The diet of king eiders wintering in Nuuk,Southwest Greenland,with reference to sympatric wintering common eiders

Coastal and offshore waters of Southwest Greenland are internationally important wintering areas for king eiders (Somateria spectabilis) breeding in eastern Canadian Arctic and in northwestern Greenland. This paper presents the first assessment of their winter diet. Based on esophageal-proventricular samples from 26 females (13 juveniles and 13 older birds) and 15 males (11 juveniles and four older birds) collected in 2000–2002 (November–May) in coastal waters of Nuuk, we identified 28 prey species. The diet consisted of almost equal proportions (aggregate fresh mass) of polychaetes, echinoderms, crustaceans and molluscs. The dominant prey species were Pectinaria spp. (26.8%), Strongylocentrotus droebachiensis (18.4%), Mya eideri (11.2%) and Hyas araneus (9.7%). The polychaetes have previously been identified as important prey for eiders in Greenland, but apparently not outside Greenland. Compared with a diet study of common eiders Somateria mollissima from the same wintering area, the king eiders consumed significantly less bivalves and significantly more echinoderms. This difference corresponded with observations that common eiders were feeding in shallow waters, while king eiders were feeding in deeper waters farther from the shore. Benthic surveys are needed to confirm that diet corresponds with prey availability.     

The diet of common eiders wintering in Nuuk,Southwest Greenland

Southwest Greenland provides wintering grounds for 70% (∼460,000) of the northern common eider (Somateria mollissima borealis) population. From 241 gullet samples (esophagus and proventriculus content) collected over three winters (1999–2002) near Nuuk, we identified 39 species consumed by the eiders. In contrast to studies elsewhere, fresh mass of the diet was dominated by soft-bottom species: the bivalve Mya eideri (32.8%) and the polychaete Pectinaria spp. (24.2%). The hard-bottom blue mussel (Mytilus edulis), usually the dominant prey of common eiders, was only the fourth most important diet species (7.5%). Overall, bivalves accounted for 56% of the diet. Twenty-seven prey species were minor foods with aggregate fresh mass of only 5.5%. Diets of males and females were similar, whereas juveniles consumed greater mass of crustaceans and less of bivalves. Diet diversity was higher in mid-winter than late winter, and higher in coastal habitats than in fjords. Within one important wintering area the results indicate that optimal size of blue mussels may be depleted over winter.     

Population dynamics of king eiders breeding in northern Alaska

The North American population of king eiders (Somateria spectabilis) has declined by more than 50% since the late 1970s for unknown reasons. King eiders spend most of their lives in remote areas, forcing managers to make regulatory and conservation decisions based on very little information. We incorporated available published estimates of vital rates with new estimates to build a female, stage-based matrix population model for king eiders and examine the processes underlying population dynamics of king eiders breeding at 2 sites, Teshekpuk and Kuparuk, on the coastal plain of northern Alaska and wintering around the Bering Sea (2001–2010). We predicted a decreasing population (λ = 0.981, 95% CI: 0.978–0.985), and that population growth was most sensitive to changes in adult female survival (sensitivity = 0.92). Low duckling survival may be a bottleneck to productivity (variation in ducking survival accounted for 66% of retrospective variation in λ). Adult survival was high (0.94) and invariant ( = 0.0002, 95% CI: 0.0000–0.0007); however, catastrophic events could have a major impact and we need to consider how to mitigate and manage threats to adult survival. A hypothetical oil spill affecting breeding females in a primary spring staging area resulted in a severe population decline; although, transient population dynamics were relatively stable. However, if no catastrophic events occur, the more variable reproductive parameters (duckling and nest survival) may be more responsive to management actions. Published 2012. This article is a U.S. Government work and is in the public domain in the USA.     

