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.     

Body mass regulation during incubation in female common eiders Somateria mollissima

We investigated changes in incubation behaviour induced by body fuel depletion in incubating female common eiders, which, in contrast to pelagic seabirds, fast despite being close to marine food sources. In the Svalbard Archipelago, electronic scales were placed under eider nests and the incubation of six birds was prolonged by using wax-filled eggs. Based on changes in the rate of body mass loss in normally incubating females and in ten captive birds that did not incubate, body reserves neared depletion on average four days after hatching. During prolonged incubation, females took more frequent and longer recesses. Nest attentiveness consequently decreased, but was still high. In contrast to recesses during normal incubation, during which body mass of the birds decreased, mass remained constant during the recesses of prolonged incubation. The body stores of female eiders seemed to enable them to complete incubation with a limited safety margin. A further drop in body mass is avoided when a critical body mass is reached, because birds then start feeding enough to maintain mass while continuing incubation. Presumably, a similar mechanism will enable eiders to continue incubation when body reserves are prematurely depleted before hatching.     

Long-Term Changes in Nutrients and Mussel Stocks Are Related to Numbers of Breeding Eiders Somateria mollissima at a Large Baltic Colony

Background

The Baltic/Wadden Sea eider Somateria mollissima flyway population is decreasing, and this trend is also reflected in the large eider colony at Christiansø situated in the Baltic Sea. This colony showed a 15-fold increase from 1925 until the mid-1990''s, followed by a rapid decline in recent years, although the causes of this trend remain unknown. Most birds from the colony winter in the Wadden Sea, from which environmental data and information on the size of the main diet, the mussel Mytilus edulis stock exists. We hypothesised that changes in nutrients and water temperature in the Wadden Sea had an effect on the ecosystem affecting the size of mussel stocks, the principal food item for eiders, thereby influencing the number of breeding eider in the Christiansø colony.

Methodology/Principal Finding

A positive relationship between the amount of fertilizer used by farmers and the concentration of phosphorus in the Wadden Sea (with a time lag of one year) allowed analysis of the predictions concerning effects of nutrients for the period 1925–2010. There was (1) increasing amounts of fertilizer used in agriculture and this increased the amount of nutrients in the marine environment thereby increasing the mussel stocks in the Wadden Sea. (2) The number of eiders at Christiansø increased when the amount of fertilizer increased. Finally (3) the number of eiders in the colony at Christiansø increased with the amount of mussel stocks in the Wadden Sea.

Conclusions/Significance

The trend in the number of eiders at Christiansø is representative for the entire flyway population, and since nutrient reduction in the marine environment occurs in most parts of Northwest Europe, we hypothesize that this environmental candidate parameter is involved in the overall regulation of the Baltic/Wadden Sea eider population during recent decades.     

Experimental evaluation of nest shelter effects on weight loss in incubating common eiders Somateria mollissima

Evaluating consequences of habitat selection is an important step in understanding life history strategies and behavioural decisions of animals. Kilpi and Lindström (1997) found that incubating common eiders Somateria mollissima on exposed, treeless islands lost weight faster than females nesting on wooded islands and proposed that this difference was due to adverse incubation conditions at exposed nests. Therefore, we tested whether common eiders gained an advantage when nesting in sheltered habitats by placing artificial shelters over randomly-selected females after the onset of incubation within an eider colony in arctic Canada. We predicted that sheltered females would be heavier on completion of incubation than control hens lacking shelters. Females nesting in artificial shelters experienced a more moderate thermal environment at both cold and warm temperature extremes. Eiders nesting in shelters were heavier than control females during mid incubation, consistent with habitat-specific rates of weight loss reported by Kilpi and Lindström (1997) . Natural overhead cover was available at potential nests but few eiders used those sites. We suspect that microclimatic advantages offered by sheltered sites may be offset by costs of increased female vulnerability to predators. Further work is needed to test this hypothesis, and to determine mechanisms responsible for lower weight loss in eiders attending well concealed nests.     

Do T3 levels in incubating eiders reflect the cost of incubation among clutch sizes?

