Seabirds reveal mercury distribution across the North Atlantic.

Mercury (Hg) is a heterogeneously distributed toxicant affecting wildlife and human health. Yet, the spatial distribution of Hg remains poorly documented, especially in food webs, even though this knowledge is essential to assess large-scale risk of toxicity for the biota and human populations. Here, we used seabirds to assess, at an unprecedented population and geographic magnitude and high resolution, the spatial distribution of Hg in North Atlantic marine food webs. To this end, we combined tracking data of 837 seabirds from seven different species and 27 breeding colonies located across the North Atlantic and Atlantic Arctic together with Hg analyses in feathers representing individual seabird contamination based on their winter distribution. Our results highlight an east-west gradient in Hg concentrations with hot spots around southern Greenland and the east coast of Canada and a cold spot in the Barents and Kara Seas. We hypothesize that those gradients are influenced by eastern (Norwegian Atlantic Current and West Spitsbergen Current) and western (East Greenland Current) oceanic currents and melting of the Greenland Ice Sheet. By tracking spatial Hg contamination in marine ecosystems and through the identification of areas at risk of Hg toxicity, this study provides essential knowledge for international decisions about where the regulation of pollutants should be prioritized.

Mercury contamination and potential health risks to Arctic seabirds and shorebirds.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of mercury (Hg) on Arctic biota in 2011 and 2018, there has been a considerable number of new Arctic bird studies. This review article provides contemporary Hg exposure and potential health risk for 36 Arctic seabird and shorebird species, representing a larger portion of the Arctic than during previous AMAP assessments now also including parts of the Russian Arctic. To assess risk to birds, we used Hg toxicity benchmarks established for blood and converted to egg, liver, and feather tissues. Several Arctic seabird populations showed Hg concentrations that exceeded toxicity benchmarks, with 50 % of individual birds exceeding the "no adverse health effect" level. In particular, 5 % of all studied birds were considered to be at moderate or higher risk to Hg toxicity. However, most seabirds (95 %) were generally at lower risk to Hg toxicity. The highest Hg contamination was observed in seabirds breeding in the western Atlantic and Pacific Oceans. Most Arctic shorebirds exhibited low Hg concentrations, with approximately 45 % of individuals categorized at no risk, 2.5 % at high risk category, and no individual at severe risk. Although the majority Arctic-breeding seabirds and shorebirds appeared at lower risk to Hg toxicity, recent studies have reported deleterious effects of Hg on some pituitary hormones, genotoxicity, and reproductive performance. Adult survival appeared unaffected by Hg exposure, although long-term banding studies incorporating Hg are still limited. Although Hg contamination across the Arctic is considered low for most bird species, Hg in combination with other stressors, including other contaminants, diseases, parasites, and climate change, may still cause adverse effects. Future investigations on the global impact of Hg on Arctic birds should be conducted within a multi-stressor framework. This information helps to address Article 22 (Effectiveness Evaluation) of the Minamata Convention on Mercury as a global pollutant.

A risk assessment of the effects of mercury on Baltic Sea, Greater North Sea and North Atlantic wildlife, fish and bivalves.

A wide range of species, including marine mammals, seabirds, birds of prey, fish and bivalves, were investigated for potential population health risks resulting from contemporary (post 2000) mercury (Hg) exposure, using novel risk thresholds based on literature and de novo contamination data. The main geographic focus is on the Baltic Sea, while data from the same species in adjacent waters, such as the Greater North Sea and North Atlantic, were included for comparative purposes. For marine mammals, 23% of the groups, each composing individuals of a specific sex and maturity from the same species in a specific study region, showed Hg-concentrations within the High Risk Category (HRC) and Severe Risk Category (SRC). The corresponding percentages for seabirds, fish and bivalves were 2.7%, 25% and 8.0%, respectively, although fish and bivalves were not represented in the SRC. Juveniles from all species showed to be at no or low risk. In comparison to the same species in the adjacent waters, i.e. the Greater North Sea and the North Atlantic, the estimated risk for Baltic populations is not considerably higher. These findings suggest that over the past few decades the Baltic Sea has improved considerably with respect to presenting Hg exposure to its local species, while it does still carry a legacy of elevated Hg levels resulting from high neighbouring industrial and agricultural activity and slow water turnover regime.

DNA damage in Arctic seabirds: Baseline, sensitivity to a genotoxic stressor, and association with organohalogen contaminants.