Foraging dynamics of muskoxen in Peary Land, northern Greenland

Muskoxen Ovibos moschatus in northern Greenland (79-83°N) are at the northern limit of their distribution and exist under seasonal extremes dominated by nearly 10 months of winter, much of which is without sunlight. The period of summer vegetative growth is less than two months. In the Kap København area (82°30'N), diversity of plant species is low (76 species of vascular plants) and forage biomass in major vegetation types in summer varies from over 40 g m^-2 in sedge-dominated fens to ≤5 g m^-2 in polar barrens. Nonetheless, 90-95% of the ice-free area consists of barren ground or sparcely vegetated polar desert. During summer, muskoxen apparently foraged opportunistically to maximize intake, with sedges the major food item in fens while willows were the major dietary component when on willow-dominated slopes. Quality of summer forage was high during its early phenological stages, with 21-28% crude protein and 60-75% in vitro digestibility. Microhistological analysis of winter feces indicated dominance by graminoids. Muskoxen spent > 50% of their daily activity feeding, which fits a cline of increasing feeding time with increasing latitude in summer. Increased feeding times at high latitudes appears to be a function of both reduced forage biomass and need to maximize forage intake during the brief summer period when forage quality is high. Movement rates in summer while foraging were inversely related to available forage biomass. Seasonal activity of muskoxen peaks during the rutting period (July-September) and then declines gradually through early winter to a low in late winter (March-April).     

Abundance and energy requirements of eiders (Somateria spp.) suggest high predation pressure on macrobenthic fauna in a key wintering habitat in SW Greenland

The number of common eiders (Somateria mollissima borealis) in west Greenland declined dramatically during the twentieth century, supposedly because of human activities. However, their sensitivity to alternative drivers of variation, such as climate conditions, diseases or food availability, remains unstudied. In this study, we describe prey availability and assess the trophic coupling between eiders and their macrobenthic prey in a shallow inlet, Nipisat Sound; a key wintering habitat in the south-west Greenland Open Water Area. Macrobenthic species abundance and biomass were studied, and annual production was estimated by an empirical model, including environmental characteristics, fauna composition and individual biomass. In spring 2008, average macrozoobenthic abundance and biomass were 6,912 ind m⁻² and 28.4 g ash-free dry mass (AFDM) m⁻² (647 kJ m⁻²), respectively. Annual production was estimated at 13.9 g AFDM m⁻² year⁻¹ (317 kJ m⁻² year⁻¹). During the winters of 2008–2010, we monitored the number of common eiders (S. mollissima borealis) and king eiders (Somateria spectabilis) and observed a distinct peak in abundance during winter with up to 15.000 birds in Nipisat Sound. Based on physiological costs of different activities in combination with the observed behavioural pattern, we obtained an estimate of the energy required for eiders to balance their costs of living, which amounted to 58% of the estimated total annual production of macrobenthos in Nipisat Sound. This result suggests that eider predation affects macrobenthic species composition and biomass and demonstrates the potential importance of variations in prey availability for the population dynamics of eiders in Greenland.     

Migration patterns, breeding and moulting locations of king eiders wintering in north-eastern Norway

The Norwegian coast is a very important winter area for king eiders (Somateria spectabilis), but their origin has been unknown. We determined spring and autumn migration routes, timing of migration and potential breeding areas of king eiders wintering in north-eastern Norway using implanted satellite transmitters. Five males and five females were equipped with transmitters in February 2008, and location data were received from six birds. All birds departed within 10 days in mid April and flew to the Pechora Sea and Kara Sea in western Russia where they staged until mid June. Subsequently, four of five birds with active transmitters (two females and two males) moved east to potential breeding locations on the Gydan and Taymyr Peninsulas. In early July, the males moved west to moult at Kolguyev Island and in the Pechora Sea. The two females departed in mid July, one probably moulting between the gulfs of the Ob and Yenisei Rivers, after which it moved to the Pechora Sea. The other female probably moulted in the eastern Taymyr, thereafter moving to Novaya Zemlya. This female returned to the north-eastern coast of Norway 1 December, while the other female returned 2 January. For the males, the transmitters stopped in December/January when they were still in the Pechora/Kolguyev area. King eiders wintering in north-eastern Norway thus originate from the western half of Arctic Russia, and the Taymyr Peninsula is probably the dividing point between the eastern and western flyways.     

