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Biological invasions by exotic species impose substantial ecological, economic and social costs worldwide, being a major threat to biodiversity conservation. Because not all individuals introduced in the new environments become successful invaders, the identification of factors underlying variation in invasion success would be essential for evaluating invasion risk. Here, we test several host–parasite hypotheses accounting for invasion success of house sparrows Passer domesticus in Peru. According to the Enemy Release Hypothesis, invasive house sparrows from Peru showed lower prevalence and genetic diversity of haemosporidian parasites than sparrows from their natural range (Spain), indicating that the release from their natural parasites may have favoured the spread of sparrows in the new area of occurrence. We also showed that Peruvian sparrows had larger uropygial glands and higher anti‐bacterial activity in its secretion than sparrows from Spain, suggesting selection in defensive mechanisms driven by pathogens when colonizing new environments. Finally, we showed that uninfected sparrows had larger uropygial glands and higher anti‐bacterial activity than malaria‐infected house sparrows, implying that uropygial gland secretions may act as a defensive mechanism against haemosporidian infections. Alternatively, a condition‐dependent trade‐off exists between synthesis of uropygial secretion and immune response. These outcomes provide essential information for identifying potential invaders and designing interventions. The release from their natural malaria parasites may have favoured the spread of sparrows in South America: invasive house sparrows from Peru showed lower prevalence and genetic diversity of haemosporidian parasites than sparrows from their natural range in Spain. Furthermore, invasive sparrows had larger uropygial glands and higher anti‐bacterial activity in their secretion than sparrows from their original range, suggesting selection in defensive mechanisms driven by pathogens when colonizing new environments. [ABSTRACT FROM AUTHOR]

Pathogens such as bacteria, fungi and malaria and related haemosporidians provoke negative effects on the fitness of their hosts. Animals have developed a range of defensive mechanisms to resist or eliminate these parasitic infections and their negative fitness costs. The uropygial gland secretion has been proposed to act as defensive barrier of skin and plumage in the fight against bacteria and fungi, and may prevent birds from acquiring haemosporidian infections. Thus, the secretion of uropygial glands of birds may favour survival of individuals. However, whether uropygial gland secretion influence survival remains unknown. Here we explore if the size of the uropygial gland and malaria infection influence survival of house martins Delichon urbica. We showed, for the first time, that the volume of the uropygial gland positively predicted survival prospects of malaria infected house martins. Malaria infected birds had the lowest probability of survival, with the effect of gland size on survival prospects depending on infection: infected house martins with larger uropygial glands were better able to survive to the next breeding season, while infected birds with small uropygial glands were not. These results highlight the importance of uropygial gland secretion in the life history of wild birds. [ABSTRACT FROM AUTHOR]

Understanding connections between breeding, stopover and wintering grounds for long-distance migratory birds can provide important insight into factors influencing demography and the strength of carry-over effects among various periods of the annual cycle. Using previously described, multi-isotope (δ13C, δ15N, δ2H) feather isoscapes for Africa, we identified the most probable wintering areas for house martins Delichon urbica breeding at Badajoz in southwestern Spain. We identified two most-probable wintering areas differing in isotopic signature in west Africa. We found that the probability to winter in the isotopic cluster two was related to age and sex of individuals. Specifically, experienced males (i.e. two years or older) winter in the isotopic cluster two with a greater probability than experienced females, whereas first-year females winter in the isotopic cluster two with a greater probability than first-year males. In addition, wintering area was correlated with breeding phenology, with individuals wintering in the isotopic cluster two initiating their clutches earlier than those wintering in the isotopic cluster one. For birds wintering in the isotopic cluster two, there was no relationship between age and clutch initiation date. In contrast, young birds wintering in the isotopic cluster one initiated their clutches earlier than experienced birds wintering in this area. There was no significant correlation between wintering area and clutch size or the number of fledglings produced. We hypothesize that the relationship among social status, population density and winter habitat quality should be the most important driver of the carry-over effect we found for this population. [ABSTRACT FROM AUTHOR]

