Tough decisions: Reproductive timing and output vary with individuals' physiology,behavior and past success in a social opportunistic breeder

This article is part of a Special Issue "SBN 2014".Photoperiod and the hormonal response it triggers are key determinants of reproductive timing in birds. However, other cues and physiological traits may permit flexibility in the timing of breeding and perhaps facilitate adaptation to global change. Opportunistic breeders are excellent models to study the adaptive significance of this flexibility, especially at the individual level. Here, we sought to quantify whether particular male physiological and behavioral traits were linked to reproductive timing and output in wild-derived zebra finches. We repeatedly assessed male stress-induced corticosterone levels (CORT), basal metabolic rate (BMR), and activity before releasing them into outdoor aviaries and quantifying each pair's breeding timing, investment, and output over a seven-month period. Despite unlimited access to food and water, the colony breeding activity occurred in waves, probably due to interpair social stimulations. Pairs adjusted their inter-clutch interval and clutch size to social and temperature cues, respectively, but only after successful breeding attempts, suggesting a facultative response to external cues. When these effects were controlled for statistically or experimentally, breeding intervals were repeatable within individuals across reproductive attempts. In addition, males' first laying date and total offspring production varied with complex interactions between pre-breeding CORT, BMR and activity levels. These results suggest that no one trait is under selection but that, instead, correlational selection acts on hormone levels, metabolism, and behavior. Together our results suggest that studying inter-individual variation in breeding strategy and their multiple physiological and behavioral underpinnings may greatly improve our understanding of the mechanisms underlying the evolution of breeding decisions.     

Should I stay or should I go? Hormonal control of nest abandonment in a long-lived bird,the Adélie penguin

According to life-history theory, long-lived birds should favor their survival over the current reproductive attempt, when breeding becomes too costly. In seabirds, incubation is often associated with spontaneous long-term fasting. Below a threshold in body reserves, hormonal and metabolic shift characteristics of a switch from lipid to protein utilization (phase III, PIII) occur. These metabolic changes are paralleled by nest abandonment and stimulation of refeeding behavior. Parental behavior is then under control of two hormones with opposite effects: corticosterone (CORT) and prolactin which stimulate foraging and incubation behavior, respectively.The aim of this study was to determine the respective role of these two hormones in nest abandonment by Adélie penguins. To this end, plasma hormone levels were measured before egg-laying and at departure from the colony (i.e. when birds were relieved by their partner or abandoned their nest), and related to nutritional state and incubation success.We found that males abandoning their nest in PIII presented high CORT levels and low prolactin levels. Interestingly, males which presented high plasma levels of prolactin in PIII did not abandon. We show that although CORT is the first hormone to be affected by prolonged energy constraints, the combined effects of high CORT and low prolactin levels are necessary for parents to favor self-maintenance and abandon the nest. We provide insights into time-course changes of the endocrine profile as PIII proceeds and report that reaching proteolytic late fasting is not sufficient to induce nest abandonment in a long-lived bird.     

Nest Crypsis, Reproductive Value of a Clutch and Escape Decisions in Incubating Female Mallards Anas platyrhynchos

In cryptically coloured birds, remaining on the nest despite predator approach (risk‐taking) may decrease the likelihood that the nest will be detected and current reproductive attempt lost. By contrast, flushing may immediately reveal the nest location to the predator. Escape decisions of incubating parents should therefore be optimized based on the risk‐to‐parent/cost of escape equilibrium. Animal prey may assess predation risk depending on a variety of cues, including the camouflage that vegetation provides against the predator. We examined interactive effects of nest crypsis and the current reproductive value of a clutch on flushing distances in incubating mallards (Anas platyrhynchos) approached by a human. Our results were consistent with predictions of parental investment theory: flushing distances were inversely correlated with measures of the reproductive value of the current clutch, namely with clutch size, stage of incubation and mean egg volume. Independently of a reproductive value of a clutch, nest concealment explained a significant portion of the variation in flushing distance among females; individual females tended to increase/decrease flushing distances according to change in nest cover. The results further suggest that vegetation concealment greatly influenced the risk of nest detection by local predators, suggesting that vegetation may act as a protective cover for incubating female. A female's ability to delay flushes according to the actual vegetation cover might thus be viewed as an antipredator strategy that reduces premature nest advertising to visually oriented predators. We argue, however, that shorter flying distances from densely covered sites might be maladaptive in areas where a predator's ability to detect incubating female does not rely on visual cues of nests.     

