Genetic evidence for kin-based female social structure in common eiders (Somateria mollissima)

Kin-based social groups are commonly studied among cooperativelybreeding species but have been less studied in "nontraditional"group breeding systems. We investigated the presence of kin-basedsociality among females in the common eider (Somateria mollisima),a colonial nesting sea duck that exhibits high levels of natalphilopatry in females. Previous studies of female socialityin common eiders have been restricted to observations duringbrood rearing. However, aggregations of female common eidersare also observed during other periods of the life cycle suchas colony arrival and nesting. Here we apply a novel, empiricalframework using molecular markers and field sampling to geneticallycharacterize female social groups at several stages of the commoneider life cycle. When compared with mean estimates of interindividualrelatedness for the entire colony, significantly higher levelsof relatedness were found between females within groups arrivingto the colony in flight, between females and nearest neighborsat the time of nest site selection, and between groups of femalesdeparting the colony with ducklings. Both full-sibling and half-siblingequivalent relationships were also found within these groups.Therefore, throughout each of several stages including in-flightcolony arrival, nesting, and brood rearing, we provide the firstgenetically confirmed evidence of female kin-based social groupsin common eiders and anseriformes in general.     

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

Common eiders, Somateria mollissima, breed on two types of island in the Northern Baltic: open, sparsely vegetated islands and wooded islands with dense mixed forests. On open islands 79.8% of the nests were on open cliffs, exposed to wind and rain whereas on wooded islands 91.7% of the nests were sheltered inside dense spruce and pine thickets. We found that clutch size on open islands was significantly smaller than on wooded islands. Females started breeding simultaneously in the two habitats and they were similar in body size as measured by the length of the radio-ulna. During incubation females on open islands lost weight at a faster rate than females on wooded islands (34 g/day and 19 g/day, respectively). Heat loss is faster on open than wooded islands and therefore we suggest that the faster weight loss of females on open islands result from thermodynamically adverse incubation conditions. Because the eider is an extreme capital breeder, energy used for egg production cannot be used for incubation. To sustain a higher incubation cost on open islands, the optimal clutch size is therefore lower than on wooded islands. Received: 27 January 1997 / Accepted: 14 March 1997     

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.     

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.     

Allometry, bilateral asymmetry and sexual differences in the vocal tract of common eiders Somateria mollissima and king eiders S. spectabilis

Intraspecific sexual differences, high variation, and positive allometry of sexually-selected external display structures are common. Many sexually-selected anatomical specializations occur in the avian vocal tract but intraspecific variation and allometry have been investigated little. The tracheal bulla bulla syringealis occurs in males of most duck species. We quantified variation and size-scaling of the bulla, plus sexual differences in size of trachea, bronchi, and vocal muscles, for 62 common eiders Somateria mollissima and 51 king eiders S. spectabilis. Trends were similar in both species. Bullar ossification and definitive size occurred early in life: bullar size did not differ between first-year and older males. Bullar size did not vary more than size of other body parts (CVs of 3.4–7.0% for bullar length and breadth). Bullar size scaled to body size with negative allometry or isometry. Vocal muscles were 10–50% thicker in males than females, a much greater sexual difference than in body size (CVs of 3–6% on linear body-size variables). Vocal muscles were larger on the left side in both sexes and bilateral asymmetry was slightly more pronounced in males. Low variation and a trend towards negative allometry suggest that bullar size is under stabilizing selection; if bullar size affects vocal attributes of voice, then the latter cannot be condition-dependent. We recommend comparative research on vocal communication, vocal individuality and vocal-tract anatomy and function in eiders and other ducks.     