Migration is a complex trait although little is known about genetic correlations between traits involved in such migration syndromes. To assess the migratory responses to climate change, we need information on genetic constraints on evolutionary potential of arrival dates in migratory birds. Using two long-term data sets on barn swallows Hirundo rustica (from Spain and Denmark), we show for the first time in wild populations that spring arrival dates are phenotypically and genetically correlated with morphological and life history traits. In the Danish population, length of outermost tail feathers and wing length were negatively genetically correlated with arrival date. In the Spanish population, we found a negative genetic correlation between arrival date and time elapsed between arrival date and laying date, constraining response to selection that favours both early arrival and shorter delays. This results in a decreased rate of adaptation, not because of constraints on arrival date, but constraints on delay before breeding, that is, a trait that can be equally important in the context of climate change. [ABSTRACT FROM AUTHOR]

Many secondary sexual characters vary in a systematic way with the age of individuals, with young and old individuals displaying at lower levels than individuals of intermediate age. Analyses quantifying the within-individual and among-individual components of phenotypic variation can help partition effects of phenotypic plasticity and selective mortality. We analysed phenotypic variation in the expression of a secondary sexual character, tail length, in male and female barn swallows Hirundo rustica from four European populations studied during 11-26 years, using linear mixed effect models to describe age-related expression. Tail length increased from yearlings to intermediate aged birds with a subsequent decrease at old age. In males, this age-related pattern was because of both within-subject and between-subject effects, with no difference among populations. Males having longer lifespan had shorter tails when young than those having shorter lifespan. Females showed similar patterns of age-related variation as males, with no difference among populations. The major difference between sexes was that the between-subject effects (i.e. disappearance effects or selection) were much more important for males compared to females for which lifetime variation in tail length was mainly because of a within-subject effect (i.e., a plastic response). These findings suggest that whereas males trade greater expression of the secondary sexual character at young age against longevity, that was not the case for females. This is consistent with tail length being more costly in males than in females, with the cost of long tails potentially being offset by elevated mating success, whereas that is not the case in females. [ABSTRACT FROM AUTHOR]

Senescence is the deterioration of the phenotype with age caused by negative effects of mutations acting late in life or the physiological deterioration of vital processes. Birds have traditionally been assumed to senescence slowly despite their high metabolic rates, high blood sugar levels and high body temperature. Here we investigate the patterns of age-related performance of sperm of a long distance migrant, the barn swallow Hirundo rustica, varying in age from 1 to 6 years, analysed by the computer-assisted sperm analysis equipment. Sperm showed deteriorating performance in terms of linear movement, track velocity, straight line velocity and reduced proportions of rapidly moving, progressive and motile sperm with age. In a second series of experiments, we assessed performance of sperm from the same males in neutral medium and in medium derived from the reproductive tract of females in an attempt to test if sperm of old males performed relatively better in female medium, as expected from extra-pair paternity being negatively related to male age, but not to female age. Older males showed consistently better performance in female medium than in neutral medium in terms of track velocity, straight line velocity and reduced proportions of rapidly moving, progressive and motile sperm, whereas young males showed better performance in neutral medium. These results provide evidence of declining sperm performance for important reproductive variables not only with age, but also with the sperm of old males performing differentially better in female medium than young males. [ABSTRACT FROM AUTHOR]

Bird song is believed to honestly reveal male quality including the ability of singers to face parasitism. In a natural population of barn swallows Hirundo rustica, we experimentally imposed a cost on song production by an immune challenge. We therefore vaccinated a group of reproducing males with an antigen (Newcastle disease virus), and injected phosphate-buffered saline to a control group. Immune challenge significantly reduced one song feature, rattle duration. This decrease was related to male quality, as measured by tail length, because males with short tails reduced the duration of their rattle significantly more than males with long tails. In addition, another song feature, strophe duration, decreased in the control group, while it remained constant in the challenged group. Duration of the rattle has previously been found to be positively related to testosterone level, and it may hence reflect male competitive ability. Thus, male barn swallows may not have the potential to produce long rattles when their immune system is challenged. By maintaining their strophe duration after an immune challenge, males may compensate for the decrease in rattle duration. Our results suggest that different song features may convey different types of information, and that barn swallow song, in particular their rattle, may reliably reveal information about activation of the immune system. [ABSTRACT FROM AUTHOR]

