Males and females often exhibit different behaviours during mate acquisition, pair‐bonding and parenting, and a convenient label to characterize these behaviours is sex role. The diverse roles that male and female shorebirds (plovers, sandpipers and allies) exhibit in mating and parenting have played a key role in advancing mainstream theories in avian ecology and behavioural biology including sexual selection, sexual conflict and parental cooperation. Recent advances in shorebird research have also highlighted the significance of the social environment in driving sex role behaviours by linking the adult sex ratio with breeding behaviour and population demography. Here we review the key advances in sex role research using shorebirds as an ecological model system. We identify knowledge gaps and argue that shorebirds have untapped potential to accelerate diverse research fields including evolutionary genomics, movement ecology, social networks and environmental changes. Future studies of sex roles will benefit from individual‐based monitoring using advanced tracking technologies, and from multi‐team collaborations that are facilitated by standardized data collection methodologies across different species in the field. These advances will not only contribute to our understanding of reproductive strategies, but they will also have knock‐on effects on predicting population resilience to environmental changes and on prioritizing species for conservation.

Migrating animals are thought to be important spillover sources for novel pathogens. Haemosporidians (malaria-related parasites) are one such group of pathogens that commonly spillover into novel host communities if competent vectors are present. In birds, shorebirds (sandpipers, plovers and allies) perform some of the longest avian migrations, yet they are traditionally perceived as relatively free from haemosporidians. Although low prevalence fits several theories, such as effective immune responses or low exposure to vectors, few studies have been carried out in freshwater inland sites, where the vectors of haemosporidians (e.g. mosquitoes) are abundant, with a mixture of actively migrating (staging) and breeding hosts. Here we report the prevalence of three haemosporidian parasites, Haemoproteus, Leucocytozoon and Plasmodium, screened in 214 shorebirds from 15 species sampled in a freshwater marshland, southern Belarus. Contrary to most previous studies, we found that haemosporidians were frequent, with an overall prevalence in the community of 16.36%, including the locally breeding shorebirds (23.13%, 134 individuals of 10 species). However, actively migrating shorebirds had much lower prevalence (0.05%, 55 individuals of five species). We suggest that blood parasite infections are more common in shorebirds than currently acknowledged. Yet, actively migrating species may be free from haemosporidians or carry suppressed infections, leading to lower prevalence or even apparent absence in some species. Taken together, we theorize that a combination of sampling biases has driven our understanding of haemosporidian prevalence in shorebirds and future studies should take the migratory status of individuals into account when reporting prevalence. Furthermore, we argue that birds undergoing active migration may be less likely sources of spillover events than previously assumed.

The insulin-like growth factor 1 (IGF-1) is a pleiotropic hormone that regulates essential life-history traits and is known for its major contribution to determining individual ageing processes. High levels of IGF-1 have been linked to increased mortality and are hypothesised to cause oxidative stress. This effect has been observed in laboratory animals, but whether it pertains to wild vertebrates has not been tested. This is surprising because studying the mechanisms that shape individual differences in lifespan is important to understanding mortality patterns in populations of free-living animals. We tested this hypothesis under semi-natural conditions by simulating elevated IGF-1 levels in captive bearded reedlings, a songbird species with an exceptionally fast pace of life. We subcutaneously injected slow-release biodegradable microspheres loaded with IGF-1 and achieved a systemic 3.7-fold increase of the hormone within the natural range for at least 24 h. Oxidative damage to lipids showed marked sexual differences: it significantly increased the day after the manipulation in treated males and returned to baseline levels four days post-treatment, while no treatment effect was apparent in females. Although there was no overall difference in survival between the treatment groups, high initial (pre-treatment) IGF-1 and low post-treatment plasma malondialdehyde levels were associated with enhanced survival prospects in males. These results suggest that males may be more susceptible to IGF-1-induced oxidative stress than females and quickly restoring oxidative balance may be related to fitness. IGF-1 levels evolve under opposing selection forces, and natural variation in this hormone’s level may reflect the outcome of individual optimization.

Most organisms have to cope with unpredictable environmental challenges such as fluctuations in nutritional resources. Insulin-like growth factor-1 (IGF-1) is an evolutionarily conserved hormone that is highly sensitive to the individual nutritional status and regulates major life-history traits including lifespan and reproduction across vertebrates. We investigated the role of IGF-1 during periods of food shortages by altering between two feeding regimes (110 and 70% of daily food intake) after a period of ad libitum feeding in captive bearded reedlings (Panurus biarmicus). Each dietary treatment was repeated twice. Birds lost mass under food restriction, but the magnitude of mass change depended on the preceding dietary conditions. Moreover, bearded reedlings showed large, repeatable individual differences in their IGF-1 reaction norms with some individuals increasing IGF-1 levels in response to a restricted diet, whereas others showed no responses or decreased IGF-1 levels. This variation was explained by differences in average body mass: heavier individuals had higher IGF-1 levels during the control treatment and were more likely to decrease IGF-1 levels in response to the dietary restriction than did lighter ones. This result uncovers an individual by environment interaction (I × E) and may have important implications for the evolution of IGF-1 related hormonal phenotypes in this species.