Consequences of past and present harvest management in a declining flyway population of common eiders Somateria mollissima

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Salt gland function in the common eider duck (Somateria mollissima)

The function of the supra-orbital salt gland was studied in the common eider duck (Somateria mollissima). The maximum salt-secreting capacity was determined in (1) wild ducks which had been living in a marine environment, (2) ducks reared in captivity on fresh water, and (3) ducks from group 2 adapted to salt water. The maximum secreting capacity was found by infusing a solution of NaCl (1000 mosmol·kg^-1) at increasing rates, from 0.691 to 1.671 mosmol·min^-1. Freshwater-adapted ducks secreted at a maximum rate of 0.785 mosmol·min^-1 (1500 mosmol·kg^-1). Adapted to salt water they increased their capacity, and the best duck secreted at a rate of 1215 mosmol·min^-1 (1600 mosmol·kg^-1). The best wild duck secreted at a rate of 1516 mosmol·min^-1. Ducks in group 3 were used to examine the response to a hyperosmotic or an isoosmotic infusion. The amount of salt (NaCl) given per unit time was the same. Given a hyperosmotic solution their salt glands secreted at a high rate: 30 min after the infusion had stopped the ducks had excreted 94% of the sodium infused, 92.9% via the salt gland. Given an isoosmotic solution they secreted at a rate about half the infusion rate: 30 min after cessation of infusion they had excreted 73% of the sodium, 42.9% via the salt gland and the rest by the kidneys.Abbreviations A II angiotensin II - AV I arginine vasotocin - ED freshwater-adapted ducks - FW fresh water - SD saltwater-adapted ducks - SW sea water - WD wild ducks     

Effects of down collection on incubation temperature, nesting behaviour and hatching success of common eiders (Somateria mollissima) in west Iceland

Incubating common eiders (Somateria mollissima) insulate their nests with down to maintain desirable heat and humidity for their eggs. Eiderdown has been collected by Icelandic farmers for centuries, and down is replaced by hay during collection. This study determined whether down collecting affected the female eiders or their hatching success. We compared the following variables between down and hay nests: incubation temperature in the nest, incubation constancy, recess frequency, recess duration, egg rotation and hatching success of the clutch. Temperature data loggers recorded nest temperatures from 3 June to 9 July 2006 in nests insulated with down (n = 12) and hay (n = 12). The mean incubation temperatures, 31.5 and 30.7°C, in down and hay nests, or the maximum and minimum temperatures, did not differ between nest types where hatching succeeded. Cooling rates in down, on average 0.34°C/min and hay nests 0.44°C/min, were similar during incubation recesses. Females left their nests 0–4 times every 24 h regardless of nest type, for a mean duration of 45 and 47.5 min in down and hay nests, respectively. The mean frequency of egg rotation, 13.9 and 15.3 times every 24 h, was similar between down and hay nests, respectively. Hatching success adjusted for clutch size was similar, 0.60 and 0.67 in down and hay nests. These findings indicate that nest down is not a critical factor for the incubating eider. Because of high effect sizes for cooling rate and hatching success, we hesitate to conclude that absolutely no effects exist. However, we conclude that delaying down collection until just before eggs hatch will minimize any possible effect of down collection on females.     

Sexual behavior and plasma androgen concentrations in the male eider duck (Somateria mollissima)

Eider ducks showed clear tidal and seasonal cycles of display when involved in pair-formation behaviour. Plasma androgen (testosterone and dihydrotestosterone) concentrations did not follow similar tidal cycles but there was a 4-fold increase of androgen in spring when rates of display increased 2-fold. There was no difference in androgen levels in blood samples taken from paired birds before and after coitus. Androgens therefore appear to be essential for the expression of sexual behaviour, but there is no apparent quantitative correlation between overt sexual activity and androgen concentrations.     

