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Lay Summary The pace-of-life syndrome hypothesis predicts that environmental differences can trigger the evolution of different life-history strategies, including differences in personality. A 5-year study reveals that blue tits breeding in different habitats differ in personality and in life-history characteristics, such as adult survival probability. This is a rare demonstration of personality differences across a small geographical scale, suggesting behavior and life-history traits have coevolved in a local adaptation process., Environmental heterogeneity can result in spatial variation in selection pressures that can produce local adaptations. The pace-of-life syndrome hypothesis predicts that habitat-specific selective pressures will favor the coevolution of personality, physiological, and life-history phenotypes. Few studies so far have compared these traits simultaneously across different ecological conditions. In this study, we compared 3 personality traits (handling aggression, exploration speed in a novel environment, and nest defense behavior) and 1 physiological trait (heart rate during manual restraint) across 3 Corsican blue tit (Cyanistes caeruleus) populations. These populations are located in contrasting habitats (evergreen vs. deciduous) and are situated in 2 different valleys 25 km apart. Birds from these populations are known to differ in life-history characteristics, with birds from the evergreen habitat displaying a slow pace-of-life, and birds from the deciduous habitat a comparatively faster pace-of-life. We expected personality to differ across populations, in line with the differences in pace-of-life documented for life-history traits. As expected, we found behavioral differences among populations. Despite considerable temporal variation, birds exhibited lower handling aggression in the evergreen populations. Exploration speed and male heart rate also differed across populations, although our results for exploration speed were more consistent with a phenotypic difference between the 2 valleys than between habitats. There were no clear differences in nest defense intensity among populations. Our study emphasizes the role of environmental heterogeneity in shaping population divergence in personality traits at a small spatial scale.

The pace-of-life syndrome (i.e., POLS) hypothesis posits that behavioral and physiological traits mediate the trade-off between current and future reproduction. This hypothesis predicts that life history, behavioral, and physiological traits will covary under clearly defined conditions. Empirical tests are equivocal and suggest that the conditions necessary for the POLS to emerge are not always met. We nuance and expand the POLS hypothesis to consider alternative relationships among behavior, physiology, and life history. These relationships will vary with the nature of predation risk, the challenges posed by resource acquisition, and the energy management strategies of organisms. We also discuss how the plastic response of behavior, physiology, and life history to changes in ecological conditions and variation in resource acquisition among individuals determine our ability to detect a fast-slow pace of life in the first place or associations among these traits. Future empirical studies will provide most insights on the coevolution among behavior, physiology, and life history by investigating these traits both at the genetic and phenotypic levels in varying types of predation regimes and levels of resource abundance. We revisit the pace-of-life syndrome hypothesis, suggesting that behaviors involving a risk of death or injury should coevolve with higher metabolic rates, higher fecundity, faster growth, and heightened mortality rates. Empirical support for this hypothesis is mixed. We show how relaxing some of the assumptions underlying the pace-of-life syndrome hypothesis allows us to consider alternative relationships among behavior, physiology, and life history, and why we fail to meet the predictions posed by the pace-of-life syndrome hypothesis in some populations. Our discussion emphasizes the need to re-integrate the role of the species’ natural history, ecological conditions, and phenotypic plasticity in shaping relationships among behavior, physiology, and life history.

While the study of the origins of biological diversity across species has provided numerous examples of adaptive divergence, the realization that it can occur at microgeographic scales despite gene flow is recent, and scarcely illustrated. We review here evidence suggesting that the striking phenotypic differentiation in ecologically relevant traits exhibited by blue tits Cyanistes caeruleus in their southern range‐edge putatively reflects adaptation to the heterogeneity of the Mediterranean habitats. We first summarize the phenotypic divergence for a series of life history, morphological, behavioural, acoustic and colour ornament traits in blue tit populations of evergreen and deciduous forests. For each divergent trait, we review the evidence obtained from common garden experiments regarding a possible genetic origin of the observed phenotypic differentiation as well as evidence for heterogeneous selection. Second, we argue that most phenotypically differentiated traits display heritable variation, a fundamental requirement for evolution to occur. Third, we discuss nonrandom dispersal, selective barriers and assortative mating as processes that could reinforce local adaptation. Finally, we show how population genomics supports isolation – by – environment across landscapes. Overall, the combination of approaches converges to the conclusion that the strong phenotypic differentiation observed in Mediterranean blue tits is a fascinating case of local adaptation.

Longitudinal multi-plot studies of nest architecture in non-domesticated birds are rare. Here we report the results of an 18-year multi-plot study that compared heights of fresh female-built nests in Corsican Great Tits ( Parus major) and Blue Tits ( Cyanistes caeruleus ogliastrae). Boxes with relatively small nest-chambers were erected in a landscape with habitat patches that were either dominated by food-rich broad-leaved deciduous oaks ( Quercus humilis) or food-poor evergreen oaks ( Quercus ilex). Nest height tended to change with oak habitat type in a species-specific manner. Great Tits built shallower nests than Blue Tits supporting the published findings from other geographic regions (Poland, UK, southern mainland France). Potential impacts of local climate factors and risks related to nest predation on the species-specific expression of the size of nests are discussed.

