One contribution of 14 to a theme issue 'Behavioural, ecological and evolutionary responses to extreme climatic events'. As global climate change progresses, the occurrence of potentially disruptive climatic events such as storms are increasing in frequency, duration and intensity resulting in higher mortality and reduced reproductive success. What constitutes an extreme climatic event? First we point out that extreme climatic events in biological contexts can occur in any environment. Focusing on field and laboratory data on wild birds we propose a mechanistic approach to defining and investigating what extreme climatic events are and how animals cope with them at physiological and behavioural levels. The life cycle of birds is made up of life-history stages such as migration, breeding and moult that evolved to match a range of environmental conditions an individual might expect during the year. When environmental conditions deteriorate and deviate from the expected range then the individual must trigger coping mechanisms (emergency life-history stage) that will disrupt the temporal progression of life-history stages, but enhance survival. Using the framework of allostasis, we argue that an extreme climatic event in biological contexts can be defined as when the cumulative resources available to an individual are exceeded by the sum of its energetic costs-a state called allostatic overload. This allostatic overload triggers the emergency life-history stage that temporarily allows the individual to cease regular activities in an attempt to survive extreme conditions. We propose that glucocorticoid hormones play a major role in orchestrating coping mechanisms and are critical for enduring extreme climatic events.This article is part of the themed issue 'Behavioural, ecological and evolutionary responses to extreme climatic events'.

There is increasing evidence that individuals in many species avoid areas exposed to chronic anthropogenic noise, but the impact of noise on those who remain in these habitats is unclear. One potential impact is chronic physiological stress, which can affect disease resistance, survival and reproductive success. Previous studies have found evidence of elevated stress-related hormones (glucocorticoids) in wildlife exposed to human activities, but the impacts of noise alone are difficult to separate from confounding factors. Here we used an experimental playback study to isolate the impacts of noise from industrial activity (natural gas drilling and road noise) on glucocorticoid levels in greater sage-grouse (Centrocercus urophasianus), a species of conservation concern. We non-invasively measured immunoreactive corticosterone metabolites from fecal samples (FCMs) of males on both noise-treated and control leks (display grounds) in two breeding seasons. We found strong support for an impact of noise playback on stress levels, with 16.7% higher mean FCM levels in samples from noise leks compared with samples from paired control leks. Taken together with results from a previous study finding declines in male lek attendance in response to noise playbacks, these results suggest that chronic noise pollution can cause greater sage-grouse to avoid otherwise suitable habitat, and can cause elevated stress levels in the birds who remain in noisy areas.

Individuals at the forefront of a range shift are likely to exhibit phenotypic traits that distinguish them from the population breeding within the historic range. Recent studies have examined morphological, physiological and behavioral phenotypes of individuals at the edge of their range. Several studies have found differences in the hypothalamic–pituitary–adrenal (HPA) axis activity in response to acute restraint stress in individuals at the range limits. HPA axis activation leads to elevations in glucocorticoids that regulate physiology and behavior. Here we compare the hormonal profiles and morphometrics from Gambel’s white-crowned sparrows (Zonotrichia leucophrys gambelii) breeding at the northern limit of the population’s range to those birds breeding within the historic population range. Birds breeding at the northern limit experienced a harsher environment with colder temperatures; however, we found no differences in arthropod prey biomass between the northern limit and more southern (historic) sites. Males at the northern limit had higher body condition scores (mass corrected for body size) compared to individuals within the historic range, but no differences were found in beak and tarsus lengths, wing chord, muscle profile or fat stores. In males during the pre-parental stage, before breeding commenced, HPA axis activity was elevated in birds at the northern limit of the range, but no differences were found during the parental or molt stages. Females showed no differences in HPA axis activity during the parental stage. This study suggests that “pioneering” individuals at the limits of their breeding range exhibit physiology and morphology that are distinct from individuals within the historic range.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-015-3447-7) contains supplementary material, which is available to authorized users.

HighlightsMany species are expanding or contracting their geographic distribution.Range changes are caused by climate change, human disturbance and invasive species.What are the characteristics of pioneers in new habitat?Individuals at the leading edge appear to be highly variable.In range expanding songbirds responses to stress are extremely variable.

The framework of allostasis, allostatic load and overload (i.e., stability through change) attempts to combine homeostasis processes in day-to-day responses of physiology and behavior. These include predictive changes in environment such as seasons, and facultative responses to perturbations. The latter can be severe, occur at any time, and may present considerable additional challenges to homeostasis. Hormonal cascades, such as the hypothalamo-pituitary-adrenal cortex (HPA) axis, play a key role in responses to perturbations across vertebrate taxa. Glucocorticoids have been implicated in these processes in relation to energy balance that plays a role in determining responses to energetic demand (allostatic load) and influencing subsequent physiology and behavior associated with coping. Circulating glucocorticoid levels are likely regulated in part based on an individual’s proximity to energetic crisis, identified as the perturbation resistance potential (PRP). In the model of allostatic load, PRP is quantified as the difference between available resources and all energetic costs of allostatic load such as daily routines, life history stages (breeding, migration, molt and so on), and the impact of environmental perturbations. PRP can change gradually or abruptly and may be reflected by spikes in blood hormone levels. The pattern of individual responsiveness to PRP may vary and has specific implications for the activation of mineralocorticoid vs glucocorticoid-type receptors, hormone metabolizing enzymes and other downstream factors in target tissues. However, PRP is a difficult metric to measure. Here, we examine the variety of cues that animals may use to inform them about the status of their PRP and probability of energetic crisis. We consider (1) elevation in glucocorticoids as an endocrine “decision,” and (2) error management strategies in evaluating responsiveness to cues that may reflect or predict an impending energetic crisis. The potential for differential receptor activation as well as further integrative “decisions” to determine the diverse and sometimes contradictory effects of receptor activation and its downstream actions are important to the consideration of error management. This perspective offers insight into the basis of intra- and inter-individual variability in responsiveness and opens an avenue toward improving compatibility of the allostasis model with more classical views on “stress”.