The occurrence and performance of leucistic Barnacle Geese Branta leucopsis

The paper reports the occurrence of whitish 'leucistic' forms in populations of Barnacle Geese Branta leucopsis. A total of 15 (including 11 in the last 21 years) have been reported in the population breeding in Svalbard and wintering in the Solway Firth, northern Britain, whereas the two other populations, though much larger, have produced only one bird of the same type between them in the last 30 years. Leucism is controlled by a single, recessive allele.
The median lifespan of leucistic birds is 2–3 years, compared to 8–10 years for normal plumaged birds. This is because they are sought out by (illegal) hunters. There is no evidence that there is any difference in pairing or breeding performance between white and normal birds, though one male, which is still alive aged 18 years, has produced 13 young; this performance is matched fay less than 2% of normal geese.
It is suggested that the Svalbard population was established recently, from a few founders from the Siberian population; if one or two of these carried the allele controlling leucism, this would explain its much higher prevalence in this population than in the other discrete groups of Barnacle Geese.     

Possible effects of local enrichment by gulls on feeding-site selection by wintering Barnacle Geese Branta leucopsis

The selection of salt-marsh feeding sites by wintering Barnacle Geese Branta leucopsis was investigated on Schiermonnikoog, The Netherlands. Use of sites by geese was estimated by counts of goose droppings. Geese spent significantly more time grazing in sites with evidence of Herring Gull Larus argentatus and Lesser Black-backed Gull Lfuscus breeding activity than in nearby non-gull sites. The nitrogen content of red fescue grass Festuca rubra was significantly greater in gull sites. We suggest that the geese preferred to feed in these locations because of the improved nutritional value of the grass. The higher nitrogen content of F. rubra in gull sites was most likely the result of nutrient inputs from gull droppings rather than goose droppings.     

Body size,fitness and compatibility in Barnacle Geese Branta leucopsis

Large-sized Barnacle Geese Branta leucopsis of both sexes had a higher probability of breeding successfully in any particular year and produced more goslings than did smaller birds. Large females paired at an earlier age, suggesting that they were preferred as mates and were likely to have entered the breeding population earlier. The relative sizes of the pair bond members also affected fitness. Most birds were able to maximize their breeding performance by mating with relatively similar sized partners; the greater the size disparity of mates, the lower the breeding performance. This supports the idea that compatibility of mates may be important in determining fitness of the pair. The success of different pair types was also affected by environmental conditions, with certain size combinations doing better in some years and poorly in other years.     

Environmentally induced morphological variation in the Barnacle Goose,Branta leucopsis

The environmental conditions to which juvenile barnacle geese (Branta leucopsis) were exposed during growth were found to affect their body size at fledging as well as their final adult body size. Small juveniles showed compensatory growth from the time of fledging up to one year of age, but this did not fully compensate the differences in body size that were established before fledging. The variation in protein content in plants eaten during growth could probably explain the observed body size differences, sometimes of more than 10%, between different categories of adult geese. Our results imply that one cannot infer selection on morphological characters from differences between samples of adult birds from different localities or from different cohorts within a population, without first showing that environmental conditions during growth do not affect the development of the characters under study.     

The development of a genome wide SNP set for the Barnacle goose Branta leucopsis

Migratory birds are of particular interest for population genetics because of the high connectivity between habitats and populations. A high degree of connectivity requires using many genetic markers to achieve the required statistical power, and a genome wide SNP set can fit this purpose. Here we present the development of a genome wide SNP set for the Barnacle Goose Branta leucopsis, a model species for the study of bird migration. We used the genome of a different waterfowl species, Mallard Anas platyrhynchos, as a reference to align Barnacle Goose second generation sequence reads from an RRL library and detected 2188 SNPs genome wide. Furthermore, we used chimeric flanking sequences, merged from both Mallard and Barnacle Goose DNA sequence information, to create primers for validation by genotyping. Validation with a 384 SNP genotyping set resulted in 374 (97%) successfully typed SNPs in the assay, of which 358 (96%) were polymorphic. Additionally, we validated our SNPs on relatively old (30 years) museum samples, which resulted in a success rate of at least 80%. This shows that museum samples could be used in standard SNP genotyping assays. Our study also shows that the genome of a related species can be used as reference to detect genome wide SNPs in birds, because genomes of birds are highly conserved. This is illustrated by the use of chimeric flanking sequences, which showed that the incorporation of flanking nucleotides from Mallard into Barnacle Goose sequences lead to equal genotyping performance when compared to flanking sequences solely composed of Barnacle Goose sequence.     

