Markus Öst

Social and maternal factors affecting duckling survival in eiders Somateria mollissima

1. With the aid of a novel survivorship model, an 8-year field study of social and maternal factors affecting duckling survival in eiders (Somateria mollissima) revealed that duckling survival probability varies in accordance with maternal brood-rearing strategy. This variability in survival provides compelling evidence of different annual fitness consequences between females that share brood-rearing and those that tend their broods alone. Consequently, as prebreeding survival is often a major source of individual variation in lifetime reproductive success, a females annual, state-dependent (e.g. condition) choice of a brood-rearing strategy can be a critical fitness decision. 2. Variance in duckling survival among lone tender broods was best explained by a model with significant interannual variability in survival, and survivorship tending to increase with increasing clutch size at hatch. Clutch size was correlated positively with female condition. Hatch date and female body condition together affected duckling survival, but their contributions are confounded. We were unable to identify a relationship between female age or experience and duckling survival. 3. Variance in duckling survival among multifemale brood-rearing coalitions was best explained by a model that included the number of tenders, the number of ducklings and interannual variation in how their ratio affected survivorship. Hatch date did not significantly influence survival. 4. Expected duckling survival is higher in early life for lone tenders when compared with multifemale brood-rearing coalitions. However, as ducklings approach 2-3 weeks of age, two or three females was the optimal number of tenders to maximize daily duckling survival. The survivorship advantage of multifemale brood-rearing coalitions was most evident in years of average survival. 5. The observed frequency distribution of female group sizes corresponds with the distribution of offspring survival probabilities for these groups. Evidence for optimal group sizes in nature is rare, but the most likely candidates may be groups of unrelated animals where entry is controlled by the group members, such as for female eiders. 6. Our study demonstrates that differences in social factors can lead to different predictions of lifetime reproductive success in species with shared parental care of self-feeding young.

Female characteristics and parental care mode in the creching system of eiders, Somateria mollissima

Eider females may abandon their young, care alone, or join in multifemale creches. We studied the characteristics of female eiders adopting these strategies in 1996-1999. Female condition at hatching varied significantly between years. Over all years, 31% of all females abandoned, 23% tended alone and 46% creched. In the year when average female condition at hatching was lowest, abandonment rate peaked (67%). Creching birds were further identified as true crechers staying more than 2 weeks with the original creche, and transient crechers leaving the creche. The condition of females shortly before hatching showed a decreasing trend, with lone tenders being in best condition, followed by true crechers, transient crechers and abandoners. Clutch size, date of hatching relative to the population median, and female body size did not differ between groups. Individual females switched between care modes between years. Females weighed significantly less when abandoning than when tending, with no significant weight difference when the females remained as tenders between 2 years. This is consistent with the energetic salvage strategy hypothesis, which states that females in poor body condition should be more prone to abandon their brood. Our results support an adaptive approach to offspring care behaviour in eiders, driven by female condition. uf6d9 2001 The Association for the Study of Animal Behaviour

Brood Parasitism in a Population of Common Eider (somateria Mollissima)

The common eider differs from many other ducks in being a colonial capital breeder, producing eggs from stored resources. These traits are expected to influence the occurrence of conspecific brood parasitism (CBP), which is particularly common in waterfowl. We analysed CBP in an eider population in the central Baltic Sea 2001-2002, using non-destructive egg albumen sampling combined with protein fingerprinting. This technique greatly increases the detection of parasitic eggs compared to more traditional methods. Parasitic eggs occurred in 20-22% of 164 nests studied, 6% of 754 eggs being laid by other than the host female. Parasitism increased with nest density, was rather evenly distributed over the laying season, and occurred both early and late in the laying sequence of the host. Protein fingerprinting showed that host females laid up to seven eggs, more than previously reported. Among 33 parasitised nests 22 had one parasitic egg, nine had two and two had three. In all but one case all parasitic eggs within a nest were laid by the same female. Although colonial breeding facilitates CBP, it is less frequent in this eider population than in several other diving ducks. Possible contributing reasons are the relatively small clutch size and start of incubation after egg 2 or 3, limiting the time window for successful parasitism.

