Maarten J.J.E. Loonen

Subordinates explore but dominants profit: resource competition in high Arctic barnacle goose flocks

Social dominance plays an important role in assessing and obtaining access to patchy or scarce food sources in group-foraging herbivores. We investigated the foraging strategies of individuals with respect to their social position in the group in a flock of nonbreeding, moulting barnacle geese, Branta leucopsis, on high Arctic Spitsbergen. We first determined the dominance rank of individually marked birds. The dominance of an individual was best described by its age and its sex-specific body mass. Mating status explained the large variation in dominance among younger birds, as unpaired yearlings ranked lowest. In an artificially created, competitive situation, subordinate individuals occupied explorative front positions in the flock and were the first to find sites with experimentally enriched vegetation. Nevertheless, they were displaced quickly from these favourable sites by more dominant geese which were able to monopolize them. The enhanced sites were subsequently visited preferentially by individuals that succeeded in feeding there when the exclosures were first opened. Data on walking speed of foraging individuals and nearest-neighbour distances in the group suggest that subordinates try to compensate for a lower energy intake by exploring and by lengthening the foraging bout. Observations of our focal birds during the following breeding season revealed that females that returned to the study area were significantly more dominant in the previous year than those not seen in the area again. Copyright 2001 The Association for the Study of Animal Behaviour.

Herbivore impact on moss depth, soil temperature and arctic plant growth

Abstract We provide evidence for a mechanism by which herbivores may influence plant abundance in arctic ecosystems. These systems are commonly dominated by mosses, the thickness of which influences the amount of heat reaching the soil surface. Herbivores can reduce the thickness of the moss layer by means of trampling and consumption. Exclusion of grazing by barnacle geese and reindeer over a period of 7 years at Ny-Ålesund, Spitsbergen, caused an increase in the thickness of the moss layer, and a reduction in soil temperature of 0.9 °C. Soil temperature was negatively correlated with moss-layer thickness across sites, with highest soil temperatures where moss layers were shallow. We found that moss growth did not respond to experimental manipulation of soil temperature, but the grass Poa arctica (arctic meadow-grass) and the dicot Cardamine nymanii (polar cress) suffered a 50% reduction in biomass when growing in chilled soils.

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, FST, 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.

Short-term and long-term facilitation of goose grazing by livestock in the Dutch Wadden Sea area

We studied the impact of livestock grazing on the distribution ofBranta bernicla bernicla (Dark-bellied Brent goose) in the Dutch Wadden Sea during spring. It was hypothesized that livestock facilitate short-term (within-season) grazing for geese as well as long-term (over years). Therefore we measured grazing pressure by geese in salt marsh and polder areas that were either grazed (spring-grazed) or ungrazed during spring (summer-grazed). Additionally, we carried out a preference experiment with captive geese to test the preference between spring-grazed and summer-grazed polder swards. We furthermore compared patterns of use by geese between long-term ungrazed and grazed salt marshes.In May, there is a difference in grazing pressure by geese between polder pastures that are grazed or ungrazed during spring. In this month, the ungrazed polder pastures are abandoned and the geese shift to either the grazed polder pastures or to the salt marsh. Vegetation in the polder that had been spring-grazed had a lower canopy height and a higher tiller density than summer-grazed vegetation. The captive geese in the preference experiment showed a clear preference for vegetation that had been spring-grazed by sheep over ungrazed vegetation. Goose grazing pressure was negatively correlated to canopy height, both on the polder and on the salt marsh. Within the plant communities dominated byFestuca rubra andPuccinellia maritima, marshes that were intensively grazed by livestock generally had higher grazing pressure by geese than long-term ungrazed or lightly grazed salt marshes.

The influence of goose grazing on the growth of Poa arctica: overestimation of overcompensation

The effect of grazing by barnacle geese (Branta leucopsis) on above-ground production was investigated on the Arctic tundra grass Poa arctica at Spitsbergen during the growing season of 1996. We applied three treatments: orated; whole season ungrazed and temporary ungrazed during the second half of the season. In each treatment individual shoots were marked and their leaves regularly measured to calculate production, Production was calculated in three different ways: Increment in Standing Crop (ISC), Above-ground Biomass Production (ABP) and leafbirths. ISC in temporary and permanent exclosures is the most commonly used method, but it neglects senescence and disappearance of biomass during the experiment. In our study, ISC of grazed Poa arctica was significantly higher than of ungrazed swards. However, the flow of senescing biomass was much lai ger in ungrazed swards compared to grazed swards. Thereby an overestimation of overcompensation was introduced. Calculation of ABP takes this flow of senescing biomass into account. In terms of ABP, Poa arctica did not overcompensate the losses due to goose grazing. Accordingly, the number of leafbirths did not differ between grazed and ungrazed swards. In several other studies, where senescence and disappearance of leaves were neglected, inclusion of these fluxes would at least decrease overcompensation. Therefore, we advocate regarding senescence and disappearance of leaves while measuring overcompensation.

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.

