asclepiadis, C. a. asclepiadis, and H. opulenta, perform better on V. nigrum and V. rossicum than on V. hirundinaria. Larval performance of the root feeder E. asclepiadeus followed a similar pattern, but adult fecundity of this insect did not vary among host plant species. Immature development time of the pre‐dispersal seed feeder E. connexa is similar among hosts, but larvae grow larger on V. nigrum and V. hirundinaria than on V. rossicum. All herbivores are promising biological control agents of Vincetoxicum in North America and we have prioritized host range testing with H. opulenta and E. asclepiadeus.     

Within seasons and among years: when are corticosterone levels repeatable?

Hormones play a central role in integrating internal and external cues to help mediate life-history decisions as well as changes in behavior and physiology of individuals. Describing the consistency of endocrine traits within and among individuals is an important step for understanding whether hormonal traits are dependable predictors of phenotypes that selection could act upon. However, few long-term field studies have investigated the individual consistency of hormonal traits. Glucocorticoid hormones mediate homeostatic responses to environmental variation as well as stress responses to acute, unpredictable disturbances. We characterized the repeatability of plasma corticosterone concentrations in two species of free-living passerines across multiple years. We found repeatability in baseline corticosterone concentrations in both sexes of great tits (Parus major) and in female tree swallows (Tachycineta bicolor) within the breeding season but no repeatability of this trait among seasons or across years. Stress-induced levels of corticosterone were only assessed in great tits and were not repeatable in either sex. Our data suggest that in line with their function in mediating responses of individuals to longer-term and acute demands, both baseline and stress-induced plasma corticosterone concentrations are rather plastic traits. However, individuals may differ in their degree of trait plasticity and hence in behavioral and physiological responses to a variety of organismal challenges.     

Impaired swimming behaviour and morphology of sperm from barn swallows Hirundo rustica in Chernobyl

We investigated the motility and morphology of live sperm from barn swallows Hirundo rustica breeding in radioactively contaminated areas around Chernobyl and control areas in Ukraine in order to test the hypothesis that swimming behaviour and morphology of sperm was impaired by radioactive contamination. We obtained sperm samples from 98% of sampled birds, thus avoiding sampling bias due to the fraction of males not producing sperm samples. Analyses of within- and between-sample repeatability revealed significant and intermediate to large estimates for all sperm parameters. There were significant differences between the Chernobyl area and the control area for two of 11 sperm behaviour parameters, and significant interactions between area and year for six of these parameters. The proportion of sperm with abnormal morphology was elevated in barn swallows from Chernobyl. A principal component (PC) analysis revealed four significant axes that explained 88% of the variance in sperm behaviour parameters. One of these principal components differed between areas, and three components showed significant year by area interactions. PC2 representing the frequency of slow sperm increased with increasing radiation in one year, but not another. PC3 representing sperm with high linearity, small amplitude of lateral head displacement and low track velocity decreased with increasing background radiation level. PC4 reflecting a large proportion of static sperm with high beat cross frequency increased with increasing background radiation level. Sperm behaviour as reflected by principal components was predictable among years from information on level of radiation, and it was predictable among sites in different years. These results are consistent with the hypothesis that sperm behaviour and morphology have been affected by radiation due to the Chernobyl accident.     

An analysis of continent-wide patterns of sexual selection in a passerine bird

Patterns of selection are widely believed to differ geographically, causing adaptation to local environmental conditions. However, few studies have investigated patterns of phenotypic selection across large spatial scales. We quantified the intensity of selection on morphology in a monogamous passerine bird, the barn swallow Hirundo rustica, using 6495 adults from 22 populations distributed across Europe and North Africa. According to the classical Darwin-Fisher mechanism of sexual selection in monogamous species, two important components of fitness due to sexual selection are the advantages that the most attractive males acquire by starting to breed early and their high annual fecundity. We estimated directional selection differentials on tail length (a secondary sexual character) and directional selection gradients after controlling for correlated selection on wing length and tarsus length with respect to these two fitness components. Phenotype and fitness components differed significantly among populations for which estimates were available for more than a single year. Likewise, selection differentials and selection gradients differed significantly among populations for tail length, but not for the other two characters. Sexual selection differentials differed significantly from zero across populations for tail length, particularly in males. Controlling statistically for the effects of age reduced the intensity of selection by 60 to 81%, although corrected and uncorrected estimates were strongly positively correlated. Selection differentials and gradients for tail length were positively correlated between the sexes among populations for selection acting on breeding date, but not for fecundity selection. The intensity of selection with respect to breeding date and fecundity were significantly correlated for tail length across populations. Sexual size dimorphism in tail length was significantly correlated with selection differentials with respect to breeding date for tail length in male barn swallows across populations. These findings suggest that patterns of sexual selection are consistent across large geographical scales, but also that they vary among populations. In addition, geographical patterns of phenotypic selection predict current patterns of phenotypic variation among populations, suggesting that consistent patterns of selection have been present for considerable amounts of time.