Environmental contaminants are found throughout Arctic marine ecosystems, and their presence in seabirds has been associated with toxicological responses. However, there are few studies of genotoxicity in Arctic avian wildlife. The purpose of the present study was to quantify deoxyribonucleic acid (DNA) damage in lymphocytes of selected seabird species and to examine whether accumulation of organohalogen contaminants (ΣOHCs) affects DNA damage. Blood was sampled from common eider (Somateria mollissima), black guillemot (Cepphus grylle), black-legged kittiwake (Rissa tridactyla), glaucous gull (Larus hyperboreus), arctic skua (Stercorarius parasiticus), and great skua (Stercorarius skua) in Kongsfjorden, Svalbard (Norway). Contaminant concentrations found in the 6 species differed, presumably because of foraging ecology and biomagnification. Despite large differences in contaminant concentrations, ranging from ΣOHCs 3.3 ng/g wet weight in the common eider to ΣOHCs 895 ng/g wet weight in the great skua, there was no strong difference among the species in baseline DNA damage or sensitivity to a genotoxic stressor (hydrogen peroxide). Baseline levels of DNA damage were low, with median values ranging from 1.7% in the common eider to 8.6% in the great skua. There were no associations between DNA damage and contaminants in the investigated species, suggesting that contaminant concentrations in Kongsfjorden are too low to evoke genotoxic effects, or possibly that lymphocytes are resistant to strand breakage. Clearly, genotoxicity is a topic for future studies of Arctic seabirds. Environ Toxicol Chem 2018;37:1084-1091. © 2017 SETAC.

Hidden survival heterogeneity of three Common eider populations in response to climate fluctuations.

Understanding how individuals and populations respond to fluctuations in climatic conditions is critical to explain and anticipate changes in ecological systems. Most such studies focus on climate impacts on single populations without considering inter- and intra-population heterogeneity. However, comparing geographically dispersed populations limits the risk of faulty generalizations and helps to improve ecological and demographic models. We aimed to determine whether differences in migration tactics among and within populations would induce inter- or intra-population heterogeneity in survival in relation to winter climate fluctuations. Our study species was the Common eider (Somateria mollissima), a marine duck with a circumpolar distribution, which is strongly affected by climatic conditions during several phases of its annual cycle. Capture-mark-recapture data were collected in two arctic (northern Canada and Svalbard) and one subarctic (northern Norway) population over a period of 18, 15, and 29 years respectively. These three populations have different migration tactics and experience different winter climatic conditions. Using multi-event and mixture modelling, we assessed the association between adult female eider survival and winter conditions as measured by the North Atlantic Oscillation (NAO) index. We found that winter weather conditions affected the survival of female eiders from each of these three populations. However, different mechanisms seemed to be involved. Survival of the two migrating arctic populations was impacted directly by changes in the NAO, whereas the subarctic resident population was affected by the NAO with time lags of 2-3 years. Moreover, we found evidence for intra-population heterogeneity in the survival response to the winter NAO in the Canadian eider population, where individuals migrate to distinct wintering areas. Our results illustrate how individuals and populations of the same species can vary in their responses to climate variation. We suspect that the found variation in the survival response of birds to winter conditions is partly explained by differences in migration tactic. Detecting and accounting for inter- and intra-population heterogeneity will improve our predictions concerning the response of wildlife to global changes.

Differences in speciation progress in feather mites (Analgoidea) inhabiting the same host: the case of Zachvatkinia and Alloptes living on arctic and long-tailed skuas.

Recent molecular phylogenetic analyses have revealed that some apparently oligoxenous feather mite species are in fact monoxenous cryptic species with little morphological differentiation. In this study we analyzed two species, Zachvatkinia isolata (Avenzoariidae) and Alloptes (Sternalloptes) stercorarii (Alloptidae) which prefer different parts of the plumage of two sister species of birds: arctic skua (Stercorarius parasiticus) and long-tailed skua (S. longicaudus) breeding on tundra in the High Arctic archipelago of Svalbard. Given that there are no reports about hybridization events between the host species, we expected that both skuas would have a species-specific acarofauna. The genetic distances among DNA-barcode sequences (COI and 28S rDNA), phylogenetic tree topologies, and haplotype networks of the COI sequences of mites suggested extensive gene flow in Z. isolata between and within populations inhabiting both skua species, whereas the Alloptes populations were host specific and sufficiently genetically separated as to warrant species-level status. The discrepancy in the genetic structure of Alloptes and Zachvatkinia populations suggests frequent but transient contacts between the two skua species in which the probability of mite exchange is much higher for Zachvatkinia, which is present in high numbers and inhabits exposed parts of primary flight feathers, than for the less abundant Alloptes that lives primarily in more protected and inaccessible parts of the plumage. We discuss the possible nature of these contacts between host species and the area(s) where they might take place. The star-like structures in the haplotype network as well as high haplotype diversity and low nucleotide diversity observed in Z. isolata are concordant with the known dispersal strategy of feather mites: vertical colonization of new host individuals followed by rapid growth of founder populations.