Complete development of avian eggs requires external heat, inducing in most species an energetic cost of incubation for the parents. Triiodothyronine (T(3)) has been implicated in the control of the metabolic rate and is decreased during fasting in most bird species. This raises the question of the regulation of T(3) during reproduction when incubation (thus heat production) is associated with fasting (and energy sparing). In this study, plasma concentrations of T(3) were studied for different clutch sizes in incubating, as well as in nonincubating, fasting female eiders. Our results show that the T(3) levels decrease during fasting in nonincubating birds, whereas they were maintained during the incubation fast. T(3) levels increased in female eiders at hatching. The plasma T(3) level did not vary among natural clutch sizes in eiders but did so when manipulated. T(3) levels increased when eggs were added (to a maximum of six eggs, i.e., the biggest natural clutch size) or removed (to two eggs, i.e., the smallest natural clutch size). Our results suggest that (1) high T(3) levels during incubation may participate to a threshold of heat production and incubation metabolic rate in eiders despite the fact that they are fasting; (2) since T(3) is associated with the energy expenditure in birds, incubating an enlarged or reduced clutch size may lead to a higher energetic cost of incubation in eiders; and (3) the energy demand of the ducklings at hatching is probably important, as the female T(3) concentrations are then at their highest levels. Thus, any modification of the natural clutch size leads to a rise in the T(3) level of the incubating female, suggesting an additional cost of incubation. Knowing that there is no variation of T(3) levels among natural clutch sizes, this study suggests that a female eider produces a number of eggs corresponding to the energy she can invest in incubation.     

Feather corticosterone reveals effect of moulting conditions in the autumn on subsequent reproductive output and survival in an Arctic migratory bird

For birds, unpredictable environments during the energetically stressful times of moulting and breeding are expected to have negative fitness effects. Detecting those effects however, might be difficult if individuals modulate their physiology and/or behaviours in ways to minimize short-term fitness costs. Corticosterone in feathers (CORTf) is thought to provide information on total baseline and stress-induced CORT levels at moulting and is an integrated measure of hypothalamic–pituitary–adrenal activity during the time feathers are grown. We predicted that CORTf levels in northern common eider females would relate to subsequent body condition, reproductive success and survival, in a population of eiders nesting in the eastern Canadian Arctic during a capricious period marked by annual avian cholera outbreaks. We collected CORTf data from feathers grown during previous moult in autumn and data on phenology of subsequent reproduction and survival for 242 eider females over 5 years. Using path analyses, we detected a direct relationship between CORTf and arrival date and body condition the following year. CORTf also had negative indirect relationships with both eider reproductive success and survival of eiders during an avian cholera outbreak. This indirect effect was dramatic with a reduction of approximately 30% in subsequent survival of eiders during an avian cholera outbreak when mean CORTf increased by 1 standard deviation. This study highlights the importance of events or processes occurring during moult on subsequent expression of life-history traits and relation to individual fitness, and shows that information from non-destructive sampling of individuals can track carry-over effects across seasons.     

Effects of implanted satellite transmitters on behavior and survival of female common eiders

Surgically implanted satellite transmitters have been widely used in studies of avian ecology, yet little is known about their potential impacts on birds. We implanted satellite transmitters with percutaneous antennae (approx. 50 g) in 17 female common eiders (Somateria mollissima) at a breeding colony in Arctic Canada. Among females implanted during incubation, 11 of 12 nests were abandoned within 1 week of being radioed. We observed no differences in the proportion of time that implanted female eiders allocated to basic behaviors. Radioed birds were more likely to pick or preen their abdominal (site of surgical incision) and posterior–dorsal (site of antenna exit) regions than unmarked females, although these behaviors were rare (approx. 0.3% of total time budget). Three of 10 females re-observed had a pronounced limp following surgery, but we observed no walking difficulties among these females in subsequent seasons, and we observed some implanted eiders nesting in subsequent years. Mark–resighting models suggest eiders with transmitters had lower apparent survival the year after implantation (67.0%; 85% CI: 47.8–81.9%) than did color-banded eiders (87.5%, 85% CI: 82.5–91.2%), but there was no model support for a survival difference in subsequent years. We conclude that transmitter implantation in common eiders leads to short-term changes in behavior and a decline in first year survival. We encourage researchers to collect similar data on their study subjects where possible and use it to determine the degree to which data are representative of the greater population. © 2011 The Wildlife Society.     