Should I stay or should I go? Hormonal control of nest abandonment in a long-lived bird, the Adélie penguin

According to life-history theory, long-lived birds should favor their survival over the current reproductive attempt, when breeding becomes too costly. In seabirds, incubation is often associated with spontaneous long-term fasting. Below a threshold in body reserves, hormonal and metabolic shift characteristics of a switch from lipid to protein utilization (phase III, PIII) occur. These metabolic changes are paralleled by nest abandonment and stimulation of refeeding behavior. Parental behavior is then under control of two hormones with opposite effects: corticosterone (CORT) and prolactin which stimulate foraging and incubation behavior, respectively.The aim of this study was to determine the respective role of these two hormones in nest abandonment by Adélie penguins. To this end, plasma hormone levels were measured before egg-laying and at departure from the colony (i.e. when birds were relieved by their partner or abandoned their nest), and related to nutritional state and incubation success.We found that males abandoning their nest in PIII presented high CORT levels and low prolactin levels. Interestingly, males which presented high plasma levels of prolactin in PIII did not abandon. We show that although CORT is the first hormone to be affected by prolonged energy constraints, the combined effects of high CORT and low prolactin levels are necessary for parents to favor self-maintenance and abandon the nest. We provide insights into time-course changes of the endocrine profile as PIII proceeds and report that reaching proteolytic late fasting is not sufficient to induce nest abandonment in a long-lived bird.     

Food supplementation experiments: A tool to reveal mechanisms that mediate timing of reproduction

Food supplementation of free-living animals has been used to address the role of availability of resources in the timing of reproduction. A meta-analysis by Schoech and Hahn suggested that responsiveness of the reproductive axis to the supplementary cue of food is lessened at higher latitudes, presumably because the brief time during which conditions are appropriate to rear offspring has led to an evolved resistance to supplementary cues with a primary reliance on photoperiod. Unfortunately, few investigators have examined the potential underlying mechanisms that mediate this differential responsiveness to supplemental food across latitudes. Considerable research, however, links nutritional state and plasma glucocorticoid levels, both of which impinge upon the reproductive axis. Long-term research on Florida scrub-jays (Aphelocoma coerulescens) in my laboratory shows that suburban birds with access to ad libitum supplemental food express early breeding and lower plasma corticosterone (CORT) levels in comparison to jays in nearby natural habitat. Furthermore, supplementation in natural habitat advances laying, with the largest effects occurring in bad years (i.e., years defined by late breeding and poor reproductive output by non-supplemented controls). Similarly, reproductive output of supplemented jays is greater and exhibits considerably less variance than do controls, suggesting fitness benefits of supplementation that are tied to advanced breeding. Generally, CORT levels in early-breeding supplemented jays are lower than are those of controls. Also, regression analysis suggests that clutch-initiation dates of non-supplemented female breeders are predicted by baseline CORT levels. Although these data are not conclusive and trends can be obscured by year-effects, they suggest a role for CORT in timing of breeding. Whether this link might help to explain the above-referenced latitudinal trends remains to be characterized.     

Food Supplementation Fails to Reveal a Trade-Off between Incubation and Self-Maintenance in Female House Wrens

Incubating birds must allocate their time and energy between maintaining egg temperature and obtaining enough food to meet their own metabolic demands. We tested the hypothesis that female house wrens (Troglodytes aedon) face a trade-off between incubation and self-maintenance by providing females with supplemental food during incubation. We predicted that food supplementation would increase the amount of time females devoted to incubating their eggs, lower their baseline plasma corticosterone levels (a measure of chronic stress), and increase their body mass, haematocrit (a measure of anaemia), and reproductive success relative to control females. As predicted, food-supplemented females spent a greater proportion of time incubating their eggs than control females. Contrary to expectation, however, there was no evidence that food supplementation significantly influenced female baseline plasma corticosterone levels, body mass, haematocrit, or reproductive success. However, females with high levels of corticosterone at the beginning of incubation were more likely to abandon their nesting attempt after capture than females with low levels. Corticosterone significantly increased between the early incubation and early nestling stages of the breeding cycle in all females. These results suggest that although food supplementation results in a modest increase in incubation effort, it does not lead to significantly lower levels of chronic stress as reflected in lower baseline corticosterone levels. We conclude that female house wrens that begin the incubation period with low levels of plasma corticosterone can easily meet their own nutritional needs while incubating their eggs, and that any trade-off between incubation and self-feeding does not influence female reproductive success under the conditions at the time of our study.     