Avian malaria parasites are supposed to exert negative effects on host fitness because these intracellular parasites affect host metabolism. Recent advances in molecular genotyping and microscopy have revealed that coinfections with multiple parasites are frequent in bird–malaria parasite systems. However, studies of the fitness consequences of such double infections are scarce and inconclusive. We tested if the infection with two malaria parasite lineages has more negative effects than single infection using 6 years of data from a natural population of house martins. Survival was negatively affected by both types of infections. We found an additive cost from single to double infection in body condition, but not in reproductive parameters (double-infected had higher reproductive success). These results demonstrate that malaria infections decrease survival, but also have different consequences on the breeding performance of single- and double-infected wild birds. [ABSTRACT FROM AUTHOR]

Variation in the expression of sexually selected traits among individuals is widely investigated on the premise that these traits evolved to signal male quality. Significant repeatabilities of sexual signals and their associations with condition, mating success, survivorship and age may be the signatures of sexual selection. However, little is known about the relationship between these sexual attributes. Here we studied 28 acoustic and visual traits in the barn swallow, Hirundo rustica, that may potentially function in sexual selection. Based on effect sizes calculated at the between-individual level, we assessed the relationship between repeatability, condition-dependence, attractiveness, age-dependence and viability indicator value of sexual traits using sexual signals as the units of analyses. Those traits that showed high within-year repeatability also showed high between-year repeatability, indicating that between-individual variation is consistent within and among seasons. In addition, age-dependence of traits, probably causing between-year variation, was negatively related to between-year repeatability. Condition-dependence was negatively correlated with effect sizes for the extent to which traits predicted viability. Therefore, traits that are positively related to immediate condition are those that are negatively related to survival, which may be the signature of a trade-off between current and future reproductive success ultimately reflecting signal reliability. No other significant relationship was found between trait attributes. We conclude that multiple sexual signals reflect different aspects of male quality in the barn swallow. [ABSTRACT FROM AUTHOR]

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 tall 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. [ABSTRACT FROM AUTHOR]

Maternal effects such as androgen in avian eggs can mediate evolutionary responses to selection, allowing manipulation of offspring phenotype and promoting trans-generational adaptive effects. We tested the predictions of two adaptive hypotheses that have been proposed to explain female variation in yolk androgen allocation in birds, using the barn swallow Hirundo rustica as a model. We found no support for the first hypothesis proposing that yolk androgen varies as a function of breeding density in order to prepare offspring for different breeding densities. However, we found experimental support for the hypothesis that female yolk androgen allocation depends on mate attractiveness and that it constitutes an example of differential allocation. Females increased the concentration of androgens in their eggs when mated to males with experimentally elongated tails. Female phenotypic quality as measured by arrival date and clutch size was positively related to egg androgen concentration, consistent with the hypothesis that this is a costly investment, constrained by female condition. We found correlative evidence of a direct relationship between egg androgen concentration and performance of offspring as measured by mass increase. [ABSTRACT FROM AUTHOR]

The T-cell mediated response to a challenge of the immune system with phytohemagglutinin (PHA) is a common measure used in ecological studies of host-parasite interactions or parasite-mediated selection. We investigated the temporal dynamics of this response in house sparrows Passer domesticus in order to determine factors that contribute to temporal and individual variation in PHA response. After an initial significant increase in response from injection to six hours post-injection, there was no further significant change in mean response after 12, 24, 36, 48 and 72 hours. Responses at night were consistently stronger than during daytime. Individuals may benefit from producing both a strong and a quick immune response to parasite attacks, although this may be impossible because of the observed positive relationship between maximum response and latency to maximum response. House sparrows with Haemoproteus infections had lower PHA responses and smaller maximum responses than individuals without infections. Individuals in prime body condition had stronger PHA responses than individuals in poor condition. House sparrows with large diurnal fluctuations in body mass had a weaker maximum response than individuals with small fluctuations. Since individuals with Haemoproteus infections have large fluctuations in body mass, this result suggests that infected individuals with large fluctuations in mass are unable to mount a strong immune response. [ABSTRACT FROM AUTHOR]

