Maternal and environmental effects can have profound effects on offspring performance by generating variation in offspring phenotypes, independent of genetic effects. Within avian broods, differential maternal investment of resources across the laying sequence is thought to be an adaptive strategy to modulate competitive hierarchies induced by hatching asynchrony. In this study, we evaluated the relative importance of maternally derived within-clutch variation and the asymmetric post-hatching environment for growth and survival of common tern (Sterna hirundo) siblings. We experimentally manipulated hatching order, resulting in chicks from last-laid eggs hatching first and vice versa. Although both initial age and size asymmetries were larger within experimental than control broods, the early survival of last-hatched chicks was similar between groups. Initial positive effects of egg size disappeared as siblings approached fledging. Ultimately at fledging, both within-brood growth and cumulative survival patterns were similar between experimental and control broods, suggesting that the effects of systematic variation of egg constituents (e.g., maternally derived yolk hormones) and egg size are too subtle and largely overwhelmed by the effects of hatching asynchrony. Therefore, we conclude that variation in offspring phenotypes is pre-dominantly determined by the social environment experienced post-hatching. Maternal effects may further fine-tune phenotypic variation in response to varying environmental conditions, but this needs to be tested through empirical studies in which multiple maternal effects are measured simultaneously under different environmental conditions.

The hormonal response to social challenges has been widely studied, however, most work focused on adult behavior in a reproductive context although developing animals also encounter important social challenges early in life. We studied the relationship between acute sibling competition and plasma corticosterone (CORT) and testosterone (T) in common tern (Sterna hirundo) chicks, a species whose young compete for access to food by scramble interactions. Blood samples were taken in nests with two and only one single chick both immediately after a feeding bout and in non-challenged controls. We found that T levels were lower in siblings challenged by a feeding bout as compared to controls, which may be explained by the fact that T suppresses begging behavior and is only elevated in response to territorial intrusion but not sibling competition in a related species. Singletons had, corrected for body condition, generally lower CORT levels than siblings suggesting that growing up with siblings creates a competitive environment in which high CORT levels are sustained irrespective of a social challenge. CORT levels were also negatively correlated with body condition and were higher in males than in females. The latter may be related to sex-specific food requirements and susceptibility to stress. Our results suggest a possible suppressive effect of acute sibling competition on T secretion, and a positive effect on CORT levels by longer term sibling competition. The degree to which these dynamics are related to begging or aggression, or both, needs further experimental work.

Although sibling competition in avian species has been extensively studied, the proximate mechanisms remain largely unknown. Recent research proposed that steroid hormones, in particular testosterone (T) and corticosterone (CORT), might play a role either in promoting competitive behavioral displays or in response to chronic nutritional stress accompanied by a sustained competitive situation. Here, we examine body condition, endogenous T levels and fecal glucocorticoid metabolites (FGM) as non-invasive measures of CORT in sibling broods of wild common tern chicks (Sterna hirundo) during three post-natal developmental stages. In this species, distinct within-brood size asymmetries are imposed by an asynchronous hatching interval, and sexes show slightly different growth patterns. First-hatched (a-)chicks were in better condition than their later-hatched sibling (b-chick). FGM levels inversely covaried with condition and were elevated at the end of pre-fledging development. T levels of a- and b-chicks changed with age, although the direction of the changes differed, with b-chicks eventually having higher levels than their older siblings. Survival to fledging was not associated with FGM but with T levels, which tended to be higher in surviving chicks. Our results are discussed with regard to how plasticity in steroid hormones could be involved in mediating sibling competition in common terns.

Once a bird has fledged it becomes hardly accessible for researchers and consequently knowledge about post-fledging ontogeny is scarce. In this study on juvenile Common Terns (Sterna hirundo) we used an automated transponder-based detection and weighing system at Banter See colony, Northern Germany, which enabled us to investigate body mass growth of post-fledglings and its consequences for their survival until first return to the natal colony when 2 years old. We analysed data from two contrasting breeding seasons, 2000 and 2001, in order to determine inter-year and inter-sex variation of post-fledging parameters assumed to potentially affect subadult survival, such as the period a juvenile is still present at colony surroundings (departure age), its fledging mass and last recorded post-fledging body mass, and hatch date. Using an information-theoretic model selection approach, neither the date of hatching nor the departure age was found to affect survival. The only predictor of survival was last post-fledging body mass whereas fledging mass itself was of minor importance. Although there was weak evidence for an interaction with year, individuals of the cohort 2000, which left the colony area on average 5 g lighter than those reared under the more favourable conditions in 2001, did not exhibit lower return probability. We suggest that under unfavourable conditions selection had eliminated weak individuals prior to fledging or during the post-fledging period. This study underlines the importance of the post-fledging period and its consequences for survival, especially in species with prolonged parental care post-fledging.

In long-lived vertebrates, individuals generally visit potential breeding areas or populations during one or more seasons before reproducing for the first time. During these years of prospecting, they select a future breeding site, colony, or mate and improve various skills and their physical condition to meet the requirements of reproduction. One precondition of successful reproduction is arrival in time on the breeding grounds. Here, we study the intricate links among the date of initial spring arrival, body mass, sex, and the age of first breeding in the common tern Sterna hirundo, a long-lived migratory colonial seabird. The study is based on a unique, individual-based, long-term dataset of sexed birds, marked with transponders, which allow recording their individual arrival, overall attendance, and clutch initiation remotely and automatically year by year over the entire lifetime at the natal colony site. We show that the seasonal date of initial arrival at the breeding grounds predicts the individual age at first reproduction, which mostly occurs years later. Late first-time arrivals remain delayed birds throughout subsequent years. Our findings reveal that timing of arrival at the site of reproduction and timing of reproduction itself are coherent parameters of individual quality, which are linked with the prospects of the breeding career and may have consequences for fitness.

Many ecological studies use stress hormones to assess the condition, health or disturbance levels of wild organisms. Common blood sampling protocols for this research involve trapping individuals and taking blood within three minutes to obtain a "baseline" for analysis of stress hormones ("conventional method"). In some situations it may be difficult to get an accurate measure of baseline values; therefore, alternative sampling techniques may be preferable. We compared corticosterone levels in samples taken via a newly developed, minimally invasive blood sampling technique with corticosterone levels in blood taken via the conventional method. We collected samples from incubating adult common terns Sterna hirundo via blood sucking bugs (Heteroptera, Triatominae) contained in "dummy eggs" ("bug method") and compared measured corticosterone concentrations to concentrations in blood taken from the same birds using the conventional method. We found no significant differences in mean or variance of baseline corticosterone levels between samples collected via the different methods. This suggests that the bug method offers a viable alternative for hormone sampling.