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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.

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.

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.

Previous studies have linked winter habitat with subsequent breeding phenology and physical condition of migratory birds, but few have found delayed effects of winter habitat on subsequent reproductive success. The aim of this study was to test if African winter habitat is related to subsequent reproductive success of house martins (Delichon urbicum) breeding at a colony in Spain. We measured stable isotope (δ2H, δ13C, δ15N) values from feathers moulted in West Africa and used confirmatory path analysis to test if isotopic values of winter-grown feathers were related to reproductive success through the mediation of breeding phenology and body condition. We conducted separate analyses for males, females and age classes (yearlings vs ≥ 2 years old). Experienced males wintering in habitats of higher rainfall (as inferred from lower feather δ2H values) were in better body condition and produced more offspring during the subsequent breeding season. In contrast, we did not find any effect of winter habitat on reproductive success of young males or females. These findings provide evidence consistent with a complex causal link between winter habitat quality and subsequent breeding success of long-distance migratory songbirds.

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle-aged females (i.e. 2-4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within-individual increase in timing of breeding and clutch size in early life and a greater within-individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle-aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor-quality members of the pairs, mainly poor-quality males. Age-related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life-history strategies in free-ranging animals from more than a single population can throw new light on life-history theory, population dynamics and evolutionary studies of senescence.

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.

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.

Protandry is the difference in arrival date between males and females, with competition among males for access to preferred territories (the rank advantage hypothesis) or mating success (the mate opportunity hypothesis) supposedly driving the evolution of protandry. The fitness costs and benefits of protandry accruing to individuals differing in degree of protandry (arrival date of a male relative to the arrival date of his partner) have never been quantified. We analyzed the fitness consequences of sex differences in arrival date in the barn swallow Hirundo rustica, in which arrival date can be precisely estimated and the fitness of pairs differing in degree of individual protandry assessed. Early arriving males had greater mating success than late arriving males. The number of extrapair offspring in own nests decreased with increasing degree of individual protandry, whereas the number of offspring fathered by a focal male was unrelated to individual protandry. There was directional selection on individual protandry as shown by pairs with a larger than average degree of protandry reproducing early and, hence, supposedly producing more recruits. There was also stabilizing selection on individual protandry as shown by pairs with an intermediate degree of protandry reproducing early. Annual production of fledglings increased with early arrival of males, but not with early arrival of females, once the effect of laying date had been considered, with no additional effect of individual protandry. Neither male nor female survival was significantly related to degree of individual protandry. These findings are consistent not only with the mate opportunity hypothesis but also with a sexual conflict hypothesis, suggesting that males and females differ in their optimal timing of arrival due to sex-specific fitness costs and benefits. Key words: barn swallow, extrapair paternity, Hirundo rustica, optimal arrival time, sex ratio. [Behav Ecol]

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.

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.

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.

Ever since the Chernobyl accident in 1986, that contaminated vast areas in surrounding countries with radiation, abnormalities and birth defects have been reported in human populations. Recently, several studies suggested that the elevated frequency of such abnormalities can be attributed to poverty and stress in affected human populations. Here, we present long-term results for a free-living population of barn swallows, Hirundo rustica , demonstrating the presence of 11 morphological abnormalities in populations around Chernobyl, but much less frequently in an uncontaminated Ukrainian control population and three more distant control populations. The presence of these abnormalities in barn swallows is associated with reduced viability. These findings demonstrate a link between morphological abnormalities and radiation in an animal population that cannot be attributed to poverty and stress. The most parsimonious hypothesis for abnormalities in animal and human populations alike is that the effects are caused by the same underlying cause, viz. radiation derived from the Chernobyl accident.

11 pages; International audience; UNCORRECTED PROOF 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 in terms of 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.

