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Recent studies have demonstrated that generic statistical signals derived from time series of population abundance and fitness-related traits of individuals can provide reliable indicators of impending shifts in population dynamics. However, how the seasonal timing of environmental stressors influences these early warning indicators is not well understood. The goal of this study was to experimentally assess whether the timing of stressors influences the production, detection and sensitivity of abundance- and trait-based early warning indicators derived from declining populations. In a multi-generation, season-specific habitat loss experiment, we exposed replicate populations of Drosophila melanogaster to one of two rates of chronic habitat loss (10% or 20% per generation) in either the breeding or the non-breeding period. We counted population abundance at the beginning of each season, and measured body mass and activity levels in a sample of individuals at the end of each generation. When habitat was lost during the breeding period, declining populations produced signals consistent with those documented in previous studies. Inclusion of trait-based indicators generally improved the detection of impending population collapse. However, when habitat was lost during the non-breeding period, the predictive capacity of these indicators was comparatively diminished. Our results have important implications for interpreting signals in the wild because they suggest that the production and detection of early warning indicators depends on the season in which stressors occur, and that this is likely related to the capacity of populations to respond numerically the following season.

Carry-over effects occur when past experience influences current individual performance. Although variation in conspecific density in one season has been shown to carry over to influence dynamics in the following season, the proximate ecological mechanisms driving these effects are unknown. One hypothesis is that high density decreases food availability, resulting in poor physiological condition, which in turn compromises performance the next season. Alternatively, high conspecific density could also lead to a high degree of antagonistic interactions, decreasing the amount of time individuals spend foraging. To investigate these hypotheses, we applied a factorial design where both conspecific density and per capita food availability during the non-breeding period were independently manipulated in seasonal populations of common fruit flies (Drosophila melanogaster Meigen, 1830). Individual condition at the beginning of the breeding period was influenced by per capita food availability but not density during the previous non-breeding period. In contrast, reproductive output was most strongly influenced by the interaction between per capita food availability and density in the previous non-breeding period, such that populations that experienced high non-breeding densities and low food availability had the lowest reproductive output. However, the strength of this effect was relatively weak. Our results demonstrate how environmental and social conditions in one part of the annual cycle can carry over to influence individual performance in subsequent periods.

Phenotypic plasticity allows organisms to adjust the timing of life-history events in response to environmental and demographic conditions. Shifts by individuals in the timing of breeding with respect to variation in age and temperature are well documented in nature, and these changes are known to scale to affect population dynamics. However, relatively little is known about how organisms alter phenology in response to other demographic and environmental factors. We investigated how pre-breeding temperature, breeding population density, age, and rainfall in the first month of life influenced the timing and plasticity of lay date in a population of Savannah sparrows (Passerculus sandwichensis) monitored over 33 years (1987-2019). Females that experienced warmer pre-breeding temperatures tended to lay eggs earlier, and also advanced their lay date as they aged, but breeding population density had no effect on lay date. Natal precipitation interacted with age to influence lay date plasticity, with females that experienced high precipitation levels as nestlings advancing lay dates more strongly over the course of their lives. We also found evidence for varied pace of life; females that experienced high natal precipitation had shorter lifespans and reduced fecundity, but more nesting attempts over their lifetimes. Rainfall during the nestling period increased through time, while population density and fecundity declined, suggesting that increased precipitation on the breeding grounds may be detrimental to breeding females and ultimately the viability of the population as a whole. Our results suggest that females adjust their laying date in response to pre-breeding temperature, and as they age, while presenting new evidence that environmental conditions during the natal period can affect phenological plasticity and generate downstream, population-level effects.

Evidence that organisms evolve rapidly enough to alter ecological dynamics necessitates investigation of the reciprocal links between ecology and evolution. Data that link genotype to phenotype to ecology are needed to understand both the process and ecological consequences of rapid evolution. Here, we quantified the suite of elements in individuals (i.e., ionome) and differences in the fluxes of key nutrients across populations of threespine stickleback. We find that allelic variation associated with freshwater adaptation that controls bony plating is associated with changes in the ionome and nutrient recycling. More broadly, we find that adaptation of marine stickleback to freshwater conditions shifts the ionomes of natural populations and populations raised in common gardens. In both cases ionomic divergence between populations was primarily driven by differences in trace elements rather than elements typically associated with bone. These findings demonstrate the utility of ecological stoichiometry and the importance of ionome-wide data in understanding eco-evolutionary dynamics.