Feather corticosterone levels on wintering grounds have no carry-over effects on breeding among three populations of great skuas (Stercorarius skua).

Environmental conditions encountered by migratory seabirds in their wintering areas can shape their fitness. However, the underlying physiological mechanisms remain largely unknown as birds are relatively inaccessible during winter. To assess physiological condition during this period, we measured corticosterone concentrations in winter-grown primary feathers of female great skuas (Stercorarius skua) from three breeding colonies (Bjørnøya, Iceland, Shetland) with wintering areas identified from characteristic stable isotope signatures. We subsequently compared winter feather corticosterone levels between three wintering areas (Africa, Europe and America). Among females breeding in 2009, we found significant differences in feather corticosterone levels between wintering areas. Surprisingly, levels were significantly higher in Africa despite seemingly better local ecological factors (based on lower foraging effort). Moreover, contrary to our predictions, females sharing the same wintering grounds showed significant differences in feather corticosterone levels depending on their colony of origin suggesting that some skuas could be using suboptimal wintering areas. Among females wintering in Africa, Shetland females showed feather corticosterone levels on average 22% lower than Bjørnøya and Iceland females. Finally, the lack of significant relationships between winter feather corticosterone levels and any of the breeding phenology traits does not support the hypothesis of potential carry-over effects of winter feather corticosterone. Yet, the fitness consequences of elevated feather corticosterone levels remain to be determined.

Dietary versus maternal sources of organochlorines in top predator seabird chicks: an experimental approach.

We examined the relative importance of dietary sources and maternal transfer on organochlorine concentrations (∑OCs) in Great skua chicks (Stercorarius skua) in Shetland by food supplementing parents with known wintering area. We predicted that experimental chicks (whose parents were supplemented) should have (i) higher growth rates and, (ii) lower ∑OCs due to growth dilution effect and/or due to being fed with less contaminated food compared to control chicks. We also predicted a significant influence of maternal wintering area on chicks' ∑OCs. Plasma ∑OCs of adults, assessed prior to the manipulation, significantly differed between wintering areas of birds. Chicks were weighed every 5 days and plasma ∑OCs were assessed at 20 days old. Based on nitrogen and carbon stable isotope analysis, the supplementary food contributed on average 20% of the dietary protein of the chicks. Although experimental chicks experienced better developmental conditions, supplementary food did not alleviate their organochlorine burden. Nevertheless, chicks whose mothers wintered in Europe showed ∑OCs 50% higher than chicks whose mothers wintered in Africa. Moreover, based on the positive relationship between ∑OCs of chicks and females, the contaminant load of Great skua chicks in Shetland appears to be more influenced by maternal transfer than by trophic transfer.

Trans-equatorial migration routes, staging sites and wintering areas of a high-Arctic avian predator: the long-tailed Skua (Stercorarius longicaudus).

The Long-tailed Skua, a small (<300 g) Arctic-breeding predator and seabird, is a functionally very important component of the Arctic vertebrate communities in summer, but little is known about its migration and winter distribution. We used light-level geolocators to track the annual movements of eight adult birds breeding in north-east Greenland (n = 3) and Svalbard (n = 5). All birds wintered in the Southern Hemisphere (mean arrival-departure dates on wintering grounds: 24 October-21 March): five along the south-west coast of Africa (0-40°S, 0-15°E), in the productive Benguela upwelling, and three further south (30-40°S, 0-50°E), in an area extending into the south-west Indian Ocean. Different migratory routes and rates of travel were documented during post-breeding (345 km d(-1) in late August-early September) and spring migrations (235 km d(-1) in late April) when most birds used a more westerly flyway. Among the different staging areas, a large region off the Grand Banks of Newfoundland appears to be the most important. It was used in autumn by all but one of the tracked birds (from a few days to three weeks) and in spring by five out of eight birds (from one to more than six weeks). Two other staging sites, off the Iberian coast and near the Azores, were used by two birds in spring for five to six weeks. Over one year, individuals travelled between 43,900 and 54,200 km (36,600-45,700 when excluding staging periods) and went as far as 10,500-13,700 km (mean 12,800 km) from their breeding sites. This study has revealed important marine areas in both the south and north Atlantic Ocean. Sustainable management of these ocean basins will benefit Long-tailed Skuas as well as other trans-equatorial migrants from the Arctic.

