Implications of migratory and exotic birds and the mosquito community on West Nile virus transmission.

BACKGROUND: Vector-borne diseases like West Nile virus (WNV) pose a global health challenge, with rising incidence and distribution. Culex mosquitoes are crucial WNV vectors. Avian species composition and bird community diversity, along with vector communities, influence WNV transmission patterns. However, limited knowledge exists on their impact in southwestern Spain, an area with active WNV circulation in wild birds, mosquitoes, and humans. METHODS: To address this, we conducted a comprehensive study investigating the contributions of migratory and exotic bird species to WNV transmission and the influence of mosquito community composition. RESULTS: Analysing 1194 serum samples from 44 avian species, we detected WNV antibodies in 32 samples from 11 species, four for the first time in Europe. Migratory birds had higher WNV exposure likelihood than native and exotic species, and higher phylogenetic diversity in bird communities correlated with lower exposure rates. Moreover, in 5859 female mosquitoes belonging to 12 species, we identified WNV competent vectors like Cx. pipiens s.l. and the Univittatus subgroup. Birds with WNV antibodies were positively associated with competent vector abundance, but negatively with overall mosquito species richness. CONCLUSIONS: These findings highlight the complex interactions between bird species, their phylogenetics, and mosquito vectors in WNV transmission. Understanding these dynamics will help to implement effective disease control strategies in southwestern Spain.

Unravelling the mosquito-haemosporidian parasite-bird host network in the southwestern Iberian Peninsula: insights into malaria infections, mosquito community and feeding preferences.

BAKGROUND: Vector-borne diseases affecting humans, wildlife and livestock have significantly increased their incidence and distribution in the last decades. Because the interaction among vectors-parasite-vertebrate hosts plays a key role driving vector-borne disease transmission, the analyses of the diversity and structure of vector-parasite networks and host-feeding preference may help to assess disease risk. Also, the study of seasonal variations in the structure and composition of vector and parasite communities may elucidate the current patterns of parasite persistence and spread as well as facilitate prediction of how climate variations may impact vector-borne disease transmission. Avian malaria and related haemosporidian parasites constitute an exceptional model to understand the ecology and evolution of vector-borne diseases. However, the characterization of vector-haemosporidian parasite-bird host assemblages is largely unknown in many regions. METHODS: Here, we analyzed 5859 female mosquitoes captured from May to November in five localities from southwestern Spain to explore the composition and seasonal variation of the vector-parasite-vertebrate host network. RESULTS: We showed a gradual increase in mosquito abundance, peaking in July. A total of 16 different haemosporidian lineages were found infecting 13 mosquito species. Of these assemblages, more than 70% of these vector-parasite associations have not been described in previous studies. Moreover, three Haemoproteus lineages were reported for the first time in this study. The prevalence of avian malaria infections in mosquitoes varied significantly across the months, reaching a maximum in November. Mosquito blood-feeding preference was higher for mammals (62.5%), whereas 37.5% of vectors fed on birds, suggesting opportunistic feeding behavior. CONCLUSION: These outcomes improve our understanding of disease transmission risk and help tovector control strategies.

The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Exploring the adjustment to parasite pressure hypothesis: differences in uropygial gland volume and haemosporidian infection in palearctic and neotropical birds.

Parasites are globally widespread pathogenic organisms, which impose important selective forces upon their hosts. Thus, in accordance with the Adjustment to parasite pressure hypothesis, it is expected that defenses among hosts vary relative to the selective pressure imposed by parasites. According to the latitudinal gradient in diversity, species richness and abundance of parasites peak near the equator. The uropygial gland is an important defensive exocrine gland against pathogens in birds. Size of the uropygial gland has been proposed to vary among species of birds because of divergent selection by pathogens on their hosts. Therefore, we should expect that bird species from the tropics should have relatively larger uropygial glands for their body size than species from higher latitudes. However, this hypothesis has not yet been explored. Here, we analyze the size of the uropygial gland of 1719 individual birds belonging to 36 bird species from 3 Neotropical (Peru) and 3 temperate areas (Spain). Relative uropygial gland volume was 12.52% larger in bird species from the tropics than from temperate areas. This finding is consistent with the relative size of this defensive organ being driven by selective pressures imposed by parasites. We also explored the potential role of this gland as a means of avoiding haemosporidian infection, showing that species with large uropygial glands for their body size tend to have lower mean prevalence of haemosporidian infection, regardless of their geographical origin. This result provides additional support for the assumption that secretions from the uropygial gland reduce the likelihood of becoming infected with haemosporidians.