Migration strategies differ greatly among and within avian populations. The associated trade‐offs and fitness consequences of diverse strategies and how they persist are pertinent questions in migration research. Migration is a costly endeavour, presumably compensated for by better survival conditions in the non‐breeding area. One way to assess the cost of alternative strategies is to investigate the investment in movement across the entire annual cycle, an assessment made increasingly feasible with improvements in tracking technology. Our study focuses on lesser black‐backed gulls, generalist seabirds that exploit a broad range of resources, exhibit diverse migration strategies and have potentially altered migration strategies in response to human activities and climate change. We used GPS tracking to quantify lesser black‐backed gulls’ movement throughout their annual cycle and compare trade‐offs among four migration strategies. The annual cumulative distance travelled by long distance migrants wintering in west Africa, over 4000 km from their breeding colony, did not differ significantly from individuals of the same breeding colony wintering only a few hundred kilometres away in Great Britain. Short distance migrants returned to the colony first, and long distance migrants returned last. Sex and wing length were not correlated with maximum range, cumulative distance travelled or timing. Individuals spent only a small proportion of their time in flight and spent on average 17% of their time at sea throughout an annual cycle, suggesting a reliance on inland resources for many individuals. Analysing movement throughout the annual cycle can change our perspective and understanding of consequences of different migration strategies. Our study shows that a range of migration strategies coexists and we propose that the long term costs and benefits of these strategies balance out. Diversity in migration strategies may contribute to the resilience of this species in the face of ongoing anthropogenic impact on the environment.

For declining wild populations, a critical aspect of effective conservation is understanding when and where the causes of decline occur. The primary drivers of decline in migratory and seasonal populations can often be attributed to a specific period of the year. However, generic, broadly applicable indicators of these season-specific drivers of population decline remain elusive. We used a multi-generation experiment to investigate whether habitat loss in either the breeding or non-breeding period generated distinct signatures of population decline. When breeding habitat was reduced, population size remained relatively stable for several generations, before declining precipitously. When non-breeding habitat was reduced, between-season variation in population counts increased relative to control populations, and non-breeding population size declined steadily. Changes in seasonal vital rates and other indicators were predicted by the season in which habitat loss treatment occurred. Per capita reproductive output increased when non-breeding habitat was reduced and decreased with breeding habitat reduction, whereas per capita non-breeding survival showed the opposite trends. Our results reveal how simple signals inherent in counts and demographics of declining populations can indicate which period of the annual cycle is driving declines.

Migratory birds undergo impressive body remodelling over the course of an annual cycle. Prior to long-distance flights, red knots (Calidris canutus islandica) reduce gizzard mass while increasing body mass and pectoral muscle mass. Although body mass and pectoral muscle mass are functionally linked via their joint effects on flight performance, gizzard and pectoral muscle mass are thought to be independently regulated. Current hypotheses for observed negative within-individual covariation between gizzard and pectoral muscle mass in free-living knots are based on a common factor (e.g. migration) simultaneously affecting both traits, and/or protein limitation forcing allocation decisions. We used diet manipulations to generate within-individual variation in gizzard mass and test for independence between gizzard and pectoral muscle mass within individuals outside the period of migration and under conditions of high protein availability. Contrary to our prediction, we observed a negative within-individual covariation between gizzard and pectoral muscle mass. We discuss this result as a potential outcome of an evolved mechanism underlying body remodelling associated with migration. Although our proposed mechanism requires empirical testing, this study echoes earlier calls for greater integration of studies of function and mechanism, and in particular, the need for more explicit consideration of the evolution of mechanisms underlying phenotypic design.

Age-related increases in the repeatable expression of labile phenotypic traits are often assumed to arise from an increase in among-individual variance due to differences in developmental plasticity or by means of state-behavior feedbacks. However, age-related increases in repeatability could also arise from a decrease in within-individual variance as a result of stabilizing trait expression, that is, canalization. Here we describe age-related changes in within-individual and among-individual variance components in two correlated traits— gizzard mass and exploration behavior—in a medium-sized shorebird, the red knot (Calidris canutus). Increased repeatability of gizzard mass came about due to an increase in among-individual variance, unrelated to differences in developmental plasticity, together with decreases in within-individual variance consistent with canalization. We also found canalization of exploration but no age-related increase in overall repeatability, which suggests that showing predictable expression of exploration behavior may be advantageous from a very young age onward. Contrasts between juveniles and adults in the first year after their capture provide support for the idea that environmental conditions play a key role in generating among-individual variation in both gizzard mass and exploration behavior. Our study shows that stabilization of traits occurs under constant conditions: with increased exposure to predictable cues, individuals may become more certain in their assessment of the environment allowing traits to become canalized.

Behavioural differences may arise in the absence of genetic or environmental variation. Chaotic dynamics may influence behavioural development, and so this among-individual variation. We discuss methods and experimental designs to test this idea. Ultimately, nonlinear and chaotic behavioural development may explain much of natural variation., Comment: Version of manuscript following initial rejection and invitation to resubmit from journal. Further revised version now published, see: https://www.sciencedirect.com/science/article/pii/S0003347218300654