In the endeavour to understand the causes and consequences of the variation in animal personality, a wide range of studies were carried out, utilising various aspects to make sense of this biological phenomenon. One such aspect integrated the study of physiological traits, investigating hypothesised physiological correlates of personality. Although many of such studies were carried out on vertebrates (predominantly on birds and mammals), studies using arthropods (mainly insects) as model organisms were also at the forefront of this area of research. In order to review the current state of knowledge on the relationship between personality and the most frequently studied physiological parameters in arthropods, we searched for scientific articles that investigated this relationship. In our review, we only included papers utilising a repeated-measures methodology to be conceptually and formally concordant with the study of animal personality. Based on our literature survey, metabolic rate, thermal physiology, immunophysiology, and endocrine regulation, as well as exogenous agents (such as toxins) were often identified as significant affectors shaping animal personality in arthropods. We found only weak support for state-dependence of personality when the state is approximated by singular elements (or effectors) of condition. We conclude that a more comprehensive integration of physiological parameters with condition may be required for a better understanding of state's importance in animal personality. Also, a notable knowledge gap persists in arthropods regarding the association between metabolic rate and hormonal regulation, and their combined effects on personality. We discuss the findings published on the physiological correlates of animal personality in arthropods with the aim to summarise current knowledge, putting it into the context of current theory on the origin of animal personality.

Environmental challenges, such as fluctuations in food availability, could influence reproductive investment. If resource availability is poor, individuals need to decide in which life‐history process they invest more energy and vital compounds, which determine the cost of reproduction. In resource allocation, the physiological pathways have important roles. The aim of our study was to examine whether food availability influenced physiological traits (insulin‐like growth factor‐1 (IGF‐1) concentration, antioxidant capacity (OXY), level of oxidative damage (ROM) and haematocrit value) prior to egg‐laying in female canaries Serinus canaria. We also tested whether these physiological traits were associated with traits reflecting the reproductive investment in egg macro‐ (egg mass and yolk mass) and micronutrient content (eggshell biliverdin‐ and protoporphyrin‐based colouration). To test these questions we conducted a food restriction experiment with control and food‐restricted (72% of control food) groups. Our study showed that food‐restricted canary females delayed egg‐laying and the physiological traits differed between the groups. At the pre‐laying period, after 9–11 days of treatment, a reduction in plasma IGF‐1 concentration, ROM level and haematocrit value was detected in the control group, whereas in the food‐restricted group plasma IGF‐1 concentration increased, while ROM level and haematocrit value did not change. Plasma level of OXY was not influenced by treatment or breeding period. Plasma concentration of IGF‐1, haematocrit value and oxidative status before egg laying did not affect the egg characteristics. Our study highlights the importance of breeding stage when studying and interpreting the effects of food restriction on physiological traits of breeding birds. Moreover, our data suggest that nutritional limitation had an effect on the timing of egg‐laying that could be mediated by changes in physiological variables.

Hormonal pathways have been proposed to be key at modulating how fast individuals grow and reproduce and how long they live (i.e., life history trajectory). Research in model species living under controlled environment is suggesting that insulin-like growth factor 1 (IGF-1), which is an evolutionarily conserved polypeptide hormone, has an important role in modulating animal life histories. Much remains, however, to be done to test the role played by IGF-1 in shaping the phenotype and life history of animals in the wild. Using a wild long-lived bird, the Alpine swift (Tachymarptis melba), we show that adults with higher levels of IGF-1 had longer wings and shorter telomeres. Hence, telomeres being a proxy of lifespan in this species, our results support a potential role of IGF-1 at shaping the life-history of wild birds and suggest that IGF-1 may influence the growth-lifespan trade-off.

Moulting is a crucial, yet often overlooked life-history stage in many animals, when they renew their integumental structures. This life-history stage is an energetically demanding somatic growth event that has particular importance in birds because feathers play a crucial role in flight, insulation and communication. Somatic growth processes are regulated by the evolutionarily conserved peptide hormone insulin-like growth factor 1 (IGF-1). However, the role of IGF-1 in feather growth remains unknown. In this study, we captured 41 juvenile free-living bearded reedlings (Panurus biarmicus) that had started their first complete moult and brought them into captivity. Then, we manipulated their circulating IGF-1 levels using poly-(lacticco-glycolid acid) microparticles (microspheres) that provide a sustained release of IGF-1. The treatment increased IGF-1 levels but did not affect the feather growth rate. However, 2 weeks after the treatment, birds in the increased IGF-1 group were moulting more feathers simultaneously than the controls and were at a more advanced stage of moult. Birds with experimentally increased IGF-1 levels had better quality feathers (measured by a lower number of fault bars) than the controls. These results suggest that an increase in IGF-1 does not speed up feather growth, but may alter moult intensity by initiating the renewal of several feathers simultaneously. This may shorten the overall moulting time but may imply costs in terms of IGF-1-induced oxidative stress.

