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.

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.

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.

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 female's 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.

Parental effort and reproductive skew in coalitions of brood rearing female common eiders.

Members of breeding groups face conflicts over parental effort when balancing antipredatory vigilance and feeding. Empirical evidence has shown disparate responses to manipulations of parental effort. We develop a model in which we determine the evolutionarily stable effort of partners given their body conditions, allowing the benefits of shared care to be unevenly divided, and we test this model's predictions with data on common eiders (Somateria mollissima). Eiders show uniparental female care; females may share brood rearing, or they may tend alone, and their body condition at hatching of the young shows large environmentally induced variation. The model predicts that parental effort (vigilance) in a coalition is lower than when tending alone, controlling for parental condition; this prediction is supported by the data. The parental effort in a coalition should be positively correlated with body condition, and this prediction is also supported. Finally, parental effort should increase when partner condition decreases and vice versa; this prediction is partially supported. The Nash bargaining game may provide promising avenues by which to determine the precise settlement of reproductive skew and effort between coalition partners in the future.

Analysis of nodularin-R in eider (Somateria mollissima), roach (Rutilus rutilus L.), and flounder (Platichthys flesus L.) liver and muscle samples from the western Gulf of Finland, northern Baltic Sea.

Nodularin (NODLN) is a cyanobacterial hepatotoxin that may cause toxic effects at very low exposure levels. The NODLN-producing cyanobacterium Nodularia spumigena forms massive blooms in the northern Baltic Sea, especially during the summer. We analyzed liver and muscle (edible meat) samples from common eider (Somateria mollissima), roach (Rutilus rutilus L.), and flounder (Platichthys flesus L.) for NODLN-R by liquid chromatography/mass spectrometry (LC-MS) and enzyme-linked immunosorbent assay (ELISA). Thirty eiders, 11 roach, and 15 flounders were caught from the western Gulf of Finland between September 2002 and October 2004. Eiders from April to June 2003 were found dead. The majority of samples were analyzed by LC-MS and ELISA from the same sample extracts (water:methanol:n-butanol, 75:20:5, v:v:v). Nodularin was detected in 27 eiders, nine roach, and eight flounders. Eider liver samples contained NODLN up to approximately 200 microg/kg dry weight and muscle samples at approximately 20 microg/kg dry weight, roach liver samples 20 to 900 microg NODLN/kg dry weight and muscle samples 2 to 200 microg NODLN/kg dry weight, and flounder liver samples approximately 5 to 1,100 microg NODLN/kg dry weight and muscle samples up to 100 microg NODLN/kg dry weight. The NODLN concentrations found in individual muscle samples of flounders, eiders, and roach (1-200 microg NODLN/kg dry wt) indicate that screening and risk assessment of NODLN in Baltic Sea edible fish and wildlife are required for the protection of consumer's health.

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.

Evidence of chromosomal damage in common eiders (Somateria mollissima) from the Baltic Sea.

Common eiders nesting in the Baltic Sea are exposed to generally high levels of contaminants including potentially genotoxic polycyclic aromatic hydrocarbons and organochlorines. Blood samples were collected from eiders at eight sites in the Baltic Sea and two sites in the Beaufort Sea. DNA content variation was estimated using the flow cytometric method, and subsequently utilized as a biomarker of genetic damage. We observed no significant differences in genetic damage among populations within either the Baltic or Beaufort Seas. However, eider populations from the Baltic Sea had significantly elevated estimates of genetic damage compared to populations from the Beaufort Sea.

An adenovirus associated with intestinal impaction and mortality of male common eiders (Somateria mollissima) in the Baltic Sea.

We examined 10 common eider (Somateria mollissima) males found dead in 1998 during a die-off in the northern Baltic Sea off the southwestern coast of Finland. We diagnosed impaction of the posterior small intestine with mucosal necrosis as the cause of death in all 10 and isolated adenoviruses from cloacal samples of six birds. The adenovirus isolates were not neutralized by reference antisera to group I, II, or III avian adenoviruses. Cloacal swabs from 22 apparently healthy eider females nesting at the mortality area were negative for viruses. An adenovirus isolated from one of the eiders caused clinical signs of illness and gastrointestinal pathology in experimentally infected mallard (Anas platyrhynchos) ducklings. These findings suggest that the adenovirus contributed to the mortality of common eider males in the Finnish archipelago.

Isolation and characterization of a reovirus from common eiders (Somateria mollissima) from Finland.

Samples of brain, intestine, liver, lung, spleen, and bursa of Fabricius were collected from five common eider (Somateria mollissima) duckling carcasses during a die-off in the western Gulf of Finland (59 degrees 50'N, 23 degrees 15'E) in June 1996. No viral activity was observed in specific-pathogen-free chicken embryos inoculated with tissue suspensions, but samples of bursa of Fabricius from three birds were positive when inoculated into Muscovy duck (Cairina moschata) embryo fibroblasts. The isolates were characterized as nonenveloped RNA viruses and possessed several characteristics of the genus Orthoreovirus. Virus particles were icosahedral with a mean diameter of 72 nm and were stable at pH 3.0; their genome was separated into 10 segments by polyacrylamide gel electrophoresis. Mallard (Anas platyrhynchos) ducklings experimentally infected with the eider reovirus showed elevated serum activities of aspartate aminotransferase, creatine kinase, and lactate dehydrogenase enzymes and focal hemorrhages in the liver, spleen, and bursa of Fabricius. During 1997-99, the prevalence of neutralizing antibodies to the isolated virus ranged from 0 to 86% in 302 serum samples collected from incubating eider hens at three nesting areas along coastal Finland. The highest seroprevalence was found in Hanko in 1999, just weeks before reports of an uninvestigated mortality event resulting in the death of an estimated 98% of ducklings at that location. These findings raise the question of potential involvement of the virus in poor duckling survival and eider population declines observed in several breeding areas along coastal Finland since the mid-1980s.

Habitat-specific clutch size and cost of incubation in common eiders, Somateria mollissima.

Common eiders, Somateria mollissima, breed on two types of island in the Northern Baltic: open, sparsely vegetated islands and wooded islands with dense mixed forests. On open islands 79.8% of the nests were on open cliffs, exposed to wind and rain whereas on wooded islands 91.7% of the nests were sheltered inside dense spruce and pine thickets. We found that clutch size on open islands was significantly smaller than on wooded islands. Females started breeding simultaneously in the two habitats and they were similar in body size as measured by the length of the radio-ulna. During incubation females on open islands lost weight at a faster rate than females on wooded islands (34 g/day and 19 g/day, respectively). Heat loss is faster on open than wooded islands and therefore we suggest that the faster weight loss of females on open islands result from thermodynamically adverse incubation conditions. Because the eider is an extreme capital breeder, energy used for egg production cannot be used for incubation. To sustain a higher incubation cost on open islands, the optimal clutch size is therefore lower than on wooded islands.