Bird populations most exposed to climate change are less sensitive to climatic variation.

The phenology of many species shows strong sensitivity to climate change; however, with few large scale intra-specific studies it is unclear how such sensitivity varies over a species' range. We document large intra-specific variation in phenological sensitivity to temperature using laying date information from 67 populations of two co-familial European songbirds, the great tit (Parus major) and blue tit (Cyanistes caeruleus), covering a large part of their breeding range. Populations inhabiting deciduous habitats showed stronger phenological sensitivity than those in evergreen and mixed habitats. However, populations with higher sensitivity tended to have experienced less rapid change in climate over the past decades, such that populations with high phenological sensitivity will not necessarily exhibit the strongest phenological advancement. Our results show that to effectively assess the impact of climate change on phenology across a species' range it will be necessary to account for intra-specific variation in phenological sensitivity, climate change exposure, and the ecological characteristics of a population.

Phenotypic plasticity drives phenological changes in a Mediterranean blue tit population.

Earlier phenology induced by climate change, such as the passerines' breeding time, is observed in many natural populations. Understanding the nature of such changes is key to predict the responses of wild populations to climate change. Genetic changes have been rarely investigated for laying date, though it has been shown to be heritable and under directional selection, suggesting that the trait could evolve. In a Corsican blue tit population, the birds' laying date has significantly advanced over 40 years, and we here determine whether this response is of plastic or evolutionary origin, by comparing the predictions of the breeder's and the Robertson-Price (STS) equations, to the observed genetic changes. We compare the results obtained for two fitness proxies (fledgling and recruitment success), using models accounting for their zero inflation. Because the trait appears heritable and under directional selection, the breeder's equation predicts that genetic changes could drive a significant part of the phenological change observed. We, however, found that fitness proxies and laying date are not genetically correlated. The STS, therefore, predicts no evolution of the breeding time, predicting correctly the absence of trend in breeding values. Our results also emphasize that when investigating selection on a plastic trait under fluctuating selection, part of the fitness-trait phenotypic covariance can be due to within individual covariance. In the case of repeated measurements, splitting within and between individual covariance can shift our perspective on the actual intensity of selection over multiple selection episodes, shedding light on the potential for the trait to evolve.

Age-dependent phenological plasticity in a wild bird.

Life-history traits are often plastic in response to environmental factors such as temperature or precipitation, and they also vary with age in many species. Trait variation during the lifetime could thus be partly driven by age-dependent plasticity in these traits. We study whether plasticity of a phenological trait-the egg-laying date-with respect to spring temperature, varies with age, and explore whether this variation relates to changes in breeding success throughout the life cycle. We use data from a four-decade long-term monitoring of a wild population of blue tits in Corsica, to estimate age-dependent plasticity of reproductive phenology and annual reproductive success. We show that both laying date plasticity and annual reproductive success vary with age: young and old females are less plastic, and fledge fewer offspring, than middle-age females. Furthermore, in contrast to young and prime-age females, in old females fledging success does not depend on laying date. Phenological plasticity is a major mechanism for coping with rapid environmental variation. Our results suggest that understanding its role in adaptation to climate change and population persistence requires integrating the age structure of the population.

Phenotypic plasticity in response to climate change: the importance of cue variation.

Phenotypic plasticity is a major mechanism of response to global change. However, current plastic responses will only remain adaptive under future conditions if informative environmental cues are still available. We briefly summarize current knowledge of the evolutionary origin and mechanistic underpinnings of environmental cues for phenotypic plasticity, before highlighting the potentially complex effects of global change on cue availability and reliability. We then illustrate some of these aspects with a case study, comparing plasticity of blue tit breeding phenology in two contrasted habitats: evergreen and deciduous forests. Using long-term datasets, we investigate the climatic factors linked to the breeding phenology of the birds and their main food source. Blue tits occupying different habitats differ extensively in the cues affecting laying date plasticity, as well as in the reliability of these cues as predictors of the putative driver of selective pressure, the date of caterpillar peak. The temporal trend for earlier laying date, detected only in the evergreen populations, is explained by increased temperature during their cue windows. Our results highlight the importance of integrating ecological mechanisms shaping variation in plasticity if we are to understand how global change will affect plasticity and its consequences for population biology. This article is part of the theme issue 'The role of plasticity in phenotypic adaptation to rapid environmental change'.

Selection on skewed characters and the paradox of stasis.

Observed phenotypic responses to selection in the wild often differ from predictions based on measurements of selection and genetic variance. An overlooked hypothesis to explain this paradox of stasis is that a skewed phenotypic distribution affects natural selection and evolution. We show through mathematical modeling that, when a trait selected for an optimum phenotype has a skewed distribution, directional selection is detected even at evolutionary equilibrium, where it causes no change in the mean phenotype. When environmental effects are skewed, Lande and Arnold's (1983) directional gradient is in the direction opposite to the skew. In contrast, skewed breeding values can displace the mean phenotype from the optimum, causing directional selection in the direction of the skew. These effects can be partitioned out using alternative selection estimates based on average derivatives of individual relative fitness, or additive genetic covariances between relative fitness and trait (Robertson-Price identity). We assess the validity of these predictions using simulations of selection estimation under moderate sample sizes. Ecologically relevant traits may commonly have skewed distributions, as we here exemplify with avian laying date - repeatedly described as more evolutionarily stable than expected - so this skewness should be accounted for when investigating evolutionary dynamics in the wild.