A multifactorial explanation for variation in breeding performance of Brent Geese Branta bernicla

The dramatic differences in annual breeding success of Dark-bellied Brent Geese Branta b. bernicla are generally interpreted as being caused by variable conditions on the arctic breeding grounds. The recently revived discussion on this issue mentions, in particular, the cyclic variation in abundance of lemmings, resulting in cyclic variation of predation pressure on bird nests by arctic foxes, as the main factor determining whether Brent Geese are able to breed successfully or not. New data on the spring condition of Brent Geese in Western Europe and wind conditions during migration from the spring staging areas to the breeding grounds show that these factors also influence breeding output.     

Shared or Unshared Parental Care in Overwintering Brent Geese (Branta bernicla hrota)

We examine brood size effects on the behaviour of wintering parent and juvenile brent geese (Branta bernicla hrota) to test predictions of shared and unshared parental care models. The behaviour of both parents and offspring appear to be influenced by declining food availability over the winter. Parental vigilance increased with brood size and may be explained by vigilance having functions in addition to antipredator behaviour where the benefits are shared among the brood. There was no increase in parental aggression with brood size and this does not fit the prediction of shared care. Nevertheless, large families are able to monopolize better feeding areas compared with smaller families and large families static feed more but walk feed less than do small families, the former apparently being the preferred mode. The presence of additional young, rather than increasing the amount of parental aggression, seems to enhance the family's competitive ability. Because parents with large broods benefit from enhanced access to resources there is likely to be no additional significant cost in the parental care of larger broods (sensu Trivers 1972 ).     

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.     

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.     

Breeding of waders (Charadrii) and Brent Geese Branta bernicla bernicla at Pronchishcheva Lake, northeastern Taimyr, Russia, in a peak and a decreasing lemming year

During summer 1991, lemmings occurred at high densities in Arctic tundra at Pronchishcheva Lake in the northeastern Taimyr Peninsula, whereas, in 1992, lemming densities were substantially lower and decreased further during the summer. In 1991, avian predators such as Snowy Owls Nyctea scandiaca, gulls and skuas bred well; Arctic foxes Alopex lagopus were rarely observed in the study area but bred in the immediate vicinity. In both years there was a late thaw, but this did not deter breeding by birds. The insect food supply for waders showed similar patterns of abundance in both years. In 1991, 73 nests of nine species of wader were found within a 14-km² study area, and Dark-bellied Brent Geese Branta bernicla bernicla nested in association with Snowy Owls. The overall density of wader nests was estimated to be 7 per km². Clutches disappeared at only two wader nests and no Brent Goose nests, and the Mayfield estimate of the daily probability of predation for waders was 0.0022. In contrast, the daily probability of predation was 0.20 in 1992, when there was a similar breeding density of waders. Arctic foxes were seen searching for food daily within the study area, and fox droppings were found associated with nests taken by predators. The predicted scenarios for peak and decreasing lemming years (the Roselaar-Summers hypothesis), i.e. low predation and high nest success in the peak year and high predation and low nest success in the decreasing year, therefore occurred.     

Dark-bellied brent geese Branta b. bernicla breeding near snowy owl Nyctea scandiaca nests lay more and larger eggs

Several studies have demonstrated that snowy owls Nyctea scandiaca defend an area around their nests against predators, hereby inadvertently creating safe havens for breeding dark-bellied brent geese Branta b. bernicla . However, studies investigating brent goose breeding ecology within the predator-exclusion zones of the snowy owls are absent. In 1999 and 2005, years of high lemming abundance Lemmus sibiricus and Dicrostonyx torquatus , brent geese were primarily breeding in association with snowy owls in the Medusa river catchment on western Taimyr, Russia. Goose nest failure, either as a result of nest abandonment by the adult birds or of nest depredation, increased with increasing distance from the owl nests. Within the brent goose colonies, clutch size as well as egg size increased with decreasing distance from the snowy owl nest, indicating an increasing adult quality closer to owl nests. However, as a result of the abandonment of eggs and goslings, the increasing clutch size did not result in a higher nest success during this study. Apparently brent geese compete for breeding sites close to owl nests, but details of this process remain unknown.     