Antiparasite treatments reduce humoral immunity and impact oxidative status in raptor nestlings.

Parasites are natural stressors that may have multiple negative effects on their host as they usurp energy and nutrients and may lead to costly immune responses that may cause oxidative stress. At early stages, animals may be more sensitive to infectious organisms because of their rapid growth and partly immature immune system. The objective of this study was to explore effects of parasites by treating chicks of two raptor species (northern goshawk Accipiter gentilis and white-tailed sea eagle Haliaeetus albicilla) against both endoparasites (internal parasites) and ectoparasites (external parasites). Nests were either treated against ectoparasites by spraying with pyrethrin or left unsprayed as control nests. Within each nest, chicks were randomly orally treated with either an antihelminthic medication (fenbendazole) or sterile water as control treatment. We investigated treatment effects on plasma (1) total antioxidant capacity TAC (an index of nonenzymatic circulating antioxidant defenses), (2) total oxidant status TOS (a measure of plasmatic oxidants), and (3) immunoglobulin levels (a measure of humoral immune function). Treatment against ectoparasites led to a reduction in circulating immunoglobulin plasma levels in male chicks. TOS was higher when not receiving any parasite reduction treatment and when receiving both endo- and ectoparasitic reduction treatment compared with receiving only one treatment. TAC was higher in all treatment groups, when compared to controls. Despite the relatively low sample size, this experimental study suggests complex but similar relationships between treatment groups and oxidative status and immunoglobulin levels in two raptor species.

Assessing the effects of climate on host-parasite interactions: a comparative study of European birds and their parasites.

BACKGROUND: Climate change potentially has important effects on distribution, abundance, transmission and virulence of parasites in wild populations of animals. METHODOLOGY/PRINCIPAL FINDING: Here we analyzed paired information on 89 parasite populations for 24 species of bird hosts some years ago and again in 2010 with an average interval of 10 years. The parasite taxa included protozoa, feather parasites, diptera, ticks, mites and fleas. We investigated whether change in abundance and prevalence of parasites was related to change in body condition, reproduction and population size of hosts. We conducted analyses based on the entire dataset, but also on a restricted dataset with intervals between study years being 5-15 years. Parasite abundance increased over time when restricting the analyses to datasets with an interval of 5-15 years, with no significant effect of changes in temperature at the time of breeding among study sites. Changes in host body condition and clutch size were related to change in temperature between first and second study year. In addition, changes in clutch size, brood size and body condition of hosts were correlated with change in abundance of parasites. Finally, changes in population size of hosts were not significantly related to changes in abundance of parasites or their prevalence. CONCLUSIONS/SIGNIFICANCE: Climate change is associated with a general increase in parasite abundance. Variation in laying date depended on locality and was associated with latitude while body condition of hosts was associated with a change in temperature. Because clutch size, brood size and body condition were associated with change in parasitism, these results suggest that parasites, perhaps mediated through the indirect effects of temperature, may affect fecundity and condition of their hosts. The conclusions were particularly in accordance with predictions when the restricted dataset with intervals of 5-15 years was used, suggesting that short intervals may bias findings.

Temporal dynamics of circulating persistent organic pollutants in a fasting seabird under different environmental conditions.