The effects of herbivory on plant fitness have traditionally been assessed by reductions in performance measured in terms of physiological, growth, and reproductive parameters. We investigated the direct effect of herbivory on a measure of foliar developmental instability (fluctuating asymmetry in leaf width) in evergreen stone oak Quercus rotundifolia. Since leaves from more than a single growth season are present simultaneously, it is possible to compare measures of developmental instability of newly formed leaves in relation to the extent of herbivory in the previous season as assessed from damage to old leaves. We found significantly increased asymmetry in leaf width when leaves from the previous year showed signs of herbivory. This result was confirmed experimentally by using leaf removal to mimic herbivory. Oak trees where many old leaves were removed had increased foliar asymmetry compared to oak trees where only few leaves had been removed anduntreated control trees. Using foliar asymmetry in old leaves from the previous year as a baseline for these comparisons confirmed that foliar asymmetry of individual trees increased as a consequence of herbivory. Oak trees with high degrees of asymmetry were more likely to die than oak trees with low asymmetry. These results suggest that foliar asymmetry is symptomatic of a stressed plant. Herbivores can bothproduce such a stress, leading to increased asymmetry, and attack differentially more asymmetric plants. In the extreme, the most asymmetric plants are those most likely to die. [ABSTRACT FROM AUTHOR]

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity. 2. The change of haematocrit of Barn Swallows ( Hirundo rustica) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed. 3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males. 4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length. 5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males. [ABSTRACT FROM AUTHOR]

1. Senescence reflects age-dependent changes in residual reproductive value. Annual survival rates of the barn swallow Hirundo rustica L. increased from 1- to 2-year-old individuals, but decreased among 5 years old or older individuals. Estimates of age-dependent reproductive value showed a similar pattern. 2. Longitudinal data from two long-term population studies were used to test whether a number of different measures of performance (condition-dependent morphological traits, migratory performance, reproductive success, intensity of parasitism) changed among individuals when reaching old age. 3. The length of the outermost tail feathers (a secondary sexual character) decreased among old individuals, while two measures of individual developmental instability increased with age. Migratory performance decreased in old barn swallows as reflected by a delay in spring arrival at the breeding grounds. Reproductive performance measured as seasonal reproductive success decreased with age. The intensity of infestations with an haematophagous mite and a mallophagous ectoparasite increased among old barn swallows. 4. These results suggest that the condition-dependent secondary sexual character, developmental stability, and measures of migratory and reproductive performance deteriorated, and the frequency of parasitism increased among old individuals. Ageing was thus associated with a general deterioration of performance. [ABSTRACT FROM AUTHOR]

Studies the ability of female barn swallows or Hirundo rustica to adjust their reproductive effort to the attractiveness of their mates. Influence of secondary sexual characters in the reproductive effort of female animals; Experimental manipulation of the male tail length.

Heat-shock proteins (HSPs) are synthesized by animals and plants in response to various stressors. The level of the HSP60 stress protein was measured from the cell fraction of peripheral blood obtained from nestling house martins ( Delichon urbica) to test whether ectoparasitism increased the concentration of stress protein. We assessed HSP from nestlings raised in nests previously treated with an insecticide or infested with 50 martin bugs ( Oeciacus hirundinis). In addition, haematozoa infections were checked in blood smears. Nestlings from parasite-infested nests, or nestlings infected with trypanosomes, had increased levels of HSP in their blood cells. Nestling growth as determined from wing length was negatively related to HSP60 levels and within-brood variation in wing length increased with increasing levels of the stress protein independently of treatment and infection by trypanosomes. These results suggest HSPs may play a role in host-parasite interactions, and that they can be used reliably for measuring physiological responses to parasites. [ABSTRACT FROM AUTHOR]

Parasites often exert strong selection pressures on their hosts that have evolved anti-parasite defences to counter the negative effects of parasites. We studied the relationship between intensity of parasitism, one aspect of host immune response, and host reproductive success, using the house martin bug Oeciacushirundinis and its house martin Delichonurbica host as a model system. Experimental manipulation of parasite load of nests during laying of the first clutch altered the intensity of parasitism. Parasites reduced the reproductive success of their hosts measured in terms of body condition and survival of nestlings. Host immune response, measured as the concentration of gammaglobulins and total plasma proteins, was positively associated with parasite reproduction, estimated as the number of juvenile parasites, but was only weakly related to the intensity of adult parasites. The concentration of gammaglobulins was negatively related to nestling body mass, implying a trade-off between immune function and body condition. Parasite reproduction thus exerts a cost on hosts by increasing the immune response. [ABSTRACT FROM AUTHOR]

Replies to the article of Klaudia Witte which commented on the authors' study on the differential allocation hypothesis which was published in the 1993 issue of the `Evolution' periodical. Earlier studies of the sexual behavior Hirundo rustica; Inability of the differential allocation hypothesis to account for differential female investment in offspring when they are mated to males with the most attractive phenotype.