Movements and mortality of common goldeneye Bucephala clangula broods in a patchy environment

We tested the effect of timing and intensity of deer browsing on compensatory response by willow grown for bioenergy. Browsing was simulated several times during summer and once during winter. The clipping was done at different intensities during the first year after establishment in a new willow plantation, and during the first year after harvest in an older willow coppice. Total aboveground biomass and biomass available for deer browsing were recorded both at the end of the first and the second growing periods after clipping. Both in the new plantation and in the older coppice, the willows fully compensated for biomass losses after winter clipping, irrespective of clipping intensity. On the other hand, total biomass production usually decreased after high-intensity summer clipping. Such a seasonal difference may be explained by the nutrition of remaining buds after clipping. In the new willow plantation, total produced biomass after summer clipping still bottomed at about 50% of the level of control stools at the end of the experiment. In the older coppice, usual browsing levels by deer in summer, roughly corresponding to the moderate clipping intensities used in this experiment, did not influence total produced biomass. The newly established willow stools reacted to summer clipping by undercompensating in terms of biomass available for deer browsing, while older stools could overcompensate. Compared to stools clipped in late summer, willow stools clipped during early summer were able to compensate earlier and stronger. The higher the proportion of twig biomass removed by summer clipping, the lower the compensatory growth by willow. Depending on the circumstances, the fast-growing willows responded within the whole continuum from under- to overcompensation, and some implications for the management of willow plantations and large herbivores are discussed.     

Conspecific nest parasitism is associated with inequality in nest predation risk in the common goldeneye (Bucephala clangula)

Previous studies of the role of nest predation in conspecificnest parasitism have not taken into account the possibilitythat predation risk may not be randomly distributed among nestsites and that breeding individuals may use different cues toassess the risk and adjust their reproductive tactic betweenyears accordingly. Especially in cavity-nesting species, therole of nest predation in conspecific nest parasitism has beendownplayed, while the role of nest site limitation has beenhighlighted. Using both observational and experimental data,I show that in the common goldeneye (Bucephala clangula), acavity-nesting species in which conspecific nest parasitism iscommon, predation risk varies considerably between nest sitesand does not follow a random expectation. The inequality inpredation risk between nest sites also showed up in the occurrenceof parasitized nests in an experimental setup. Nests parasitizedin year t were more frequent in those nest sites that were notdepredated during the previous nesting attempt in year t - nthan in nest sites that were depredated and in control nestsites that had not been used for nesting before. A nest siteaddition experiment revealed that conspecific nest parasitismwas not associated with nest site limitation. My findings givesupport for the hypothesis that nest predation is an importantecological factor explaining conspecific nest parasitism ingoldeneyes.     

Survival of common goldeneye Bucephala clangula ducklings in relation to weather, timing of breeding, brood size, and female condition

The survival of common goldeneye Bucephala clangula ducklings during their first week of life was studied in relation to hatching date, brood size, female condition, and weather (temperature and precipitation) during the first week post-hatch by using data from radio-marked females and their broods. Also, age-specific variation in the survival of the young was determined until fledging (over 50 d of age). Survival was lowest in the first week after hatching. Hatching date, brood size, and first week temperature and precipitation were poor predictors of duckling survival during the first week after nest exodus. Instead, the ducklings of females in a better body condition survived better in their first week of life. The results suggest that weather does not have a direct effect on downy ducklings' survival, but the condition of the female seems to be an important determinant of the survival of common goldeneye ducklings.     

Decline of Canada geese (Branta canadensis) and common goldeneye (Bucephala clangula) associated with a collapse of eelgrass (Zostera marina) in a Nova Scotia estuary

