Summary1. The role of individual experience vs. the use of conspecific cues on breeding dispersal decisions have seldom been determined in colonial birds. We studied causes of breeding dispersal in the lesser kestrel ( Falco naumanni ), a species that breeds in colonies of variable size as well as solitarily. During a 6-year study in Spain, we gathered information on 486 subsequent breeding attempts and on 26 explanatory variables which evaluated individual experience, conspecific cues in terms of breeding performance and colony size, and different ecological and populational characteristics. 2. Two decisions were separately analysed: whether or not to disperse, and how far to move. Generalized Linear Mixed Models (GLMMs) allowed us to identify the relative contribution of each explanatory variable while controlling for the non-independence of individual dispersal decisions across years. 3. Females seemed to disperse more often than males (34% vs. 19%), and both sexes apparently dispersed less with age. However, a GLMM showed that experience (i.e. the number of years a bird bred in a particular colony) was the only factor influencing breeding dispersal. Birds showed higher site fidelity the greater their experience in a colony, which could be related to benefits derived of increased local familiarity. A second GLMM showed that, before birds acquired experience in a particular colony, individual nest failure due to predation and proximity to other colonies increased the probability of dispersal, dispersal being also higher in colonies with poor conspecific breeding success. Furthermore, solitary nesting birds were more prone to disperse and dispersal probability decreased the larger the colony of origin, according to fitness expectations associated with colony size. 4. A GLMM explaining dispersal distances retained two variables -birds dispersed farther the lower the breeding density in the surroundings, and the larger the distance to the nearest colony. Dispersing birds tended to settle within their previous foraging areas (median dispersal distance = 1·6 km), being constrained by the availability of nearby colonies. 5. Lesser kestrels mainly cue on their own breeding performance and experience in a particular colony at the time of taking a dispersal decision. However, inexperienced birds also partially cue on the size and breeding success of their own colonies ( but not on the size or breeding performance of other colonies), and birds moved larger distances when dispersing from areas of low populational density. These results support some degree of conspecific attraction.

Understanding the selective forces that shape genetic variation in natural populations remains a high priority in evolutionary biology. Genes at the major histocompatibility complex (MHC) have become excellent models for the investigation of adaptive variation and natural selection because of their crucial role in fighting off pathogens. Here we present one of the first data sets examining patterns of MHC variation in wild populations of a bird of prey, the lesser kestrel, Falco naumanni. We report extensive polymorphism at the second exon of a putatively functional MHC class II gene, Fana‐DAB*1. Overall, 103 alleles were isolated from 121 individuals sampled from Spain to Kazakhstan. Bayesian inference of diversifying selection suggests that several amino acid sites may have experienced strong positive selection (ω = 4.02 per codon). The analysis also suggests a prominent role of recombination in generating and maintaining MHC diversity (ρ = 4Nc = 0.389 per codon, θ = 0.017 per codon). Both the Fana‐DAB*1 locus and a set of eight polymorphic microsatellite markers revealed an isolation‐by‐distance pattern across the Western Palaearctic (r = 0.67; P = 0.01 and r = 0.50; P = 0.04, respectively). Nonetheless, geographical variation at the MHC contrasts with relatively uniform distributions in the frequencies of microsatellite alleles. In addition, we found lower fixation rates in the MHC than those predicted by genetic drift after controlling for neutral mitochondrial sequences. Our results therefore underscore the role of balancing selection as well as spatial variations in parasite‐mediated selection regimes in shaping MHC diversity when gene flow is limited.

Despite its key role in population dynamics and evolutionary ecology, little is known about factors shaping survival in long-lived territorial species. Here, we assessed several hypotheses that might explain variability in survival in a migratory Spanish population of a long-lived territorial species, the Egyptian vulture Neophron percnopterus, using a 16-year monitoring period and live-encounter histories of 835 individually marked birds. Cormack-Jolly-Seber captureÁrecapture models showed no evidence for effects of sex or nestling body condition on survival. However, the normalized difference vegetation index (NDVI; an indicator of primary productivity) of natal territories had positive effects on juvenile survival, indicating that environmental conditions experienced early in life can determine survival prospects. Survival increased with age (0.7390.02 in the first 2 years to 0.7890.03 in years 3 and 4) to later decrease when birds were five years old (0.6090.05), the age at which they acquire the adult plumage, abandon the communal lifestyle of juveniles, and may look for a breeding territory. At older ages, survival was higher for non-breeding (0.7590.02) and breeding adults (0.8390.02). Among the latter, birds that recruited into better territories had higher survival prospects. Age-specific variation in survival in this species may be related to behavioural changes linked to dispersal and recruitment into the breeding population, while survival prospects of adult birds strongly depend on breeding territory selection. These results suggest a tradeoff between recruiting soon, and thus reducing mortality costs of a long and extensive dispersal period, and trying to recruit into a good quality territory. Finally, annual survival rates for birds of all age classes were positively related with the NDVI in their African wintering grounds. Although this relationship was probably mediated by food availability, further research is needed to properly identify the limiting factors that are affecting trans-Saharan migrants, especially in light of global climate change.

