Survival of Svalbard pink-footed geese Anser brachyrhynchus in relation to winter climate, density and land-use

1. Global change may strongly affect population dynamics, but mechanisms remain elusive. Several Arctic goose species have increased considerably during the last decades. Climate, and land-use changes outside the breeding area have been invoked as causes but have not been tested. We analysed the relationships between conditions on wintering and migration staging areas, and survival in Svalbard pink-footed geese Anser brachyrhynchus. Using mark-recapture data from 14 winters (1989-2002) we estimated survival rates and tested for time trends, and effects of climate, goose density and land-use. 2. Resighting rates differed for males and females, were higher for birds recorded during the previous winter and changed smoothly over time. Survival rates did not differ between sexes, varied over time with a nonsignificant negative trend, and were higher for the first interval after marking (0.88-0.97) than afterwards (0.74-0.93). Average survival estimates were 0.967 (SE 0.026) for the first and 0.861 (SE 0.023) for all later survival intervals. 3. We combined 16 winter and spring climate covariates into two principal components axes. F1 was related to warm/wet winters and an early spring on the Norwegian staging areas and F2 to dry/cold winters. We expected that F1 would be positively related to survival and F2 negatively. F1 explained 23% of survival variation (F1,10=3.24; one-sided P=0.051) when alone in a model and 28% (F1,9=4.50; one-sided P=0.031) in a model that assumed a trend for survival. In contrast, neither F2 nor density, land-use, or scaring practices on important Norwegian spring staging areas had discernible effects on survival. 4. Climate change may thus affect goose population dynamics, with warmer winters and earlier springs enhancing survival and fecundity. A possible mechanism is increased food availability on Danish wintering and Norwegian staging areas. As geese are among the main herbivores in Arctic ecosystems, climate change, by increasing goose populations, may have important indirect effects on Arctic vegetation. Our study also highlights the importance of events outside the breeding area for the population dynamics of migrant species.     

Snow-patch foraging by pink-footed geese Anser brachyrhynchus in south Iceland

The pre-nesting feeding behaviour of pink-footed geese was studied in hayfields in southern Iceland during the late spring of 1989. Persistent snow-patches protected underlying grass from the effects of severe night-time frost. Areas within 1 m of snow-patches had significantly greater amounts of green material than those further away; green material contained more than double the protein of brown, dead material, which predominated in open fields. Geese spent nearly 60 times more time feeding within 1 m of snow patches than expected by chance, and 20 times more time within 2–5 m. Their feeding rates here were faster and their step rate slower than further away. In this way, the geese selected the prime forage as soon as it became available.     

Genetic consequences of breaking migratory traditions in barnacle geese Branta leucopsis

Cultural transmission of migratory traditions enables species to deal with their environment based on experiences from earlier generations. Also, it allows a more adequate and rapid response to rapidly changing environments. When individuals break with their migratory traditions, new population structures can emerge that may affect gene flow. Recently, the migratory traditions of the Barnacle Goose Branta leucopsis changed, and new populations differing in migratory distance emerged. Here, we investigate the population genetic structure of the Barnacle Goose to evaluate the consequences of altered migratory traditions. We used a set of 358 single nucleotide polymorphism (SNP) markers to genotype 418 individuals from breeding populations in Greenland, Spitsbergen, Russia, Sweden and the Netherlands, the latter two being newly emerged populations. We used discriminant analysis of principal components, F_ST, linkage disequilibrium and a comparison of geneflow models using migrate ‐n to show that there is significant population structure, but that relatively many pairs of SNPs are in linkage disequilibrium, suggesting recent admixture between these populations. Despite the assumed traditions of migration within populations, we also show that genetic exchange occurs between all populations. The newly established nonmigratory population in the Netherlands is characterized by high emigration into other populations, which suggests more exploratory behaviour, possibly as a result of shortened parental care. These results suggest that migratory traditions in populations are subject to change in geese and that such changes have population genetic consequences. We argue that the emergence of nonmigration probably resulted from developmental plasticity.     