Adaptive responses of animals to climate change are most likely insufficient.

Biological responses to climate change have been widely documented across taxa and regions, but it remains unclear whether species are maintaining a good match between phenotype and environment, i.e. whether observed trait changes are adaptive. Here we reviewed 10,090 abstracts and extracted data from 71 studies reported in 58 relevant publications, to assess quantitatively whether phenotypic trait changes associated with climate change are adaptive in animals. A meta-analysis focussing on birds, the taxon best represented in our dataset, suggests that global warming has not systematically affected morphological traits, but has advanced phenological traits. We demonstrate that these advances are adaptive for some species, but imperfect as evidenced by the observed consistent selection for earlier timing. Application of a theoretical model indicates that the evolutionary load imposed by incomplete adaptive responses to ongoing climate change may already be threatening the persistence of species.

Host-parasite interaction explains variation in the prevalence of avian haemosporidians at the community level.

Parasites are a selective force that shape host community structure and dynamics, but host communities can also influence parasitism. Understanding the dual nature from host-parasite interactions can be facilitated by quantifying the variation in parasite prevalence among host species and then comparing that variation to other ecological factors that are known to also shape host communities. Avian haemosporidian parasites (e.g. Plasmodium and Haemoproteus) are abundant and widespread representing an excellent model for the study of host-parasite interactions. Several geographic and environmental factors have been suggested to determine prevalence of avian haemosporidians in bird communities. However, it remains unknown whether host and parasite traits, represented by phylogenetic distances among species and degree of specialization in host-parasite relationships, can influence infection status. The aims of this study were to analyze factors affecting infection status in a bird community and to test whether the degree of parasite specialization on their hosts is determined by host traits. Our statistical analyses suggest that infection status is mainly determined by the interaction between host species and parasite lineages where tolerance and/or susceptibility to parasites plays an essential role. Additionally, we found that although some of the parasite lineages infected a low number of bird individuals, the species they infected were distantly related and therefore the parasites themselves should not be considered typical host specialists. Infection status was higher for generalist than for specialist parasites in some, but not all, host species. These results suggest that detected prevalence in a species mainly results from the interaction between host immune defences and parasite exploitation strategies wherein the result of an association between particular parasite lineages and particular host species is idiosyncratic.

Volume and antimicrobial activity of secretions of the uropygial gland are correlated with malaria infection in house sparrows.

BACKGROUND: Animals have developed a wide range of defensive mechanisms against parasites to reduce the likelihood of infection and its negative fitness costs. The uropygial gland is an exocrine gland that produces antimicrobial and antifungal secretions with properties used as a defensive barrier on skin and plumage. This secretion has been proposed to affect the interaction between avian hosts and their ectoparasites. Because uropygial secretions may constitute a defense mechanism against ectoparasites, this may result in a reduction in prevalence of blood parasites that are transmitted by ectoparasitic vectors. Furthermore, other studies pointed out that vectors could be attracted by uropygial secretions and hence increase the probability of becoming infected. Here we explored the relationship between uropygial gland size, antimicrobial activity of uropygial secretions and malaria infection in house sparrows Passer domesticus. METHODS: A nested-PCR was used to identify blood parasites infection. Flow cytometry detecting absolute cell counting assessed antimicrobial activity of the uropygial gland secretion RESULTS: Uninfected house sparrows had larger uropygial glands and higher antimicrobial activity in uropygial secretions than infected individuals. We found a positive association between uropygial gland size and scaled body mass index, but only in uninfected sparrows. Female house sparrows had larger uropygial glands and higher antimicrobial activity of gland secretions than males. CONCLUSION: These findings suggest that uropygial gland secretions may play an important role as a defensive mechanism against malaria infection.

Do malaria parasites manipulate the escape behaviour of their avian hosts? An experimental study.

