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Individual heterogeneity in diet and foraging behaviour is common in wild animal populations, and can be a strong determinant of how populations respond to environmental changes. Within populations, variation in foraging behaviour and the occurrence of individual tactics in relation to resources distribution can help explain differences in individual fitness, and ultimately identify important factors affecting population dynamics. We examined how foraging behaviour and habitat during the breeding period related to the physiological state of a long‐ranging seabird adapted to sea ice, the Antarctic petrel Thalassoica antarctica.Firstly, using GPS tracking and state‐switching movement modelling (hidden Markov models) on 124 individual birds, we tested for the occurrence of distinct foraging tactics within our study population. Our results highlight a large variation in the movement and foraging behaviour of a very mobile seabird, and delineate distinct foraging tactics along a gradient from foraging in dense pack ice to foraging in open water.Secondly, we investigated the effects of these foraging tactics on individual state at return from a foraging trip. We combined movement data with morphometric and physiological measurements of a suite of plasma metabolites that provided a general picture of a bird's individual state. Foraging in denser sea ice was associated with lower gain in body mass during brooding, as well as lower level of energy acquisition (plasma triacylglycerol) during both brooding and incubation. We found no clear relationship between the foraging tactic in relation to sea ice and the energetic stress (changes in plasma corticosterone), energetic balance (β‐hydroxybutyrate) or trophic level (δ15N). However, a shorter foraging range was related to both the energetic balance (positively) and the trophic level (negatively).Our results highlight a diverse range of foraging tactics in relation to sea ice in Antarctic petrels. While the various foraging tactics do not seem to strongly alter energetic balance, they may affect other aspects of Antarctic petrels' physiology. Future changes in sea‐ice habitats can thus be expected to have an impact on the individual state of seabirds such as Antarctic petrels, which could ultimately affect their population dynamics. Nonetheless, strong individual heterogeneity in the use of sea‐ice habitats by a typical pagophilic species might strengthen its resilience to environmental changes and in particular to forecasted sea‐ice loss. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

1. A combination of timing of and body condition (i.e., mass) at arrival on the breeding grounds interact to influence the optimal combination of the timing of reproduction and clutch size in migratory species. This relationship has been formalized by Rowe et al. in a condition-dependent individual optimization model (American Naturalist, 1994, 143, 689-722), which has been empirically tested and validated in avian species with a capital-based breeding strategy. 2. This model makes a key, but currently untested prediction; that variation in the rate of body condition gain will shift the optimal combination of laying date and clutch size. This prediction is essential because it implies that individuals can compensate for the challenges associated with late timing of arrival or poor body condition at arrival on the breeding grounds through adjustment of their life history investment decisions, in an attempt to maximize fitness. 3. Using an 11-year data set in arctic-nesting common eiders (Somateria mollissima), quantification of fattening rates using plasma triglycerides (an energetic metabolite), and a path analysis approach, we test this prediction of this optimization model; controlling for arrival date and body condition, females that fatten more quickly will adjust the optimal combination of lay date and clutch size, in favour of a larger clutch size. 4. As predicted, females fattening at higher rates initiated clutches earlier and produced larger clutch sizes, indicating that fattening rate is an important factor in addition to arrival date and body condition in predicting individual variation in reproductive investment. However, there was no direct effect of fattening rate on clutch size (i.e., birds laying on the same date had similar clutch sizes, independent of their fattening rate). Instead, fattening rate indirectly affected clutch size via earlier lay dates, thus not supporting the original predictions of the optimization model. 5. Our results demonstrate that variation in the rate of condition gain allows individuals to shift flexibly along the seasonal decline in clutch size to presumably optimize the combination of laying date and clutch size. [ABSTRACT FROM AUTHOR]