Snow conditions as an estimator of the breeding output in high-Arctic pink-footed geese Anser brachyrhynchus

The Svalbard-breeding population of pink-footed geese Anser brachyrhynchus has increased during the last decades and is giving rise to agricultural conflicts along their migration route, as well as causing grazing impacts on tundra vegetation. An adaptive flyway management plan has been implemented, which will be based on predictive population models including environmental variables expected to affect goose population development, such as weather conditions on the breeding grounds. A local study in Svalbard showed that snow cover prior to egg laying is a crucial factor for the reproductive output of pink-footed geese, and MODIS satellite images provided a useful estimator of snow cover. In this study, we up-scaled the analysis to the population level by examining various measures of snow conditions and compared them with the overall breeding success of the population as indexed by the proportion of juveniles in the autumn population. As explanatory variables, we explored MODIS images, satellite-based radar measures of onset of snow melt, winter NAO index, and the May temperature sum and May thaw days. To test for the presence of density dependence, we included the number of adults in the population. For 2000–2011, MODIS-derived snow cover (available since 2000) was the strongest indicator of breeding conditions. For 1981–2011, winter NAO and May thaw days had equal weight. Interestingly, there appears to have been a phase shift from density-dependent to density-independent reproduction, which is consistent with a hypothesis of released breeding potential due to the recent advancement of spring in Svalbard.     

Colonization history of the high-arctic pink-footed goose Anser brachyrhynchus

Population structure and phylogeography of the pink-footed goose, Anser brachyrhynchus Baillon 1833, was studied using mtDNA control region sequences (221 bp) from 142 individuals. Present breeding areas of the species in Greenland, Iceland, and Svalbard were largely covered by ice during the late Pleistocene. In pairwise comparisons phiST estimates showed significant differentiation among eastern and western populations, whereas sampling localities within both areas were not differentiated. The mtDNA data indicate that the populations have separated recently (less than 10 000 years ago) and present breeding areas were colonized from one refugial population. The levels of haplotype and nucleotide diversity were approximately five times higher for the eastern population compared to the western population and suggest that the latter was colonized by a subset of eastern birds. Time to the most recent common ancestor of the species is 32 000-46 000 years, i.e. the present mtDNA variation of the pink-footed goose has accumulated during the last 0.1 My. Estimates of the long-term female effective population size (5400-7700 for the eastern population) imply that the refugial population of the pink-footed goose has been large. Tundra habitats were more extensive in cold periods of the late Pleistocene than today and may have sustained population sizes that allowed the accumulation of extant genetic polymorphism. It is not probable that the postulated small refugial areas in the high latitudes had a significant role in maintaining this diversity.     

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.     

Assessing body condition and energy budget components by scoring abdominal profiles in free-ranging pink-footed geese Anser brachyrhynchus

An abdominal profile index (API) was developed for pink-footed geese Anser brachyrhynchus as a measure of body condition. On basis of carcass analysis of 56 adult geese with known API prior to collection, we found significant linear relationships between API against body mass, abdominal fat and total energy content. Hence, changes in API reflect net energy intake rates. As an example of the applicability of the calibration, we compared APIs of individually marked geese before and after long migration episodes and estimated the cost of flight at 8.9 kJ/km. In addition we estimated gain rates at three major staging sites along the spring flyway indicating an increase in fueling rates with latitude. Calibration of APIs and energy contents offers new opportunities for field studies of waterfowl energetics.     

Minimal intra-seasonal dietary overlap of barnacle and pink-footed geese on their breeding grounds in Svalbard

We analysed barnacle Branta leucopsis and pink-footed goose Anser brachyrhynchus summer diets (May–July 2003) based on the proportions of different plant constituents in the faecal material of adult breeding birds in Sassendalen, Svalbard to assess potential inter-specific competition. Diets were highly restricted and overlapped little during pre-nesting and post hatch. During incubation both species showed greatest variety in their diet, reflecting site-specific differences in local food abundance. However, locally the diets of pink-footed and barnacle geese resembled each other most at this time (although still differing significantly). The conflicting needs of nest defence and maintenance of body condition constrains the extent of the feeding resource utilised by nesting pairs and explains slightly greater dietary overlap at this time. Hence, there is little evidence of inter-specific competition (interference or depletion) at present, but this is most likely to be manifest during the incubation period in the future if goose numbers continue to increase. More detailed investigations of the degree of spatial overlap of the two species and their effects on plant structure, quality and community composition are necessary to predict likely outcomes of expected increases in numbers of both goose species.     