Escape behaviour is the behaviour that birds and other animals display when already caught by a predator. An individual exhibiting higher intensity of such anti-predator behaviour could have greater probabilities of escape from predators. Parasites are known to affect different aspects of host behaviour to increase their own fitness. Vector-transmitted parasites such as malaria parasites should gain by manipulating their hosts to enhance the probability of transmission. Several studies have shown that malaria parasites can manipulate their vectors leading to increased transmission success. However, little is known about whether malaria parasites can manipulate escape behaviour of their avian hosts thereby increasing the spread of the parasite. Here we used an experimental approach to explore if Plasmodium relictum can manipulate the escape behaviour of one of its most common avian hosts, the house sparrow Passer domesticus. We experimentally tested whether malaria parasites manipulate the escape behaviour of their avian host. We showed a decrease in the intensity of biting and tonic immobility after removal of infection with anti-malaria medication compared to pre-experimental behaviour. These outcomes suggest that infected sparrows performed more intense escape behaviour, which would increase the likelihood of individuals escaping from predators, but also benefit the parasite by increasing its transmission opportunities.

Detecting transmission areas of malaria parasites in a migratory bird species.

The identification of the regions where vector-borne diseases are transmitted is essential to study transmission patterns and to recognize future changes in environmental conditions that may potentially influence the transmission areas. SGS1, one of the lineages of Plasmodium relictum, is known to have active transmission in tropical Africa and temperate regions of Europe. Nuclear sequence data from isolates infected with SGS1 (based on merozoite surface protein 1 (MSP1) allelic diversity) have provided new insights on the distribution and transmission areas of these allelic variants. For example, MSP1 alleles transmitted in Africa differ from those transmitted in Europe, suggesting the existence of two populations of SGS1 lineages. However, no study has analysed the distribution of African and European transmitted alleles in Afro-Palearctic migratory birds. With this aim, we used a highly variable molecular marker to investigate whether juvenile house martins become infected in Europe before their first migration to Africa. We explored the MSP1 allelic diversity of P. relictum in adult and juvenile house martins. We found that juveniles were infected with SGS1 during their first weeks of life, confirming active transmission of SGS1 to house martins in Europe. Moreover, we found that all the juveniles and most of adults were infected with one European transmitted MSP1 allele, whereas two adult birds were infected with two African transmitted MSP1 alleles. These findings suggest that house martins are exposed to different strains of P. relictum in their winter and breeding quarters.

Assessing multivariate constraints to evolution across ten long-term avian studies.

BACKGROUND: In a rapidly changing world, it is of fundamental importance to understand processes constraining or facilitating adaptation through microevolution. As different traits of an organism covary, genetic correlations are expected to affect evolutionary trajectories. However, only limited empirical data are available. METHODOLOGY/PRINCIPAL FINDINGS: We investigate the extent to which multivariate constraints affect the rate of adaptation, focusing on four morphological traits often shown to harbour large amounts of genetic variance and considered to be subject to limited evolutionary constraints. Our data set includes unique long-term data for seven bird species and a total of 10 populations. We estimate population-specific matrices of genetic correlations and multivariate selection coefficients to predict evolutionary responses to selection. Using Bayesian methods that facilitate the propagation of errors in estimates, we compare (1) the rate of adaptation based on predicted response to selection when including genetic correlations with predictions from models where these genetic correlations were set to zero and (2) the multivariate evolvability in the direction of current selection to the average evolvability in random directions of the phenotypic space. We show that genetic correlations on average decrease the predicted rate of adaptation by 28%. Multivariate evolvability in the direction of current selection was systematically lower than average evolvability in random directions of space. These significant reductions in the rate of adaptation and reduced evolvability were due to a general nonalignment of selection and genetic variance, notably orthogonality of directional selection with the size axis along which most (60%) of the genetic variance is found. CONCLUSIONS: These results suggest that genetic correlations can impose significant constraints on the evolution of avian morphology in wild populations. This could have important impacts on evolutionary dynamics and hence population persistence in the face of rapid environmental change.

Assessing the effects of climate on host-parasite interactions: a comparative study of European birds and their parasites.

