Nocturnal feeding by Dark-bellied Brent Geese Branta bernicla bernicla

The occurrence and extent of nighttime feeding by Dark-bellied Brent Geese Branta bernicla bernicla on salt marshes adjacent to their tidal roost site were assessed on the north Norfolk coast using position-sensitive radio transmitters. Nine birds were monitored on 143 bird-nights, and feeding was recorded on 87. The frequency of feeding by night varied widely between individuals. The mean proportion of the night spent feeding for one bird was 19.7 ± 8%. Of this, 59% occurred within ± 1.5 h of high tide. Neither the proportion of the whole night spent feeding nor the intensity of feeding around high tide varied with the length of the night or with time after sunset at which high tide occurred. The extent of nocturnal feeding was also independent of the brightness and duration of moonlight. The birds fed for significantly longer on colder nights, and there was a significant positive relationship between the proportion of the night spent feeding and maximum temperature the preceding day. Analysis of daytime activity budgets on inland pastures showed that the proportion of time engaged in nonfeeding, energy-expensive activities was also positively related to maximum daytime temperature. Increases in the proportion of the night spent feeding after warmer days were achieved by increasing the period of time spent feeding on either side of the high tide, not by increasing feeding intensity at high tide. It is concluded that the geese compensate for increased energy expenditure during the day by increasing the extent to which they feed at night. Implications of increasing energy expenditure as a result of disturbance during the day are discussed.     

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.     

Dark-bellied Brent Geese Branta bernicla bernicla, as recorded by satellite telemetry, do not minimize flight distance during spring migration

Nine Dark-bellied Brent Geese Branta bernicla bernicla were equipped with satellite transmitters during spring staging in the Dutch Wadden Sea in 1998 and 1999. The transmitters (in all cases less than 3% of body mass) were attached to the back by a flexible elastic harness. One juvenile female was tracked to the Yamal peninsula in 1998. Eight adult males were selected from a single catch of 75 to span the range of body mass observed on the date of capture (11 May 1999) and all but the lightest individual completed the first lap of the migratory flight to the White Sea, Russia, according to the time schedule normal for the species. Six birds were successfully tracked to Taymyr for a total distance averaging 5004 km (range 45775164) but judging from later movements none bred (although 1999 was a breeding year). Although the routes chosen during spring migration were closely similar, none of the tagged birds migrated together. On average the geese used 16 flights to reach their summer destinations on Taymyr. The longest uninterrupted flights during the first half of the journey (Wadden Sea to Kanin) covered 1056 km (mean of seven adult males, range 7681331), while the corresponding value for the second half of the migration (KaninTaymyr) was only 555 km (mean of six adult males). Only 7% of total time during spring migration was spent in active flight, as contrasted to c.  80% at long-term stopovers. Overall average travelling speed was 118 km/day (range 97148). Including fattening prior to departure the rate of travel falls to 62 km/day (range 4970), in keeping with theoretical predictions. Routes followed deviated from the great circle route, adding at least 700 km (16%) to the journey from Wadden Sea to Taymyr, and we conclude that the coastal route is chosen to facilitate feeding, drinking and resting en route instead of minimizing total flight distance.     

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 ).     

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.     

Incubation capacity contributes to constraints on maximal clutch size in Brent Geese Branta bernicla nigricans

Lack ( 1967 ) proposed that clutch size in species with precocial young was determined by nutrients available to females at the time of egg formation; since then others have suggested that regulation of clutch size in these species may be more complex. We tested whether incubation limitation contributes to ultimate constraints on maximal clutch size in Black Brent Geese (Black Brant) Branta bernicla nigricans. Specifically, we investigated the relationship between clutch size and duration of the nesting period (i.e. days between nest initiation and the first pipped egg) and the number of goslings leaving the nest. We used experimental clutch manipulations to assess these questions because they allowed us to create clutches that were larger than the typical maximum of five eggs in this species. We found that the per‐capita probability of egg success (i.e. the probability an egg hatched and the gosling left the nest) declined from 0.81 for two‐egg clutches to 0.50 for seven‐egg clutches. As a result of declining egg success, clutches containing more than five eggs produced, at best, only marginally more offspring. Manipulating clutch size at the beginning of incubation had no effect on the duration of the nesting period, but the nesting period increased with the number of eggs a female laid naturally prior to manipulation, from 25.4 days (95% CI 25.1–25.7) for three‐egg clutches to 27.7 days (95% CI 27.3–28.1) for six‐egg clutches. This delay in hatching may result in reduced gosling growth rates due to declining forage quality during the brood rearing period. Our results suggest that the strong right truncation of Brent clutches, which results in few clutches greater than five, is partially explained by the declining incubation capacity of females as clutch size increases and a delay in hatching with each additional egg laid. As a result, females laying clutches with more than five eggs would typically gain little fitness benefit above that associated with a five‐egg clutch.     

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.     

Energetic consequences of a major change in habitat use: endangered Brent Geese Branta bernicla hrota losing their main food resource

Coastal seagrasses are declining at increasing rates worldwide, forcing herbivores previously reliant on these habitats to abandon them in search of alternative ways to fulfil their daily energy budgets. After two decades of declining seagrass abundance in Mariager Fjord, Denmark, the Svalbard breeding population of Light‐bellied Brent Geese Branta bernicla hrota has experienced substantial changes in habitat use at this traditional autumn staging area. Declines in seagrasses have caused birds to depend increasingly on Sea Lettuce Ulva lactuca in recent years, and forced birds into terrestrial habitats such as saltmarsh and winter wheat. In contrast to those birds exploiting aquatic habitats, birds relying on these new habitats showed higher energy expenditure and failed to balance their energy budget. Eelgrass (Zostera) was energetically superior to other food resources, with marine Ulva being second best. Predicted body mass development under two different scenarios indicate that present habitat use resulted in a midwinter body mass around 122 g lower than just 20 years ago, equivalent to c. 9.4% of Brent Goose body weight. Even after controlling for inter‐annual differences in thermoregulatory costs, the effect of changes in habitat use translated into a body mass reduction of c. 56 g, which could adversely affect survival and future reproduction. Flyway‐wide declines in Zostera abundance and further reductions in traditional habitats due to climate change give cause to reassess projected population trends and consequent management implications for the East Atlantic flyway population of Light‐bellied Brent Geese.     

