Endocrine flexibility can facilitate or constrain the ability to cope with global change.

Global climate change has increased average environmental temperatures world-wide, simultaneously intensifying temperature variability and extremes. Growing numbers of studies have documented phenological, behavioural and morphological responses to climate change in wild populations. As systemic signals, hormones can contribute to orchestrating many of these phenotypic changes. Yet little is known about whether mechanisms like hormonal flexibility (reversible changes in hormone concentrations) facilitate or limit the ability of individuals, populations and species to cope with a changing climate. In this perspective, we discuss different mechanisms by which hormonal flexibility, primarily in glucocorticoids, could promote versus hinder evolutionary adaptation to changing temperature regimes. We focus on temperature because it is a key gradient influenced by climate change, it is easy to quantify, and its links to hormones are well established. We argue that reaction norm studies that connect individual responses to population-level and species-wide patterns will be critical for making progress in this field. We also develop a case study on urban heat islands, where several key questions regarding hormonal flexibility and adaptation to climate change can be addressed. Understanding the mechanisms that allow animals to cope when conditions become more challenging will help in predicting which populations are vulnerable to ongoing climate change. This article is part of the theme issue 'Endocrine responses to environmental variation: conceptual approaches and recent developments'.

Repeatable negotiation rules? Only females show repeatable responses to partner removal in a brood-provisioning songbird.

Theoretical models indicate that the evolution of biparental care depends on how parents behaviourally negotiate their level of care in response to those of their partner and whether sexes and individuals consistently vary in their response (compensatory response). While the compensatory response has been widely investigated empirically, its repeatability has rarely been assessed. In this study, we used a reaction norm approach to investigate the repeatability of the compensatory offspring provisioning of a parent after temporary removal of its partner in the pied flycatcher (Ficedula hypoleuca) across different breeding seasons and partners. We found that only females partially compensated for the short-term removal of the partner and their response was significantly repeatable across years while breeding with different partners. This study highlights the importance of considering among individual differences in negotiation rules to better understand the role of negotiation mechanisms in the evolution of parental care strategies.

Songbird parents coordinate offspring provisioning at fine spatio-temporal scales.

For parents, rearing offspring together is far from a purely cooperative exercise, as a conflict of interest ('sexual conflict') exists over their optimum level of care. Recent theory emphasizes that sexual conflict can be evolutionarily resolved, and complete parental cooperation can occur, if parents directly respond ('negotiate') to each other and coordinate their level of care. Despite numerous experiments showing that parents are responsive to each other, we still lack empirical evidence of the behavioural mechanisms by which this negotiation occurs. In this study, we investigated the spatio-temporal coordination of parental provisioning behaviour as a possible mechanism of negotiation over parental care. We deployed an automated radiotracking technology to track the provisioning activity of wild great tit Parus major pairs during chick rearing. Our analyses represent the first detailed spatial and temporal description of foraging coordination in songbird parents in a natural context. We demonstrate that the foraging behaviour of the two parents is highly coordinated in space and time, with parents changing their foraging locations in conjunction with their partners' movements. Therefore, foraging coordination could be a mechanism by which parents directly monitor and respond to each other's level of investment.

Are individuals consistent? Endocrine reaction norms under different ecological challenges.

Quantifying organismal capacity for compensatory mechanisms is essential to forecast responses to environmental change. Despite accumulating evidence for individual variation in physiological plasticity, the causes and consequences of this variation remain unclear. An outstanding question is whether individual reaction norms are consistent across different environmental challenges, i.e. whether an individual that is responsive to one environmental variable will be equally responsive to a different environmental variable. Additionally, are these reaction norms themselves consistent over time, i.e. repeatable? Here, we quantified individual baseline glucocorticoid responses in house sparrows, Passer domesticus, to sequential manipulations of temperature, wind speed and food unpredictability that were repeated in discrete blocks of sampling under both control and stressor-exposed conditions. Individuals significantly decreased their baseline corticosterone levels and increased their mass during treatment exposure. This response was consistent across environmental challenge types. There was high repeatability in the intercept and slope of the baseline corticosterone reaction norm between environmental challenges but broad credible intervals in the repeatability of the reaction norm slope, suggesting that although glucocorticoid levels during baseline conditions are repeatable, among-individual variation in the shape of the glucocorticoid response may be higher than within-individual variation. Within-subject variation in baseline corticosterone levels was mainly explained by within-individual variation in body mass during stressor exposure. Despite the high lability in physiological traits, endocrine plasticity is repeatable across environmental challenges and may be able to evolve as a result of genetic accommodation, in which selection acts on genetic variation of reaction norms.

Optimizing livestock carrying capacity for wild ungulate-livestock coexistence in a Qinghai-Tibet Plateau grassland.

Wild ungulates are an important part of terrestrial ecosystems and play a critical role in maintaining ecosystem health and integrity. In many grassland ecosystems that are habituated by wild ungulates, the coexistence of domestic ungulates has created a conflict over grazing resources. Solving this conflict requires a balanced and sustainable policy that satisfies both the needs of wildlife protection and food production. Here, we assess the optimal grassland livestock carrying capacity of an alpine grassland on the Qinghai-Tibet Plateau, given the coexistence of wild populations of kiangs (Equus kiang) and Tibetan gazelles (Procapra picticaudata), two key species grazing in this region. We use kriging and the MaxEnt method to estimate the population sizes of kiangs and Tibetan gazelles in Maduo County, Qinghai Province. We then convert the estimated population size of the two species into sheep units and calculate the residual carrying capacity for livestock grazing. We show that after accounting for the grazing need for kiangs and Tibetan gazelles, grassland in Maduo is capable of supporting 420,641 sheep units, which is slightly more than the current livestock population. However, the residual carrying capacity is highly uneven across the region, and overgrazing is found in many areas of Maduo, especially in northern Maduo. This research provides a useful framework for planning sustainable livestock farming for the Qinghai-Tibet Plateau and other regions facing wildlife-livestock conflict.

