Drivers of within- and among-individual variation in risk-taking behaviour during reproduction in a long-lived bird.

Plastic and selective mechanisms govern parental investment adjustments to predation threat. We investigated the relative importance of plasticity and selection in risk-taking propensity of incubating female common eiders Somateria mollissima facing unprecedented predation in SW Finland, Baltic Sea. Using a 12-year individual-based dataset, we examined within- and among-individual variation in flight initiation distance (FID), in relation to predation risk, nest detectability, individual traits and reproductive investment (NFID = 1009; Nindividual = 559). We expected females nesting in riskier environments (higher predation risk, lower nest concealment) to mitigate environmentally imposed risk by exhibiting longer FIDs, and females investing more in current reproduction (older, in better condition or laying larger clutches) to display shorter FIDs. The target of predation-adult or offspring-affected the mechanisms adapting risk-taking propensity; females plastically increased their FID under higher adult predation risk, while risk-avoiding breeders were predominant on islands with higher nest predation risk. Risk-taking females selected thicker nest cover, consistent with personality-matching habitat choice. Females plastically attenuated their anti-predator response (shorter FIDs) with advancing age, and females in better body condition were more risk-taking, a result explained by selection processes. Future research should consider predator type when investigating the fitness consequences of risk-taking strategies.

Mitigating impacts of invasive alien predators on an endangered sea duck amidst high native predation pressure.

Anthropogenically introduced invasive species represent a major threat to global biodiversity by causing population declines and extinctions of native species. The negative impacts of introduced predators are well documented, yet a fundamental knowledge gap exists regarding the efficiency of potential mitigation methods to restore the ecosystem. Other understudied aspects concern prey behavioural antipredator responses and the historical context of native predator-prey interactions, which may moderate invasion impacts on native prey. Invasion impacts of American mink (Neovison vison) and raccoon dog (Nyctereutes procyonoides) into the Baltic Sea archipelago are poorly understood, and the efficiency of removal efforts as a means to alleviate depredation pressure on native prey is debated. Here, we examine the effectiveness of invasive predator removal on ground-nesting female common eider (Somateria mollissima) mortality, breeding success and breeding propensity over a 9-year period, while controlling for predation risk imposed by the main native predator, the white-tailed eagle (Haliaeetus albicilla). Our results clearly show that intensified removal of American minks and raccoon dogs decreased the number of female eiders killed during nesting, while improving both nesting success and breeding propensity. Such obvious positive effects of invasive predator removal are particularly noteworthy against the backdrop of a soaring eagle population, indicating that the impacts of invasives may become accentuated when native predators differ taxonomically and by hunting mode. This study shows that invasive alien predator removal is an effective conservation measure clearly aiding native fauna even under severe native predation pressure. Such cost-effective conservation actions call for governmental deployment across large areas.

Sex-specific effects of the in ovo environment on early-life phenotypes in eiders.

Maternal effects affect offspring phenotype and fitness. However, the roles of offspring sex-specific sensitivity to maternal glucocorticoids and sex-biased maternal investment remain unclear. It is also uncertain whether telomere length (a marker associated with lifespan) depends on early growth in a sex-specific manner. We assessed whether maternal traits including corticosterone (CORT; the main avian glucocorticoid) and in ovo growth rate are sex-specifically related to offspring CORT exposure, relative telomere length (RTL) and body condition in eiders (Somateria mollissima). We measured feather CORT (fCORT), RTL and body condition of newly hatched ducklings, and growth rate in ovo was expressed as tarsus length at hatching per incubation duration. Maternal traits included baseline plasma CORT, RTL, body condition and breeding experience. We found that fCORT was negatively associated with growth rate in daughters, while it showed a positive association in sons. Lower offspring fCORT was associated with higher maternal baseline plasma CORT, and fCORT was higher in larger clutches and in those hatching later. The RTL of daughters was negatively associated with maternal RTL, whereas that of males was nearly independent of maternal RTL. Higher fCORT in ovo was associated with longer RTL at hatching in both sexes. Duckling body condition was mainly explained by egg weight, and sons had a slightly lower body condition. Our correlational results suggest that maternal effects may have heterogeneous and even diametrically opposed effects between the sexes during early development. Our findings also challenge the view that prenatal CORT exposure is invariably associated with shorter telomeres.