Mean numbers of migrant Canada geese (Branta canadensis) in Antigonish Harbour in the southern Gulf of St. Lawrence (Canada) during October to December were similar (approx. 450–500 birds) for the period 1998–2000. Similarly, during this period, geese used two foraging sites. However, in 2001, the average number of birds decreased by half and the primary foraging sites were used only rarely. This coincided with a decline of about 95% in the biomass of roots and rhizomes of eelgrass (Zostera marina) that occurred between October 2000 and 2001. Eelgrass is the principal food of geese in this estuary. In addition, there was a reduction of around 50% in the numbers of common goldeneye (Bucephala clangula), which feed on invertebrates associated with eelgrass. Lower than usual weekly abundances of geese and goldeneye are probably the result of an unusually short residence time in the estuary, rather than a decline in the total number of visiting migrants. We attribute these changes in the distribution and abundance of geese and goldeneyes to the dramatic decline in eelgrass. Electronic Publication     

Selection of nest sites by a hole-nesting duck, the Goldeneye Bucephala clangula

Nest site preferences were examined for a population of Goldeneye Ducks breeding in nest boxes in Värmland, central Sweden. Some nest boxes were occupied more often than others even if females returning to the same nest box were excluded from the analysis. Nest boxes located higher up trees were occupied more often than those close to the ground and some spatial 'cluster groups' of boxes were occupied more often than others. Otherwise nest site prefernces were not related to any measured physical attributes of the boxes. Prefernces for nest boxes seemed to be based mainly on a tendency for females to select those that had been occupied by other females in the preceding year, especially if they had bred successfully. As a result of this, the occupancy of nest boxes was not random over years but rather progressed in a series of runs; a period of consecutive years in which a box was occupied was followed by a period of years in which it was empty.
There were reproductive consequences for these prefernces in that females occupying preferred boxes were less likely to lose their clutch to a predator. These females also bred earlier in the year and produced larger clutches and broods than females breeding in other boxes.     

Female wing plumage reflects reproductive success in Common Goldeneye Bucephala clangula

Recent studies on the function of female plumage characteristics have yielded ambiguous results. Some studies have found an association between different physiological, ecological or behavioural traits and female plumage, while others have found no association and interpret female plumage as neutral in function. We observed a high variance among females in both wing plumage and breeding success in female Common Goldeneyes Bucephala clangula , a sexually plumage-dimorphic diving duck. We studied the association between female wing plumage and hatching date. Principal component analysis of four wing patch area measurements derived a single factor describing wing plumage. Wing plumage was strongly associated with hatching date, which is the most important determinant of goldeneye recruit production; irrespective of age, females with more white in the wing bred earlier than individuals with more black in the wing. We propose that the wing pattern in Common Goldeneye females reflects individual quality.     

Isolation and characterization of 22 polymorphic microsatellite loci for the Barrow's goldeneye (Bucephala islandica)

We describe primers and polymerase chain reaction conditions to amplify 22 microsatellite loci from the Barrow's goldeneye (Bucephala islandica). The primers were tested on 27 individuals from a population breeding in British Columbia, Canada. The developed primer pairs yielded an average of 6.11 alleles per locus (range 2-12), an average observed heterozygosity of 0.70 (range 0.07-0.96) and a polymorphic information content of 0.07-0.88.     

Reliability of egg morphology to detect conspecific brood parasitism in goldeneyes Bucephala clangula examined using protein fingerprinting

Eadie (1989) developed a method based on variation between females in egg length, width and weight to detect conspecific brood parasitism in the field: using these three egg measures, Euclidean distance between all pairs of eggs within a clutch is calculated, and if maximum Euclidean distance (MED) between any two eggs exceeds a threshold value the nest is considered parasitized. The MED method has been tested in Finnish and Scottish common goldeneye Bucephala clangula populations but the results have been contradicting. Here we use protein fingerprinting to assess the validity of the MED method. Data comprised 35 clutches of which we knew, based on protein fingerprinting, how many different females laid the clutch (range 1–5 females). The mean MED of non-parasitized clutches (laid by 1 female only) was 1.470 (95% CI: lower 1.169, upper 1.771; n=21) and that of parasitized clutches (laid by 2 or more females) was 3.654 (95% CL: lower 3.083, upper 4.225; n=14). Using a MED>3.0 as a criterion to identify parasitized clutches 89% of all clutches were classified correctly either parasitized or non-parasitized when compared to the identification based on protein fingerprinting. Clutch size and the number of females (beyond 2 females) did not affect the clutch MED, whereas the status of parasitism did. Repeatability of egg length, width and weight were: 0.63, 0.76 and 0.80, respectively, implying that, variation in these egg measures occurs among rather than within females. Our new results confirm that the MED method is reliable enough to detect parasitism in common goldeneye.     