Red-winged blackbirds (Agelaius phoeniceus) with higher baseline glucocorticoids also invest less in incubation and clutch mass

Glucocorticoid hormones facilitate responses to environmental challenges by mediating diverse physiological and behavioral changes, including resource mobilization and altered reproductive effort. Elevated glucocorticoids might indicate that an individual is facing high levels of environmental challenges and thus, elevated concentrations might be associated with reduced fitness (CORT-fitness hypothesis). Alternatively, the energetic demands of reproduction might be a challenge that requires elevated glucocorticoids to mobilize resources to support reproductive effort, ultimately increasing reproductive investment and fitness (CORT-adaptation hypothesis). Investigations of glucocorticoid-fitness relationships have yielded mixed results. Variation in the direction of this relationship could be caused in part by differences in the contexts in which the relationship was assessed. Incorporating context, such as life history stage, could be key to understanding the role of glucocorticoids in influencing fitness outcomes. We investigated the relationship between corticosterone and reproductive effort and success within a single life history stage: incubation of eggs. In an observational study, we measured baseline corticosterone in incubating female red-winged blackbirds (Agelaius phoeniceus), monitored incubation behavior, and determined hatching success for each nest. Incubating birds with higher baseline corticosterone concentrations had more frequent, shorter incubation bouts and spent less time overall incubating their clutches of eggs than birds with lower corticosterone concentrations. Elevated corticosterone was also associated with lower clutch mass, but neither corticosterone nor incubation effort were correlated with hatching success. Although experimental tests are needed to establish causation, these results suggest that during the incubation period, corticosterone might shift resource investment towards self-maintenance, and away from current reproductive effort.     

Incubation patterns in a central-place forager affect lifetime reproductive success: scaling of patterns from a foraging bout to a lifetime

Background

Long-lived seabirds face a conflict between current and lifelong reproductive success. During incubation shifts, egg neglect is sometimes necessary to avoid starvation, but may compromise the current reproductive attempt. However, factors underlying this decision process are poorly understood. We focus on the ancient murrelet, Synthliboramphus antiquus, an alcid with exceptionally long incubation shift lengths, and test the impact of environmental factors on incubation shift length in relation to reproductive success.

Methodology/Principal Findings

Using an information theoretic approach, we show that incubation shift length was a strong predictor of reproductive success for ancient murrelets at Reef Island, Haida Gwaii, British Columbia, Canada during the 2007 and 2008 breeding seasons. The most important factors explaining an individual''s shift length were egg size, wind speed and the length of the mate''s previous shift. Wind speed and tide height were the two most important factors for determining foraging behavior, as measured by dive frequency and depth.

Conclusions/Significance

Our study demonstrates that (i) species-specific reproductive strategies interact with environmental conditions such as wind speed to form multiple incubation patterns and (ii) maintaining regular incubation shifts is an essential component of reproductive success.     

Incubation and brooding performance of the Antarctic Petrel Thalassoica antarctica at Svarthamaren, Dronning Maud Land

Antarctic Petrel Thalassoica antarctica incubation and brooding effort was studied at Svarthamaren, Dronning Maud Land, during the austral summer of 1991–1992. The females probably left the nest site shortly after egg laying. The duration of incubation and brooding shifts as well as the daily weight loss (absolute and proportionate) were comparable with those of other similar-sized procellariform species. Males spent more time incubating and brooding than did females, suggesting higher female energy stress due to egg laying. Incubating birds which were below average weight were likely to desert the nests before their mates returned from feeding trips. Both males and females lost approximately one-fifth of their body-weight during their first incubation shifts. Nevertheless, they increased their initial weights from egg laying to hatching and had their highest initial weights when they returned to start the shift during which the egg hatched. No factors related to adult body-weight explained the duration of the incubation shifts. Both males and females gained weight at a higher rate when at sea than they lost it during incubation, and it is suggested that factors unrelated to food availability or individual feeding skills may be important in regulating the duration of the incubation shifts and the stay at sea.     

