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.     

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.     

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.     

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.     

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.     

Bryophyte DNA sequences from faeces of an arctic herbivore, barnacle goose (Branta leucopsis)

We tested DNA extraction methods and PCR conditions for the amplification of bryophyte DNA from barnacle goose (Branta leucopsis) faeces collected from Spitsbergen (Svalbard). Both the Qiagen stool kit and a silica-based extraction method received sufficient DNA from fresh and older droppings, as indicated by successful amplification of the plastid psbA-trnH spacer. Standard Taq polymerase outperformed two hot start polymerases. Sequencing of cloned PCR products revealed at least ten moss and two angiosperm sequences. This first example of identifying bryophyte DNA from faeces will allow analysing moss diets of arctic herbivores with a DNA barcoding approach.     

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 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]     

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.     

吉林向海发现白颊黑雁

正2018年5月3日傍晚在位于吉林省白城市通榆县的吉林向海国家级自然保护区鹤类核心区青年坝附近(44°59′51.5″N,122°17′52.1″E,海拔168 m)使用NIKON-D4相机800 mm定焦镜头拍摄到1只黑白相间雁类,拍摄距离约120 m。经查阅相关文献(段文科等2017)确定本次发现的鸟类为白颊黑雁(Branta leucopsis)。     

Site fidelity and range size of wintering Barnacle Geese Branta leucopsis

CapsuleBarnacle Geese restrict their movements to relatively few key sites and exhibit considerable variation in ranging behaviour.

Aims To examine individual and seasonal variation in site fidelity, habitat use, range size and foraging strategies of Barnacle Geese Branta leucopsis.

Methods The movements of 18 male Barnacle Geese tagged in two discrete areas were tracked for 3–6 months from late autumn until departure on the spring migration.

Results Tagged geese concentrated their feeding in a relatively small proportion of apparently suitable habitat. Geese moved increasingly further afield in midwinter, and there was a clear predeparture shift to the largest area of relatively undisturbed, and possibly more nitrogen-rich, saltmarsh on the Solway. Birds from one of the two capture sites tended to be more sedentary and have smaller home ranges.

Conclusion There is considerable between-site and inter-individual variation in the degree of site-faithfulness of Barnacle Geese.     

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.     

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.     

The morphological development of the locomotor and cardiac muscles of the migratory barnacle goose (Branta Leucopsis)

The masses of the locomotor and acardiac muscles of wild barncale goose gollings, from migratory population, were examined systematically during development and their values compared to those of pre-migratory geese. Pre-flight development was typified by approximately linear increases of body, leg, and heart ventricular mass with respect to age. Flight muscle showed an exponential increase in mass. Pectoralis muscle mass was 14.2 ± 0.3% of body mass (1297 ± 73g, n=7) in early flying goslings compared to 16.6 ± 0.3% of body mass (2318 ± 109g, n=8) in pre-migratory geese. Post-flight development was typified by stasis of leg muscle mass but hypertrophy of Ventricular and pectoralis muscle mass in proportion to body mass. Ventricular mass relative to body mass showed the lowest values at 5 weeks of age (0.62 ± 0.01%) with peak values at 1 week of age (10.4 ± 0.04%). The latter may be associated with both requirements of thermoregualation in these precocial, arctic breeding geese and the need to forage approximately 24 hours post-hatch. Peak values for leg muscle mass, relative to body mass, were found at 3 weeks of age (12.7 ± 0.36%), with lowest values in the pre-migratory geese (6.7 ± 0.21%), while peak values for pectoralis muscle mass were expressed in the premigratory geese with lowest values at 1 week of age (0.94 ± 0.07%). Ventricular mass was proportional to leg muscle mass up to 5 weeks of age (M_v= 0.38M_t^0.68, r²=0.95), but subsequent increase in ventricular mass was proportional to pectoralis muscle mass (M_v= 0.25M_p^0.73, r²= 0.81).     

Natal and breeding dispersal of Canada geese Branta canadensis

Natal dispersal in Canada Geese Branta canadensis breeding in the English midlands is male-biased; males are more likely to change sites between birth and breeding, and move greater distances when they do so. Breeding dispersal is confined to within-site movements, and the evidence for a sex bias is equivocal. It is suggested that the ability to exploit information on brood rearing areas may be an important selection pressure favouring female philopatry in wildfowl.     

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.