Temporal dynamics of persistent organic pollutants (POPs) were examined in fasting common eider (Somateria mollissima) females in one subarctic (68° N; over 5 years) and one high arctic colony (78° N; 3 years). Blood concentrations of polychlorinated biphenyl (PCB)-153; 1-dichloro-2,2-bis (p-chlorophenyl) ethylene (p,p'-DDE), and hexachlorobenzene (HCB) were measured twice each season in eider females (total n = 162) during incubation (at day 5 and day 20). The mean wet weight concentrations of PCB-153 were ~3-5 times higher in the subarctic colony, whereas p,p'-DDE and HCB concentrations tended to be higher in high arctic than in subarctic eiders late in the incubation period. All POPs increased during incubation fast, but the relative increase in mean concentration varied more among years in high arctic than in subarctic eiders. In the high arctic, both lipid-metabolism and the increase in circulating POP concentrations were highest in the year when the mean ambient temperature was lowest. Moreover, females with low body condition and high lipid metabolism (body mass loss) had stronger increase in circulating concentrations of p,p'-DDE and HCB; the effect size being within the same order of magnitude in the two colonies. Hence, since eiders at high latitudes metabolized relatively more lipids, they experienced higher exposure of p,p'-DDE and HCB over the incubation period than birds inhabiting the more benign subarctic region.

Individual variation in biomarkers of health: influence of persistent organic pollutants in Great skuas (Stercorarius skua) breeding at different geographical locations.

Persistent organic pollutants (POPs) have been shown to cause adverse effects on a number of biomarkers of health in birds. POPs may impair immune function and alter the stress response, defined as a suite of behavioral and physiological responses to environmental perturbations. Recent studies have also proposed that POPs can induce oxidative stress. Nevertheless, there is a lack of studies simultaneously assessing the potential damaging effects of POPs on the latter biomarkers. In this study, we examined the contribution of legacy (organochlorines; (OCs)) and emerging (flame retardants; PBDEs) POPs to individual variations in stress levels (feather corticosterone), humoral immunity (plasma immunoglobulin Y levels) and oxidative stress occurring in three breeding colonies of a top predator seabird, the Great skua (Stercorarius skua), distributed from temperate regions to the high Arctic: Shetland (60°N), Iceland (63°N) and Bjørnøya (74°N). Our results demonstrated that plasma concentrations of OCs in Great skuas from Bjørnøya are among the highest in North Atlantic seabirds, with up to 7900 µg/kg (ww) ∑OCs. Yet, a latitudinal gradient in POP levels was observed with all compounds being significantly higher in Bjørnøya than in Iceland and Shetland (on average 4-7 fold higher for OCs and 2.5-4.5 for PBDEs, respectively). Contrary to our predictions, skuas breeding at the least contaminated site (i.e., Shetland) experienced the poorest physiological condition; i.e., the highest levels of stress hormones (25% higher) and oxidative stress (50% higher) and the lowest immunoglobulin levels (15% lower) compared to the two other colonies. Finally, our results failed to point out consistent within-colony relationships between biomarkers of health and POPs. Overall, it is suggested that other ecological factors such as food availability could constrain physiological indicators more than anthropogenic contaminants.

A first evaluation of the usefulness of feathers of nestling predatory birds for non-destructive biomonitoring of persistent organic pollutants.

In previous studies, feathers of adult predatory birds have been evaluated as valid non-destructive biomonitor matrices for persistent organic pollutants (POPs). In this study, we assessed for the first time the usefulness of nestling raptor feathers for non-destructive biomonitoring of POPs. For this purpose, we collected body feathers and blood of nestlings from three avian top predators from northern Norway: northern goshawks (Accipiter gentilis), white-tailed eagles (Haliaeetus albicilla) and golden eagles (Aquila chrysaetos). We were able to detect a broad spectrum of legacy POPs in the nestling feathers of all three species (Σ PCBs: 6.78-140ng g(-1); DDE: 3.15-145ng g(-1); Σ PBDEs: 0.538-7.56ng g(-1)). However, these concentrations were lower compared to other studies on raptor species, probably due to the aspect of monitoring of nestlings instead of adults. Besides their analytical suitability, nestling feathers also appear to be biologically informative: concentrations of most POPs in nestling feathers showed strong and significant correlations with blood plasma concentrations in all species (p<0.050; 0.775

Indeterminate laying and flexible clutch size in a capital breeder, the common eider.

Clutch size control in capital breeders such as large waterfowl has been much debated. Some studies have concluded that clutch size in ducks is determined before the start of laying and does not change in response to egg additions or removals. The response, however, may depend on the timing of tests, and experiments may have been too late for females to alter the number of eggs. We here study clutch size responses to predation of first and second eggs in the common eider, using protein fingerprinting of egg albumen to verify that the same female continues laying in the nest after predation. Sixty of 79 females with early egg predation (one or both of the two first eggs) deserted the nest. Among the 19 females that stayed and continued laying, the mean number of eggs produced was 4.4, significantly higher than the 3.7 in non-predated nests. The staying females had similar egg size and clutch initiation date as females that deserted, and their body mass and clutch initiation date was similar to that of females whose clutches were not predated. Even capital-breeding common eiders may therefore be indeterminate layers, as many females in which early eggs are removed lay more eggs than others. A previous study has shown that they can reduce their laying if eggs are added. Our results add to increasing evidence that ducks have more flexible egg production than previously thought.