Annual variation in predation risk is related to the direction of selection for brain size in the wild.

The direction of predator-mediated selection on brain size is debated. However, the speed and the accuracy of performing a task cannot be simultaneously maximized. Large-brained individuals may be predisposed to accurate but slow decision-making, beneficial under high predation risk, but costly under low risk. This creates the possibility of temporally fluctuating selection on brain size depending on overall predation risk. We test this idea in nesting wild eider females (Somateria mollissima), in which head volume is tightly linked to brain mass (r2 = 0.73). We determined how female relative head volume relates to survival, and characterized the seasonal timing of predation. Previous work suggests that relatively large-brained and small-brained females make slow versus fast nest-site decisions, respectively, and that predation events occur seasonally earlier when predation is severe. Large-brained, late-breeding females may therefore have higher survival during high-predation years, but lower survival during safe years, assuming that predation disproportionately affects late breeders in such years. Relatively large-headed females outsurvived smaller-headed females during dangerous years, whereas the opposite was true in safer years. Predation events occurred relatively later during safe years. Fluctuations in the direction of survival selection on relative brain size may therefore arise due to brain-size dependent breeding phenology.

To breed or not to breed: drivers of intermittent breeding in a seabird under increasing predation risk and male bias.

Intermittent breeding may be adaptive for long-lived species subjected to large accessory reproductive costs, but it may also reflect reduced adaptation to the environment, reducing population growth. Nevertheless, environmental influences on breeding propensity, particularly that of predation risk, remain poorly understood and difficult to study, because non-breeders are typically not identified. Female eiders Somateria mollissima from the Baltic Sea provide an excellent testbed, because nesting females have been exposed to intensifying predation and growing male bias that may increase female harassment. We based our study on long-term data (14 years) on females captured and marked at the nest, and females individually identified at sea irrespective of capture status. We hypothesized that breeding propensity decreases with increasing predation risk and male bias, and increases with breeder age. Consistent with our hypotheses, females nesting on islands with higher nest predation risk were more likely to skip breeding, and breeding probability increased with age. In contrast, the steep temporal decline in breeding propensity could not be reliably attributed to annual adult sex ratio or to the abundance of white-tailed sea eagles (Haliaeetus albicilla), the main predator on females, at the nearby Hanko Bird Observatory. Breeding probability showed significant consistent individual variation. Our results demonstrate that spatiotemporal variation in predation risk affects the decision to breed and high incidence of non-breeding was associated with low fledging success. The increased frequency of intermittent breeding in this declining population should be explicitly considered in demographic models, and emphasis placed on understanding the preconditions for successful reproduction.

Increased male bias in eider ducks can be explained by sex-specific survival of prime-age breeders.

In contrast to theoretical predictions of even adult sex ratios, males are dominating in many bird populations. Such bias among adults may be critical to population growth and viability. Nevertheless, demographic mechanisms for biased adult sex ratios are still poorly understood. Here, we examined potential demographic mechanisms for the recent dramatic shift from a slight female bias among adult eider ducks (Somateria mollissima) to a male bias (about 65% males) in the Baltic Sea, where the species is currently declining. We analysed a nine-year dataset on offspring sex ratio at hatching based on molecularly sexed ducklings of individually known mothers. Moreover, using demographic data from long-term individual-based capture-recapture records, we investigated how sex-specific survival at different ages after fledgling can modify the adult sex ratio. More specifically, we constructed a stochastic two-sex matrix population model and simulated scenarios of different survival probabilities for males and females. We found that sex ratio at hatching was slightly female-biased (52.8%) and therefore unlikely to explain the observed male bias among adult birds. Our stochastic simulations with higher survival for males than for females revealed that despite a slight female bias at hatching, study populations shifted to a male-biased adult sex ratio (> 60% males) in a few decades. This shift was driven by prime reproductive-age individuals (≥5-year-old), with sex-specific survival of younger age classes playing a minor role. Hence, different age classes contributed disproportionally to population dynamics. We argue that an alternative explanation for the observed male dominance among adults-sex-biased dispersal-can be considered redundant and is unlikely, given the ecology of the species. The present study highlights the importance of considering population structure and age-specific vital rates when assessing population dynamics and management targets.