Diet and body condition of spectacled eiders wintering in pack ice of the Bering Sea

Spectacled eiders (Somateria fischeri) winter among leads in the Bering Sea pack ice, where they dive 40-70 m for benthic prey. During the first icebreaker cruises into that area, esophagi of collected eiders contained only clams, mostly Nuculana radiata, with no trace of the once-dominant Macoma calcarea. Alternative prey used elsewhere (snails, amphipods, other bivalves) were available but not eaten. Eiders ate mainly N. radiata 18-24 mm long, although M. calcarea of this length contained 62% more energy. Percent body lipid of eiders averaged 12Dž% (SD) for 26 adult males and 14Dž% for 12 adult females. Mean body mass (-SE) of these males in late March (1,688ᆩ g) was higher than reported for 53 males after arriving at breeding areas in late May (1,494ᆢ g). Body mass of these females (1,550ᆷ g) was lower (but not significantly) than reported for 11 females upon arrival at breeding sites (1,623ᇂ g). In 1999, the last spectacled eiders left the wintering area on 21 April, 4-8 weeks before their typical arrival at breeding sites. Their location is unknown in the interim, when habitats used appear critical to acquiring reserves for reproduction.     

Reduced immunocompetence and cost of reproduction in common eiders

Immunocompetence may be especially important in long-lived species where infectious organisms may have detrimental effects upon future reproductive value of hosts. The resource demand for immunocompetence may reduce resource availability for reproduction and a trade-off between these traits has therefore been proposed. Capital breeders, such as the common eider (Somateria mollissima), rely upon accumulated body reserves during reproduction. Eiders lose more than 40% of pre-breeding body mass during egglaying and incubation and many females abandon their ducklings to other females after hatching. Results from our observational study show that levels of leukocytes (i.e., lymphocytes, heterophils and heterophil/lymphocyte ratio) are not related to body mass early in the incubation period. However, eider females with low initial body mass showed signs of immunosuppression (i.e., decreased late levels of lymphocytes) and increased response towards stressors (i.e., increased heterophil/lymphocyte ratio) later in the incubation period. Moreover, females with low lymphocyte levels more frequently abandoned their brood, and females abandoning young had an increased return rate to the next breeding season. However, among brood abandoning females return rate was lower for the females with low lymphocyte levels. These results suggest that immunosuppression, as indicated by low lymphocyte levels, is a reproductive cost that may be partly compensated for by abandoning young.     

Winter climate affects subsequent breeding success of common eiders

The phenology of spring migration depends on the severity of the preceding winter and approaching spring. This severity can be quantified using the North Atlantic Oscillation (NAO) index; positive values indicate mild winters. Although milder winters are correlated with earlier migration in many birds in temperate regions, few studies have addressed how climate‐induced variation in spring arrival relates to breeding success. In northern Europe, the NAO‐index correlates with ice cover and timing of ice break‐up of the Baltic Sea. Ice cover plays an important role for breeding waterfowl, since the timing of ice break‐up constrains both spring arrival and onset of breeding. We studied the effects of the winter‐NAO‐index and timing of ice break‐up on spring migration, laying date, clutch size, female body condition at hatching and fledging success of a short‐distance migrant common eider (Somateria mollissima) population from SW Finland, the Baltic Sea, 1991–2004 (migration data 1979–2004). We also examined the correlation between the NAO‐index and the proportion of juvenile eiders in the Danish hunting bag, which reflects the breeding success on a larger spatial scale. The body condition of breeding females and proportion of juveniles in the hunting bag showed significant positive correlations with the NAO, whereas arrival dates showed positive correlations and clutch size and fledging success showed negative correlations with the timing of ice break‐up. The results suggest that climate, which also affects ice conditions, has an important effect on the fledging success of eiders. Outbreaks of duckling disease epidemics may be the primary mechanism underlying this effect. Eider females are in poorer condition after severe winters and cannot allocate as much resources to breeding, which may impair the immune defense of ducklings. Global climate warming is expected to increase the future breeding success of eiders in our study population.     

