Parenting in a changing environment: A long-term study of prolactin, parental effort and reproductive success in common eiders.

Parental care is regulated by multiple endocrine mechanisms. Among these hormones, prolactin (PRL) is involved in the expression of parental behaviors. Despite the consensus that PRL mediates variation in parental effort with age and body condition, its role in the adjustment of parental effort to fluctuating environmental conditions, including changing predation pressure, still awaits further investigation. To shed light on this knowledge gap, we relied on a long-term monitoring of female common eiders Somateria mollissima (n = 1277 breeding attempts, 2012-2022) incubating under fluctuating predation risk to investigate the link between baseline PRL levels and female minimum age, body condition, clutch size, environmental parameters (predation pressure, climate, nest microhabitat) and hatching success. We predicted that PRL would be higher in older females, those in better condition or incubating larger clutches. We also predicted that females would reduce parental effort when nesting under challenging environmental conditions (high predation pressure or poor climatic conditions), translated into reduced baseline PRL levels. We also explored how variation in PRL levels, female characteristics and environmental parameters were related to hatching success. Following our predictions, PRL levels were positively associated with body condition and female age (before showing a senescent decline in the oldest breeders). However, we did not observe any population-level or individual-level reduction in PRL levels in response to increasing predation pressure. Population-level baseline PRL levels instead increased over the study period, coincident with rising predation threat, but also increasing female body condition and age. While we did not provide evidence for a direct association between baseline PRL levels and predation risk, our results support the idea that elevated baseline PRL levels promote hatching success under internal constraints (in young, inexperienced, breeders or those incubating a large clutch) or constraining environmental conditions (during years of high predation pressure or poor climatic and foraging conditions). Finally, the low repeatability of baseline PRL levels and high interannual variability highlight considerable within-individual flexibility in baseline PRL levels. Further research should explore flexibility in parental effort to changing environmental conditions, focusing on both baseline and stress-induced PRL levels.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Adult predation risk drives shifts in parental care strategies: a long-term study.

1. Grouping provides antipredatory benefits, and therefore aggregation tendencies increase under heightened predation risk. In socially breeding groups, however, conflicts over reproductive shares or safety tend to disintegrate groups. Group formation thereby involves a balance between the antipredatory benefits of aggregation and the destabilizing effect of reproductive conflict. 2. We study the grouping behaviour of a facultatively social precocial sea duck with uniparental female care, the eider (Somateria mollissima Linnaeus). Females tend their young solitarily or in groups of 2-5 females. Here, we focus on the effect predation on adults has on group-formation decisions of brood-caring females. 3. By modifying an existing bidding game over care, we model the effects of predation risk on the width of the window of selfishness, which delimits the reproductive sharing allowing cooperation within brood-rearing coalitions, and generate predictions about the relative frequencies of solitary versus cooperative parental care modes. Furthermore, we model the dilution effect as a function of female group size and predation risk. 4. The window of selfishness widens with increasing predation risk, and the dilution of predation risk increases with both female group size and increasing predation risk, yielding the following predictions: (i) cooperative brood care becomes more prevalent and, conversely, solitary brood care less prevalent under heightened predation risk and (ii) group sizes increase concomitantly. 5. We tested these predictions using 13 years of data on female grouping decisions and annual predation rates, while controlling for the potentially confounding effect of female body condition. 6. Our data supported both predictions, where heightened predation risk of nesting females, but not changes in their condition, increased the relative frequency of cooperative brood care. Increased female nesting mortality also resulted in larger groups of cooperative females. 7. The predation risk of incubating females has long-term implications for later parental care decisions. We discuss the potential consequences of predation-induced shifts in group size on per capita fitness and population-wide productivity.

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.

Do female ornaments indicate quality in eider ducks?

The fitness consequences of female ornamentation remain little studied and the results are often contradictory. Female ornamentation may be an artefact of a genetic correlation with male ornamentation, but this possibility can be disregarded if the ornament only occurs in females. Female-specific white wing bars in eiders (Somateria mollissima) have been suggested to indicate individual quality, and we studied size variation in this trait in relation to key fitness components and quality attributes. We found that clutch size, body condition, female age, hatching date and success were unrelated to female ornament size; ornament size was explained by its size in the previous year. In contrast, good body condition was associated with hatching success. These results suggest that the breadth of the white wing bars does not indicate individual quality in our study population.

Age-specific nest-site preference and success in eiders.

Variation in nest concealment is puzzling given the expected strong selection for safe nest sites. Selecting a concealed nest may decrease the risk of clutch predation but hinder parents from escaping predators, providing a possible solution to this paradox. Because the relative value of current versus future reproduction may vary with breeder age or state, nest concealment may also vary as a function of these attributes. We tested four predictions of the female and clutch safety trade-off hypothesis in eiders (Somateria mollissima): (1) nest concealment is negatively related to escape possibilities, (2) our capture rate of females is higher in covered nests, (3) egg predation is higher in open nests, and (4) overall nest success is unrelated to nest habitat. We also analysed nest microhabitat preferences and nest success relative to breeder age and body condition, controlling for nest spatial centrality. As expected, nest concealment and potential escape angle were negatively related, and capture by us, indicating female predation vulnerability, increased with nest cover. Clutch size was smaller in open nests, suggesting higher partial clutch predation, while it was larger among experienced and good-condition breeders. The probability of successful hatching was unrelated to nest habitat, positively associated with breeder experience, and negatively associated with hatching date. Experienced females selected more concealed and centrally located nests without sacrificing potential escape angles. The age-specific spatial distribution of nests on islands was unrelated to nest initiation dates, indicating no apparent competition. The age-specific preference of eiders for concealed nests may reflect declining reproductive value with age or confidence in surviving despite selecting a concealed nest. The apparently positive relationship between female age and survival and fecundity in eiders refutes the former alternative. Individual improvement in choosing safe nest sites, coupled with differential survival of individuals performing well, most likely explains age-specific nest-site preference and success.

Relatedness and spatial proximity as determinants of host-parasite interactions in the brood parasitic Barrow's goldeneye (Bucephala islandica).

Recent studies, which have found evidence for kin-biased egg donation, have sparked interest in re-assessing the parasitic nature of conspecific brood parasitism (CBP). Since host-parasite kinship is essential for mutual benefits to arise from CBP, we explored the role of relatedness in determining the behaviour of conspecific nest parasites and their hosts in nesting female Barrow's goldeneyes (Bucephala islandica), a duck in which CBP is common. The results revealed that the amount of parasitism increased with host-parasite relatedness, the effect of which was independent of geographical proximity of host and parasite nests. Proximity per se was also positively associated with the amount of parasitism. Furthermore, while hosts appeared to reduce their clutch size as a response to the presence of parasitic eggs, the magnitude of host clutch reduction also tended to increase with increasing relatedness to the parasite. Hence, our results indicate that both relatedness and spatial proximity are important determinants of CBP, and that host clutch reduction may be an adaptation to nest parasitism, modulated by host-parasite relatedness. Taken together, the results provide a demonstration that relatedness influences host and parasite behaviour in Barrow's goldeneyes, resulting in kin-biased egg donation.