Blood and feather concentrations of toxic elements in a Baltic and an Arctic seabird population.

We report blood and feather concentrations of elements in the Baltic Sea and Arctic population of common eiders (Somateria mollissima). The endangered Baltic Sea population of eiders was demonstrably affected by element pollution in the 1990s. While blood concentrations of Hg were higher in Baltic breeding eiders, blood Se, As and Cd concentrations were higher in Arctic eiders. Blood concentrations of Pb, Cr, Zn and Cu did not differ between the two populations. While blood Pb concentrations had declined in Baltic eiders since the 1990s, Hg concentrations had not declined, and were above concentrations associated with adverse oxidative effects in other bird species. Inconsistent with blood concentrations, feather concentrations suggested that Pb, Zn, and Cd exposure was higher in Baltic eiders, and that Hg exposure was higher in Arctic eiders. Our study thus emphasizes the need for comprehensive evaluation of toxic element status, covering the annual cycle of a species.

DNA double-strand breaks in incubating female common eiders (Somateria mollissima): Comparison between a low and a high polluted area.

Alterations in the genetic material may have severe consequences for individuals and populations. Hence, genotoxic effects of environmental exposure to pollutants are of great concern. We assessed the impact of blood concentrations of persistent organic pollutants (POPs) and mercury (Hg) on DNA double-strand break (DSB) frequency, in blood cells of a high-exposed Baltic, and lower exposed Arctic population of common eiders (Somateria mollissima). Furthermore, we examined whether the genotoxic response was influenced by antioxidant concentration (plasma total glutathione (tGSH) and total antioxidant capacity) and female body mass. The DNA DSB frequency did not differ between the two populations. We found significant positive relationships between Hg and DNA DSB frequency in Baltic, but not in Arctic eiders. Although both p,p'-DDE and PCB 118 had a lesser effect than Hg, they exhibited a positive association with DNA DSB frequency in Baltic eiders. Antioxidant levels were not important for the genotoxic effect, suggesting alternative mechanisms other than GSH depletion for the relationship between Hg and DNA DSBs. Hence, the Baltic population, which is considered to be endangered and is under the influence of several environmental stressors, may be more susceptible to genotoxic effects of environmental exposure to Hg than the Arctic population.

State-dependent capital and income breeding: a novel approach to evaluating individual strategies with stable isotopes.

BACKGROUND: Species-specific strategies for financing the costs of reproduction are well understood, forming a continuum ranging from high to low reliance on stored nutrients. Animals relying mostly on stored reserves are termed 'capital breeders', whereas 'income breeders' rely mostly on concurrent intake when financing the costs of reproduction. The role and adaptive value of individual variation in these strategies remain elusive. Life-history theory posits that capital breeding should be favoured when offspring reproductive value peaks, typically occurring early in the season, and that current income should increasingly be used with progressing season. Because resource limitation may hamper flexible resource allocation, a corollary prediction is that only good-condition individuals may show the expected seasonal shift in resource use. To test this prediction, we examined stable isotopes (δ(13)C and δ(15)N) in blood and lipid-free egg yolk of breeding eider females (Somateria mollissima) from the Baltic Sea to assess the role of individual variation in the use of proteins from local diet vs. stored reserves. RESULTS: We show for the first time that individuals from a single population differ in their utilization of stored reserves and concurrent intake to finance the costs of reproduction. Consistent with our prediction, heavy females predominantly used stored reserves for producing egg yolks early in the season, increasingly relying on local feeding with later onset of breeding, whereas light females showed no seasonal change in allocation strategy. CONCLUSIONS: Stable isotope profiling at the individual level is a powerful tool for monitoring relative changes in investment strategies through time, showing promise as an early warning indicator of ecological change in food webs.

