Incubation behavior of king eiders on the coastal plain of northern Alaska

Incubating birds balance their energetic demands during incubation with the needs of the developing embryos. Incubation behavior is correlated with body size; larger birds can accumulate more endogenous reserves and maintain higher incubation constancy. King eiders (Somateria spectabilis) contend with variable and cold spring weather, little nesting cover, and low food availability, and thus are likely to rely heavily on endogenous reserves to maintain high incubation constancy. We examined the patterns of nest attendance of king eiders at Teshekpuk and Kuparuk, Alaska (2002–2005) in relation to clutch size, daily temperature, and endogenous reserves to explore factors controlling incubation behavior. Females at Kuparuk had higher constancy (98.5 ± 0.2%, n = 30) than at Teshekpuk (96.9 ± 0.8%, n = 26), largely due to length of recesses. Mean recess length ranged from 21.5 to 23.7 min at Kuparuk, and from 28.5 to 51.2 min at Teshekpuk. Mean body mass on arrival at breeding grounds (range; Teshekpuk 1,541–1,805, Kuparuk 1,616–1,760), and at the end of incubation (Teshekpuk 1,113–1,174, Kuparuk 1,173–1,183), did not vary between sites or among years (F < 1.1, P > 0.3). Daily constancy increased 1% with every 5°C increase in minimum daily temperature (β _min = 0.005, 95% CI 0.002, 0.009). Higher constancy combined with similar mass loss at Kuparuk implies that females there met foraging requirements with shorter recesses. Additionally, females took more recesses at low temperatures, suggesting increased maintenance needs which were potentially ameliorated by feeding during these recesses, indicating that metabolic costs and local foraging conditions drove incubation behavior.     

Use of serum biochemistry to evaluate nutritional status and health of incubating common eiders (Somateria mollissima) in Finland

During 1997-1999, we collected serum samples from 156 common eider (Somateria mollissima) females incubating eggs in the Finnish archipelago of the Baltic Sea. We used serum chemistry profiles to evaluate metabolic changes in eiders during incubation and to compare the health and nutritional status of birds nesting at a breeding area where the eider population has declined by over 50% during the past decade, with birds nesting at two areas with stable populations. Several changes in serum chemistries were observed during incubation, including (1) decreases in serum glucose, total protein, albumin, beta-globulin, and gamma-globulin concentrations and (2) increases in serum uric acid, creatine kinase, and beta-hydroxybutyrate concentrations. However, these changes were not consistent throughout the 3-yr period, suggesting differences among years in the rate of carbohydrate, lipid, and protein utilization during incubation. The mean serum concentrations of free fatty acids, glycerol, and albumin were lowest and the serum alpha- and gamma-globulin levels were highest in the area where the eider population has declined, suggesting a role for nutrition and diseases in the population dynamics of Baltic eiders.     

Factors Influencing Nesting Success of King Eiders on Northern Alaska's Coastal Plain

Abstract: King eider (Somateria spectabilis) populations have declined markedly in recent decades for unknown reasons. Nest survival is one component of recruitment, and a female's chance of reproductive success increases with her ability to choose an appropriate nesting strategy. We estimated variation in daily nest survival of king eiders at 2 sites, Teshekpuk and Kuparuk, Alaska, USA, 2002–2005. We evaluated both a priori and exploratory competing models of nest survival that considered importance of nest concealment, seclusion, and incubation constancy as strategies to avoid 2 primary egg predators, avian (Larus spp., Stercorarius spp., and Corvus corax) and fox (Alopex lagopus). We used generalized nonlinear techniques to examine factors affecting nest survival rates and information-theoretic approaches to select among competing models. Estimated nest survival, accounting for a nest visitation effect, varied considerably across sites and years (0.21–0.57); however, given our small sample size, much of this variation may be attributable to sampling variation (s̀²_process = 0.007, 95% CI: 0.003–0.070). Nest survival was higher at Kuparuk than Teshekpuk in all years; however, due to the correlative nature of our data, we cannot determine the underlying causes with any certainty. We found mixed support for the concealed breeding strategy; females derived no benefit from nesting in areas with more willow (Salix spp.; measure of concealment) except that the observer effect diminished as willow cover increased. We suggest these patterns are due to conflicting predation pressures. Nest survival was not higher on islands (measure of seclusion) or with increased incubation constancy but was higher post-fox removal, indicating that predator control on breeding grounds could be a viable management option. Nest survival was negatively affected by our nest visitations, most likely by exposing the nest to avian scavengers. We recommend precautions be taken to limit the effects of nest visits in future studies and to consider them as a possible negative bias in estimated nest survival. Future models of the impacts of development within the breeding grounds of king eider should consider the influence of humans in the vicinity of nests.     