BACKGROUND: Climate change potentially has important effects on distribution, abundance, transmission and virulence of parasites in wild populations of animals. METHODOLOGY/PRINCIPAL FINDING: Here we analyzed paired information on 89 parasite populations for 24 species of bird hosts some years ago and again in 2010 with an average interval of 10 years. The parasite taxa included protozoa, feather parasites, diptera, ticks, mites and fleas. We investigated whether change in abundance and prevalence of parasites was related to change in body condition, reproduction and population size of hosts. We conducted analyses based on the entire dataset, but also on a restricted dataset with intervals between study years being 5-15 years. Parasite abundance increased over time when restricting the analyses to datasets with an interval of 5-15 years, with no significant effect of changes in temperature at the time of breeding among study sites. Changes in host body condition and clutch size were related to change in temperature between first and second study year. In addition, changes in clutch size, brood size and body condition of hosts were correlated with change in abundance of parasites. Finally, changes in population size of hosts were not significantly related to changes in abundance of parasites or their prevalence. CONCLUSIONS/SIGNIFICANCE: Climate change is associated with a general increase in parasite abundance. Variation in laying date depended on locality and was associated with latitude while body condition of hosts was associated with a change in temperature. Because clutch size, brood size and body condition were associated with change in parasitism, these results suggest that parasites, perhaps mediated through the indirect effects of temperature, may affect fecundity and condition of their hosts. The conclusions were particularly in accordance with predictions when the restricted dataset with intervals of 5-15 years was used, suggesting that short intervals may bias findings.

Malaria infection and feather growth rate predict reproductive success in house martins.

Carry-over effects take place when events occurring in one season influence individual performance in a subsequent season. Blood parasites (e.g. Plasmodium and Haemoproteus) have strong negative effects on the body condition of their hosts and could slow the rate of feather growth on the wintering grounds. In turn, these winter moult costs could reduce reproductive success in the following breeding season. In house martins Delichon urbica captured and studied at a breeding site in Europe, we used ptilochronology to measure growth rate of tail feathers moulted on the winter range in Africa, and assessed infection status of blood parasites transmitted on the wintering grounds. We found a negative association between haemosporidian parasite infection status and inferred growth rate of tail feathers. A low feather growth rate and blood parasite infections were related to a delay in laying date in their European breeding quarters. In addition, clutch size and the number of fledglings were negatively related to a delayed laying date and blood parasite infection. These results stress the importance of blood parasites and feather growth rate as potentially mechanisms driving carry-over effects to explain fitness differences in wild populations of migratory birds.

Diversity, loss, and gain of malaria parasites in a globally invasive bird.

Invasive species can displace natives, and thus identifying the traits that make aliens successful is crucial for predicting and preventing biodiversity loss. Pathogens may play an important role in the invasive process, facilitating colonization of their hosts in new continents and islands. According to the Novel Weapon Hypothesis, colonizers may out-compete local native species by bringing with them novel pathogens to which native species are not adapted. In contrast, the Enemy Release Hypothesis suggests that flourishing colonizers are successful because they have left their pathogens behind. To assess the role of avian malaria and related haemosporidian parasites in the global spread of a common invasive bird, we examined the prevalence and genetic diversity of haemosporidian parasites (order Haemosporida, genera Plasmodium and Haemoproteus) infecting house sparrows (Passer domesticus). We sampled house sparrows (N = 1820) from 58 locations on 6 continents. All the samples were tested using PCR-based methods; blood films from the PCR-positive birds were examined microscopically to identify parasite species. The results show that haemosporidian parasites in the house sparrows' native range are replaced by species from local host-generalist parasite fauna in the alien environments of North and South America. Furthermore, sparrows in colonized regions displayed a lower diversity and prevalence of parasite infections. Because the house sparrow lost its native parasites when colonizing the American continents, the release from these natural enemies may have facilitated its invasion in the last two centuries. Our findings therefore reject the Novel Weapon Hypothesis and are concordant with the Enemy Release Hypothesis.

Individual responses in spring arrival date to ecological conditions during winter and migration in a migratory bird.

1. We studied lifetime arrival patterns in the barn swallow (Hirundo rustica L.) in relation to variation in ecological conditions, as reflected by the normalized difference vegetation index (NDVI) in the Sub-Saharan winter quarters and at stopover sites in North Africa. 2. Migratory birds have recently advanced their arrival dates, but the relative role of microevolution and phenotypic plasticity as mechanisms of response to changing environmental conditions remains unknown. To distinguish between these two possibilities, we investigated the change in the arrival date using cross-sectional and longitudinal analyses. 3. We predicted that the effect (i.e. slopes) of environmental conditions in stopover or winter areas on arrival date should be similar using cross-sectional and longitudinal analyses in case phenotypic plasticity is the underlying mechanism, or they should differ in case microevolution is the mechanism. 4. As expected according to a previous cross-sectional study, we found an advance in the arrival date when ecological conditions improve in stopover areas and a delay in the arrival date when ecological conditions improve in the winter quarters. 5. Change in the arrival time at the breeding grounds due to ecological conditions found en route and, in the winter areas, was mainly due to phenotypic plasticity as shown by similarities in the slopes found in these relationships using cross-sectional and longitudinal analyses. 6. We also investigated sex and age of barns swallows as sources of variation in the arrival time with respect to conditions experienced in winter and stopover areas. We found that earlier arrival at the breeding grounds due to prevailing ecological conditions found en route in North Africa was similar for males and females of all age-classes. In contrast, individuals tended to delay departure when ecological conditions improved in the winter quarters, but this delay differed among age classes, with old individuals delaying departure more than middle-aged and yearling birds. 7. The migratory response of individuals to changing climatic conditions experienced during different parts of their life provides evidence for individuals responding differently to prevailing conditions in the winter quarters depending on their age, but not to conditions experienced en route during spring migration.