Predation risk may affect the allocation priorities of limiting resources by potential prey. Investment in immune function should receive reduced priority, when hosts are exposed to predators because of the costs of immune function. We tested this hypothesis by randomly exposing adult house sparrows, Passer domesticus, to either a cat, Felis catus, or a rabbit, Oryctolagus cuniculus, for 6 h while assessing their ability to raise a T-cell--mediated immune response to a challenge with phytohemagglutinin. Sparrows exposed to a cat had a significant reduction of, on average, 18% and 36% in T-cell response in two different experiments compared with sparrows that were exposed to a rabbit. In a field experiment with a barn owl, Tyto alba, or a rock dove, Columba livia, placed next to a nest-box during laying, we found a mean reduction in T-cell--mediated immune response of 20%. In males, the reduction in cell-mediated immune response owing to cat exposure increased with increasing size of the badge, which is a secondary sexual character, but only during the breeding season. In a third experiment, house sparrows were either exposed to a barn own, T. alba, or a rock dove, C. livia, and development of malarial infections was recorded during the following 6 weeks. Individual sparrows exposed to a predator had a higher prevalence and intensity of Haemoproteus malarial infection than did control individuals. Therefore, exposure to predators reduced their ability of hosts to cope with parasitism mediated through effects on immune function. Copyright 2004.

in relation to time of day,body condition and blood parasite infection. – Oikos 101: 291–298.The T-cell mediated response to a challenge of the immune system with phytohemag-glutinin (PHA) is a common measure used in ecological studies of host-parasiteinteractions or parasite-mediated selection. We investigated the temporal dynamics ofthis response in house sparrows

We tested the novel hypothesis that arrival date in migratory birds represents a reliable indicator of male quality that can be used by females as a cue in extrapair mating decisions. Secondary sexual characters are often condition-dependent, and competition for early arrival leads to condition-dependent migration. Hence, both secondary sexual characters and arrival date are predicted to be condition-dependent indicators of male phenotypic quality. We studied the relationship between expression of a secondary sexual character, arrival date, and condition, respectively, and extrapair paternity in a Spanish population of barn swallows, Hirundo rustica. By using microsatellite markers to determine paternity, we showed that 17.8% of all offspring (N = 674) and 32.4% of all broods (N = 170) were due to extrapair paternity. Quasi-parasitism (in which the male nest owner fathered the offspring, but the eggs were laid by another female) occurred in 2.6% of all nestlings and 2.9% of all broods. Individuals were consistent in the frequency of extrapair paternity among first, second, and third broods. Males with long outermost tail feathers, arriving early and in prime body condition, had little extrapair paternity in their nests. This was also the case when controlling for the confounding effects of male age. Partial correlation analysis was used to investigate the direct and indirect effects of tail length, arrival date, and body condition on extrapair paternity. Body condition accounted for most of the variance in extrapair paternity, whereas tail length and arrival date accounted for a smaller proportion of the variance. Body condition was strongly correlated with tail length and arrival date. However, because females cannot directly assess condition or arrival date (males arrive before females), females may obtain an indirect measure of condition and migration ability from tail length and other phenotypic traits of males. This suggests that extrapair paternity depends on the effects of condition, through its indirect effects on arrival date, tail length, and other variables. Copyright 2003.

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.

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.

The functional signi¢cance of elongated, narrow tips of the tail feathers of certain birds, so-called tail streamers, has recently been discussed from an aerodynamic point of view, and the e¡ects of sexual selection on such traits have been questioned. We review our long-term ¢eld studies using observational and experimental approaches to investigate natural and sexual selection in the barn swallow, Hirundo rustica, which has sexually size-dimorphic outermost tail feathers. Experimental manipulation of the length of the outermost tail feathers has demonstrated sexual selection advantages of tail elongation and disadvantages of tail shortening, with opposite e¡ects for natural selection in terms of foraging e¤ciency, haematocrit and survival. These ¢ndings are contrary to the prediction of a general deterioration from both shortening and elongation, if the tail trait was determined solely by its e¡ects on aerodynamic e¤ciency and £ight manoeuvrability. Patterns of sexual selection in manipulated birds conform with patterns in unmanipulated birds, and selection di¡erentials for di¡erent components of sexual selection in manipulated birds are strongly positively correlated with di¡erentials in unmanipulated birds. Age and sex di¡erences in tail length, and geographical patterns of sexual size dimorphism, are also consistent with sexual selection theory, but inconsistent with a purely natural selection advantage of long outermost tail feathers in male barn swallows., A.P.M. was supported by the Danish Natural Sciences Research Council, S.M. by a postdoctoral grant fromthe Spanish Ministry of Education, A.B. by a Marie Curie postdoctoral grant of the European Communities, and F.deL. by Spanish DGICYT (PB 95020).