Corticosterone alone does not trigger a short term behavioural shift in incubating female common eiders Somateria mollissima, but does modify long term reproductive success

The trade-off between reproductive effort and adult survival in birds is modulated by several factors. Corticosterone and prolactin have additive effects on reproductive behaviour by stimulating foraging and parental behaviours, respectively. When incubation is associated with fasting, nest desertion is supposed to be activated by an unknown refeeding signal when body condition becomes critically deteriorated. The concomitant rise in corticosterone levels has been suggested to be the triggering factor. We tested the role of corticosterone on reproductive success by observing the effect of corticosterone implants on reproductive success and on plasma prolactin concentration in female common eiders Somateria mollissima . Implanted females showed a significant increase in corticosterone and a decrease in prolactin levels. Despite their enhanced daily body mass loss, females did not abandon incubation nor did they start to refeed in the four days following implantation. These data show that the experimentally induced rise in plasma corticosterone concentration alone does not trigger nest desertion. However, after 25 days of incubation, implanted females displayed a higher rate of egg loss, suggesting lower nest attentiveness towards the end of incubation. We suggest that the short-term effects of corticosterone may be dependent on the energy state of the bird. However, the late-induced change in reproductive success is indirectly linked to corticosterone, and we suggest that either a prolactin decrease, or a depletion in protein body reserves, may participate in the long-term adjustment of incubation behaviour in female eiders.     

Blood metabolite levels in normal and handicapped pied flycatchers rearing broods of different sizes

We measured levels of select metabolites (glucose, triglycerides, free fatty acids, glycerol, uric acid) and corticosterone in the blood plasma of adult pied flycatchers Ficedula hypoleuca while they were rearing broods whose sizes were modified experimentally. We also made it more difficult than normal for some pairs of birds to forage by removing certain wing and tail feathers (handicapping them). Both procedures have been shown previously to change parental workload. We did this in order to determine if the birds alter their use of nutrients in response to differences in their workload. Metabolite levels were not influenced by handicapping or brood size. However, the concentration of free fatty acids in the plasma of females and of triglycerides in the plasma of males was directly related to the frequency with which the adults fed their nestlings. These findings suggest that the two sexes have different ways of coping with the work associated with rearing the brood: females apparently undergo brief daily fasts while feeding their chicks, whereas males take more time to feed themselves while providing food for their young, and spend more time doing so as their workload increases. The flycatchers exhibited high concentrations of uric acid and corticosterone in the blood plasma; corticosterone and glycerol were positively correlated in females; and corticosterone and triglyceride levels were negatively correlated in males; all of which suggest that gluconeogenesis provides some of the energy required for their parental activities.     

Body mass and clutch size may modulate prolactin and corticosterone levels in eiders

Altered body condition, increased incubation costs, and egg loss are important proximate factors modulating bird parental behavior, since they inform the adult about its remaining chances of survival or about the expected current reproductive success. Hormonal changes should reflect internal or external stimuli, since corticosterone levels (inducing nest abandonment) are known to increase while body condition deteriorates, and prolactin levels (stimulating incubation) decrease following egg predation. However, in a capital incubator that based its investment on available body reserves and naturally lost about half of its body mass during incubation, corticosterone should be maintained at a low threshold to avoid protein mobilization for energy supply. This study focused on the regulation of corticosterone and prolactin release in such birds during incubation, when facing egg manipulation (control, reduced, or increased) or a stressful event. Blood samples were taken before and after clutch manipulation and at hatching. Corticosterone levels were determined before and after 30 min of captivity. Female eiders exhibited a high hypothalamic-pituitary-adrenal sensitivity, plasma concentration of corticosterone being increased by four- to fivefold following 30 min of captivity. The adrenocortical response was not modified by body mass loss but was higher in birds for which clutch size was increased. In the same way, females did not show different prolactin levels among the experimental groups. However, when incubation started, prolactin levels were correlated to body mass, suggesting that nest attendance is programmed in relation to the female initial body condition. Moreover, due to an artifactual impact of bird manipulation, increased baseline corticosterone was associated with a prolactin decrease in the control group. These data suggest that, in eiders, body mass and clutch size modification can modulate prolactin and corticosterone levels, which cross-regulate each other in order to finely control incubation behavior.     