Reproductive investment is connected to innate immunity in a long-lived animal.

Life-history theory predicts that organisms optimize their resource allocation strategy to maximize lifetime reproductive success. Individuals can flexibly reallocate resources depending on their life-history stage, and environmental and physiological factors, which lead to variable life-history strategies even within species. Physiological trade-offs between immunity and reproduction are particularly relevant for long-lived species that need to balance current reproduction against future survival and reproduction, but their underlying mechanisms are poorly understood. A major unresolved issue is whether the first-line innate immune function is suppressed by reproductive investment. In this paper, we tested if reproductive investment is associated with the suppression of innate immunity, and how this potential trade-off is resolved depending on physiological state and residual reproductive value. We used long-lived capital-breeding female eiders (Somateria mollissima) as a model. We showed that the innate immune response, measured by plasma bacteria-killing capacity (BKC), was negatively associated with increasing reproductive investment, i.e., with increasing clutch size and advancing incubation stage. Females in a better physiological state, as indexed by low heterophil-to-lymphocyte (H/L) ratios, showed higher BKC during early incubation, but this capacity decreased as incubation progressed, whereas females in poorer state showed low BKC capacity throughout incubation. Although plasma BKC generally declined with increasing H/L ratios, this decrease was most pronounced in young females. Our results demonstrate that reproductive investment can suppress constitutive first-line immune defence in a long-lived bird, but the degree of immunosuppression depends on physiological state and age.

Antioxidant Responses in Relation to Persistent Organic Pollutants and Metals in a Low- and a High-Exposure Population of Seabirds.

Oxidative stress occurs when there is an imbalance between the production of reactive oxygen species (ROS) and antioxidant defense. Exposure to pollutants may increase ROS and affect antioxidant levels, and the resulting oxidative stress may negatively affect both reproduction and survival. We measured concentrations of 18 persistent organic pollutants (POPs) and 9 toxic elements in blood, as well as total antioxidant capacity (TAC), total glutathione (tGSH), and carotenoids in plasma of Baltic and Arctic female common eiders (Somateria mollissima) (N = 54) at the end of their incubation-related fasting. The more polluted Baltic population had higher TAC and tGSH concentrations compared to the Arctic population. Carotenoid levels did not differ between populations. The effect of mixtures of pollutants on the antioxidants was assessed, and the summed molar blood concentrations of 14 POPs were positively related to TAC. There was no significant relationship between the analyzed pollutants and tGSH concentrations. The adaptive improvement of the antioxidant defense system in the Baltic population may be a consequence of increased oxidative stress. However, both increased oxidative stress and energy allocation toward antioxidant defense may have adverse consequences for Baltic eiders at the incubation stage, when energy resources reach an annual minimum due to incubation-related fasting.

Persistent organic pollutant levels and the importance of source proximity in Baltic and Svalbard breeding common eiders.