Re‐colonization by common eiders Somateria mollissima in the Aleutian Archipelago following removal of introduced arctic foxes Vulpes lagopus

Islands provide refuges for populations of many species where they find safety from predators, but the introduction of predators frequently results in elimination or dramatic reductions in island‐dwelling organisms. When predators are removed, re‐colonization for some species occurs naturally, and inter‐island phylogeographic relationships and current movement patterns can illuminate processes of colonization. We studied a case of re‐colonization of common eiders Somateria mollissima following removal of introduced arctic foxes Vulpes lagopus in the Aleutian Archipelago, Alaska. We expected common eiders to resume nesting on islands cleared of foxes and to re‐colonize from nearby islets, islands, and island groups. We thus expected common eiders to show limited genetic structure indicative of extensive mixing among island populations. Satellite telemetry was used to record current movement patterns of female common eiders from six islands across three island groups. We collected genetic data from these and other nesting common eiders at 14 microsatellite loci and the mitochondrial DNA control region to examine population genetic structure, historical fluctuations in population demography, and gene flow. Our results suggest recent interchange among islands. Analysis of microsatellite data supports satellite telemetry data of increased dispersal of common eiders to nearby areas and little between island groups. Although evidence from mtDNA is suggestive of female dispersal among island groups, gene flow is insufficient to account for recolonization and rapid population growth. Instead, near‐by remnant populations of common eiders contributed substantially to population expansion, without which re‐colonization would have likely occurred at a much lower rate. Genetic and morphometric data of common eiders within one island group two and three decades after re‐colonization suggests reduced movement of eiders among islands and little movement between island groups after populations were re‐established. We predict that re‐colonization of an island group where all common eiders are extirpated could take decades.     

Occurrence and food consumption of the common eider,Somateria mollissima,in the Wadden Sea of Schleswig-Holstein

The number of eider in the Wadden Sea of Schleswig-Holstein was counted by aerial surveys during 1986 and 1987. The highest number occurred during migration in October 1987 with 151 000 ducks, the lowest number during the breeding time in May 1987 with 6000 ducks. About 100 000–120 000 eiders moult in July/August in the Wadden Sea of Schleswig-Holstein, but only 30 000–40 000 stay over winter. The average number was 62 000 ducks. Eider have increased in number since the seventies, when the average population size was estimated to be only 23 000. The increase referred mainly to moulting and migrating eider, whereas numbers in winter remained constant. There are substantial changes in the spatial distribution over the year. In most areas sites used during moult, migration and winter can be clearly separated, although so far no obvious differences in the morphology of these areas could be found. The annual food consumption was calculated to be 3.1×10⁶ kg AFDW or 1.3 g AFDW×m⁻²×year⁻¹, which is about 5% of the average biomass of macrozoobenthos. The increase in the number of eider has led to a significant increase in total food consumption of carnivorous birds, which was estimated at 7.1×10⁶ kg AFDW × year⁻¹ in the seventies and now reaches 9.0×10⁶ kg AFDW×year⁻¹, of which the eider takes 34%. The reasons for and consequences of the increase of the eider are discussed in context with the eutrophication of the North Sea and possible competition with shellfishery. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Oxidative capacity of tissues contributing to thermogenesis in eider (Somateria mollissima) ducklings: changes associated with hatching

1. The development of liver and skeletal muscle oxidative capacities during hatching of the common eider (Somateria mollissima) in the Arctic has been investigated by monitoring tissue cytochrome c oxidase activity. 2. The specific activity of the liver enzyme did not change as the embryo underwent hatching, nor during subsequent growth of the duckling into adulthood. 3. Thigh muscle enzyme specific activity increased by a factor of 3.4 during the 24 h prehatching period, remained elevated for at least 48 h after hatching, and then returned to the embryonic (-24 h) level in adults. 4. Histochemically visualized NADH-tetrazolium reductase of a typical red thigh muscle, M. vastus lateralis, showed a distinct increase in activity as the hatching process progressed to completion. 5. Electron microscopy of sectioned M. vastus lateralis revealed a dramatic increase in the density of the myofibrillar structure (number of mitochondrial profiles per unit area), and in the surface area of mitochondrial crista membranes in the course of the 48 h interval from 1 day prehatching to 1 day after hatching. 6. The significance of these changes for the scaling of thermoregulatory heat generation in the newly hatched eider duckling is discussed.     