Summary 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.

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.

Secondary sexual characters are assumed to be costly to produce and maintain, and this will select for morphological modifications that reduce the magnitude of such costs. Here we test whether a feather ornament, the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject to a directional female mate preference, and other aspects of the morphology used for flight have been modified to increase aerodynamic performance. This was done by making comparisons among sexes within populations, among individuals varying in tail length within populations, and among populations from different parts of Europe. Male barn swallows experienced reduced drag from their elongated tail feathers by morphological modifications of the ornamental feathers as compared to females. Morphological features of the outermost tail feathers were unrelated to tail length in both males and females within populations. Wing and tail morphology (length of central tail feathers and wings, wing span, wing area, wing loading, and aspect ratio) was modified in males compared to females. Barn swallows with long tails had morphological tail and wing modifications that reduced the cost of a large ornament, and similar modifications were seen among populations. The costs of the exaggerated secondary sexual character were therefore reduced by the presence of cost-reducing morphological modifications. The assumptions of reliable signalling theory, that signals should be costly, but more so to low than to high quality individuals, were not violated because long-tailed male barn swallows had the largest cost-reducing morphological characters.

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.

Postjuvenile moult in Azure‐winged Magpies Cyanopica cyanea cooki was studied in free‐living and captive birds. Most natal primary (99.7%) and secondary (81.2%) feathers were retained, but only 50% of rectrices. Estimates of individual moult duration in free living birds in the following feather tracts gave: 65 days for tertials, 13 days for greater coverts, 97 days for rectrices, and 30 days for the alula. Wing coverts were the first to begin moult, followed by tertials. Alula feathers were the last to start moult. Tertial moult commenced at an average age of 97 days, regardless of hatching date. In captive birds, body moult was initiated in dorsal and upper tail coverts at the end of July and culminated in the interramal region by the end of November, with a mean moult duration of 122 days. Data on plumage growth and development are presented, and factors influencing the extent, timing and duration of moult are discussed.

A sexual dimorphism in the postjuvenile moult of the Azure‐winged Magpie Cyanopica cyanea cooki is described. Males retain significantly fewer primaries, secondaries, upper greater primary coverts ...

Although Chernobyl is perhaps the largest environmental disaster ever, there has been minimal monitoring of the status of free-living organisms or humans in stark contrast to Hiroshima and Nagasaki, where careful monitoring has continued for over 60 years. Public health issues are traditionally

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.

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.

In this paper, we present an analysis of genetic variation in three wild populations of the barn swallow, Hirundo rustica. We estimated the P, E, and G matrices for six linear morphological measurements and tested for variation among populations using the Flury hierarchical method and the jackknife followed by MANOVA method. Because of nonpositive-definite matrices, we had to employ ‘bending’ to analyse the G and E matrices with the Flury method. Both statistical methods agree in finding that the P and G matrices are significantly different but comparison between the analysis of the P matrices and pairwise analyses of the P, E, and G matrices suggests caution in interpreting the Flury results concerning differences in matrix structure. The significant variation among the populations in the G matrices appears to be due in large measure to the most geographically distant population.

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., This study was supported by grants from the Spanish Ministry of Education (S.M.), a Marie Curie grant of the European Community (A.B.), Danish Natural Sciences Research Council (A.P.M.), Spanish DGICYT project PB 95020 (F.d.L.) and Universidad de Alcala de Henares project EO31/97 (J.M., J.P. and F.R.-C.).