Understanding the causes and consequences of population phenotypic divergence is a central goal in ecology and evolution. Phenotypic divergence among populations can result from genetic divergence, phenotypic plasticity or a combination of the two. However, few studies have deciphered these mechanisms for populations geographically close and connected by gene flow, especially in the case of personality traits. In this study, we used a common garden experiment to explore the genetic basis of the phenotypic divergence observed between two blue tit (Cyanistes caeruleus) populations inhabiting contrasting habitats separated by 25 km, for two personality traits (exploration speed and handling aggression), one physiological trait (heart rate during restraint) and two morphological traits (tarsus length and body mass). Blue tit nestlings were removed from their population and raised in a common garden for up to five years. We then compared adult phenotypes between the two populations, as well as trait‐specific Qst and Fst. Our results revealed differences between populations similar to those found in the wild, suggesting a genetic divergence for all traits. Qst ‐ Fst comparisons revealed that the traits divergences likely result from dissimilar selection patterns rather than from genetic drift. Our study is one of the first to report a Qst ‐ Fst comparison for personality traits and adds to the growing body of evidence that population genetic divergence is possible at a small scale for a variety of traits including behavioural traits.

Standardized long-term multi-plot investigations of variation in nest size in free-ranging model species are rare, despite their value for understanding how the environment influences plastic traits such as nest size. Here, we report the results of an 18-yr descriptive study of nest size in first clutches produced by secondary-cavity nesting great tits (Parus major) and blue tits (Cyanistes caeruleus) in the Mediterranean, whilst also taking individual (lay-date, clutch size, age, species) and environmental characteristics (e.g. weather, oak habitat, region) into account. Nests of both species were built in relatively small standardized boxes erected in habitat patches that differed in the presence of the dominant oak species which was either summergreen deciduous or evergreen sclerophyllous. Nest size showed strong differences between species, and also in relation to environmental factors. Great tits built smaller nests than blue tits, Corsican birds had larger nests than mainland ones, nests tended to be larger in evergreen oak-habitat, and larger nests were built during drier weather. None of the individual-specific traits most often considered in long-term studies (lay-date, clutch size, and female age) were related to nest size in either species. Experimental approaches will be required to identify the underlying mechanisms that caused the observed phenotypic diversity for nest size in our study system.

The study of the spatial distribution of relatives in a population under contrasted environmental conditions provides critical insights into the flexibility of dispersal behaviour and the role of environmental conditions in shaping population relatedness and social structure. Yet few studies have evaluated the effects of fluctuating environmental conditions on relatedness structure of solitary species in the wild. The aim of this study was to determine the impact of interannual variations in environmental conditions on the spatial distribution of relatives [spatial genetic structure (SGS)] and dispersal patterns of a wild population of eastern chipmunks (Tamias striatus), a solitary rodent of North America. Eastern chipmunks depend on the seed of masting trees for reproduction and survival. Here, we combined the analysis of the SGS of adults with direct estimates of juvenile dispersal distance during six contrasted years with different dispersal seasons, population sizes and seed production. We found that environmental conditions influences the dispersal distances of juveniles and that male juveniles dispersed farther than females. The extent of the SGS of adult females varied between years and matched the variation in environmental conditions. In contrast, the SGS of males did not vary between years. We also found a difference in SGS between males and females that was consistent with male-biased dispersal. This study suggests that both the dispersal behaviour and the relatedness structure in a population of a solitary species can be relatively labile and change according to environmental conditions.

Summary Understanding the causes of animal personality (i.e. consistent behavioural differences) is a major aim of evolutionary studies. Recent theoretical work suggests that major personality traits may contribute to evolutionary trade-offs. However, such associations have only been investigated in a few study systems, and even less so in free ranging animal populations. Eastern chipmunks exhibit consistent individual differences in exploration, ranging from slow to fast. Birth cohorts also experience dramatic differences in age at first breeding opportunity due to annual differences in beech mast. Individuals may breed for the first time at 24, 33 or 50% of their average life span, depending on year of birth. Here, we used data from a long-term survey on a wild population to investigate the relationship between reproductive life history and consistent individual differences in exploration. We determined whether predictable differences in age at first breeding opportunity among birth cohorts were associated with exploration differences and favoured individuals with different exploration. Birth cohorts with a predictably earlier age at first breeding opportunity were faster explorers on average. Slower explorers displayed their highest fecundity (females) or highest fertilization success (males) later in their life compared with faster explorers. Overall, slow explorers attained a higher lifetime reproductive success than fast explorers when given an opportunity to reproduce later in their life. Our results suggest that the timing of mating seasons, associated with fluctuating food abundance, may favour individual variation in exploration and maintain population variation through its effects on reproductive life history. Together, our result shed light on how fluctuation in ecological conditions may maintain personality differences and on the nature of the relationships between animal personality and life history.