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.     

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.     

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.     

Recess behaviour of the incubating Common Eider Somateria mollissima

Female eiders show the highest incubation constancy among ducks. However, detailed data on their behaviour during the short recesses were not available. In this study, activities during recesses were observed in a colony of female eiders breeding in the high arctic to define what the principal purposes of these temporary nest desertions are. Incubating eiders do not feed during their recess periods. Drinking salt or fresh water or eating snow (28.7%) and preening (38.0%) were the two main activities observed. After snow melting, the time spent drinking decreased (14.3%) while the time spent preening was unchanged (38.1%). Moreover, despite the fact that females started to fly longer distances to reach fresh water sources, the recess duration did not change significantly (401.5 vs 431.0 s). Since recess during incubation increases the risk of breeding failure, drinking presumably has an important survival value for female eiders. In this context, our data suggest that searching for fresh water at a greater distance is favoured despite the higher energetic cost for a fasting bird. Accepted: 5 February 2000     

Do purely capital layers exist among flying birds? Evidence of exogenous contribution to arctic-nesting common eider eggs

The strategy of relying extensively on stored resources for reproduction has been termed capital breeding and is in contrast to income breeding, where needs of reproduction are satisfied by exogenous (dietary) resources. Most species likely fall somewhere between these two extremes, and the position of an organism along this gradient can influence several key life-history traits. Common eiders (Somateria mollissima) are the only flying birds that are still typically considered pure capital breeders, suggesting that they depend exclusively on endogenous reserves to form their eggs and incubate. We investigated the annual and seasonal variation in contributions of endogenous and exogenous resources to egg formation in eiders breeding at the East Bay colony in the Canadian Arctic. We collected prey items along with females and their eggs during various stages of breeding and used two complementary analytical approaches: body reserve dynamics and stable isotope [δ(13)C, δ(15)N] mixing models. Indices of protein reserves remained stable from pre-laying to post-laying stages, while lipid reserves declined significantly during laying. Similarly, stable isotope analyses indicated that (1) exogenous nutrients derived from marine invertebrates strongly contributed to the formation of lipid-free egg constituents, and (2) yolk lipids were constituted mostly from endogenous lipids. We also found evidence of seasonal variation in the use of body reserves, with early breeders using proportionally more exogenous proteins to form each egg than late breeders. Based on these results, we reject the hypothesis that eiders are pure capital layers. In these flying birds, the fitness costs of a strict capital breeding strategy, such as temporary loss of flight capability and limitation of clutch and egg size, may outweigh benefits such as a reduction in egg predation rate.     

Radio‐tags have no behavioral or physiological effects on a migratory songbird during breeding and molt

ABSTRACT Radio‐tags are commonly used in studies of avian behavior and ecology, even though many studies have revealed negative impacts of radio‐tags. Other studies of songbirds have revealed no negative effects of radio‐tags on parental care, nesting success, or annual return rates. However, such studies have not addressed the potential physiological costs of carrying radio‐tags or determined if such costs affect birds during the energetically costly molt period. Simultaneously investigating the behavioral, physiological, and fitness consequences of radio‐tags is important for evaluating possible costs. We studied Wood Thrushes (Hylocichla mustelina) in northwestern Pennsylvania and used a repeated‐measures design to determine if radio‐tags affected parental care behavior and nesting success, and measured levels of plasma lipid metabolites (β‐hydroxybutyrate, free glycerol, and triglycerides) to evaluate physiological condition. We also examined the possible physiological effects of radio‐tags during the energetically demanding period of molt in late summer. Radio‐tags did not affect male or female feeding rates, nesting success, number of nesting attempts, number of eggs laid, fledging success, or hatching success, and we found no significant effects of radio‐tag use on levels of plasma lipid metabolites of nesting females. During molt, Wood Thrushes with radio‐tags had lower levels of β‐hydroxybutyrate and higher levels of trigylcerides, suggesting they were in better, not worse, energetic condition. Our results suggest that radio‐tags do not have negative effects on the behavior or physiology of Wood Thrushes. Similar studies on a wider range of songbirds are needed before concluding radio‐tags have little or no impact during the molting period.     