Effects of Supplementary Feeding on the Breeding Ecology of the Buff-Throated Partridge in a Tibetan Sacred Site,China

Our goal was to document effects of year-round supplemental feeding on breeding ecology of the Buff-throated Partridge, Tetraophasis szechenyii, within a Tibetan sacred site. We evaluated effects of supplemental feeding used as religious/cultural practices which could potentially aid conservation of endangered phasianids. We compared fed breeding groups to neighboring nonfed groups. Fed groups initiated first clutches significantly earlier than nonfed groups. Earlier laying groups within fed and nonfed groups showed significantly lower hatching rates than later groups; however, fed groups showed significantly higher hatching rates than nonfed groups laying in the same period. Earlier laying increased opportunities to renest. All six fed groups with clutch failures renested compared to only one of five nonfed groups with clutch failures. Fed female breeders showed significantly greater investment in their young with larger clutches and larger eggs, which likely increased survivability of early hatchlings. We observed no predation on birds at feeding sites and recorded only four cases of predation on incubating females, which showed no detectable difference between fed and nonfed groups. Ground-nesting birds typically face high risks of predation. Ten of the 48 groups nested in trees, which occurs in few phasianid species. Tree nests showed significantly higher hatching rates compared to ground nests; however, we found no significant difference in tree nesting between fed and nonfed groups. This partridge is one of four gallinaceous species with cooperative breeding. Breeding groups with helpers had significantly greater reproductive success than single pairs, and fed female breeders with helpers laid bigger eggs than single pairs. Comparing annual reproductive output per group, fed groups not only produced significantly more independent young (≥150 days post-hatching), their young hatched significantly earlier, which likely have greater reproductive value over later hatched young of nonfed groups. Supplemental feeding year-round is likely what enabled the successes of the fed partridges.     

Divorce in the socially monogamous zebra finch: Hormonal mechanisms and reproductive consequences

Up to 80% of all bird species are socially monogamous. Divorce (switching partners) or pair disruption (due to the death of a partner) has been associated with decreased reproductive success, suggesting social monogamy is a strategy that may maximize fitness via coordination between partners. Previous studies have demonstrated the effects of divorce and pair disruption on immediate reproductive success. Here, we used a paired experimental design in the zebra finch (Taeniopygia guttata) to examine the hormonal mechanisms that modulate parental behavior and reproductive success in response to a partnership change (hereafter divorce). Specifically, we examined the effects of divorce on the avian stress hormone corticosterone (CORT) in both parents and nestlings, parental behaviors (incubation and nestling provisioning), prolactin (PRL), and reproductive success. We found that divorce resulted in delayed clutch initiation, reduced clutch mass, and an increase in nestling CORT response to a standardized stressor. These effects on reproductive investment and chick CORT response were not clearly determined by parental endocrine responses. Divorce had no effect on the level of parental CORT. PRL levels were highly correlated within a pair regardless of treatment, were negatively related to the investment that males made in incubation, and increased in experimental males as a result of pair disruption. This study demonstrates the fundamental impact which divorce has not only on reproduction, but also the physiological stress responses of offspring and suggests that in socially monogamous animals the maintenance of a stable partnership over time could be advantageous for long term fitness.     

Hypothalamic monoamine turnover in ring doves (streptopelia risoria), courting,incubating and brooding young

1. The turnovers of hypothalamic 5-hydroxytryptamine (5HT), dopamine (DA) and noradrenaline (NE) were measured in male and female ring doves (Streptopelia risoria) at three stages of the breeding cycle: courtship, 3 days after pairing; early incubation, 1–2 days after egg laying; and brooding, 1–3 days after the squabs had hatched.2. In both sexes plasma LH decreased progressively from courtship through incubation to brooding young. Crop sacs were fully developed in doves brooding young but not at other stages of the reproductive cycle, indicating increased concentrations of plasma prolactin.3. The turnovers of 5HT and DA in both sexes were significantly higher (P<0.001) in doves brooding young than in birds incubating eggs or nest building. The turnover of DA was higher in females than in males at the onset of incubation. The turnover of NE was lower (P > 0.01) in females at the onset of incubation than during courtship or brooding.4. Increased turnover of hypothalamic DA may be more closely related to brooding behaviour than to changes in prolactin or LH secretion.5. Increased hypothalamic 5HT turnover in brooding doves appears to be more directly related to crop sac development, and by inference increased prolactin secretion, than to depressed plasma LH concentrations.     