An experimental study on the effects of polymorphiasis in common eider ducklings.

Eight common eider (Somateria mollissima) ducklings were experimentally infected from 1 June through 13 June, 1995 with acanthocephalans (Polymorphus minutus) by allowing the birds to feed on Gammarus spp. (Gammarus oceanicus, G. salinus, G. zaddachi, and G. lacustris) containing acanthocephalan cystacanths. Uninfected Gammarus spp. were fed to a control group of seven ducklings. No mortality of ducklings occurred during the experiment. However, the infected ducklings gained weight more slowly than the control birds. After the 2 wk study period, the mean serum concentrations of total protein, albumin, beta-globulin, gamma-globulin, fructosamine and creatine kinase were lower in the infected group than in the controls. The mean (+/-SE) number of acanthocephalans in the intestine of the infected ducklings was 21 (+/-4). The parasites were attached to the mucosa of the posterior small intestine of the infected ducklings with a mixed inflammatory reaction consisting of heterophils and mononuclear lymphocytes surrounding the attachment sites.     

Factors affecting nest site selection and nesting success in the Common Eider Somateria mollissima

Nesting site selection and nesting success in Common Eiders Somateria mollissima were studied over a 3-year period (1991–1993) in the Mast River delta (58^o24'N, 94^o24'W), 40 km east of Churchill, Manitoba, Canada. Eiders preferentially nested on islands that had incubating Lesser Snow Geese Anser caerulescens caerulescens on them; this effect was also seen between years on the same islands. Eiders which nested on islands with geese had a reduced chance of having eggs taken by predators during egg-laying and had a greater chance of hatching once incubation had begun, independent of the number of other eiders nesting on the island. The distance to a goose nest was less in nests which did not lose eggs prior to incubation than in those which did, but there was no difference in the distance to a goose nest in eider clutches which did or did not hatch. Artificial eider eggs placed closer to goose nests had a lower probability of being predated than those placed farther away. Nests on islands farther up the river and farther from the mainland had higher nesting success, presumably because these nesting islands were inaccessible to Arctic Foxes Alopex lagopus.     

Does winter region affect spring arrival time and body mass of king eiders in northern Alaska?

Events during the non-breeding season may affect the body condition of migratory birds and influence performance during the following breeding season. Migratory birds nesting in the Arctic often rely on endogenous nutrients for reproductive efforts, and are thus potentially subject to such carry-over effects. We tested whether king eider (Somateria spectabilis) arrival time and body mass upon arrival at breeding grounds in northern Alaska were affected by their choice of a winter region in the Bering Sea. We captured birds shortly after arrival on breeding grounds in early June 2002–2006 at two sites in northern Alaska and determined the region in which individuals wintered using satellite telemetry or stable isotope ratios of head feathers. We used generalized linear models to assess whether winter region explained variation in arrival body mass among individuals by accounting for sex, site, annual variation, and the date a bird was captured. We found no support for our hypothesis that either arrival time or arrival body mass of king eiders differed among winter regions. We conclude that wintering in different regions in the Bering Sea is unlikely to have reproductive consequences for king eiders in our study areas.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

Use of the Beaufort Sea by King Eiders Breeding on the North Slope of Alaska

Abstract: We estimated areas used by king eiders (Somateria spectabilis) in the Alaskan Beaufort Sea, how distributions of used areas varied, and characteristics that explained variation in the number of days spent at sea, to provide regulatory agencies with baseline data needed to minimize impacts of potential offshore oil development. We implanted sixty king eiders with satellite transmitters at nesting areas on the North Slope of Alaska, USA, in 2002-2004. More than 80% of marked eiders spent >2 weeks staging offshore prior to beginning a postbreeding molt migration. During postbreeding staging and migration, male king eiders had much broader distributions in the Alaskan Beaufort Sea than female eiders, which were concentrated in Harrison and Smith Bays. Distribution did not vary by sex during spring migration in the year after marking. Shorter residence times of eiders and deeper water at locations used during spring migration suggest the Alaskan Beaufort Sea might not be as critical a staging area for king eiders during prebreeding as it is postbreeding. Residence time in the Beaufort Sea varied by sex, with female king eiders spending more days at sea than males in spring and during postbreeding. We conclude the Alaskan Beaufort Sea is an important staging area for king eiders during postbreeding, and eider distribution should be considered by managers when mitigating for future offshore development. We recommend future studies examine the importance of spring staging areas outside the Alaskan Beaufort Sea.     