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No abstract

Causes of natal dispersal were studied in an isolated population of the migratory, facultatively colonial Lesser Kestrel (Falco naumanni) in northeastern Spain. During a seven‐year study, we gathered information on natal dispersal of 751 individuals and on 24 explanatory variables that evaluated individual traits, conspecific cues in terms of colony size, and different ecological and populational features. We examined separately whether or not individuals changed colonies and how far they moved. Dispersal from the natal colony was high (83%), and dispersers covered a median distance of 7225 m (range 112–136 500 m). The probability of natal dispersal decreased with the size of the natal colony and with the distance to the nearest colony, but not with the distance to unoccupied buildings, in the year of recruitment. Moreover, the shorter the distance to the nearest colony, the shorter the distances that individuals dispersed, particularly to large colonies (>10 pairs). These results support the conspecific attraction hypothesis. Accordingly, the probability of recruiting in the natal colony increased with the proportion of philopatric adults, although beyond a threshold individuals were prevented from recruiting by the more dominant adults in a despotic way. The timing of arrival from the wintering grounds was positively related to the probability of colony change, especially in males. To the extent that this variable is related to the phenotypic quality of individuals, it reinforces the idea that natal dispersal is shaped by intraspecific competition in crowded colonies. Males showed lower frequencies of colony change and dispersed shorter distances than did females, as explained by the different role of the sexes in nest acquisition and defense. Our results indicate that natal dispersal is an evolutionarily labile trait simultaneously influenced by ecological, social, and individual features both within and between populations.

Avian coloniality traditionally has been investigated by examining how breeding success varies with colony size, but other crucial fitness components rarely have been examined. This may lead to wrong conclusions because unmeasured parameters may change the final fitness balance. We used multistate capture-recapture models to investigate adult survival and dispersal in relation to colony size within a long-term monitored population of lesser kestrels (Falco naumanni). Nest predation probability decreases with colony size, and adult survival is predicted to show the same trend because adults are exposed to the same suite of predators. As expected, survival probability was higher in large colonies (0.72+/-0.015; mean+/-SE) than in medium or small colonies (0.65+/-0.02). Additionally, dispersal probabilities were higher going from small to large colonies (0.20+/-0.01) than from large to small (0.08+/-0.01), as predicted by theory of habitat selection shaped by fitness maximization. These asymmetries are likely to generate size-specific colony population dynamics, so they should be taken into account in studies of colonial birds and other metapopulation-like systems. Allee effects, that is, positive density dependence, appear to be the cause of the evolution of dispersal behavior and may explain the maintenance of coloniality in this species.

Summary1. Factors governing dispersal rates have seldom been examined in spatially structured populations of vertebrates. We gathered information during 8 years on transfer rates between subpopulations in a spatially structured, growing population of colonial lesser kestrels Falco naumanni , and analysed the contribution of several variables related to spatial isolation and characteristics of both subpopulation of origin and destination on probabilities of dispersal. 2. Lesser kestrels were highly philopatric to their subpopulations, but first-breeders dispersed more often than adults (26% vs. 4%, n = 1706) because adults were reluctant to move from familiar areas. Frequency of subpopulation change was higher in females than in males (first-recruiters: 30% vs. 22%, n = 987; adults 5% vs. 1%, n = 719), according to their different breeding roles. However, all populational factors had an equal effect on individuals of different sex and dispersal status. 3. Movement rates decreased with inter-subpopulation distance -indicating that travelling to distant subpopulations may impose costs in terms of breeding prospectsand with the number of subpopulations, which increased during our study period. 4. Conspecific attraction strongly influenced the probability of dispersal: it was relatively higher in largely populated subpopulations, and individuals of large subpopulations were reluctant to change to others. These results were neither influenced by the size and breeding density of the subpopulations nor by habitat quality in terms of food availability or risk of predation, as indicated by breeding success of kestrels at each subpopulation. The number of conspecifics could be used by the kestrels as a cue of patch quality in terms of settlement options, and large subpopulations could be more easily detected by prospecting birds. 5. Our study highlights the fact that several assumptions of theoretical metapopulation modelling are often not fulfilled in nature. Both theoretical models and management strategies on spatially structured populations or metapopulations should thus consider the number, population size, and spatial distribution of local populations, as well as their relationships with the dispersal ability of the species.