Events in one part of the annual cycle often affect the performance (and subsequently fitness) of individuals later in the season (carry‐over effects). An important aspect of this relates to the timing of activities. For example, many studies on migratory birds have shown that relatively late‐spring arrival in the breeding area reduces both the likelihood of getting a mate or territory and reproductive success. In contrast, relatively little is known about the movements of individuals in non‐migratory populations during the non‐breeding season. Few studies have investigated the timing of arrival at the breeding area in such species, possibly due to the assumption that most individuals remain in the area during the non‐breeding season. In this study, we used 4 years of data from a transponder‐based automated recording system set up in a non‐migratory population of blue tits Cyanistes caeruleus to describe individual variation in arrival at the breeding site. We investigated whether this variation can be explained by individual characteristics (sex, body size or status), and we assessed its effect on aspects of reproductive success in the subsequent breeding season. We found substantial variation in arrival date and demonstrate that this trait is individual‐specific (repeatable). Females arrived later than males, but the arrival dates of social pair members were more similar than expected by chance, which suggests that individuals may mate assortatively depending on their arrival in the breeding area. Arrival predicted both whether an individual would end up breeding that season and several aspects of its breeding success. Our study suggests that individuals of non‐migratory species leave the breeding area during the non‐breeding season. Hence, it may be useful to consider variation in the scale of movements between breeding and non‐breeding sites, rather than using a simple dichotomy between ‘resident’ and ‘migratory’ species. We conclude that the timing of pre‐breeding events, in particular arrival date, may be an overlooked, but important, fitness‐relevant trait in non‐migratory species.
The evolutionary consequences of individual genetic diversity are frequently studied by assessing heterozygosity–fitness correlations (HFCs). The prevalence of positive and negative HFCs and the predominance of general versus local effects in wild populations are far from understood, partly because comprehensive studies testing for both inbreeding and outbreeding depression are lacking. We studied a genetically diverse population of blue tits in southern Germany using a genome‐wide set of 87 microsatellites to investigate the relationship between proxies of reproductive success and measures of multilocus and single‐locus individual heterozygosity (MLH and SLH). We used complimentary measures of MLH and partitioned markers into functional categories according to their position in the blue tit genome. HFCs based on MLH were consistently negative for functional loci, whereas correlations were rather inconsistent for loci found in nonfunctional areas of the genome. Clutch size was the only reproductive variable showing a general effect. We found evidence for local effects for three measures of reproductive success: arrival date at the breeding site, the probability of breeding at the study site and male reproductive success. For these, we observed consistent, and relatively strong, negative effects at one functional locus. Remarkably, this marker had a similar effect in another blue tit population from Austria (~400 km to the east). We suggest that a genetic local effect on timing of arrival might be responsible for most negative HFCs detected, with carry‐over effects on other reproductive traits. This effect could reflect individual differences in the distance between overwintering areas and breeding sites.
Understanding the genomic landscape of adaptation is central to understanding microevolution in wild populations. Genomic targets of selection and the underlying genomic mechanisms of adaptation can be elucidated by genome‐wide scans for past selective sweeps or by scans for direct fitness associations. We sequenced and assembled 150 haplotypes of 75 blue tits (Cyanistes caeruleus) of a single Central European population by a linked‐read technology. We used these genome data in combination with coalescent simulations (i) to estimate an historical effective population size of ~250,000, which recently declined to ~10,000, and (ii) to identify genome‐wide distributed selective sweeps of beneficial variants probably originating from standing genetic variation (soft sweeps). The genes linked to these soft sweeps, but also those linked to hard sweeps based on new beneficial mutants, showed a significant enrichment for functions associated with gene expression and transcription regulation. This emphasizes the importance of regulatory evolution in the population’s adaptive history. Soft sweeps were further enriched for genes related to axon and synapse development, indicating the significance of neuronal connectivity changes in the brain potentially linked to behavioural adaptations. A previous scan of heterozygosity–fitness correlations revealed a consistent negative effect on arrival date at the breeding site for a single microsatellite in the MDGA2 gene. Here, we used the haplotype structure around this microsatellite to explain the effect as a local and direct outbreeding effect of a gene involved in synapse development.
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