Aggressive neighbors and dense nesting: nest site choice and success in high-Arctic common eiders

Minimizing the risk of nest predation has led some bird species to exploit the nest defense behavior of other species. At Nasaruvaalik Island, Nunavut, Canada, some common eiders (Somateria mollissima borealis) nest within the boundaries of Arctic tern (Sterna paradisaea) colonies, while others nest elsewhere on the island, away from the terns. We tested the effects of location (within vs. outside the tern colonies), density of common eider nests, and annual variation on the nesting parameters of common eiders. Our results suggest that nesting in association with Arctic terns does not confer an obvious benefit to eiders. Such associative nesting of eiders and terns may be the result of overlapping habitat preferences between the two species, or a general scarcity of suitable nesting habitat for ground-nesting species in the high Arctic. However, eiders nesting in higher densities with other eiders had greater nest success and lower total clutch predation, indicating a positive correlation between nest density and success.     

Cost of reproduction in a long-lived bird: incubation effort reduces immune function and future reproduction

Life-history theory predicts that increased current reproductive effort should lead to a fitness cost. This cost of reproduction may be observed as reduced survival or future reproduction, and may be caused by temporal suppression of immune function in stressed or hard-working individuals. In birds, consideration of the costs of incubating eggs has largely been neglected in favour of the costs of brood rearing. We manipulated incubation demand in two breeding seasons (2000 and 2001) in female common eiders (Somateria mollissima) by creating clutches of three and six eggs (natural range 3-6 eggs). The common eider is a long-lived sea-duck where females do not eat during the incubation period. Mass loss increased and immune function (lymphocyte levels and specific antibody response to the non-pathogenic antigens diphtheria and tetanus toxoid) was reduced in females incubating large clutches. The increased incubation effort among females assigned to large incubation demand did not lead to adverse effects on current reproduction or return rate in the next breeding season. However, large incubation demand resulted in long-term fitness costs through reduced fecundity the year after manipulation. Our data show that in eiders, a long-lived species, the cost of high incubation demand is paid in the currency of reduced future fecundity, possibly mediated by reduced immune function.     

Endogenous reserve dynamics of northern common eiders wintering in Greenland

Endogenous reserves influence both survival and reproduction of many waterfowl species, but little is known about reserve levels of most species during the nonbreeding season, particularly those wintering at high latitudes. We investigated whether age, sex, and season were related to carcass composition of northern common eiders (Somateria mollissima borealis) wintering in southwest Greenland during 1999–2002. Adults carried more lipid and protein than juveniles during all winters. Among both age classes, males and females had similar fat levels but males carried slightly more protein. There was no dramatic seasonal variation in lipid or protein content. This suggests that during the period of this study, these eiders did not experience large-scale nutritional shortfalls. As predicted, Greenlandic eiders carried more lipid reserves than eider populations wintering in more temperate environments. Contrary to prediction, there was little relation between reserve levels and photoperiod, ambient temperature, or hunting disturbance intensity. Our results suggest that both sexes are equally capable of dealing with nutritional deficits, and that juvenile birds are more prone to nutritional stress as evidenced by their consistently poorer body condition.     

Avian cholera in eider ducks in Maine

Outbreaks of avian cholera (Pasteurella multocida) occur frequently in common eiders (Somateria mollissima dresseri) in Maine during early summer. Studies over a seven year period show that over 90% of the loss occurred in incubating females and might be associated with their weakened condition because females do not feed during the incubation period. High nesting densities also may contribute to the losses. The exact source of P. multocida is unknown although carrier birds were found.     

