Javier Balbontín

Effects of malaria double infection in birds: one plus one is not two

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.

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.

Variations in the age of mates as an early warning signal of changes in population trends? The case of Bonelli's eagle in Andalusia

Methods to evaluate population trends have recently received particular attention because of the decreasing patterns shown by several species since the nineteenth century. We have studied demographic traits of the Bonellis eagle (Hieraaetus fasciatus) population of Andalusia (southern Spain) for 20 years (1980–2000). This species is endangered and is suffering a rapid decline in most of its distribution range. Although our study population has remained stable for the last 10 years, both age at first breeding and productivity have decreased during the last 5 years. We propose that age at first breeding could be used as a potential early warning signal to detect possible future changes in population trends of long-lived species with deferred maturation age. In particular, in this species spatial variation in demographic traits could help to detect whether unnatural events such as shooting are decreasing age at first breeding and productivity. If occupation of territories by non-adults is caused by human persecution, conservation efforts in the particular case of Bonellis eagle have to be directed to protect those breeding territories where an increase in the percentage of pairs containing at least one non-adult individual would be detected.

The proportion of immature breeders as a reliable early warning signal of population decline: evidence from the Spanish imperial eagle in Doñana.

Abstract Methods to evaluate population trends have recently received particular attention because of perceived declines in several species during the 20th century. We investigated whether age at first breeding could be used as an “early warning signal” to detect possible changes in population trends in long-lived species with deferred maturity using data from the Spanish imperial eagle (Aquila adalberti) population in Donana National Park (Spain). This bird of prey is an endangered species that has suffered a rapid decline in this population during the last 10 years. As a result of our 27-year monitoring (1976–2002) study, we detected that an increase in immature breeding birds occurred before population decline became evident. The proportion of immature-plumaged breeders in the population was significantly higher during the period of decline than during the period of stability. In our case, more than 10% of immature breeders can be considered as an “early warning signal” that anticipates population decline. Owing to the ignorance of this warning signal, urgent actions for the recovery of this eagle population started 10 years later than necessary, and when population size had been reduced.

Quantitative genetics of migration syndromes: a study of two barn swallow populations

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.

Senescent sperm performance in old male birds.

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.

From local monitoring to a broad-scale viability assessment: a case study for the Bonelli's Eagle in western Europe

Population viability analysis (PVA) has become a basic tool of current conservation practice. However, if not accounted for properly, the uncertainties inherent to PVA predictions can decrease the reliability of this type of analysis. In the present study, we performed a PVA of the whole western European population (France, Portugal, and Spain) of the endangered Bonellis Eagle (Aquila fasciata), in which we thoroughly explored the consequences of uncertainty in population processes and parameters on PVA predictions. First, we estimated key vital rates (survival, fertility, recruitment, and dispersal rates) using monitoring, ringing, and bibliographic data from the period 1990-2009 from 12 populations found throughout the studied geographic range. Second, we evaluated the uncertainty about model structure (i.e., the assumed processes that govern individual fates and population dynamics) by comparing the observed growth rates of the studied populations with model predictions for the same period. Third, using the model structures suggested in the previous step, we assessed the viability of both the local populations and the overall population. Finally, we analyzed the effects of model and parameter uncertainty on PVA predictions. Our results strongly support the idea that all local populations in western Europe belong to a single, spatially structured population operating as a source- sink system, whereby the populations in the south of the Iberian Peninsula act as sources and, thanks to dispersal, sustain all other local populations, which would otherwise decline. Predictions regarding population dynamics varied considerably, and models assuming more constrained dispersal predicted more pessimistic population trends than models assuming greater dispersal. Model predictions accounting for parameter uncertainty revealed a marked increase in the risk of population declines over the next 50 years. Sensitivity analyses indicated that adult and pre-adult survival are the chief vital rates regulating these populations, and thus, the conservation efforts aimed at improving these survival rates should be strengthened in order to guarantee the long-term viability of the European populations of this endangered species. Overall, the study provides a framework for the implementation of multi-site PVAs and highlights the importance of dispersal processes in shaping the population dynamics of long-lived birds distributed across heterogeneous landscapes.

Plasma chemistry in booted eagle (Hieraaetus pennatus) during breeding season.

Most studies that have examined raptor plasma chemistry have been conducted on birds living in captivity. In this study, we describe typical plasma chemistry values indicators of body condition in free-living Booted Eagles, Hieraaetus pennatus, from Doñana National Park (Spain). Values are compared with those of other raptors. Mean concentrations of creatinine, uric acid and urea were lower in adults than in nestlings, while glucose, DAT and AAT were lower in nestlings than in adults. Interactions of age/sex affected plasma mean levels of creatine kinase, glucose, AAT, uric acid and urea. Adult females showed significantly lower levels of creatine kinase, uric acid and urea than adult males and nestlings. Adult males had significantly higher levels of AAT than the other groups. The lowest levels of glucose and the highest levels of uric acid were found in nestling females. We think the differences in blood parameters can be explained by differences in size of species, of individuals (because of both body condition and sexual dimorphism) and diet.

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.

Environmental conditions during early life accelerate the rate of senescence in a short-lived passerine bird

Environmental conditions experienced in early life may shape subsequent phenotypic traits including life history. We investigated how predation risk caused by domestic cats (Felis silvestris catus) and local breeding density affected patterns of reproductive and survival senescence in Barn Swallows (Hirundo rustica) breeding semicolonially in Denmark. We recorded the abundance of cats and the number of breeding pairs at 39 breeding sites during 24 years and related these to age-specific survival rate and reproductive senescence to test predictions of the life history theory of senescence. We found evidence for actuarial senescence for the first time in this species. Survival rate increased until reaching a plateau in midlife and then decreased later. We also found that survival rate was higher for males than females. Local breeding density or predation risk did not affect survival as predicted by theory. Barn Swallows with short lives did not invest more in reproduction in early life, inconsistent with expectations for trade-offs between reproduction and survival as theory suggests. However, we found that the rate of reproductive decline during senescence was steeper for individuals exposed to intense competition, and predation pressure accelerated the rate of reproductive senescence, but only in sites with many breeding pairs. These latter results are in accordance with one of the predictions suggested by the life history theory of aging. These results emphasize the importance of considering intraspecific competition and interspecific interactions such as predation when analyzing reproductive and actuarial senescence.