There are multiple observations around the globe showing that in many avian species, both the timing of migration and breeding have advanced, due to warmer springs. Here, we review the literature to disentangle the actions of evolutionary changes in response to selection induced by climate change versus changes due to individual plasticity, that is, the capacity of an individual to adjust its phenology to environmental variables. Within the abundant literature on climate change effects on bird phenology, only a small fraction of studies are based on individual data, yet individual data are required to quantify the relative importance of plastic versus evolutionary responses. While plasticity seems common and often adaptive, no study so far has provided direct evidence for an evolutionary response of bird phenology to current climate change. This assessment leads us to notice the alarming lack of tests for microevolutionary changes in bird phenology in response to climate change, in contrast with the abundant claims on this issue. In short, at present we cannot draw reliable conclusions on the processes underlying the observed patterns of advanced phenology in birds. Rapid improvements in techniques for gathering and analysing individual data offer exciting possibilities that should encourage research activity to fill this knowledge gap.

The rate of adaptive evolution, the contribution of selection to genetic changes that increase mean fitness, is determined by the additive genetic variance in individual relative fitness. To date, there are few robust estimates of this parameter for natural populations, and it is therefore unclear whether adaptive evolution can play a meaningful role in short-term population dynamics. We developed and applied quantitative genetic methods to long-term datasets from 19 wild bird and mammal populations and found that, while estimates vary between populations, additive genetic variance in relative fitness is often substantial and, on average, twice that of previous estimates. We show that these rates of contemporary adaptive evolution can affect population dynamics and hence that natural selection has the potential to partly mitigate effects of current environmental change.

Apparent changes in breeding performance with age measured at the population level can be due to changes in individual capacity at different ages, or to the differential survival of individuals with different capabilities. Estimating the relative importance of the two is important for understanding ageing patterns in natural populations, but there are few studies of such populations in which these effects have been disentangled. We analysed laying date and clutch size as measures of individual performance in a population of mute swans (Cygnus olor) studied over 25 years at Abbotsbury, UK. On both measures of breeding performance, individuals tended to improve up to the age of 6 or 7, and to decline after about the age of 12. Individuals with longer lifespans performed better at all ages (earlier laying, larger clutches) than animals that ceased breeding earlier. We conclude that the apparent mean increase in performance with age in mute swans is due to both individual improvement and differential survival of individuals who perform well, while the decline in older age groups is due to individual loss of function. Our results underline the need to take individual differences into account when testing hypotheses about life histories in wild populations.

Significance: Many ecological and evolutionary processes strongly depend on the way natural selection varies over time. However, a gap remains when trying to connect theoretical predictions to empirical work on this question: Most theory assumes that adaptation involves tracking a moving optimum phenotype through time, but this is seldom estimated empirically. Here, we have assembled a large database of wild bird and mammal populations, to estimate patterns of fluctuations in the optimum breeding date and its influence on the variability of natural selection. We find that optimum fluctuations are prevalent. However, their influence on temporal variance in natural selection is partly buffered by tracking of the optimum phenotype through individual phenotypic plasticity.

As an extension of the classic life history theory, the recently highlighted pace-of-life syndrome hypothesis predicts the coevolution of behavioural, physiological and life-history traits. For instance, bolder and shyer individuals do not only differ in personality profiles, but also in neuro-endocrinology and breeding patterns. While theory predicts that bolder (i.e. proactive), more aggressive individuals should colonize more rapidly urbanized habitats than shyer (i.e. reactive), less aggressive individuals, it is also predicted that across generations, adaptive selection processes could favour shyer individuals that are more sensitive to novel environmental cues. Here we compared two personality traits (handling aggression, exploration score in a novel environment), one physiological trait related to stress response (breath rate) and four breeding traits (lay date, clutch size, hatching success and fledging success) in a rural and an urban study population of Mediterranean great tits Parus major. Mixed models revealed strong phenotypic divergence between forest and city in most traits explored, in particular in personality, whereby urban great tits were more reactive to stress and faster explorers compared to rural birds (yet not more aggressive). Urban birds also laid smaller broods earlier in spring compared to their rural conspecifics, and city broods resulted in lower hatching success yet interestingly fledging success was similar. Nest-box centered measures of anthropogenic (artificial light, pedestrians and cars) perturbation and resource abundance allowed us to go beyond the classical forest/city comparison by exploring the phenotypic variation across an urbanization gradient. This revealed that high urbanisation in nest-box surroundings was associated overall with earlier breeding and smaller clutches, but also with faster breath rate, although these trends showed strong annual variation. Ongoing rapid urbanization and non-random gene flow between rural and urban great tits could both contribute to the high prevalence of bold breeders in the city. Our study suggests the existence of urban and rural great tit ecotypes with different pace-of-life, but also a finer-scale divergence along the degree of urbanisation within the city. Future studies are required to determine whether this phenotypic variation at different spatiotemporal scales is adaptive and whether it has a genetic basis or results from phenotypic plasticity.