Intraclutch egg-size variation in the Barnacle Goose Branta leucopsis: an egg-removal experiment

We carried out an egg-removal experiment on Barnacle Geese Branta leucopsis to test the hypothesis that tactile contact between the brood patch and eggs accumulating in the nest during laying was causally related to intraclutch egg-size variation, acting through the development of incubation behaviour. Egg removal had no effect on mean egg mass, relaying interval or clutch size: mean (±s.e.) clutch sizes for first clutches were 4.9 ± 0.9 and 4.7 ± 0.9 eggs for control and experimental females, respectively. We therefore conclude that Barnacle Geese have a determinate pattern of egg laying. Daily removal of all eggs from the first-laid egg did not affect the within-clutch pattern of egg-size variation: in both control and experimental clutches the first egg was relatively small, the second or third egg was largest and there was a linear decline in egg size to the smallest, last-laid egg. There was no significant difference in either the absolute or relative size of the last-laid eggs in control and experimental clutches. Tactile contact with eggs in the nest, therefore, is not required for the expression of intraclutch egg-size variation in Barnacle Geese. In this respect, determinate-laying Barnacle Geese differ from species with indeterminate laying patterns (e.g. gulls) where egg removal does affect the normal pattern of within-clutch egg-size variation. If hormonal changes (e.g. prolactin) associated with onset of incubation are causally related to the intraclutch decrease in egg size, then the stimuli involved appear to differ between determinate and indeterminate layers.     

Barnacle Goose (Branta leucopsis) feeding ecology and trophic relationships on Kolguev Island: The usage patterns of nutritional resources in tundra and seashore habitats

Insight into colonization of new areas by invasive species and their relationships with aboriginal species is a major challenge in ecology, allowing for prediction of the outcome of biological invasions. In this context, the current expansion of the Barnacle Goose (Branta leucopsis) is of great interest. The seasonal dynamics of Barnacle Goose feeding and the structure of its diet on Kolguev Island during the nesting and postnesting periods (May 27–July 29) have been examined. The diet structures of the invasive Barnacle Goose and the aboriginal species White-fronted goose (Anser albifrons) and Bean goose (A. fabalis) have been compared. The probability of interspecific competition and the ecological advantages in the diet patterns of these species have been estimated. If the Barnacle Goose population on Kolguev Island continues to increase, this may led to a decline in the population of aboriginal species of geese on the island.     

The energetics of barnacle geese (Branta leucopsis) flying in captive and wild conditions

Experimental data on the relationship between mean heart rate (f(H)) and mean rate of oxygen consumption (VO(2)) of captive barnacle geese during flights in a wind tunnel are assessed in terms of their capacity to predict the typical VO(2) of wild barnacle geese, based on the recordings of their f(H), while undertaking autumn migratory flights between Spitsbergen (78 degrees N) and Caerlaverock, Scotland (55 degrees N). A significant linear relationship has been demonstrated between the f(H) and simultaneously recorded VO(2) of a single barnacle goose (B-B) flying in the wind tunnel (VO(2)=1.42 f(H)-304, r(2)=0.82, P<0.001, N=12 flights). Data recorded from three additional geese (N=4 flights), lay within the 95% prediction intervals of the relationship for goose B-B. When these geese (mean body mass=2.06 kg, n=4) were flown in the wind tunnel (WT) without the mask, they had a mean f(H) of 451+/-23 beats min(-1), yielding an estimate for VO(2) of 336+/-33 ml min(-1). However, f(H) has also been recorded from wild barnacle geese (mean migratory f(H) of 253 beats min(-1)), and substitution of this value into the above calibration equation results in an unrealistically low value for mean migratory VO(2) of only 55 ml min(-1). Various factors, such as differences in heart mass, selective tissue perfusion, environmental temperature and flock formation, which might account for some of the difference in f(H) between the captive and wild geese are discussed. Comparison with other WT studies shows that inter-species minimum mass-specific VO(2) declines with increasing body mass (M(b); range 0.035-2.8 kg) as 173 M(b)(-0.224), r(2)=0.848.     