Limited genetic differentiation among breeding,molting, and wintering groups of the threatened Steller’s eider: the role of historic and contemporary factors

Due to declines in the Alaska breeding population, the Steller’s eider (Polysticta stelleri) was listed as threatened in North America in 1997. Periodic non-breeding in Russia and Alaska has hampered field-based assessments of behavioral patterns critical to recovery plans, such as levels of breeding site fidelity and movements among three regional populations: Atlantic-Russia, Pacific-Russia and Alaska. Therefore, we analyzed samples from across the species range with seven nuclear microsatellite DNA loci and cytochrome b mitochondrial (mt)DNA sequence data to infer levels of interchange among sampling areas and patterns of site fidelity. Results demonstrated low levels of population differentiation within Atlantic and Pacific nesting areas, with higher levels observed between these regions, but only for mtDNA. Bayesian analysis of microsatellite data from wintering and molting birds showed no signs of sub-population structure, even though band-recovery data suggests multiple breeding areas are present. We observed higher estimates of F-statistics for female mtDNA data versus male data, suggesting female-biased natal site fidelity. Summary statistics for mtDNA were consistent with models of historic population expansion. Lack of spatial structure in Steller’s eiders may result largely from insufficient time since historic population expansions for behaviors, such as natal site fidelity, to isolate breeding areas genetically. However, other behaviors such as the periodic non-breeding observed in Steller’s eiders may also play a more contemporary role in genetic homogeneity, especially for microsatellite loci.     

Does the cost of incubation set limits to clutch size in common eiders Somateria mollissima?

We examined the effect of natural clutch size on the cost of incubation in a population of common eiders Somateria mollissima nesting in Tromsø, northern Norway. The body condition of females at day 5 in the incubation period was not related to clutch size (3–6 eggs), but females incubating large clutches lost more mass and had a lower body condition at day 20 in the incubation period than females incubating small clutches. Females incubating large clutches had a slightly shorter incubation period and a lower egg predation rate. The results do not support the hypothesis that the female's ability to produce eggs is the only ultimate control of clutch size in eider. Instead the results suggest that there may be an interaction between the allocation of body reserves to eggs and incubation, and that females producing large clutches allocate more of their body reserves to incubation than females producing small clutches, in order to shorten the incubation period and to minimise the risk of predation on eggs.     

International importance of the eastern Chukchi Sea as a staging area for migrating king eiders

The evaluation of habitats used by arctic birds on migration is crucial for their conservation. We explored the importance of the eastern Chukchi Sea (ECS) as a staging area for king eiders (Somateria spectabilis) migrating between breeding areas in Siberia and western North America and wintering areas in the Bering Sea. We tracked 190 king eiders with satellite transmitters between 1997 and 2007. In late summer, 74% of satellite-tracked king eiders migrating south staged in the ECS for 13 ± 13 (SD) days between late June and early November. During spring migration, king eiders staged in the ECS between mid-April and early June for 21 ± 10 days. All instrumented birds migrating to breeding grounds in western North America (n = 62), and 6 of 11 males migrating to breeding grounds in Siberia, used this area for at least 1 week during spring migration. The importance of this staging area renders it possible that industrial development could adversely affect king eider populations in both Siberia and North America.     

Breeding and moulting locations and migration patterns of the Atlantic population of Steller's eiders Polysticta stelleri as determined from satellite telemetry

This study was designed to determine the spring, summer, autumn, and early winter distribution, migration routes, and timing of migration of the Atlantic population of Steller's eiders Polysticta stelleri. Satellite transmitters were implanted in 20 eiders captured in April 2001 at Vadsø, Norway, and their locations were determined from 5 May 2001 to 6 February 2002. Regions where birds concentrated from spring until returning to wintering areas included coastal waters from western Finnmark, Norway, to the eastern Taymyr Peninsula, Russia. Novaya Zemlya, Russia, particularly the Mollera Bay region, was used extensively during spring staging, moult, and autumn staging; regions of the Kola, Kanin, and Gydanskiy peninsulas, Russia, were used extensively during spring and moult migrations. Steller's eiders migrated across the Barents and Kara seas and along the Kara Sea and Kola Peninsula coastal waters to nesting, moulting, and wintering areas. The majority of marked eiders (9 of 15) were flightless in near‐shore waters along the west side of Novaya Zemlya. Eiders were also flightless in northern Norway and along the Kanin and at Kola Peninsula coasts. We compare and contrast natural history characteristics of the Atlantic and Pacific populations and discuss evolutionary and ecological factors influencing their distribution.