Maturation costs affect maturation timing: sexual reproduction in a heterogonic hydra

If maturation is more costly for females, they may need more distinct environmental cues to induce sexual reproduction than males. We verified this hypothesis by comparing the indirect costs of maturation to males and females of the heterogonic Hydra oligactis, reproducing both asexually and sexually. The laboratory experiments revealed that males mature 2 weeks earlier than the first females at falling temperatures simulating the natural conditions that precede sexual reproduction. The difference between the energy costs of maturation for males versus females has been considered a likely factor responsible for the observed difference in maturation time. Available food supply positively affected the percentage of sexually mature females, indicating that females are more sensitive to food limitation than males. The number of gonads was correlated positively with the size of mature hydra for both males and females. However, males produced twice as many testes as ovaries produced by females. We postulate that females are induced later than males in order to prevent gonadal development after an unseasonable drop in temperature. As sexual reproduction in H. oligactis interferes with asexual budding, under favorable conditions for asexual proliferation unnecessary gonadal development decreases an individual’s fitness through reduction of the number of produced offspring.     

Costs and benefits of female-biased natal philopatry in the common goldeneye

Sex-biased natal dispersal in long-lived species may resultin interactions between parents and mature young of the philopatricsex. To investigate the evolutionary basis of natal philopatryin a noncooperative species, the common goldeneye Bucephalaclangula, we studied possible costs and benefits of simultaneousbreeding of females and philopatric daughters. We did not find any fitness consequences of a daughter's breeding on their mother'sbreeding in terms of nest-site selection, body weight, clutchsize, hatching date, or hatching success. Our results, therefore,did not support the assumption of the local resource competitionhypothesis, that the natally philopatric sex should be morecostly to a breeding parent. As possible benefits for daughters returning to their natal area, we tested inheritance of nestsites from mothers and explored whether daughters utilize thepresence of their mother by parasitically sneaking into hermother's nest. Daughters' nest-site selection was not associatedwith the presence of their mothers. A comparison between daughtersand control females revealed that daughters chose their nestsite closer to their natal nest than expected by nest-siteavailability alone. Daughters could not expect to inherit anest site from their mother, and we did not find other indicationsof cooperation between relatives either. The mother's clutchsize did not increase in the year breeding with the daughter, indicating daughters do not parasitize their mother's nest.We suggest that benefits such as decreased nest predation riskassociated with nesting close to the natal nest site may beimportant in the natal philopatric behavior of the species.     

Feeding ecology of the diving ducks pochard (Aythya ferina), tufted duck (A. fuligula), scaup (A. mania) and goldeneye (Bucephala clangula) overwintering on Lough Neagh, Northern Ireland

1. The diets of pochard (Aythya ferina), scaup (A. marila) and goldeneye (Bucephala clangula) overwintering on Lough Neagh are dominated by chironomid larvae, while molluscs are more important in that of tufted duck (A. fuligula). 2. Inshore areas of Lough Neagh offer poor feeding conditions for these diving ducks because chironomid larvae and molluscs are of small individual body size or low abundance. These factors lead to all four ducks foraging at least in part at depths greater than those usually exploited. 3. Due to their common consumption of molluscs, the diet of tufted duck shows a higher overlap with that of an introduced roach (Rutilus rutilus) population than with any other duck or fish species. 4. The feeding ecology of tufted duck and roach in Lough Neagh may form an example of distant competition and be at least partly responsible for recent fluctuations in the numbers of tufted duck.     