Stress responsiveness, age and body condition interactively affect flight initiation distance in breeding female eiders

Predation may drive differential selection among personality types, but the mechanism linking personality with predation risk is poorly understood. One such mechanism may be provided by stress hormones (corticosterone in birds), which are linked to boldness towards predators. However, because of feedbacks between boldness and future fitness expectations, the relationship between boldness and stress physiology may be modulated by individual quality. We investigated flight initiation distances (FIDs) of incubating eider, Somateria mollissima, females in relation to handling-induced corticosterone concentrations and individual quality (female breeding experience, body condition). We investigated whether FIDs were repeatable, and whether stress responsiveness and individual quality attributes, either independently or interactively, affected FIDs. We also analysed whether incubation duration, a period of peak predation pressure on females, would depend on female boldness towards predators (FID), controlling for individual quality. FIDs were repeatable within and between seasons, and females with higher handling-induced corticosterone concentrations generally had longer FIDs. However, this relationship was modulated by interactions between stress responsiveness and individual quality. High stress responsiveness was associated with longer FIDs in younger females, while the opposite was found for the oldest females. Furthermore, the FIDs of females in good body condition increased less strongly with increasing stress-induced circulating corticosterone. Shy females (long FIDs) and those in poor body condition had shorter incubation periods. Boldness is thus linked to stress responsiveness, helping to explain why stress responses may be correlated with survival. However, physiological stress effects on boldness cannot be understood in isolation from effects of individual quality.

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.

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 (Φxa0=xa00.720), and survival on open islands was significantly lower (Φxa0=xa00.679) than on forested islands (Φxa0=xa00.761). Nonetheless, only 0.7xa0% 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.

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.

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 Barrows 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 Barrows goldeneyes, resulting in kin‐biased egg donation.

Large-scale change in the sex ratio of a declining eider Somateria mollissima population

Abstract The breeding potential of a monogamous animal population should be maximal during equal operational sex ratio, and empirical evidence suggests that the population-wide sex ratio may be linked to population density. We studied the sex ratio of eiders Somateria mollissima migrating into the Gulf of Finland, the Baltic Sea, in nine years during 1979–2005 (1979–1980, 1982–1983 and 2001–2005), and the sex ratio of birds collected by Danish hunters during 1982–2004. In two decades, the sex ratio during peak migration has reversed from female bias to male bias, and hunting statistics have shown a significantly increasing adult male bias. Also the proportion of juvenile males has shown a significant increase (Danish hunting statistics 1982–2004), which indicates either that the primary sex ratio of ducklings is exceedingly male biased, or that the mortality of female ducklings has increased. This shift in sex ratio is paralleled by a dramatic decrease in the Baltic eider population which started in the early 1990s. The proportion of juveniles in the hunting bag, an indicator of breeding success in the Baltic, significantly decreased during our study period. The sex ratio of migrating eiders showed seasonal fluctuations, the pattern of which has changed during the study period. Particularly the proportion of late-migrating females has decreased dramatically since the early 1980s, suggesting a declining influx of subadult females. Both the increased male bias and the decreased breeding success are likely to be linked with the population decline. A primary contributor to the shift in sex ratio and the declining trend in breeding success and population size is possibly differential mortality of the sexes during breeding, as the mortality of breeding females has increased sharply in the western Gulf of Finland, mainly due to predation by white-tailed sea eagles Haliaeetus albicilla and American minks Mustela vison, the former of which has recently increased in numbers. It is unlikely that differential winter mortality of the sexes can explain our results, as the wintering area of eiders from the Gulf of Finland has remained the same, and the Danish hunting bag reflects the existing sex ratio. Our study highlights the need for future empirical and theoretical work on the relationship between population sex ratio and population density.

Eider females form non‐kin brood‐rearing coalitions

Kin selection is a powerful tool for understanding cooperation among individuals, yet its role as the sole explanation of cooperative societies has recently been challenged on empirical grounds. These studies suggest that direct benefits of cooperation are often overlooked, and that partner choice may be a widespread mechanism of cooperation. Female eider ducks (Somateria mollissima) may rear broods alone, or they may pool their broods and share brood‐rearing. Females are philopatric, and it has been suggested that colonies may largely consist of related females, which could promote interactions among relatives. Alternatively, shared brood care could be random with respect to relatedness, either because brood amalgamations are accidental and nonadaptive, or through group augmentation, assuming that the fitness of all group members increases with group size. We tested these alternatives by measuring the relatedness of co‐tending eider females in enduring coalitions with microsatellite markers. Females formed enduring brood‐rearing coalitions with each other at random with respect to relatedness. However, based on previous data, partner choice is nonrandom and dependent on female body condition. We discuss potential mechanisms underlying eider communal brood‐rearing decisions, which may be driven by the specific ecological conditions under which sociality has evolved in this species.