Preservation of winter social dominance status in Brent Geese Branta bernicla bernicla within and across winters

Dominant and subordinate individuals in a group may benefit from the stability of the social dominance organisation, avoiding excessive waste of time and energy in aggressive interactions and reducing injury risks. Nevertheless, the likely evolutionary incentive for individuals to become, and furthermore to stay, dominant may destabilise such dominance hierarchies. In this context, the relative importance of fixed (e.g. sex, morphological size) and fluctuating (e.g. body condition, mating status, reproductive success, social unit size) traits influencing the establishment and preservation of dominance relationships could play a key role in group structure. We investigated the relative role of fixed and fluctuating traits on social status in Dark-bellied Brent Geese Branta bernicla bernicla which form large fairly unstable groups both within and across winters. We compared individual dominance scores of ringed Brent Geese during four consecutive winters. Brent Geese conserved their dominance score within a given winter irrespective of their age but were generally unable to conserve it across consecutive winters. As winter dominance scores correlated best with social unit size, dominance status thus appeared to be mostly a by-product of a fluctuating trait: breeding success in the previous summer. When we considered only adults that had the same social unit size during two consecutive winters, we observed a significant preservation of dominance scores. This result suggests that a fixed trait such as sex or morphological size may still play a role in setting dominance status.     

Utilization, diet and diet selection by brent geese Branta bernicla bernicla on salt-marshes in Norfolk

The diet and utilization by brent geese of two plant communities ( Limonium/Armeria marsh—a short sward community in the upper mid salt-marsh and composed of several species, and Salicornia marsh—a community of the lower salt-marsh dominated by Salicornia europuea agg. and Aster tripolium ) were studied on the north Norfolk coast. The biomass on these communities was highest in autumn and declined markedly in November owing to the senescence and die-off of the succulent species. This decline was not significantly precipitated by the grazing of the geese but coincided with their switch to inland habitats where the biomass m^-2 and total food available on fields of winter wheat and grassland within the population's home range was 20–30 times greater. Brent geese continued to graze salt-marsh at a low intensity through the winter, feeding largely on Puccinellia maritima. In spring they returned largely to Limonium/Armeria marsh where they significantly reduced the regrowth of Puccinellia maritima.
The main plant species ealen was Puccinellia maritima , especially in mid winter. Salicornia maritima agg. leaves and seeds formed much of the diet in autumn, whereas Triglochin maritima, Plantago maritima and Aster tripolium were important foods on Limonium/Armeria marsh in spring. All these species were selected at some part of the year. Selection was related to the proline (an imino acid used by salt-marsh plants as an osmoregulator) and chloride content and avoidance related to tannins. Two species which contained condensed tannins. Limonium vulgare and Armeria maritima , were rarely eaten. As well as being a nutrient, proline may help to neutralize the effects of tannins in the diet if incorporated into proline-rich proteins.     

DHEA levels and social dominance relationships in wintering brent geese (Branta bernicla bernicla)

After testosterone, dehydroepiandrosterone (DHEA) is the main hormone involved in aggressive behaviour in birds. While the role of DHEA has been verified for wintering territorial passerines, it has not been shown for gregarious species. In wintering geese species, both sexes present very low testosterone levels and aggression in a non-sexual context is not testosterone-related. Therefore, testosterone does not seem to be responsible for aggressive behaviour by geese during winter and the role of DHEA must be explored. We used brent geese (Branta bernicla bernicla) to examine the roles of testosterone and DHEA in dominance relationships. For the first time, we highlighted the presence of plasma DHEA in free-living geese. As the level of DHEA was lower than that of testosterone, and there was no obvious impact of DHEA level on dominance status, our results failed to confirm the role of plasma DHEA in the social hierarchies of this species during winter. Nevertheless, because DHEA levels were greater in singletons than in paired birds, we discuss the need to explore hormonal and/or behavioural mechanisms implicated within dominance status acquisition and maintenance within each reproductive status class, to underline the role of the presence of relatives as a signal of dominance abilities. We also acknowledge and discuss the possibility that the long handling time may have affected DHEA levels and masked subtle differences between individuals.     