We measured the energy cost of ectoparasitism in an experimental study of the house martin bug Oeciacus hirundinis and its main host, nestlings of the house martin Delichon urbica. Nests were randomly assigned to inoculation with 0 (control) 10, or 100 bugs during egg laying, and this resulted in statistically significant differences in parasite loads following fledging of nestlings. Parasite loads negatively affected house martin nestlings as estimated from their body mass at age 16 days and from mass loss estimated over 1 day late in the nestling period. Daily energy expenditure (KJ/d), average daily metabolic rate (ml CO2/g h), and mass independent daily energy expenditure (kJ/mass0.67d) did not differ significantly between experimental treatments. However, average daily metabolic rate increased with increasing intensities of ectoparasitism. Mass independent daily energy expenditure also increased with higher levels of parasite infestation. These results demonstrate that the bug imposes an energy cost on its host by elevating the level of metabolism.

On a etudie le developpement du crâne du chat sauvage sur 75 echantillons (45 crânes de mâles et 30 de femelles). L'âge a ete determine en comptant les lignes de croissance du cement dentaire sur les canines. Dix-neuf mesures et indices differents ont egalement ete pris sur chacun des crânes. Ces 19 variables, avec les donnees de l'âge, furent soumises a une analyse multivariee, avec comme resultat la separation de certains des âges dans les deux sexes. On etudie et on discute de l'importance des muscles du crâne impliques dans la capture de proies et dans la mastication, en soulignant l'existence du dimorphisme sexuel et de ses consequences biologiques

Parasites are usually assumed to negatively affect fitness components of their hosts, particularly during adverse environmental conditions. We experimentally tested whether the ectoparasitic house martin bug (Oeciacus hirundinis) had larger negative effects on its house martin (Delichon urbica) host during second clutches. Conditions for reproduction decreased during the season, being less favourable for the second clutch than the first clutch. This was demonstrated by smaller clutch sizes, lower breeding success and smaller and lighter nestlings during the second clutch, as compared to the first, in the absence of parasites. Experimental manipulation of the intensity of parasitism resulted in the reproductive success of the host being reduced with increasing numbers of house martin bugs. This effect was much stronger during the second, as compared to the first, clutch. The house martin bug therefore had a greater detrimental effect on the reproduction of its host when conditions for reproduction were poor.

Sexual selection theory assumes that secondary sexual characters do not influence female reproductive effort. Female animals may invest relatively more in reproduction if they acquire mates of high phenotypic quality, because offspring sired by preferred males may be relatively more viable than offspring sired by less preferred males. Here we report for the first time in a field study that females of the monogamous barn swallow Hirundo rustica adjust their reproductive effort to the attractiveness of their mates. Experimental manipulation of male tail length, which is a trait currently subject to a directional female mating preference, affected the reproductive effort by females in single broods as well as their decision on the seasonal number of clutches. These results, and those of previous experiments, demonstrate that female barn swallows assess the quality of their mates throughout the reproductive season and adjust their reproductive decisions accordingly. This result has important implications for the theory of sexual selection and for the possibility of testing current models of female mate preferences, because the viability of offspring will be confounded by differential reproductive effort.

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.

The effects of herbivory on plant fitness have traditionally been assessed by reductions in performance measured in terms of physiologi .owth, 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 and untreated 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 inereased 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 both produce 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.

We studied the selective advantage of nest and egg crypsis in the Stone Curlew Burhinus oedicnemus. Experimental removal of camouflaging nest material resulted in replacement of a similar amount and size of nest material for the specific nest site. The match between the colour of the eggs and the ground was usually perfect and mismatches in coloration increased the predation rate by visually searching avian nest predators. Both nest building and choice of nest substrate appeared to increase crypsis and improved hatching success.

Many birds are negatively affected in terms of reproductive success by the presence of ectoparasites in their nests. We hypothesized that the negative effects of ectoparasitism on reproductive success would be a direct function of the number of clutches laid because of a seasonal increase in ectoparasite populations. Ectoparasites would negatively affect each clutch and the total effect of parasitism on annual reproductive success should therefore be relatively larger with an increasing number of clutches. This prediction was tested by fumigation of nests of swallows Hirundo rustica either following completion of the first clutch, completion of all clutches or by keeping nests as controls