Interactions between rate processes with different timescales explain counterintuitive foraging patterns of arctic wintering eiders

To maximize fitness, animals must respond to a variety of processes that operate at different rates or timescales. Appropriate decisions could therefore involve complex interactions among these processes. For example, eiders wintering in the arctic sea ice must consider locomotion and physiology of diving for benthic invertebrates, digestive processing rate and a nonlinear decrease in profitability of diving as currents increase over the tidal cycle. Using a multi-scale dynamic modelling approach and continuous field observations of individuals, we demonstrate that the strategy that maximizes long-term energy gain involves resting during the most profitable foraging period (slack currents). These counterintuitive foraging patterns are an adaptive trade-off between multiple overlapping rate processes and cannot be explained by classical rate-maximizing optimization theory, which only considers a single timescale and predicts a constant rate of foraging. By reducing foraging and instead digesting during slack currents, eiders structure their activity in order to maximize long-term energetic gain over an entire tide cycle. This study reveals how counterintuitive patterns and a complex functional response can result from a simple trade-off among several overlapping rate processes, emphasizing the necessity of a multi-scale approach for understanding adaptive routines in the wild and evaluating mechanisms in ecological time series.     

Costs of parasites in common eiders: effects of antiparasite treatment

Parasites are exploiting their host for the resources required for the parasite's reproduction and survival. The effects of antiparasitic treatment during incubation on body mass and return rate in female common eiders Somateria mollissima were examined in a two year study (1998 and 1999) . There was no effect of treatment on nest success, but unsuccessful females had lower body mass. Treatment had no effect on incubation time, mass-loss during incubation or return rate among successful birds. However, treatment lead to higher return rate among unsuccessful birds (treated=69%, n=13; controls=18%, n=11). This pattern remained significant also after including the birds that returned two years after the experiment. We suggest that the unsuccessful females are of lower quality and had higher costs from parasite infections than birds which completed incubation, and that the higher return rate for treated unsuccessful females may mean that costs of parasites are substantial, and that the potential fitness gain from investment in parasite defense and avoidance is large.     

Handling stress of female common eiders during avian cholera outbreaks

Researchers often consider the importance of minimizing holding time during research activities; however, the long-term costs of such handling stress is rarely measured explicitly. As part of an ongoing study of common eiders (Somateria mollissima) at a breeding colony in East Bay, Southampton Island, Nunavut, we recorded duration of restraint for females captured during avian cholera epizootics (2007 and 2008) and monitored female fates (breeding probability, onset of laying, and survival) relative to holding time. Probability of death increased with holding time in 2007 from an estimated 0.05 for females held 20 min to 0.33 for females held for 150 min. In 2008, we responded by limiting holding time to <90 min and mortality was no longer positively correlated with holding time, although total mortality was greater due to increased severity of avian cholera. In both years, longer restraint durations delayed onset of egg-laying after capture by 0.5 days for each 10 min of additional restraint but did not prevent breeding. This delay of nest initiation did not enhance survival probability. Our results show that prolonged holding time can exacerbate mortality during epizootics and emphasize the importance of minimizing restraint time in wild birds, especially in the presence of diseases. © 2011 The Wildlife Society.     

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.     

Characterizing the nutritional strategy of incubating king eiders Somateria spectabilis in northern Alaska

We measured plasma concentrations of variables associated with lipid metabolism (free fatty acids, glycerol, triglyceride, and β‐hydroxybutyrate), protein metabolism (uric acid), and baseline corticosterone to characterize the nutritional state of incubating king eiders Somateria spectabilis and relate this to incubation constancy at two sites, Kuparuk and Teshekpuk, in northern Alaska. King eiders at both sites appeared to employ a partial‐income incubation strategy, relying on both endogenous and exogenous energy resources. Females maintained high invariant levels of free fatty acids, β‐hydroxybutyrate, and glycerol throughout incubation, indicating that fat reserves were a major energy source, and not completely depleted during incubation. Similarly, uric acid did not increase, suggesting effective protein sparing or protein ingestion and adequate lipid reserves throughout incubation. Baseline corticosterone and triglyceride levels increased during incubation, indicative of an increase in foraging during late stages of incubation. Incubating females at Kuparuk had higher triglyceride concentrations but also had higher β‐hydroxybutyrate concentrations than females at Teshekpuk. This dichotomy may reflect a short‐term signal of feeding overlaying the longer‐term signal of reliance on endogenous lipid reserves due to higher food intake yet higher metabolic costs at Kuparuk because of its colder environment. Incubation constancy was not correlated with plasma concentrations of lipid or protein metabolites.