Divergent patterns of impact of environmental conditions on life history traits in two populations of a long-distance migratory bird.

Some areas have experienced recent dramatic warming due to climate change, while others have shown no change at all, or even recent cooling. We predicted that patterns of selection on life history would differ between southern and northern European populations of a long-distance migratory bird, the barn swallow Hirundo rustica, because global patterns of weather as reflected by large-scale weather phenomena such as the North Atlantic Oscillation (NAO) and the El Niño-Southern Oscillation (ENSO) have different effects on environmental conditions in different parts of the world frequented during the annual cycle. We investigated relationships between mean arrival date, dispersal rate and yearling survival rate among years, using two long-term population studies in Spain and Denmark. We found evidence of a difference in the effects of normalized difference vegetation index in North and West Africa on mean arrival date of male barn swallows, with the effect differing significantly between populations. Second, there was a significant interaction between ENSO and population on dispersal rate, showing that conditions in Africa during winter differentially affected dispersal in the two populations. Finally, the NAO index in winter had an effect on yearling survival that differed between populations. These findings highlight the divergent patterns of response to climate change among populations, and they suggest that climate change can differentially affect important life history traits with potential implications for maintenance of viable populations and gene flow among populations.

Age-related change in breeding performance in early life is associated with an increase in competence in the migratory barn swallow Hirundo rustica.

1. We investigated age-related changes in two reproductive traits (laying date and annual fecundity) in barn swallows Hirundo rustica L. using a mixed model approach to di-stinguish among between- and within-individual changes in breeding performance with age. 2. We tested predictions of age-related improvements of competence (i.e. constraint hypothesis) and age-related progressive disappearance of poor-quality breeders (i.e. selection hypothesis) to explain age-related increase in breeding performance in early life. 3. Reproductive success increased in early life, reaching a plateau at middle age (e.g. at 3 years of age) and decreasing at older age (> 4 years). Age-related changes in breeding success were due mainly to an effect of female age. 4. Age of both female and male affected timing of reproduction. Final linear mixed effect models (LME) for laying date included main and quadratic terms for female and male age, suggesting a deterioration in reproductive performance at older age for both males and females. 5. We found evidence supporting the constraints hypothesis that increases in competence within individuals, with ageing being the most probable cause of the observed increase in breeding performance with age in early life. Two mechanisms were implicated: (1) advance in male arrival date with age provided middle-aged males with better access to mates. Yearling males arrived later to the breeding grounds and therefore had limited access to high-quality mates. (2) Breeding pairs maintaining bonds for 2 consecutive years (experienced pairs) had higher fecundity than newly formed inexperienced breeding pairs. 6. There was no support for the selection hypothesis because breeding performance was not correlated with life span. 7. We found a within-individual deterioration in breeding and migratory performance (arrival date) in the oldest age-classes consistent with senescence in these reproductive and migratory traits.

Malarial parasites decrease reproductive success: an experimental study in a passerine bird.

Malarial parasites are supposed to have strong negative fitness consequences for their hosts, but relatively little evidence supports this claim due to the difficulty of experimentally testing this. We experimentally reduced levels of infection with the blood parasite Haemoproteus prognei in its host the house martin Delichon urbica, by randomly treating adults with primaquine or a control treatment. Treated birds had significantly fewer parasites than controls. The primaquine treatment increased clutch size by 18%; hatching was 39% higher and fledging 42% higher. There were no effects of treatment on quality of offspring, measured in terms of tarsus length, body mass, haematocrit or T-cell-mediated immune response. These findings demonstrate that malarial parasites can have dramatic effects on clutch size and other demographic variables, potentially influencing the evolution of clutch size, but also the population dynamics of heavily infected populations of birds.