The reliance on distant resources for egg formation in high Arctic breeding barnacle geese Branta leucopsis

Breeding in the high Arctic is time constrained and animals should therefore start with their annual reproduction as early as possible. To allow for such early reproduction in migratory birds, females arrive at the breeding grounds either with body stores or they try to rapidly develop their eggs after arrival using local resources. Svalbard breeding barnacle geese Branta leucopsis have to fly non‐stop for about 1100 km from their last continental staging site to the archipelago making the transport of body stores costly. However, environmental conditions at the breeding grounds are highly unpredictable favouring residual body stores allowing for egg production after arrival on the breeding grounds. We estimated the reliance on southern continental resources, i.e. body stores for egg formation, in barnacle geese using stable isotope ratios in the geese's forage along the flyway and in their eggs. Females adopted mixed breeding strategies by using southern resources as well as local resources to varying extents for egg formation. Southern capital in lipid‐free yolk averaged 41% (range: 23–65%), early laid eggs containing more southern capital than eggs laid late in the season. Yolk lipids and albumen did not vary over time and averaged a southern capital proportion of 54% (range: 32–73%) and 47% (range: 25–88%), respectively. Our findings indicate that female geese vary the use of southern resources when synthesising their eggs and this allocation also varies among egg tissues. Their mixed and flexible use of distant and local resources potentially allows for adaptive adjustments to environmental conditions encountered at the archipelago just before breeding.     

Incidence of embedded shotgun pellets and inferred hunting kill amongst Russian/Baltic barnacle geese Branta leucopsis

X-rays of wild caught barnacle geese from the Russian/Baltic population were taken in Denmark in the springs of 2009 and 2011 to determine the incidence of embedded shotgun pellets and to estimate the annual hunting kill. On average, 13 % of adult geese (n?=?212) and 6 % of first year geese (n?=?35) examined contained pellets in their tissue. Assuming that these birds represented a random sample, extrapolation to the entire population of c. 770,000 individuals indicates that 96,000 barnacle geese carry embedded pellets. Based on the assumption that the ratio between the number of birds with embedded shot and the total number of birds harvested per season is similar to that found in pink-footed geese Anser brachyrhynchus (recorded as 3.6:1 during 1990–1996 and 1.7:1 during 2009–2011), the annual kill of barnacle geese was estimated at 26,300–58,300 birds. The Russian/Baltic barnacle geese are protected from hunting on their winter quarters (The Netherlands, Germany and Denmark), but are quarry in Russia and are shot under license to protect agricultural crops in Estonia, Germany, Sweden and Denmark (amounting to a total of c. 1,800–3,000 geese shots per year). Barnacle geese are known to be shot illegally (accidentally and deliberately) along the migration route, but in the absence of Russian bag statistics, the contribution of illegal hunting to the overall harvest cannot be substantiated. Although the population is currently increasing at an annual rate of 8 %, the indirectly estimated hunting pressure (3–7 %) is not insubstantial and should be taken into account in future population management models.     

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.     

Variation in daily and seasonal foraging routines of non-breeding barnacle geese (Branta leucopsis): working harder does not overcome environmental constraints

This study describes the changes in stored body fat in Svalbard barnacle geese Branta leucopsis over the non-breeding period, and uses behavioural patterns to explain the variation in body fat stores. It was predicted that foraging effort would: (1) increase at low food densities; (2) increase when days were short; (3) decrease in smaller flocks; (4) be bimodally distributed throughout the day for long days, but constant for short days. Time constraints were found to be the major driving force behind foraging decisions during the shortest days of mid-winter, whereas food density was found to drive decisions during longer days. Field observations of fat stores showed that fat was rapidly accumulated at the start and end of the non-breeding period, but that fat stores remained constant during mid-winter. It is suggested that day length prevented a positive fat storage rate in mid-winter through the limitation of foraging time, even though foraging effort was high during this period. During a single day, evidence for a bimodal foraging routine was found, where feeding activity was concentrated in the early morning and late afternoon periods. This pattern was found in the full range of day lengths, which suggests that even for short days, feeding must be interrupted so that other essential activities can be conducted. It is concluded that the behavioural choices of barnacle geese were constrained by environmental conditions, and that these behavioural patterns allow the variation in fat stores to be explained.     

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.     

The onset of spring and timing of migration in two arctic nesting goose populations: the pink‐footed goose Anser bachyrhynchus and the barnacle goose Branta leucopsis