Clutch predation in great tinamous Tinamus major and implications for the evolution of egg color

Egg camouflage has been found to reduce predation in several ground‐nesting species. Therefore, the evolution of eggs that lack camouflage in ground nesting birds is puzzling. Even though clutch predation in the tropics is high, tinamous are the only tropical ground‐nesting birds that do not build a nest and do not lay cryptic eggs. I studied predation of great tinamou clutches in a lowland tropical forest and found that risk of predation was higher during incubation when the eggs are covered by the parent, than during laying when they are exposed, suggesting that predators primarily use cues from the incubating males to locate the clutch and not cues from the eggs. Clutch size had no effect on predation rate, even though larger clutches are more conspicuous to a human observer. Predation by visual cues is likely reduced during incubation by the camouflaged plumage and high nest attendance of males. If most predators use cues from the incubating male and not the eggs to locate clutches, then conspicuous egg color may have evolved in great tinamous as an intra‐specific signal. I evaluate hypotheses that may explain the maintenance of conspicuous egg color in tinamous.     

Corticosterone manipulations alter morph-specific nestling provisioning behavior in male white-throated sparrows,Zonotrichia albicollis

In the polymorphic white-throated sparrow (Zonotrichia albicollis), tan-striped males provision nestlings at higher rates than do white-striped males. In a previous study, we found that tan-striped males had lower baseline corticosterone levels than white-striped males during the nestling stage. To determine if this variation in corticosterone influences morph-specific differences in nestling provisioning behavior, we used intraperitoneal osmotic pumps to increase baseline corticosterone levels in tan-striped males (TS CORT) and administer RU486, a glucocorticoid receptor antagonist, in white-striped males (WS RU486). These manipulations essentially reversed morph-specific nestling provisioning behavior in males. TS CORT males fed nestlings at lower rates than TS controls (vehicle-only implant), and at similar rates to WS controls (vehicle-only implant), while WS RU486 males fed nestlings at higher rates than WS controls, and at similar rates to TS controls. These results demonstrate that (1) increases in baseline corticosterone (i.e., below concentrations associated with the adrenocortical response to stress) can directly or indirectly inhibit nestling provisioning behavior, and (2) corticosterone influences morph-specific variation in parental behavior in male white-throated sparrows. This study contributes to the growing evidence that modulating baseline CORT mediates parental care and self-maintenance activities in birds, and thus may serve as a mechanism for balancing current reproductive success with survival.     

Elevated plasma corticosterone increases metabolic rate in a terrestrial salamander

Plasma glucocorticoid hormones (GCs) increase intermediary metabolism, which may be reflected in whole-animal metabolic rate. Studies in fish, birds, and reptiles have shown that GCs may alter whole-animal energy expenditure, but results are conflicting and often involve GC levels that are not physiologically relevant. A previous study in red-legged salamanders found that male courtship pheromone increased plasma corticosterone (CORT; the primary GC in amphibians) concentrations in males, which could elevate metabolic processes to sustain courtship behaviors. To understand the possible metabolic effect of elevated plasma CORT, we measured the effects of male courtship pheromone and exogenous application of CORT on oxygen consumption in male red-legged salamanders (Plethodon shermani). Exogenous application of CORT elevated plasma CORT to physiologically relevant levels. Compared to treatment with male courtship pheromone and vehicle, treatment with CORT increased oxygen consumption rates for several hours after treatment, resulting in 12% more oxygen consumed (equivalent to 0.33 J) during our first 2 h sampling period. Contrary to our previous work, treatment with pheromone did not increase plasma CORT, perhaps because subjects used in this study were not in breeding condition. Pheromone application did not affect respiration rates. Our study is one of the few to evaluate the influence of physiologically relevant elevations in CORT on whole-animal metabolism in vertebrates, and the first to show that elevated plasma CORT increases metabolism in an amphibian.     

Risk-taking by incubating rufous bush robins Cercotrichas galactotes: season-dependent incubation stage effect

Risk-taking by incubating birds is commonly assumed to increase with reproductive stage due to increased nest value, and also with time of the breeding season due to decreased renesting opportunity. Nonetheless, the potential for these two factors to interact has not been given much importance. However, the extent to which risk-taking may increase with reproductive stage could be expected to vary with season. We investigated this issue using data on rufous bush robins (Cercotrichas galactotes) that we experimentally flushed from their nests at different stages of incubation and at different dates in the breeding season. We found that flushing distance of incubating birds decreased significantly with both incubation stage and season. More interestingly, we also found that incubation stage and season interact in shaping risk-taking the behaviour of incubating birds. As the breeding season progresses, and renesting opportunity declines, the effect of incubation stage on incubating parent risk-taking decreases. Overall, our work once again underlines the great complexity of behavioural and ecological factors and processes that affect risk-taking by nesting birds. In particular, it stresses the need to consider the interaction between reproductive stage and season when dealing with nest-defence behaviour in birds.     