Despite the widely used application of standardized capture-handling protocols to collect blood and assess the physiological stress response, the actual sampling design (e.g., timing and the number of blood samples) often differs between studies, and the potential implications for the measured physiological endpoints remain understudied. We, therefore experimentally tested the effects of repeated handling and multiple blood sampling on the stress response in wintering free-living great tits (Parus major). We modified a well-established sampling protocol of avian studies by adding either an additional blood sample or a "sham-manipulation" (i.e., handling associated with the blood sampling procedure without venepuncture), to disentangle the effects of handling stress and blood loss. We combined three different stress metrics along the endocrine-immune interface to investigate the acute short-term stress response: total corticosterone concentrations (Cort), the heterophil/ lymphocyte ratio (H:L), and the Leucocyte Coping Capacity (LCC). Our study provided three key results: i) no relationship between Cort levels, LCC and H:L, confirming that these three parameters represent different physiological endpoints within the stress response; ii) contrasting dynamics in response to stress by the measured parameters and iii) no difference in physiological stress levels 30 min after capture due to one additional blood sampling or handling event. By optimising the sampling design, our results provide implications for animal welfare and planning experimental procedures on stress physiology in passerine species.

Lifespan evolves as a compromise between antagonistic selection forces. Insulin-like growth factor 1 (IGF-1) is a pleiotropic hormone that regulates several life-history traits. High levels of IGF-1 have been linked to increased mortality, partly by causing oxidative stress. However, these effects have no experimental evidence in wild animals. We implanted microspheres loaded with exogenous IGF-1 into bearded reedlings, a common short-lived Eurasian songbird. The treatment elevated plasma IGF-1 levels for at least 24 h. Oxidative damage to lipids significantly increased the day after the manipulation in treated birds, but returned to baseline levels four days post-treatment. The treatment had no effect on survival over 16 months; however, birds with higher pre-treatment (baseline) IGF-1 levels had better survival prospects. These results suggest that, although high IGF-1 levels may induce oxidative damage, natural variation in this hormone’s level may reflect the outcome of individual optimization.

Avian plumage colors and ornaments are excellent models to study the endocrine mechanisms linking sexually selected traits and individual parameters of quality and condition. Insulin-like growth factor 1 (IGF-1) is an evolutionarily highly conserved peptide hormone. Its regulatory role in cell proliferation and differentiation and its high sensitivity to the nutritional state of individuals suggest it as an interesting candidate, possibly providing a link between body condition and individual capacity to grow elaborated ornamental features. We investigated whether IGF-1 levels during molting correlate with the expression of multiple ornaments in a sexually dichromatic passerine species, the bearded reedling ( Panurus biarmicus ). We collected blood samples of males and females shortly before the molting completed and measured the size and colors of ornamental traits. Our results indicate that in males, structural plumage colors, the size of the melanin-based ornament (beard), and tail length are independent traits. IGF-1 levels are associated with the length of the tail and the expression of male structural plumage components (UV coloration), but not the melanin-based ornament. In females, plumage color and tail length were independent traits, which were not related to IGF-1 levels. To the best of our knowledge, this study provides the first evidence that IGF-1 could play a role in the development of secondary sexual characters in a bird species. Significance statement IGF-1 is an evolutionarily highly conserved peptide hormone, which recently entered the center stage of research enquiry in evolutionary biology. It is considered as one of the key factors shaping individual life histories, but little is known about its effects on sexually selected traits. We investigated whether IGF-1 levels during molting predict the elaboration of multiple ornamental plumage traits in male and female bearded reedlings ( Panurus biarmicus ). Our results indicate that higher IGF-1 levels had positive effects on male structural plumage colors and tail feather length. This is the first study, bringing indication for a potential role of IGF-1 in the expression of plumage ornaments in a bird species. Our findings suggest that IGF-1 might serve as an ideal candidate to study the mechanisms linking condition and the capacity to develop sexually selected ornaments.