Relatedness and spatial proximity as determinants of host–parasite interactions in the brood parasitic Barrow's goldeneye (Bucephala islandica)

Recent studies, which have found evidence for kin-biased egg donation, have sparked interest in re-assessing the parasitic nature of conspecific brood parasitism (CBP). Since host–parasite kinship is essential for mutual benefits to arise from CBP, we explored the role of relatedness in determining the behaviour of conspecific nest parasites and their hosts in nesting female Barrow's goldeneyes ( Bucephala islandica ), a duck in which CBP is common. The results revealed that the amount of parasitism increased with host–parasite relatedness, the effect of which was independent of geographical proximity of host and parasite nests. Proximity per se was also positively associated with the amount of parasitism. Furthermore, while hosts appeared to reduce their clutch size as a response to the presence of parasitic eggs, the magnitude of host clutch reduction also tended to increase with increasing relatedness to the parasite. Hence, our results indicate that both relatedness and spatial proximity are important determinants of CBP, and that host clutch reduction may be an adaptation to nest parasitism, modulated by host–parasite relatedness. Taken together, the results provide a demonstration that relatedness influences host and parasite behaviour in Barrow's goldeneyes, resulting in kin-biased egg donation.     

Measuring the costs of reproduction

The measurement of costs of reproduction is of interest because such costs are generally assumed by life history theory. There is some controversy concerning how to measure costs: common methods include experimental manipulations of life history, such as preventing some individuals from reproducing, or estimates of genetic correlations. These two methods often yield similar results, suggesting that one can serve as a substitute for the other. There are now experiments which demonstrate that there are different mechanisms underlying the response to an experimental manipulation versus a genetic correlation, so the two methods are not equivalent in estimating costs.     

Brood size and immunity costs in zebra finches Taeniopygia guttata

Birds rearing experimentally enlarged broods have lower antibody responses to a novel antigen, and we tested three hypotheses that could explain this result. We used zebra finches Taeniopygia guttata inoculated with sheep red blood cells (SRBC) as a study system, for which this trade-off was previously demonstrated. 1. Compensatory cellular immunity: The humoral immune response is slow, and removal of SRBC through up-regulated cellular immunity could pre-empt an antibody response. However, cellular immune response to PHA decreased with increasing brood size, allowing rejection of this hypothesis. 2. Costs of antibody-production: Chicks in large broods grow less well, and birds with large broods may allocate resources to chicks instead of antibodies when these are costly. Compared to saline controls, SRBC suppressed metabolic rate in the hours following immunisation, but there was no effect in the following night, or at any time 4 and 8 days later. Fitness costs were measured by repeatedly immunising parents with SRBC while rearing young. Chick growth, parental condition, and subsequent reproduction of the parents were not affected by SRBC. We conclude that the costs of antibody formation cannot explain the trade-off between brood size and antibody responsiveness. 3. Costs of immune system maintenance: Maintaining a system enabling antibody-formation may be very costly, and birds rearing large broods may have down-regulated this system. Based on this hypothesis we predicted that antibody formation would still be reduced in parents rearing large broods when immunised after rearing the chicks. Our results confirmed this prediction, and we suggest that birds rearing large broods have lower antibody responses because they economised on the maintenance costs of the immune system.     

Brood parasitism,female condition and clutch reduction in the common eider Somateria mollisima

Conspecific brood parasitism (CBP) is an alternative reproductive tactic found in many animals with parental care. Parasitizing females lay eggs in the nests of other females (hosts) of the same species, which incubate and raise both their own and the foreign offspring. The causes and consequences of CBP are debated. Using albumen fingerprinting of eggs for accurately detecting parasitism, we here analyse its relation to female condition and clutch size in High Arctic common eiders Somateria mollissima borealis. Among 166 clutches in a Svalbard colony, 31 (19%) contained eggs from more than one female, and 40 of 670 eggs (6%) were parasitic. In 6 cases an active nest with egg(s) was taken over by another female. Many suitable nest sites were unoccupied, indicating that CBP and nest takeover are reproductive tactics, not only consequences of nest site shortage. Similarity in body mass between female categories suggests that condition does not determine whether a nesting female becomes parasitised. There was no evidence of low condition in parasites: egg size was similar in hosts and parasites, and parasitism was equally frequent early and late in the laying season. Meta‐analysis of this and 3 other eider studies shows that there is a cost of being parasitised in this precocial species: host females laid on average 7% fewer eggs than other females.