Urban resources limit pair coordination over offspring provisioning.

The amount of care parents provide to the offspring is complicated by an evolutionary conflict of interest ('sexual conflict') between the two parents. Recent theoretical models suggest that pair coordination of the provisioning may reduce this conflict and increase parent and offspring fitness. Despite empirical studies showing that pair coordination is common in avian species, it remains unclear how environmental and ecological conditions might promote or limit the ability of parents to coordinate care. We compared the level of pair coordination, measured as alternation and synchrony of the nest visits, of house wrens Troglodytes aedon pairs breeding in a rural (10 nests) and a suburban (9 nests) site and investigated how differences in parental behaviours were related to habitat composition, prey abundance and how they ultimately related to reproductive success. We found that parents alternated and synchronized their nest visits more in the rural site compared to the suburban one. The suburban site is characterized by a more fragmented habitat with more coniferous trees and less caterpillar availability. Offspring from the rural site were heavier at fledging than at the suburban site. Taken together, these results suggest that environmental conditions play an important role on the emergence of coordinated parental care and that considering environmental variables is pivotal to assess the fitness consequences of parental strategies.

Across time and space: Hormonal variation across temporal and spatial scales in relation to nesting success.

There is a renewed interest in investigating individual variation in hormone levels in relation to fitness metrics, as hormones act as mediators of life-history trade-offs. Hormone concentrations, however, are labile, responding to both internal and external stimuli, so the relationship between hormones and fitness can be non-consistent. One explanation of this inconsistent relationship is that a single hormone sample may not be representative of individual phenotypes in a free-living species. We addressed this issue by repeatedly sampling a free-living population of mountain white-crowned sparrows, Zonotrichia leucophrys oriantha, for baseline and stress-induced corticosterone (cort) and testosterone (T) across different stages of the breeding season. We measured (co)variation using three different methods, taking into account inter- and intra-individual variances, to determine whether hormone levels and the stress response are repeatable. We documented the temporal (over 3 months) and spatial (home-range) variation of individual hormone phenotypes and investigated how these components related to nesting success. At the population level, we found significant repeatability in male stress-induced cort concentrations but no repeatability in male or female baseline cort or male T concentrations. Using a new metric of intra-individual variance focusing on the stress response (profile repeatability), we found a wide range of variance scores, with most individuals showing high variation in their stress response. Similarly, we found a low level of repeatability of the reaction norm intercept and slope for the stress response across different life-history stages. Males with higher concentrations of stress-induced cort had more central home-ranges. Males with higher body condition had larger home-ranges; however, home-range size did not relate to male hormone concentrations or nesting success. We also did not find any significant relationship between variation in hormone levels and nesting success. We recommend that future studies combine both physiological and environmental components to better understand the relationship between hormones and fitness.

Genetic inheritance and environment determine endocrine plasticity to urban living.

As urban areas continue to expand globally, understanding how and why species respond to novel habitats becomes increasingly important. Knowledge of the mechanisms behind observed phenotypic changes in urban animals will enable us to better evaluate the impact of urbanization on current and future generations of wildlife. Physiological changes, such as those involved in the endocrine stress response, may allow individuals to inhabit and thrive in urbanized areas, but it is currently unknown how these changes arise in natural populations. In this study, we performed a four-way cross-foster experiment in free-living house wren chicks, Troglodytes aedon, to disentangle whether differences in baseline corticosterone between urban and rural individuals are a result of genetic and/or plastic mechanisms during development. We found that urban chicks already had higher corticosterone levels than their rural counterparts on the day they hatched, which suggests a possible genetic component to the corticosterone phenotype. However, rural offspring that were moved to an urban environment significantly increased their corticosterone levels, mimicking those of urban offspring. Our findings suggest that, although differences in baseline corticosterone concentrations between urban and rural individuals may have a genetic component, plasticity plays a pivotal role and can modify the corticosterone phenotype in response to the environment experienced in the first two weeks of life.

The effect of social connections on the discovery of multiple hidden food patches in a bird species.

Social foraging is thought to provide the possibility of information transmission between individuals, but this advantage has been proved only in a handful of species and contexts. We investigated how social connections in captive flocks of house sparrows (Passer domesticus) affected the discovery of (i.e. feeding for the first time from) two hidden food patches in the presence of informed flock-mates. At the first-discovered and most-exploited food patch social connections between birds affected the order of discovery and presumably contributed to a greater exploitation of this patch. However, social connections did not affect discovery at the second food patch despite its close spatial proximity. Males discovered the food sources sooner than females, while feeding activity was negatively related to patch discovery. Age had no effect on the order of discovery. Birds that first discovered and fed at the food patches were characterized by higher level of social indifference, i.e. followed others less frequently than other birds in an independent context. Our findings provide experimental evidence for the importance of variable social connections during social foraging in house sparrow flocks, and suggest that social attraction can contribute differently to the exploitation of different patches when multiple food sources are present.