The distance to sources and the long-range transport potential of persistent organic pollutants (POPs) are important in understanding the impact of anthropogenic pollution on natural seabird populations. The present study documented blood concentrations of POPs in the Baltic Sea (Tvärminne, Finland) population of common eiders (Somateria mollissima) in 2009 and in 2011 and compared the concentrations with the presumably less exposed Arctic population in Svalbard (Kongsfjorden, Norway). The Baltic population had 26, 10, and 5 times greater concentrations of hexachlorocyclohexane, polychlorinated biphenyls, and p,p'-dichlorodiphenyldichloroethylene than the Svalbard population. Unexpectedly, concentrations of chlordanes were higher in Svalbard eiders, whereas concentrations of hexachlorobenzenes (HCBs) did not differ between the 2 populations. Although the similar HCB levels may partly be explained by the high transport potential of HCBs, unknown factors may have been more important than distance to sources and long-range transport potential for the chlordanes. One plausible explanation may be that the fasting-related redistribution of POPs from fat to blood was greater throughout the incubation in Arctic eiders, causing them to have higher blood levels of these POPs at the end of incubation. The blood concentrations of POPs in Baltic eiders were higher than documented in any other eider population and were comparable to levels in seabirds feeding at higher trophic positions in the food chain. Environ Toxicol Chem 2016;35:1526-1533. © 2015 SETAC.

Brain size-related breeding strategies in a seabird.

The optimal compromise between decision speed and accuracy may depend on cognitive ability, associated with the degree of encephalization: larger brain size may select for accurate but slow decision-making, beneficial under challenging conditions but costly under benign ones. How this brain size-dependent selection pressure shapes avian breeding phenology and reproductive performance remains largely unexplored. We predicted that (1) large-brained individuals have a delayed breeding schedule due to thorough nest-site selection and/or prolonged resource acquisition, (2) good condition facilitates early breeding independent of relative brain size, and (3) large brain size accrues benefits mainly to individuals challenged by environmental or intrinsic constraints. To test these predictions, we examined how the relative head volume of female eiders (Somateria mollissima) of variable body condition correlated with their breeding schedule, hatching success and offspring quality. The results were consistent with our predictions. First, large head size was associated with a progressively later onset of breeding with increasing breeding dispersal distance. Second, increasing body condition advanced the timing of breeding, but this effect was significantly weaker in large-brained females. Third, larger head volume was associated with increased hatching success mainly among late breeders and those in poor body condition, and duckling body condition was positively related to maternal head volume, but only in poor-condition mothers. Our study is, to our knowledge, the first to demonstrate the presence of brain size-related differences in reproductive strategies within a single natural population.

Context dependency of baseline glucocorticoids as indicators of individual quality in a capital breeder.

Identifying markers of individual quality is a central goal of life-history theory and conservation biology. The 'corticosterone (CORT)-fitness hypothesis' postulates that low fitness signals impaired ability to cope with the environment, resulting in elevated baseline CORT levels. CORT can, however, be negatively, positively or neutrally related to fitness, depending on the context. In order to clarify this controversial issue, we elucidate the utility of using baseline CORT as a correlate of individual fitness in incubating female eiders across variable environments. An increase in serum CORT with decreasing body condition was evident in older, more experienced breeders, while increased clutch mass was associated with elevated serum CORT in females breeding late in the season. For faecal CORT, the expected negative association with body condition was observed only in early breeders. We found a strong increase in faecal CORT with increasing baseline body temperature, indicating the utility of body temperature as a complementary stress indicator. Females in good body condition had a lower baseline body temperature, but this effect was only observed on open islands, a harsher breeding habitat less buffered against weather variability. Females with higher reproductive investment also maintained a lower baseline body temperature. Nest success strongly decreased with increasing serum and faecal CORT concentrations, and individual stress hormone and body temperature profiles were repeatable over years. Although our data support the tenet that baseline CORT is negatively related to fitness, the complex context-dependent effects call for cautious interpretation of relationships between stress physiology and phenotypic quality.

Relative importance of social status and physiological need in determining leadership in a social forager.