Experimental glucocorticoid ( GC) manipulations can be useful for identifying the mechanisms that drive life-history and fitness variation in free-living animals, but predicting the effects of GC treatment can be complicated. Much of the uncertainty stems from the multi-faceted role of GCs in organismal metabolism, and their variable influence with respect to life-history stage, ecological context, age, sex and individual variation., Glucocorticoid hormones have been implicated in the regulation of parental care in many vertebrate taxa but in two seemingly contradictory ways, which sets up a potential GC-induced 'reproductive conflict'. Circulating GCs mediate adaptive physiological and behavioural responses to stressful events, and elevated levels can lead to trade offs between reproductive effort and survival (e.g. the current reproduction vs. survival hypothesis). The majority of studies examining the fitness effects of GC manipulations extend from this hypothesis. However, when animals are not stressed (likely most of the time) baseline GCs act as key metabolic regulators of daily energy balance, homoeostasis, osmoregulation and food acquisition, with pleiotropic effects on locomotor activity or foraging behaviour. Slight increases in circulating baseline levels can then have positive effects on reproductive effort (e.g. the 'cort' fitness/adaptation hypotheses), but comparatively few GC manipulation studies have targeted these small, non-stress induced increases., We review studies of GC manipulations and examine the specific hypotheses used to predict the effects of manipulations in wild, breeding vertebrates. We argue that given the dichotomous function of GCs the current 'reproduction vs. survival' paradigm is unnecessarily restrictive and predicts only deleterious GC effects on fitness. Therefore, a broader set of hypotheses should be considered when testing the fitness effects of GC manipulations., When framing experimental manipulation studies, we urge researchers to consider three key points: life-history context (e.g. long vs. short lived, semelparous vs. iteroparous, etc.), ecological context and dose delivery. [ABSTRACT FROM AUTHOR]

Biomedical researchers have long appreciated that maternal stressors can induce preparative and adaptive programming in offspring via exposure to maternal Glucocorticoids ( GCs). However, few ecologists are aware of the capacity for maternal GC exposure to translate ecological and environmental stressors into preparative and adaptive programmed offspring responses in free-living systems. We review a growing body of experimental work indicating that circulating maternal GCs link ecological stressors with adaptive programming of the stress axis. Throughout, we emphasise that natural and human-induced ecological stressors play a fundamental role in programming the capacity of individuals, populations and communities to respond to both predictable and unpredictable ecological change via translating maternal adversity into responsive programming of the vertebrate stress axis., To encourage rigorous testing of this paradigm in a broad range of ecological systems, we introduce the principal extrinsic stressors with a recognised potential to alter maternal circulating GC levels. We then review from the biomedical literature regarding the underlying physiological and epigenetic mechanisms of stress-induced programming of individual phenotypes to predict how variation in ecological stressors can produce individual variation in stress axis management., To appreciate the potential evolutionary inertia (i.e. adaptive value) of maternally programmed individual variation, we review key recent studies in free-living systems that test its adaptive function, and then discuss how variation in stress-axis programming may scale up to influence populations and ecological communities., Given the huge potential of this field, it is encouraging that ecologists are beginning to examine how and why maternal GCs translate ecological and environmental stressors into preparative stress axis programming in free-living systems. [ABSTRACT FROM AUTHOR]

Optimality theory predicts that both timing of arrival and arrival state on the breeding area will determine reproductive timing and investment in migratory organisms. We tested this idea using a condition-dependent individual optimization model ( Ardea, 1980, 225 and The American Naturalist, 1994, 698) in common eider ducks through descriptive data, path analyses and experimental manipulation. Our results support the causal pathways drawn from the optimization model indicating that individuals adjust their reproductive decisions as a function of their arrival date and body condition at arrival. Independent of body condition, early-arriving females had a longer pre-laying period, but still initiated their nests earlier, and produced larger clutches than late-arriving birds. Independent of arrival date, females in good condition laid earlier than those in poor condition. Manipulation of pre-laying female body condition confirmed that the relationship between condition and laying date was causal. Female common eiders appear to optimize reproductive decisions in response to both their external (i.e. environmental conditions affecting the egg-value) and internal (i.e. body condition) states. These adjustments seem to minimize the fitness costs of reproduction, in which higher clutch size is not associated with an apparent lower survival or future breeding probability. Our study emphasizes the importance of (i) simultaneously considering the timing of migration, the state of individuals and the seasonal change in egg-value to understand clearly birds' breeding decisions and (ii) appreciating the potential proximate and ultimate factors explaining why some individuals delay breeding and/or produce small clutches. [ABSTRACT FROM AUTHOR]