Regulation of carbohydrate metabolism in lymphoid tissue. Nature of the endogenous substrates and their contribution to the respiratory fuel of the sliced rat spleen in vitro.

1. Tissue glycogen contributes, maximally, only 10% of the respiratory fuel of the rat spleen slice in the absence of an added carbon source, and makes no significant contribution when glucose (3mM) is added. 2. The reserves of fatty acid in the form of triglyceride (35.5mumol of fatty acid/g dry wt. of tissue) fall by approx. 25% after incubation of spleen slices with or without added glucose for 2h, and , on this basis, account for 32% of the oxidative fuel. 3. In contrast, the total oxidative contribution of fatty acid reserves to the respiratory fuel, determined on the basis of inhibiton of respiration by 2-bromostearate, is 42-52%. This range includes tissue from both starved and well-fed animals and is not significantly altered by the presence of added glycose (3mM). 4. Large quantities of NH3 (31-35mumol//h per g dry wt. of tissue) are produced by spleen slices incubated in the absence of added substrates, and this value is suppressed by approx. 50% on incubation with glucose (3mM). Adenine nucleotide breakdown can account for only 17% of the total ammonia produced. 5. Individual free amino acid concentrations in spleen were determined, both in vivo and in slices before and after 60 min of incubation. Although the total free amino acid pool size increases by 45% during incubation, owing to protein breakdown, the tissue concentrations of aspartate, glutamate, glutamine and alanine do not increase. It is suggested that these amino acids areoxidized in a net sense to CO2 and water with the liberation of free NH3 via transamination reactions, glutaminase, the purine nucleotide cycle and the tricarboxylic acid cycle. 6. It is concluded that the normal endogenous metabolism of sliced rat spleen (43-52% due to lipids, 30% due to amino acids and 10% due to glycogen) is modified by added glycose only to the extent that glycogen oxidation and 50% of the contribtion made by ino acids are suppressed; endogenous lipid metabolism is unaffected.     

Patterns of lipid storage and mobilisation in the female green sea turtle ( Chelonia mydas)

Reproductive data from southern Queensland indicate that vitellogenesis in female Chelonia mydas takes approximately 8 months and is followed by a migration to a breeding area. At Heron Island, females lay multiple clutches over approximately 3 months. To investigate how females mobilise and store lipid during the breeding season we collected plasma, yolk, and fat tissue samples from females at a variety of stages during the nesting season. In breeding females, concentrations of plasma triglyceride increased seasonally. They reached peak concentrations during vitellogenesis and courtship, remained high throughout the nesting season, and then declined to a nadir after the last clutch. Plasma protein concentration increased throughout the breeding season, peaking following the last clutch for the season. Yolk lipids were highest during courtship and were similar throughout the nesting season, suggesting that uptake of lipid by ovarian follicles is completed prior to the beginning of the nesting season. Plasma triglyceride decreases in females with prolonged periods of unsuccessful nesting, and total lipid levels in adipose tissue and follicle yolks were significantly lower in atretic females. It appears that: (1) endogenous energy reserves can be reduced by stochastic environmental events (such as those reducing nesting success), and (2) a metabolic shift signalling the end of the nesting season is characterised by a drop in plasma triglycerides and slight increase in total plasma protein.     

Metabolism of endogenous sterol ester by the superovulated rat ovary in vitro

Sterol, glyceride and phospholipid were found to account for more than 90% (w/w) of the lipid extracted from whole superovulated rat ovaries. These lipids, together with non-esterified fatty acids, were assayed in slices of the tissue after incubation for various times. Whereas the concentrations of triglyceride, diglyceride and phospholipid did not change significantly during incubation, that of sterol ester markedly decreased and those of free sterol, monoglyceride and non-esterified fatty acid increased. Evidence is presented that in this tissue (in contrast with other mammalian tissues) the main endogenous substrate for respiration is fatty acid derived from sterol ester.     