The effect of mate change and new partner's age on reproductive success in the barnacle goose, Branta leucopsis

Mate retention frequencies and correlations between mate changeand reproductive performance were estimated in a populationof barnacle geese, Branta leucopsis, breeding on the islandof Gotland in the Baltic. About 90% of the pairs remained togetherfrom one breeding season to the next. Only 2.4% of the pairsdivorced, most mate changes being consequences of the deathof one partner. Divorces were not forecast by low reproductivesuccess, and seemed to be accidental. In the season before matechange, there was no difference in reproductive performance,measured as clutch size, hatching date, and number of fledgedyoung, between faithful pairs and pairs where one partner wassubsequently changed. However, in the first season with a newpartner, clutch size and number of fledged young decreased onaverage. Hence, because mate change led to a reduction in reproductivesuccess, it was concluded that mate retention is advantageous.Our results suggest that this reduction is more likely due tothe lower average age or breeding experience of new partnersthan to the benefits of breeding experience with one particularpartner. [Behav Ecol 1991 ;2:116–122]     

Conservation of migrants on their wintering grounds: an overview

Most research on the behaviour and ecology of wintering migrants has been undertaken in the Neotropics. In Africa, conservation biologists are faced with a comparative paucity of baseline data to assess the significance of environmental change, in contrast to the comparatively well-developed monitoring programmes in place on the breeding grounds. A programme of monitoring, established through international collaboration and matched with further ecological research, is required. Given the 'dispersed' distribution of most passerine migrants, determination of the significance of apparent detrimental effects can be hard to ascertain. However, migrants may provide conservationists with an effective vehicle for the provision of input into region-wide development policy. It is only through influencing regional development policies within the continent that large-scale and long-term conservation of important bird habitats will be achieved.     

Vegetation growth and a seasonal habitat shift of the barnacle goose (Branta leucopsis)

Summary Barnacle geese (Branta leucopsis) wintering on the island of Schiermonnikoog in the Netherlands abruptly switch all their foraging activities from a dairy pasture (a polder) to an adjacent salt-marsh during the early spring. We present evidence to show that this shift is related to changes in the quality of the diet available in these different habitats. Barnacle geese shift from polder to salt-marsh at the precise time that these are equal in dietary protein availability, which occurs as the food plants on the salt-marsh undergo a sudden spring growth. The dairy pasture undergoes its own spring growth shortly afterwards, and more dietary protein is available there for the rest of the year. We suggest that the salt-marsh is a more preferred habitat, but that low dietary protein during the winter prevents its use by barnacle geese. We hypothesize that the salt-marsh may be more preferred due to a lower level of disturbance which permits geese to graze more slowly, improving the utilization of food plants.     

Phenology of migration and use of wintering sites by the increasing population of dark-bellied brent geese Branta bernicla bernicla

During the recovery of the population of dark-bellied brent geese Branta bernicla bernicla , traditional wintering sites in Britain have experienced increases in numbers and new sites have been colonized. Goose counts were used to describe the migration phenology and winter use of Scolt Head, north Norfolk (a traditional site) and the Burry Inlet, South Wales (a relatively new site) over the period of re-colonization and colonization, respectively. During the 1950s, only a few hundred birds wintered at Scolt Head, and most stayed for only a short period. By the late 1980s/early 1990s, numbers had reached several thousands, their period of stay was longer and there was little variation between years in median arrival (28 October) and departure (11 March) dates. A similar process was observed during the entire colonization phase at the Burry Inlet; initially birds arrived late and departed early, but arrived progressively earlier and departed later as wintering numbers increased. The autumn arrival rates at both sites were slower than the spring departure rates. During years of good breeding productivity, those adults with young arrived slightly later in Norfolk than failed breeders and non-breeders, and a higher proportion of young birds remained in spring after the main departure. However, the overall pattern of arrival and departure did not vary according to whether there had been a good or poor breeding season. Some adults of breeding age also remained until May. The effects of food depletion and use of novel feeding habitats in relation to migration phenology and winter use of sites are discussed.     

Heart rate and rate of oxygen consumption during flight of the barnacle goose, Branta leucopsis

We report the first measurements of heart rate (f(H)) and the rate of oxygen consumption (V(O(2))) during flights from a species of bird larger than 500 g. V(O(2))was obtained from nine forward flapping flights of 8.9 min mean duration at a mean speed of 13.2 m s(-1) performed by three barnacle geese of mean mass 1.68 kg. Mean V(O(2))was 332 ml min(-1)or 201 ml min(-1) kg(-1). Sixteen flights were obtained from two of these birds equipped with heart rate data loggers, both when they were wearing a V(O(2)) mask and when they were not. During flights with the mask (mean duration 7.4 min), mean f(H) was 472 beats per min and during flights without the mask (mean duration 8.0 min) it was 391 beats per min. Heart rate was also measured in another goose flying without a respiratory mask and mean f(H) for all the three birds (mean mass 1.7 kg) flying without a mask for an average of 7.9 min at 13 m s(-1) was 378 beats per min. Resting f(H) for these three birds was 79 beats per min. The values of f(H) during flight are greater than those obtained from the same species during their autumn migration from Spitsbergen to southern Scotland. The possible reasons for this are discussed.     