Group decisions on the timing of mutually exclusive activities pose a dilemma: monopolized decision-making by a single leader compromises the optimal timing of activities by the others, while independent decision-making by all group members undermines group coherence. Theory suggests that initiation of foraging should be determined by physiological demand in social foragers, thereby resolving the dilemma of group coordination. However, empirical support is scant, perhaps because intrinsic qualities predisposing individuals to leadership (social status, experience or personality), or their interactions with satiation level, have seldom been simultaneously considered. Here, we examine which females initiated foraging in eider (Somateria mollissima) brood-rearing coalitions, characterized by female dominance hierarchies and potentially large individual differences in energy requirements due to strenuous breeding effort. Several physiological and social factors, except for female breeding experience and boldness towards predators, explained foraging initiation. Initiators spent a larger proportion of time submerged during foraging bouts, had poorer body condition and smaller structural size, but they were also aggressive and occupied central positions. Initiation probability also declined with female group size as expected given random assignment of initiators. However, the relative importance of physiological predictors of leadership propensity (active foraging time, body condition, structural size) exceeded those of social predictors (aggressiveness, spatial position) by an order of magnitude. These results confirm recent theoretical work suggesting that 'leading according to need' is an evolutionary viable strategy regardless of group heterogeneity or underlying dominance structure.

Brood size matching: a novel perspective on predator dilution.

A primary benefit of grouping is diluting the individual risk of attack by predators. However, the fact that groups are formed not always by solitary adults but also by subgroups (e.g., families) has been overlooked. The subgroup-specific benefit of predator dilution depends on its relative contribution to total group size. Therefore, the willingness of a subgroup to merge with others should increase the less it contributes to total group size, but the conflicting preferences of partners may result in the preferential merger of similar-sized subgroups. Here, we evaluate how the proportional contribution of subgroups to diluting risk affects group formation. We generate predictions using a bidding game over parental care and test them using data on common eiders (Somateria mollissima), in which females with variable-sized broods may form brood-rearing coalitions. The predictions (1) that size-matched subgroups should have a higher propensity to merge, (2) that predation should increase group formation propensity, and (3) that increased bargaining power, as proxied by female body condition, should increase the time needed to establish partnerships were all supported. Partners do negotiate over their relative contributions to predator dilution, accepting or rejecting partnerships on the basis of this criterion. Our results show that consideration of the size of subgroups before merger is critical in understanding the process of group formation under the threat of predation.

Philopatric predisposition to predation-induced ecological traps: habitat-dependent mortality of breeding eiders.

Because population size is sensitive to changes in adult survival, adult survival may be buffered against environmental variability. Philopatry may be adaptive in changing environments, but it could also constrain breeding habitat selection under changing conditions such as shifting predation regimes. Habitat preference and quality could become decoupled in long-lived philopatric species that evolved in stable environments when suddenly faced by increased adult predation risk, as dispersal may be triggered by past reproductive failure. We evaluated whether the Baltic eider (Somateria m. mollissima) population may currently face a predation-induced ecological trap. Eiders are philopatric and nest on open and forested islands. We hypothesized that open-nesting females would be disproportionately affected by increased predation. We compared female annual survival in these two habitats in 1996-2010. We also tested for effects of time trends, winter severity (NAO), female body condition, and habitat-specific predation pressure on survival. Our results revealed the lowest survival recorded for this species (Φ = 0.720), and survival on open islands was significantly lower (Φ = 0.679) than on forested islands (Φ = 0.761). Nonetheless, only 0.7 % of females changed breeding habitat type despite ample availability of alternative islands, and breeding phenology in both habitats was similar. Female survival increased with body condition, while it was unrelated to winter climate and stable over time. Open islands had a higher predation pressure on incubating females. Breeding philopatry results in a predator-mediated ecological trap for open-nesting eiders. Our results contribute to explaining the drastic decline of the Baltic eider population.

Kin association during brood care in a facultatively social bird: active discrimination or by-product of partner choice and demography?