The control of fatty acid and triglyceride synthesis in rat epididymal adipose tissue. Roles of coenzyme A derivatives, citrate and l-glycerol 3-phosphate

1. Methods are described for the extraction and assay of acetyl-CoA and of total acid-soluble and total acid-insoluble CoA derivatives in rat epididymal adipose tissue. 2. The concentration ranges of the CoA derivatives in fat pads incubated in vitro under various conditions were: total acid-soluble CoA, 0.20-0.59mm; total acid-insoluble CoA, 0.08-0.23mm; acetyl-CoA, 0.03-0.14mm. 3. An investigation was made of some postulated mechanisms of control of fatty acid and triglyceride synthesis in rat epididymal fat pads incubated in vitro. The concentrations of intermediates of possible regulatory significance were measured at various rates of fatty acid and triglyceride synthesis produced by the addition to the incubation medium (Krebs bicarbonate buffer containing glucose) of insulin, adrenaline, albumin, palmitate or acetate. 4. The whole-tissue concentrations of glucose 6-phosphate, l-glycerol 3-phosphate, citrate, acetyl-CoA, total acid-soluble CoA and total acid-insoluble CoA were assayed after 30 or 60min. incubation. The rates of fatty acid and triglyceride synthesis, calculated from the incorporation of [U-(14)C]glucose into fatty acids and glyceride glycerol respectively, and the rates of glucose uptake, lactate plus pyruvate output and glycerol output were measured over a 60min. incubation. 5. The rate of triglyceride synthesis could not be correlated with the concentrations of either l-glycerol 3-phosphate or long-chain fatty acyl-CoA (measured as total acid-insoluble CoA). Factor(s) other than the whole-tissue concentrations of these recognized precursors appear to be involved in the determination of the rate of triglyceride synthesis. 6. No relationship was found between the rate of fatty acid synthesis and the whole-tissue concentrations of the intermediates, citrate or acetyl-CoA, or with the two proposed effectors of acetyl-CoA carboxylase, citrate (as activator) or long-chain fatty acyl-CoA (as inhibitor). The control of fatty acid synthesis appears to reside in additional or alternative factors.     

Fatty acid signatures of female Brünnich's guillemots Uria lomvia suggests reliance on local prey for replacement egg production

The laying of smaller replacement eggs has been described as a time-saving adaptation because chicks generally grow faster once hatched than inside the egg. However, chicks hatched from smaller eggs have, potentially, lower survival. Consequently, the smaller replacement egg represents a benefit to the female in terms of preserving its own condition at a cost to their offspring. We test these ideas by measuring adult mass changes and plasma lipid concentration changes in male and female Brünnich's guillemots Uria lomvia breeding on Coats Island, Nunavut. Though males lost more mass than females, these differences were not significant. Between laying the first and replacement egg, plasma fatty acid concentrations declined in females and increased in males, suggesting that females mobilise less lipid to preserve their condition after laying the replacement egg. In females, plasma lipid concentrations of the dominant fatty acids found in the eggs (16:0 and 18:1) declined between the laying of first and replacement eggs while plasma concentrations of 20:1 increased in both males and females. We compared the fatty acid signatures of first and replacement egg to look for evidence of differences between the lipid sources for their production. Principal component and discriminant function analyses showed that the fatty acid signatures of replacement eggs were closer to the signatures of the local prey than those of first eggs. We suggest that females rely on local sources of energy to a greater degree for the production of the replacement egg than the first egg, but that endogenous reserves of certain nutrients are important for the production of both eggs.     

Phytohemagglutinin response and immunoglobulin index decrease during incubation fasting in female common eiders

To maximize their fitness, long-lived species face trade-offs between survival and reproduction. The cost of reproduction, which is defined as the negative effect of current parental investment on chances of adult survival and future reproduction, may affect immune function, possibly through hormonal changes. In this study, components of acquired immunity and plasma corticosterone levels of female eiders (Somateria mollissima) have been measured throughout the incubation period as a function of clutch size. These precocial birds lay up to six eggs and fast completely during incubation. Birds were sampled early and late in the incubation period, with clutches ranging from one to four eggs. T-cell-mediated immune response and humoral immunity were assessed by phytohemagglutinin (PHA) skin tests (a challenging method) and measurements of serum immunoglobulins (a monitoring method), respectively. During incubation, responses to PHA injection and immunoglobulin index significantly decreased, by about 40% and 25%, respectively. These observed decreases occurred independently of the number of eggs laid by the females. Corticosterone did not vary significantly during incubation, whatever the clutch size. We conclude that female eiders seem to reallocate their resources from immune function to reproductive effort independently of clutch size or corticosterone levels.     

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.     

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.