The diet of common eiders wintering in Nuuk,Southwest Greenland

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

The diet of a declining Red-billed Chough Pyrrhocorax pyrrhocorax population on Islay,Scotland

Capsule: We quantify Red-billed Chough Pyrrhocorax pyrrhocorax diet from faecal samples collected on Islay. Dung invertebrates formed the majority of prey biomass in dune pasture and Tipulid larvae in mixed pasture but Aphodius larvae were scarce in the diet, whereas they had been a major component in the 1980s. There are management implications from the indication of a reduction in availability of preferred food.     

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

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

Lack of competition between barnacle geese Branta leucopsis and pink-footed geese Anser brachyrhynchus during the pre-breeding period in Svalbard

The feeding ecology of barnacle geese and pink-footed geese was studied in Sassendalen, Svalbard during the pre-nesting period (late May) to assess the potential for inter-specific competition. Barnacle geese fed almost exclusively (97%) by grazing above-ground plant material, mostly (79%) along snow edges in moss-mat habitats. Pink-footed geese fed mostly (93%) by excavating below-ground parts of plants, mostly (56%) away from snow and were more evenly distributed between habitat types. Barnacle goose faeces contained mostly (62%) moss, that of pink-footed geese mostly (48%) below-ground plant storage organs (especially Bistorta viviparum L.). Principal components analysis of dropping contents showed no overlap in species diet in allopatry or sympatry. There was little overlap in diet and feeding ecology of the two species at this pre-nesting feeding site. Hence, unless increased goose feeding densities affect future vegetation density and composition, under present circumstances, increasing numbers of either species is unlikely to affect foraging conditions for the other at this important stage in the annual cycle. However, such changes could have local density-dependent intra-specific effects.     

Brent Geese Branta bernicla and Zostera; factors affecting the exploitation of a seasonally declining food resource

Brent Geese Branta bernicla wintering at Lindisfarne, northeastern England, fed almost exclusively on intertidal habitats. Their main food supply was two species of Eelgrass Zostera noltii and Zostera angustifolia. Although abundant when the birds arrived in September, this Zostera was rapidly depleted during the period October-December. Brent Goose food intake rate declined with the decreasing food supply, and the birds responded by extending the time that they spent feeding. When it was no longer possible to extend the time spent feeding (i.e. they were feeding for all of the time that the food supply was available to them), they moved away from the site. The geese fed extensively at night in order to achieve their daily feeding requirements, especially later in the season. Conversion of daily food intake to energetic intake suggested that there may have been an energetic trigger acting: the geese left the site when they were unable to satisfy their basic energy demand. No evidence was found for direct interference competition between Brent Geese and the other grazer in the system, Wigeon Anas penelope: the two species showed no spatial segregation in their feeding areas at the scale investigated nor any temporal avoidance of each other.     

Spring staging in Brent Geese Branta bernicla: feeding constraints and the impact of diet on the accumulation of body reserves

Summary The diet composition of Brent Geese Branta bernicla on a salt-marsh was quantified. Puccinellia maritima was the principal food species, while Plantago maritima and Triglochin maritima were less commonly taken. Festuca rubra only acted as a substitute for Puccinellia when production of the latter species dropped. The metabolizable energy of the food plants ranged from 5 to 11 kJ·g^–1. By assessing the ingestion rates of geese feeding on different food species, the net intake rate could be derived. Plantago and Triglochin appeared to be the most profitable plants to eat. The proportion of these species in the diet was restricted by (1) the capacity of the alimentary tract, since high intake rates combined with high water contents of the food plants easily led to overfill; and (2) the limited distribution of these plants, in combination with their rapid depletion by grazing geese. These latter factors led to an unequal allocation among individual geese. Most Plantago and Triglochin was obtained by dominant pairs within the flocks. The high quality of Puccinellia allowed geese to gain mass in spring, but the metabolizable energy of this plant species declined during the staging period, and Plantago and Triglochin increased in importance in supplying the geese with components with which to build their body reserves. The timing of the onset of spring growth of the various food species differed between years, and plant phenology was shown to have a profound effect on the final body reserves of the geese.