Neither season nor sex affects the cost of terrestrial locomotion in a circumpolar diving duck: the common eider (Somateria mollissima)

Locomotion accounts for a significant proportion of the energy budget in birds, and selection is likely to act on its economy, particularly where energy conservation is essential for survival. Birds are capable of different forms of locomotion, such as walking/running, swimming, diving and flying, and adaptations for these affect the energetic cost [cost of locomotion (CoL)] and kinematics of terrestrial locomotion. Furthermore, seasonal changes in climate and photoperiod elicit physiological and behavioural adaptations for survival and reproduction, which also influence energy budget. However, little is understood about how this might affect the CoL. Birds are also known to exhibit sex differences in size, behaviour and physiology; however, sex differences in terrestrial locomotion have only been studied in two cursorially adapted galliform species in which males achieved higher maximum speeds, and in one case had a lower mass-specific CoL than females. Here, using respirometry and high-speed video recordings, we sought to determine whether season and sex would affect the CoL and kinematics of a principally aquatic diving bird: the circumpolar common eider (Somateria mollissima). We demonstrate that eiders are only capable of a walking gait and exhibit no seasonal or sex differences in mass-specific CoL or maximum speed. Despite sharing identical limb morphometrics, the birds exhibited subtle sex differences in kinematic parameters linked to the greater body mass of the males. We suggest that their principally aquatic lifestyle accounts for the observed patterns in their locomotor performance. Furthermore, sex differences in the CoL may only be found in birds in which terrestrial locomotion directly influences male reproductive success.     

Differential investment and costs during avian incubation determined by individual quality: an experimental study of the common eider (Somateria mollissima)

Individuals of different quality may have different investment strategies, shaping responses to experimental manipulations, thereby rendering the detection of such patterns difficult. However, previous clutch-size manipulation studies have infrequently incorporated individual differences in quality. To examine costs of incubation and reproductive investment in relation to changes in clutch size, we enlarged and reduced natural clutch sizes of four and five eggs by one egg early in the incubation period in female common eiders (Somateria mollissima), a sea duck with an anorectic incubation period. Females that had produced four eggs (lower quality) responded to clutch reductions by deserting the nest more frequently but did not increase incubation effort in response to clutch enlargement, at the cost of reduced hatch success of eggs. Among birds with an original clutch size of five (higher quality), reducing and enlarging clutch size reduced and increased relative body mass loss respectively without affecting hatch success. In common eiders many females abandon their own ducklings to the care of other females. Enlarging five-egg clutches led to increased brood care rate despite the higher effort spent incubating these clutches, indicating that the higher fitness value of a large brood is increasing adult brood investment. This study shows that the ability to respond to clutch-size manipulations depends on original clutch size, reflecting differences in female quality. Females of low quality were reluctant to increase investment at the cost of lower hatch success, whereas females of higher quality apparently have a larger capacity both to increase incubation effort and brood care investment.     

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.     

Small-scale spatial and temporal variation in the demographic processes underlying the large-scale decline of eiders in the Baltic Sea

The application of uniform conservation schemes often fails to account for small-scale spatial variation in the drivers of population decline. Demographic comparisons of imperilled populations across locations are therefore crucial for successful conservation, but progress is hampered by lack of long-term data from more than a single population. The recent large-scale decline of eider ducks (Somateria mollissima) in the Baltic Sea is ideal for determining to what extent mechanisms underlying population decline can be extrapolated over larger areas. We utilized stochastic demographic methods incorporating both environmental and sampling variation to assess small-scale spatial and temporal variation in the population dynamics of eiders at Söderskär (eastern range-margin) and Tvärminne (core breeding area), situated 130 km apart. The stochastic growth rate models accurately predicted the observed differences in the rate of decline between sites and time periods. At Söderskär, established breeder survival had by far the greatest elasticity, whereas elasticity was more evenly distributed among vital rates at Tvärminne. Although the study sites showed the single largest difference in fecundity, stochastic life table response experiment analyses revealed that reduced adult female survival at Tvärminne mainly determined the observed difference in growth rates between sites. In contrast, reduced fecundity primarily differentiated the past population increase from the present population decline at Söderskär. Our results demonstrate that different mechanisms may be associated with population decline across adjacent geographic locations, and indicate that dispersal of first-time breeders may be important for population dynamics. Safeguarding adult female survival and/or fecundity should be prioritized in management efforts.     