Reproduction and migration in relation to senescence in the barn swallow Hirundo rustica: A study of avian 'centenarians'.

A. P. Moller, F. de Lope and N. Saino: Reproduction and migration in relation to senescence in the barn swallow Hirundo rustica: A study of avian 'centenarians'. Senescence reflects the decrease in age-dependent residual reproductive value, and a previous study of a cohort of migratory barn swallows (Hirundo rustica) that had reached advanced ages for this species (at least five years old) provided evidence consistent with senescence such as reduced reproductive performance and increased abundance of parasites. We studied a population of migratory barn swallows over a number of years. Several questions were asked including 1) how do longer lived birds compare to shorter lived individuals of the same species, 2) is there a difference in individuals of the same species that live in different geographical regions, and 3) how do patterns of reproduction change during aging? We compared the phenotypes of 87 individuals from three populations in Denmark, Italy and Spain that had reached at least five years old with those of control individuals matched for sex, arrival date and breeding site. These controls only became one year old, i.e., they were not found thereafter and both groups of individuals were compared in their first year of life. Results showed that there were no significant differences in any of 11 morphological characters or in the abundance of three ectoparasites between the two groups of birds. Females that eventually became very old had relatively smaller first clutches as one year olds than did one year old controls, while males that eventually became very old had mates that laid clutches that were relatively larger than those of controls. Annual fecundity showed a similar pattern, with an increased sex difference in fecundity between birds from Spain than in Italy and in Denmark. Danish barn swallows that achieved older ages had relatively lower haematocrits than control individuals of the same sex, age and arrival date captured at the same site, while the difference in haematocrits between these two categories of individuals was small for Italian birds. Spring arrival date was relatively earlier in Danish males that eventually became very old compared to control males of the same age and tail length (a correlate of fitness) that arrived at the same site, while the two age categories of females did not differ in arrival date. In the Spanish population there was no difference in arrival date between the birds that aged most successfully and controls for either sex. We will discuss these findings in relation to current senescence theories.

Vane emargination of outer tail feathers improves flight manoeuvrability in streamerless hirundines, Hirundinidae.

Recent studies have suggested that the proximal part of the swallow (Hirundo rustica) tail streamer appears to aid turning flight, as expected if streamers evolved initially purely through natural selection for enhanced manoeuvrability. However, the evolution of slender aerodynamically advantageous streamers is also predicted by an alternative hypothesis, which suggests that such a trait could develop primarily to ameliorate the aerodynamic cost of a long size-dimorphic tail. To distinguish between these hypotheses, we have investigated for the effect on manoeuvrability of trimming the tips of the outer tail feathers into short streamers, without lengthening these feathers, in two streamerless hirundine species--the house martin (Delichon urbica) and the sand martin (Riparia riparia). This allowed us to examine the aerodynamic costs and benefits of streamers at an early evolutionary stage that predates elongation of the outermost tail feathers through female choice. We showed that such initial streamers enhance manoeuvrability in streamerless hirundines, confirming the findings of recent studies. However, in contrast to these studies, we showed that improved manoeuvrability resulting from streamers could arise before the outermost tail feathers have become elongated (e.g. owing to female choice). The occurrence of such an aerodynamic advantage depends on the ancestral shape of a forked tail. This provides support for the hypothesis that streamers, like those in the barn swallow, might evolve initially purely through natural selection for enhanced manoeuvrability.

The influence of environmental conditions on immune responses, morphology and recapture probability of nestling house martins (Delichon urbica).

Nestling birds produced later in the season are hypothesized to be of poor quality with a low probability of survival and recruitment. In a Spanish population of house martins (Delichon urbica), we first compared reproductive success, immune responses and morphological traits between the first and the second broods. Second, we investigated the effects of an ectoparasite treatment and breeding date on the recapture rate the following year. Due probably to a reverse situation in weather conditions during the experiment, with more rain during rearing of the first brood, nestlings reared during the second brood were in better condition and had stronger immune responses compared with nestlings from the first brood. Contrary to other findings on house martins, we found a similar recapture rate for chicks reared during the first and the second brood. Furthermore, ectoparasitic house martin bugs had no significant effect on the recapture rate. Recaptured birds had similar morphology but higher immunoglobulin levels when nestlings compared with non-recaptured birds. This result implies that a measure of immune function is a better predictor of survival than body condition per se.