Incubation behavior adjustments,driven by ambient temperature variation,improve synchrony between hatch dates and caterpillar peak in a wild bird population

For organisms living in seasonal environments, synchronizing the peak energetic demands of reproduction with peak food availability is a key challenge. Understanding the extent to which animals can adjust behavior to optimize reproductive timing, and the cues they use to do this, is essential for predicting how they will respond to future climate change. In birds, the timing of peak energetic demand is largely determined by the timing of clutch initiation; however, considerable alterations can still occur once egg laying has begun. Here, we use a wild population of great tits (Parus major) to quantify individual variation in different aspects of incubation behavior (onset, duration, and daily intensity) and conduct a comprehensive assessment of the causes and consequences of this variation. Using a 54‐year dataset, we demonstrate that timing of hatching relative to peak prey abundance (synchrony) is a better predictor of reproductive success than clutch initiation or clutch completion timing, suggesting adjustments to reproductive timing via incubation are adaptive in this species. Using detailed in‐nest temperature recordings, we found that postlaying, birds improved their synchrony with the food peak primarily by varying the onset of incubation, with duration changes playing a lesser role. We then used a sliding time window approach to explore which spring temperature cues best predict variance in each aspect of incubation behavior. Variation in the onset of incubation correlated with mean temperatures just prior to laying; however, incubation duration could not be explained by any of our temperature variables. Daily incubation intensity varied in response to daily maximum temperatures throughout incubation, suggesting female great tits respond to temperature cues even in late stages of incubation. Our results suggest that multiple aspects of the breeding cycle influence the final timing of peak energetic demand. Such adjustments could compensate, in part, for poor initial timing, which has significant fitness impacts.     

Exogenous corticosterone and nest abandonment: A study in a long-lived bird, the Adélie penguin

Breeding individuals enter an emergency life-history stage when their body reserves reach a minimum threshold. Consequently, they redirect current activity toward survival, leading to egg abandonment in birds. Corticosterone (CORT) is known to promote this stage. How and to what extent CORT triggers egg abandonment when breeding is associated with prolonged fasting, however, requires further investigation.We manipulated free-living male Adélie penguins with CORT-pellets before their laying period. We then examined their behavioral response with respect to nest abandonment in parallel with their prolactin levels (regulating parental care), and the subsequent effects of treatment on breeding success in relieved birds.Exogenous CORT triggered nest abandonment in 60% of the treated penguins ~ 14 days after treatment and induced a concomitant decline in prolactin levels. Interestingly, prolactin levels in treated penguins that did not abandon their nest were higher at the point of implantation and also after being relieved by females, when compared with abandoning penguins. Among successful birds, the treatment did not affect the number of chicks, nor the brood mass.Our results show the involvement of CORT in the decision-making process regarding egg abandonment in Adélie penguins when incubation is associated with a natural long fast. However, we suggest that CORT alone is not sufficient to trigger nest abandonment but that 1) prolactin levels need to reach a low threshold value, and 2) a rise in proteolysis (i.e. utilization of protein as main energy substrate) seems also to be required.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

Stress hormones in relation to breeding status and territory location in colonial king penguin: a role for social density?

Because glucocorticoid (stress) hormones fundamentally affect various aspects of the behaviour, life history and fitness of free-living vertebrates, there is a need to understand the environmental factors shaping their variation in natural populations. Here, we examined whether spatial heterogeneity in breeding territory quality affected the stress of colonial king penguin (Aptenodytes patagonicus). We assessed the effects of local climate (wind, sun and ambient temperature) and social conditions (number of neighbours, distance to neighbours) on the baseline levels of plasma total corticosterone (CORT) in 77 incubating and 42 chick-brooding birds, breeding on territories of central or peripheral colony location. We also assessed the oxidative stress status of a sub-sample of central vs. peripheral chick-brooders to determine whether chronic stress arose from breeding on specific territories. On average, we found that brooders had 55 % higher CORT levels than incubators. Regardless of breeding status, central birds experienced greater social density (higher number of neighbours, shorter distance between territories) and had higher CORT levels than peripheral birds. Increasing social density positively explained 40 % of the variation in CORT levels of both incubators and brooders, but the effect was more pronounced in brooders. In contrast, climate was similar among breeding territories and did not significantly affect the CORT levels of breeding birds. In brooders, oxidative stress status was not affected by local density or weather conditions. These results highlight that local heterogeneity in breeding (including social) conditions may strongly affect the stress levels of breeding seabirds. The fitness consequences of such variation remain to be investigated.