An earlier onset of spring has been recorded for many parts of Eurasia in recent decades. This has consequences for migratory species, both in changing the conditions encountered by individuals on reaching migratory sites and in affecting cues regulating the timing of migration where decisions to migrate are influenced by local environmental variables. Here we examine the timing of spring migration for two arctic goose populations, the pink‐footed goose Anser brachyrhynchus (during 1990–2003) and barnacle goose Branta leucopsis (during 1982–2003), which both breed on Svalbard. The satellite‐derived Normalised Difference Vegetation Index (NDVI) was used to express the onset of spring at their wintering and spring staging sites. Pink‐footed geese use several sites during spring migration, ranging from the southernmost wintering areas in Belgium to two spring staging areas in Norway, and distances between sites used along the flyway are relatively short. There was a positive correlation in the onset of spring between neighbouring sites, and the geese migrated earlier in early springs. Barnacle geese, on the other hand, have a long overseas crossing from their wintering grounds in Britain to spring staging areas in Norway. Although spring advanced in both regions, there was no corresponding correlation in the timing of onset of spring between their wintering and spring staging sites, and little evidence for barnacle geese migrating earlier over the whole study period. Hence, where geese can use spring conditions at one site as an indicator of the conditions they might encounter at the next, they have responded quickly to the advancement of spring, whereas in a situation where they cannot predict, they have not yet responded, despite the advancement of spring in the spring staging area.     

Finding food in a highly seasonal landscape: where and how pink footed geese Anser brachyrhynchus forage during the Arctic spring

Food accessibility and availability in the highly seasonal Arctic landscape can be restricted by snow cover and frozen substrate, particularly in early spring. Therefore, to determine how a long distance migratory herbivore forages in such a landscape, pink‐footed geese Anser brachyrhynchus at an early spring feeding area in Svalbard were studied. Birds arrived in mid May when extensive snow cover restricted habitat availability. Geese fed in all habitats, but the highest densities occurred in wet tundra. However, prolonged snow lie restricted access to wet areas compared to dry and mesic habitats. Above ground biomass was very low in all habitats; yet sizeable amounts occurred below ground. In line with this, the majority of birds (86%) grubbed for below ground plant storage organs such as stem bases and rhizomes. Wet habitat contained greater quantities of edible and lower amounts of inedible below ground material (ratio 1:0.3) than dry (ratio 1:9) or mesic (ratio 1:4) areas. Although foraging in wet habitat prevented geese from encountering high proportions of inedible plant parts, forage species characteristic of this habitat, such as Dupontia grasses and the rush Eriophorum scheuchzeri, were more difficult to extract than food plants typical of drier habitats such as the forb Bistorta vivipara. Hence, we suggest that wet areas are preferred by pink‐footed geese, but the prolonged snow lie there made it necessary to use less preferred but much more abundant drier habitats, which experienced earlier snowmelt and indeed accommodated more than half of all goose foraging recordings.     

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]     

Modelling pink-footed goose (Anser brachyrhynchus) wintering distributions for the year 2050: potential effects of land-use change in Europe

Feeding on farmland by overwintering populations of pink-footed geese ( Anser brachyrhynchus ) conflicts with agricultural interests in Northern Europe. In order to forecast the potential future of this conflict, we used generalized linear models to relate the presence and absence of pink-footed geese to variables describing the contemporary landscape, and predicted their future distributions in relation to two land-use scenarios for the year 2050. One future scenario represented a global, economically orientated world (A1) and the other represented a regional, environmentally concerned world (B2). The probability of goose occurrence increased within cropland and grassland, and could be explained by their proximity to coast, elevation, and the degree of habitat closure. Predictions to future scenarios revealed noticeable shifts in the suitability of goose habitat evident at the local and regional scale in response to future shifts in land use. In particular, as grasslands and croplands give way to unsuitable land-use types (e.g. woody biofuel crops, increased urbanization, and forest) under both future scenarios, our models predicted a decrease in habitat suitability for geese. If coupled with continued goose population expansion, we expect that the agricultural conflict will intensify under the A1 and particularly the B2 scenarios.     

Causes of the redistribution of Pink-footed Geese Anser brachyrhynchus in Britain

The numbers of Pink-footed Geese Anser brachyrhynchus wintering in Britain have been increasing exponentially since the 1950s. The numbers of geese at virtually all roost sites have increased, but the most dramatic increase has been in Norfolk, the southernmost region occupied in Britain. This redistribution was most likely caused either by the increase in the population forcing greater numbers of geese towards more southerly sites as the result of a buffer effect, by a decrease in the quality of sites elsewhere in the range or by an increase in the attractiveness of Norfolk. Co-ordinated counts of geese around Britain and historical data of changes in agricultural practice in Norfolk were used to examine these possibilities. No evidence was found for buffer effects, degradation of sites elsewhere or an increase in food availability in Norfolk. The major difference between Norfolk and other Pink-footed Goose sites is in the food type available in Norfolk, namely harvested Sugar Beet Beta vulgaris remains. This food is of high quality and the geese are not disturbed whilst feeding on it because of its limited commercial value. The pattern of redistribution of Pink-footed Geese may therefore be the result of cultural learning of the benefits of feeding on Sugar Beet.