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.     

Dark-bellied Brent geese aggregate to cope with increased levels of primary production

We report on an aggregative response of Dark-bellied Brent geese to increased productivity of the vegetation during the growing season on agricultural fields on the island of Schiermonnikoog, the Netherlands. Plant standing crop was found to be maintained at low levels in the fields where geese activity focussed, whereas the remainder of the fields escaped herbivore control and developed a high standing crop. This pattern can be explained by a decreased efficiency of grazing in vegetation with a high standing crop. In other words, the functional response of the geese is not monotonically increasing but dome-shaped. As a consequence, continuously grazed swards are more suitable for feeding than temporarily ungrazed swards. We present a model showing that, for a dome-shaped functional response, optimal foraging under increasing primary productivity leads to spatial heterogeneity in standing crop. Beyond a certain threshold value, a further increase in productivity leads to a progressive release of vegetation from herbivore control and to the development of a high standing crop. Interestingly, our model suggests that only in a stable and predictable environment the aggregative behaviour of herbivores is able to maintain the intake rate close to its potential maximum. Misjudgement of patch quality by the herbivore or any other process disrupting the match between local primary production and consumption leads to a less than optimal intake, as suitable vegetation becomes depleted. This has important implications for ecological inferences, such as the prediction of carrying capacities in herbivore-dominated ecosystems.     

An outbreak of schistosomiasis in Atlantic brant geese, Branta bernicla hrota

A heavy infection with schistosomes of the genera Trichobilharzia and Dendritobilharzia was considered the cause of 90% mortality in a group of 40 wild-caught Atlantic brant geese (Branta bernicla hrota) that were maintained in captivity on a fresh-water pond in Aurora, Ontario. Numerous adult worms and eggs were disseminated in many organs throughout the body of all birds examined. The main pathological findings, attributed to both eggs and adults, included emaciation, thrombosis of the caudal mesenteric vein and its branches, fibrinohemorrhagic colitis, and in some birds, heptomegaly. Translocation of brant geese from their natural marine environment to a fresh-water pond may have caused them to be exposed to parasites which they would not normally encounter.     

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.     

Factors affecting nest defence in female Canada Geese Branta canadensis

The effect of various characteristics of Canada Goose Branta canadensis females and clutch size on their nest defence intensity during incubation was studied in the field during five nesting seasons. Nests were approached by a human, and the leaving distance was measured. Risk-taking in post-encounter behaviour was categorized at three levels. Nesting success was significantly associated with nest tenacity, a variable derived from leaving distance, but not with post-encounter behaviour. Nest tenacity, as well as post-encounter behaviour, differed significantly between individuals. Intensity of post-encounter behaviour, but not nest tenacity, increased with increasing age of the female. Nest tenacity, standardized over the incubation period, increased with increasing clutch size, but post-encounter behaviour intensity did not. None of the nest defence variables was correlated with the structural size of the female. Nest tenacity, but not post-encounter behaviour intensity, increased during incubation. Nest tenacity did not differ between early and late breeders, but post-encounter behaviour was more intense in late breeders. There was no evidence of positive reinforcement of nest defence through repeated visits. Nest defence did not differ between lakes, lake types or nest site types, nor was it affected by the distance to the nearest neighbouring nest. The results supported several of the predictions extracted from parental investment theory.     

Changes in body mass and hormone levels between wintering and spring staging areas in dark-bellied brent geese Branta bernicla bernicla

We document seasonal changes in body mass and plasma hormone levels (testosterone and triiodothyronine) between winter and spring quarters in dark-bellied brent geese Branta bernicla bernicla. Body mass increased between winter and spring, whereas body size was the same for the birds sampled in winter and those in spring. Adults and males were larger and heavier than juveniles and females. Plasma testosterone levels did not differ significantly between winter and spring, and did not vary with sex, age and morphometry. Plasma triiodothyronine (T₃) levels were significantly higher in spring than in winter, and were higher for males than for females. However, T₃ levels did not vary significantly with age and body measurements. To explain the findings, we discuss potential impacts of social dominance.