Nitrogen transfer between herbivores and their forage species

Herbivores may increase the productivity of forage plants; however, this depends on the return of nutrients from faeces to the forage plants. The aim of this study was to test if nitrogen (N) from faeces is available to forage plants and whether the return of nutrients differs between plant species using 15N natural abundance in faeces and plant tissue. To investigate the effect of grazing on N transfer, we carried out a grazing experiment in wet and mesic tundra on high Arctic Spitsbergen using barnacle geese (Branta leucopsis) as the model herbivore. N inputs (from faeces) increased with grazing pressure at both the wet and mesic sites, with the greatest N input from faeces at the wet site. The δ15N ratio in plant tissue from grazed plots was enriched in mosses and the dwarf shrub species, reflecting the δ15N signature of faeces-derived N, but no such pattern was observed in the dominant grasses. This study demonstrates that the δ15N signature of faeces and forage species is a useful tool to explore how grazing impacts on N acquisition. Our findings suggest that plant species which acquire their N close to the soil surface (e.g. mosses) access more of the N from faeces than species with deeper root systems (e.g. grasses) suggesting a transfer of N from the preferred forage species to the mosses and dwarf shrubs, which are less preferred by the geese. In conclusion, the moss layer appears to disrupt the nitrogen return from herbivores to their forage species.

Latitudinal variability in the seroprevalence of antibodies against Toxoplasma gondii in non-migrant and Arctic migratory geese

Toxoplasma gondii is an intracellular coccidian parasite found worldwide and is known to infect virtually all warm-blooded animals. It requires a cat (family Felidae) to complete its full life cycle. Despite the absence of wild felids on the Arctic archipelago of Svalbard, T. gondii has been found in resident predators such as the arctic fox and polar bear. It has therefore been suggested that T. gondii may enter this ecosystem via migratory birds. The objective of this study was to identify locations where goose populations may become infected with T. gondii, and to investigate the dynamics of T. gondii specific antibodies. Single blood samples of both adults and juveniles were collected from selected goose species (Anser anser, A. brachyrhynchus, Branta canadensis, B. leucopsis) at Arctic brood-rearing areas in Russia and on Svalbard, and temperate wintering grounds in the Netherlands and Denmark (migratory populations) as well as temperate brood-rearing grounds (the Netherlands, non-migratory populations). A modified agglutination test was used on serum, for detection of antibodies against T. gondii. Occasional repeated annual sampling of individual adults was performed to determine the antibody dynamics. Adults were found seropositive at all locations (Arctic and temperate, brood-rearing and wintering grounds) with low seroprevalence in brood-rearing birds on temperate grounds. As no juvenile geese were found seropositive at any brood-rearing location, but nine month old geese were found seropositive during spring migration we conclude that geese, irrespective of species and migration, encounter T. gondii infection in wintering areas. In re-sampled birds on Svalbard significant seroreversion was observed, with 42% of seropositive adults showing no detectable antibodies after 12 months, while the proportion of seroconversion was only 3%. Modelled variation of seroprevalence with field data on antibody longevity and parasite transmission suggests seroprevalence of a population within a range of 5.2-19.9%, in line with measured values. The high occurrence of seroreversion compared to the low occurrence of seroconversion hampers analysis of species- or site-specific patterns, but explains the absence of an increase in seroprevalence with age and the observed variation in antibody titre. These findings imply that even though infection rate is low, adults introduce T. gondii to the high Arctic ecosystem following infection in temperate regions.

Keeping the clock set under the midnight sun : diurnal periodicity and synchrony of avian Isospora parasites cycle in the High Arctic

For Isospora (Protozoa: Eimeriidae) parasites of passerine birds, diurnal periodicity of oocyst output is a well-described phenomenon. From the temporal zone to the tropics, oocyst production is correlated with the light-dark cycle, peaking in the afternoon hours. However, nothing is known about the existence of diurnal periodicity of these parasites in the birds of High Arctic environments, under permanent light during summer. We sampled free-ranging Snow Bunting (Aves: Passeriformes), on Svalbard in summer and tested oocysts output of Isospora plectrophenaxia. Here we show that under the permanent light conditions of Arctic summer in the wild, Isospora plectrophenaxia, a parasite of the Snow Bunting, still keeps the 24-h rhythm of oocyst output with the peak in the post-meridiem hours, despite the absence of diurnal periodicity in hosts activity. Our findings prove the ability of avian Isospora to invoke alternative cues for synchronizing the circadian rhythms. Possible cues and adaptive significance of diurnal periodicity of parasite output in High Arctic are discussed. The maintenance of synchronization and timing of the parasite life-cycle stages is under positive selection pressure even in permanent daylight in the Arctic.

Intensive grazing by Barnacle geese depletes High Arctic seed bank

Studies in the Canadian Arctic show dramatic effects of increased goose grazing on vegetation structure and soil conditions, but little is known of the role of goose grazing in the European Arctic. We focused on how geese might affect plant recruitment via effects on seed production and soil seed bank in High Arctic Svalbard. Experimental grazing by captive Barnacle geese (Branta leucopsis (Bechstein, 1803)) decreased flower densities both at normal and at high graz- ing pressure. Geese showed a clear preference for reproductive rather than vegetative shoots. Soil samples collected inside and outside 7-year-old exclosures in an intensively goose-grazed area revealed significant effects on the germinable soil seed bank. The density of viable seeds in the top soil layer inside exclosures was six times higher than in grazed plots. Lower densities of viable seeds occurred in the basal than in the top layer but there was no difference in basal layer seed density between exclosed and grazed plots. This study shows that geese have strong effects on floral abundance and conse- quently on the seed bank. We argue that goose grazing in these systems influences the potential for recovery after a dis- turbance event and thus the long-term plant species diversity and dynamics.