Strongly increasing blood concentrations of lipid-soluble organochlorines in high arctic common eiders during incubation fast.

Female common eiders (Somateria mollissima) starve during the nesting stage and may loose 30-45% of their initial body mass, mostly through lipid mobilization. In this study, the effects of fasting on the blood concentrations of three lipid-soluble organochlorines (OCs: polychlorinated biphenyl [PCB]-153; 1-dichloro-2,2-bis (p-chlorophenyl) ethylene [p,p'-DDE]; and hexachlorobenzene [HCB]) were examined in eiders breeding in the high Arctic. Blood samples were taken from females (n=47) at day 5 and day 20 of the incubation period. The mean wet weight concentrations of PCB-153 and p,p'-DDE increased strongly between day 5 and day 20 (3.6 and 8.2-fold, respectively), while HCB increased less (1.7-fold). There was a strong negative association between daily increase in PCB-153 and clutch size, and a weaker relationship for p,p'-DDE, suggesting that maternal transfer to the eggs is a significant pathway of elimination of OCs in eiders. Moreover, poor body condition (body mass controlled for body size) late in the incubation period was associated with strong daily increase of both p,p'-DDE and PCB-153, which may suggest that the release of these compounds increases when lipid reserves become depleted. For HCB, the increase was mainly associated with increase in blood lipid concentrations, and weakly to the amount of burned lipids. The causes for the differences between the compounds are, however, poorly understood. Although the absolute levels of OCs in eiders were relatively low, their rapid build up during incubation is worrying as it coincides with poor body condition and weakened immune systems.

Costs of an immune challenge and terminal investment in a long-lived bird.

An induced immune challenge can have two counteracting effects on an individual's reproductive investment. (1) The resource demand could increase to "fuel" the immunologic reaction, which in turn can lead to an adaptive decrease in investment in resource-costly activities, such as reproduction. One the other hand, (2) the individual could assume that the immune activity it experiences is indicative of a serious infection. The latter can lead to an adaptive increase in reproductive investment in response to the reduced prospects of survival and future reproduction, so called "terminal investment." To measure such life-history-related consequences of increased immune activity, one group of incubating female Common Eiders (Somateria mollissima) was injected with a nonpathogenic antigen (sheep red blood cells, SRBC) while controls were injected with sterile saline. The eider is a long-lived sea-duck. Females, who incubate the eggs and care for young without assistance from the male, engage in facultative anorexia during incubation leading to a large reduction in body mass. Eiders can abandon their young to other females at the cost of reduced young survival. The immune challenge resulted in a larger mass loss, a prolonged incubation period, and reduced return rate, demonstrating both short- and long-term costs of immune challenge. Additionally, in response to what might have been interpreted as reduced survival chances in immune-challenged females, these females more often tended their own brood after hatching, despite having suffered higher costs during incubation.

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.

Costs of immunity: immune responsiveness reduces survival in a vertebrate.

Immune defences are undoubtedly of great benefit to the host, reducing the impact of infectious organisms. However, mounting immune responses also entails costs, which may be measured by inducing immune responses against artificial infections. We injected common eider (Somateria mollissima) females with three different non-pathogenic antigens, sheep red blood cells (SRBC), diphtheria toxoid and tetanus toxoid, early in their incubation period. In the group of females that mounted a humoral immune response against SRBC, the return rate was only 27%, whereas the group of females that did not mount a response against SRBC had a return rate of 72%. Moreover, responding against diphtheria toxoid when also responding against SRBC led to a further reduction in return rate. These results are repeatable, as the same effect occurred independently in two study years. The severely reduced return rate of females producing antibodies against SRBC and diphtheria toxoid implies that these birds experienced considerably impaired long-term survival. This study thus documents severe costs of mounting humoral immune responses in a vertebrate. Such costs may explain why many organisms suppress immunity when under stress or when malnourished, and why infections may sometimes be tolerated without eliciting immune responses.

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.