Intra-group relatedness does not necessarily imply kin selection, a leading explanation for social evolution. An overlooked mechanism for generating population genetic structure is variation in longevity and fecundity, referred to as individual quality, affecting kin structure and the potential for cooperation. Individual quality also affects choosiness in partner choice, a key process explaining cooperation through direct fitness benefits. Reproductive skew theory predicts that relatedness decreases with increasing group size, but this relationship could also arise because of quality-dependent demography and partner choice, without active kin association. We addressed whether brood-rearing eider (Somateria mollissima) females preferentially associated with kin using a 6-year data set with individuals genotyped at 19 microsatellite loci and tested whether relatedness decreased with increasing female group size. We also determined the relationship between local relatedness and indices of female age and body condition. We further examined whether the level of female intracoalition relatedness differed from background relatedness in any year. As predicted, median female intra-group relatedness decreased with increasing female group size. However, the proportion of related individuals increased with advancing female age, and older females prefer smaller brood-rearing coalitions, potentially producing a negative relationship between group size and relatedness. There were considerable annual fluctuations in the level of relatedness between coalition-forming females, and in 1year this level exceeded that expected by random association. Thus, both passive and active mechanisms govern kin associations in brood-rearing eiders. Eiders apparently can discriminate between kin, but the benefits of doing so may vary over time.

Differential responses to related hosts by nesting and non-nesting parasites in a brood-parasitic duck.

Host-parasite relatedness may facilitate the evolution of conspecific brood parasitism, but empirical support for this contention remains inconclusive. One reason for this disparity may relate to the diversity of parasitic tactics, a key distinguishing feature being whether the parasite has a nest of her own. Previous work suggests that parasites without nests of their own may be of inferior phenotypic quality, but because of difficulties in identifying these parasitic individuals, little is known about their host selection criteria. We used high-resolution molecular maternity tests to assign parasitic offspring to known parasites with and without their own nests in a population of Barrow's goldeneyes (Bucephala islandica). We determined whether parasite nesting status, host-parasite relatedness and distance between host and parasite nests affected the probability of parasitizing a host and the number of eggs laid per host. We also investigated whether nesting parasites, conventionally nesting females and non-nesting parasites differed regarding their age, structural size, body condition, nesting phenology or total brood size. The probability of engaging in parasitism increased with host-parasite relatedness and spatial proximity to host nests for nesting and non-nesting females alike. However, nesting parasites increased the number of eggs donated with relatedness to the host, while non-nesting parasites did not do so. Non-nesting parasites laid fewer eggs in total, but did not differ by any of the other quality measures from conventional nesters or nesting parasites. Our study provides the first demonstration that nesting and non-nesting parasites from the same population may use different host selection criteria.

Causes and consequences of fine-scale breeding dispersal in a female-philopatric species.

The potentially confounded effects of factors affecting breeding dispersal have rarely been simultaneously examined. The consequences of breeding dispersal are even less studied, presenting a paradox: breeding dispersal seldom seems to improve breeding success, despite its presumed adaptiveness. We studied the causes and consequences of breeding dispersal in female-philopatric eiders (Somateria mollissima) in relation to the spatiotemporal predictability of nest success. Previous nest fate, breeding experience, and breeding density simultaneously affected breeding dispersal. Dispersal distances were longer among inexperienced breeders and after failed breeding. Individual dispersal distances decreased with increasing nest-site-specific breeding density, whereas island-specific nesting success peaked at intermediate densities. The fate of neighbouring nests ('public information') did not influence dispersal. Breeding dispersal was unrelated to subsequent hatching success, controlling for individual quality (body condition, breeding experience, previous nest fate), while it delayed hatch date, which is likely to impair reproductive success. This delay may result from the loss of acquired information of local breeding conditions, prolonging nest prospecting and establishment, also helping explain why breeding dispersal did not increase at high breeding densities, despite a potential reduction in nesting success. In long-lived species, however, dispersal-induced reductions in reproductive output in one season could be offset by improved parental survival prospects. Careful nest prospecting may be profitable, because overall nest success had a strong island-specific component but showed weak temporal variation, and successive individual nest fates were predictable between years. Once a safe nest site is found, females may breed at the same place successfully for many years.