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.     

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.     

Pre‐moult patterns of habitat use and moult site selection by Brent Geese Branta bernicla nigricans: individuals prospect for moult sites

In environments where habitat quality varies, the mechanism by which individuals assess and select habitats has significant consequences on their spatial distribution and ability to respond to environmental change. Each year, thousands of Black Brent Geese Branta bernicla nigricans migrate to the Teshekpuk Lake Special Area (TLSA), Alaska, to undergo a flightless wing‐moult. Over the last three decades, moulting Brent Geese have changed their distribution within the TLSA, redistributing from inland, freshwater wetlands towards coastal, brackish wetlands. To understand better the mechanism by which Brent Geese select a moult site, as well as reasons behind the long‐term shift of moulting distributions, we examined movements and habitat use of birds marked with GPS‐transmitters during the pre‐moult period. Brent Geese did not generally migrate directly to their moulting site during the pre‐moult period, defined as the time from arrival at the moulting grounds to the onset of flightlessness. Rather, individuals used an average of 3.7 ± 0.6 (se) wetland complexes and travelled a minimum of 95.14 ± 15.84 km during the pre‐moult period. Moreover, 69% of Brent Geese visited their final moult site only to leave and visit other sites before returning for the flightless moult. Brent Geese spent significant time in both inland freshwater and coastal estuarine habitats during the pre‐moult, irrespective of the habitat in which they ultimately moulted. Whereas previous research suggested that Brent Geese choose moult sites based largely upon the experience of previous years, our observations suggest a mechanism of moult site selection whereby Brent Geese ‘prospect’ for moult sites, visiting multiple potential moult sites across varied habitat types, presumably gathering information from each site and correspondingly using this information to choose an appropriate moult site. By allowing individuals to adjust their distributions in response to habitat quality cues that may change annually, such as forage type and availability, prospecting may have influenced the long‐term shift in moulting distributions of Brent Geese in the TLSA.     

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.     

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.     

Habitat switching by dark-bellied brent geese Branta b. bernicla (L.) in relation to food depletion

Seasonal changes in the distribution and feeding behaviour of dark-bellied brent geese Branta b. bernicla (L.) and the biomass of their food plants were studied in three successive winters on the Norfolk coast. The data was used, in conjunction with published information, to show how depletion, productivity and mortality of food plants drive the pattern of habitat switching in this species. It is then possible to explain the habitat shifts observed over the last 35 years and predict future changes. On arrival, geese fed first on algal beds and then on salt marsh, grass and arable fields before returning to feed entirely on the salt marsh in spring. The biomass of green algae, and subsequently the salt marsh vegetation, declined during the autumn and this could be attributed to depletion through goose grazing and natural mortality. As depletion occurred the geese fed more intensively, for a greater percentage of time and with an increasing pace rate, the net result, however, was a declining intake rate (as measured by defaecation rate). The algal biomass at which the geese switched from the algal beds to salt marsh was consistent between years, with heavy storm-induced loss of algae in one year resulting in an earlier switch. That the timing of habitat switches may be explained by depletion of food plants was further supported by historical data: the number of brent geese wintering at the site has increased dramatically over the last 30–35 years and the time of switching from algal beds to salt marsh and from salt marsh to salt marsh and fields has become progressively earlier, as expected from the increased depletion. The expected further increase in brent goose numbers will increase the rate of depletion of intertidal vegetation so that the switches between habitats will be more rapid and the geese will move inland earlier and remain inland longer. The expected increase in the brent goose population will thus result in a disproportionate increase in the levels of conflict between brent geese and agriculture.     

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.     

Fidelity and breeding probability related to population density and individual quality in black brent geese Branta bernicla nigricans

1. Patterns of temporary emigration (associated with non-breeding) are important components of variation in individual quality. Permanent emigration from the natal area has important implications for both individual fitness and local population dynamics. 2. We estimated both permanent and temporary emigration of black brent geese (Branta bernicla nigricans Lawrence) from the Tutakoke River colony, using observations of marked brent geese on breeding and wintering areas, and recoveries of ringed individuals by hunters. We used the likelihood developed by Lindberg, Kendall, Hines & Anderson 2001 (Combining band recovery data and Pollock's robust design to model temporary and permanent emigration. Biometrics, 57, 273-281) to assess hypotheses and estimate parameters. 3. Temporary emigration (the converse of breeding) varied among age classes up to age 5, and differed between individuals that bred in the previous years vs. those that did not. Consistent with the hypothesis of variation in individual quality, individuals with a higher probability of breeding in one year also had a higher probability of breeding the next year. 4. Natal fidelity of females ranged from 0.70 +/- 0.07-0.96 +/- 0.18 and averaged 0.83. In contrast to Lindberg et al. (1998), we did not detect a relationship between fidelity and local population density. Natal fidelity was negatively correlated with first-year survival, suggesting that competition among individuals of the same age for breeding territories influenced dispersal. Once females nested at the Tutakoke River, colony breeding fidelity was 1.0. 5. Our analyses show substantial variation in individual quality associated with fitness, which other analyses suggest is strongly influenced by early environment. Our analyses also suggest substantial interchange among breeding colonies of brent geese, as first shown by Lindberg et al. (1998).     

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.     

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.