Habitat used by common and king eiders in spring in the southeast Beaufort Sea and overlap with resource exploration

The southeast Beaufort Sea is a critical spring staging area for common and king eiders (Somateria mollissima v-nigrum, S. spectabilis), and is for many the final stop before they reach their breeding grounds throughout the western Canadian Arctic. The region also has significant oil and gas potential, and the recent approval of a pipeline through the Mackenzie Valley may make development of these resources economically viable. We used satellite telemetry to determine the distribution and habitat use of eiders staging in the southeast Beaufort Sea in spring, and the overlap of eiders with oil and gas exploration. From 2004 to 2009, we monitored 51 eiders equipped with platform terminal transmitters (PTTs) throughout spring migration (May–June). We compared the marine habitats used by each species, and evaluated habitat preferences using resource selection functions. The location and extent of the flaw lead (open water along the interface between mobile pack ice and stationary landfast ice) at the time when eiders were staging varied among years, but both species showed a strong preference for use of flaw lead habitats. This preference was stronger for common eiders than for king eiders, which also used pack ice extensively. Common eiders generally occurred near the landfast ice edge, whereas king eiders were just as often nearer to the pack ice edge of the flaw lead. Average water depth (±SE) for common eiders was 22 ± 2 m compared to 30 ± 1 m for king eiders. Kernel density estimators showed that eiders generally occurred in lower densities in areas of otherwise suitable habitat off the Mackenzie River delta. We suggest that this is a result of the highly turbid water discharged by the Mackenzie River, which limits visibility. Oil and gas exploration overlapped significantly with the areas used by eiders. The high density of birds using the restricted and ice-rich flaw lead habitats indicates that an accidental spill in the region could be catastrophic for Canada's western Arctic eider populations. © 2013 The Wildlife Society.     

Reproductive characteristics of the male herring in the northern Baltic Sea

The gonad weight and gonadosomatic index of the male Baltic herring Clupea harengus membras L., were highest at the beginning of the reproductive season (April/May), the values decreasing towards the end of it (July/August) during 1988–1991. The decline could not be explained by fish size but may have been due to fish condition. A high individual variation was typical for both gonad weights and gonadosomatic indices in fish of the same size and maturity stage. The mean density of sperm cells was significantly higher in June (34·9 × 10⁹ ml⁻¹) than in July (19·2 × 10⁹ ml⁻¹, Mann-Whitney U= 17; P<0·05), the variation among the males being high in both groups. Electron microscope analysis showed a severe disruption of the mitochondrial elements in males spawning at 22°C.     

Heterogeneity in blood proteins and egg albumen proteins of the eiderduck,Somateria M. mollissima (L.)

• 1.1. The heterogeneity of blood proteins (including haemoglobin) of adult and subadult male and female Eiderducks, and the egg albumen proteins was studied, in relation with some behavioural properties, as nest-site preference and being migratory or sedentary.
• 2.2. The blood proteins demonstrate a high degree of multiplicity, with two different patterns in the pH range of 5.3–5.7; half the number of blood samples showed two “single” bands. The other half two “double” bands.
• 3.3. The egg albumen proteins show three different patterns, indicating three phenotypes (aa, ab and bb), occurring in four habitat types.
• 4.4. No relation exists between blood protein or egg albumen protein patterns and behavioural aspects of habitat preference for nesting sites.
• 5.5. The Eiderducks of the Vlieland colony are residents, with a high frequency of b-allele, compared to the b-frequency of the Forvie colony birds.