Avian malaria (caused by Plasmodium spp.) and avian malaria-like infections (caused by Haemoproteus spp.) are widespread and can seriously affect the health of their bird hosts, especially of immunologically naïve individuals. Therefore, these parasites have long been in the focus of bird-parasite studies. However, the species richness and diversity of these protozoan species have only been revealed since the use of molecular techniques. Diversity and prevalence of these parasites among different bird species and even between populations of a species show a large variation. Here, we investigated prevalence of avian malaria and avian malaria-like parasites in two distant populations of a non-migratory wetland specialist passerine, the bearded reedling (Panurus biarmicus). While previous studies have shown that reed-dwelling bird species often carry various blood parasite lineages and the presence of the vectors transmitting Plasmodium and Haemoproteus species has been confirmed from our study sites, prevalence of these parasites was extremely low in our populations. This may either suggest that bearded reedlings may avoid or quickly clear these infections, or these parasites cause high mortality in this species. The remarkably low prevalence of infection in this species is consistent with earlier studies and makes bearded reedlings a possible model organism for investigating the genetic or behavioural adaptations of parasite resistance.

Physiological state is an emergent property of the interactions among physiological systems within an intricate network. Understanding the connections within this network is one of the goals in physiological ecology. Here, we studied the relationship between body condition, two neuroendocrine hormones (corticosterone and insulin-like growth factor 1 [IGF-1]) as physiological regulators, and two physiological systems related to resource metabolism (glucose) and oxidative balance (malondialdehyde). We measured these traits under baseline and stress-induced conditions in free-living house sparrows (Passer domesticus). We used path analysis to analyze different scenarios about the structure of the physiological network. Our data were most consistent with a model in which corticosterone was the major regulator under baseline conditions. This model shows that individuals in better condition have lower corticosterone levels; corticosterone and IGF-1 levels are positively associated; and oxidative damage is higher when levels of corticosterone, IGF-1, and glucose are elevated. After exposure to acute stress, these relationships were considerably reorganized. In response to acute stress, birds increased their corticosterone and glucose levels and decreased their IGF-1 levels. However, individuals in better condition increased their corticosterone levels more and better maintained their IGF-1 levels in response to acute stress. The acute stress–induced changes in corticosterone and IGF-1 levels were associated with an increase in glucose levels, which in turn was associated with a decrease in oxidative damage. We urge ecophysiologists to focus more on physiological networks, as the relationships between physiological traits are complex and dynamic during the organismal stress response.

Background Physiological regulators of life history trade-offs need to be responsive to sudden changes of resource availability. When homeostasis is challenged by unpredictable stressors, vertebrates respond through a set of physiological reactions, which can promote organismal survival. Glucocorticoids have been traditionally recognized as one of the main regulators of the physiological stress response, but the role of an evolutionarily more conserved pathway, the hypothalamic-pituitary-somatotropic (HPS) axis producing insulin-like growth factor-1 (IGF-1) has received much less attention. Although IGF-1 is known to affect several life history traits, little is known about its role in the physiological stress response and it has never been studied directly in adult wild animals. Methods In this study, we combined field observations with a controlled experiment to investigate how circulating levels of IGF-1 change in response to stress and whether this change is due to concomitant change in glucocorticoids in a free-living songbird, the bearded reedling Panurus biarmicus . We used a standard capture-restraint protocol in field observation, in which we took first and second (stress induced: 15 minutes later) samples. In a follow-up experiment, we used a minimally invasive oral corticosterone manipulation. Results We showed that corticosterone levels significantly increased while IGF-1 levels significantly decreased during capture and handling stress. However, change in corticosterone levels were not related to change in IGF-1 levels. We found that experimentally elevated corticosterone levels did not affect IGF-1 levels. Discussion Our results are the first to highlight that circulating IGF-1 levels are responsive to stress independently from glucocorticoids and suggest that the HPS axis is an autonomous physiological pathway that may play an important role as regulator of life-history decisions.

Defensive toxins are widespread in nature, yet we know little about how various environmental factors shape the evolution of chemical defense, especially in vertebrates. In this study we investigated the natural variation in the amount and composition of bufadienolide toxins, and the relative importance of ecological factors in predicting that variation, in larvae of the common toad, Bufo bufo , an amphibian that produces toxins de novo . We found that tadpoles’ toxin content varied markedly among populations, and the number of compounds per tadpole also differed between two geographical regions. The most consistent predictor of toxicity was the strength of competition, indicating that tadpoles produced more compounds and larger amounts of toxins when coexisting with more competitors. Additionally, tadpoles tended to contain larger concentrations of bufadienolides in ponds that were less prone to desiccation, suggesting that the costs of toxin production can only be afforded by tadpoles that do not need to drastically speed up their development. Interestingly, this trade-off was not alleviated by higher food abundance, as periphyton biomass had negligible effect on chemical defense. Even more surprisingly, we found no evidence that higher predation risk enhances chemical defenses, suggesting that low predictability of predation risk and high mortality cost of low toxicity might select for constitutive expression of chemical defense irrespective of the actual level of predation risk. Our findings highlight that the variation in chemical defense may be influenced by environmental heterogeneity in both the need for, and constraints on, toxicity as predicted by optimal defense theory.