Extrapolating herbivore-induced carbon loss across an arctic landscape

A high proportion of the global soil carbon stock is stored in tundra soils. However, populations of arctic-breeding migratory geese including pink-footed geese, Anser brachyrhynchus, are increasing due to agricultural changes and conservation measures in their wintering grounds. Foraging by these geese, which is widespread in extent, reduces the quantity of carbon stored in arctic tundra ecosystems. Here, the potential carbon loss caused by foraging pink-footed geese is modelled across the high-arctic archipelago of Svalbard, combining field experiments, habitat maps and published spatial models of foraging. The carbon loss caused by foraging geese was estimated three growing seasons following perturbation allowing for some recovery to take place. The carbon loss caused by 1-year worth of grubbing was estimated to be 1,700 tonnes, or 37 kg per goose. A total of over 340,000 tonnes of carbon could be affected given an unlimited increase in goose population. Estimated losses were mostly from wetter habitats, which are both carbon rich and highly selected for by foraging geese. The across-landscape carbon loss caused by geese is not great in magnitude in comparison to expected climate-driven carbon losses; however, it is locally severe, and demonstrates how migratory connectivity links processes, such as agricultural change and conservation measures in temperate Europe with carbon dynamics in the high arctic.     

Spring feeding by pink-footed geese reduces carbon stocks and sink strength in tundra ecosystems

Tundra ecosystems are widely recognized as precious areas and globally important carbon (C) sinks, yet our understanding of potential threats to these habitats and their large soil C store is limited. Land‐use changes and conservation measures in temperate regions have led to a dramatic expansion of arctic‐breeding geese, making them important herbivores of high‐latitude systems. In field experiments conducted in high‐Arctic Spitsbergen, Svalbard, we demonstrate that a brief period of early season belowground foraging by pink‐footed geese is sufficient to strongly reduce C sink strength and soil C stocks of arctic tundra. Mechanisms are suggested whereby vegetation disruption due to repeated use of grubbed areas opens the soil organic layer to erosion and will thus lead to progressive C loss. Our study shows, for the first time, that increases in goose abundance through land‐use change and conservation measures in temperate climes can dramatically affect the C balance of arctic tundra.     

Prevalence of pink-footed goose grubbing in the arctic tundra increases with population expansion

In the high-arctic archipelago of Svalbard, the pink-footed goose (Anser brachyrhynchus) population has increased dramatically over the last decades. The population increase and the corresponding range expansion suggest a substantial increase in the potential for disturbance of the tundra caused by goose herbivory. In this study, we used surveys of pink-footed goose grubbing from two separate years (2007 and 2012) to examine the temporal changes in the prevalence of grubbing in central Spitsbergen. During this time period, the Svalbard-breeding pink-footed goose population had increased from 56,400 to 80,000 individuals. We compared grubbing prevalence between the 2 years, standardising the comparison by using a published model of habitat suitability for grubbing (Speed et al. in Ecosystems 12:349-359, 2009) as a covariate. Habitat suitability was a significant predictor of grubbing prevalence across both years, with higher grubbing probability in habitats predicted to be more suitable for grubbing. The probability of grubbing was on average 4 times higher in 2012 than in 2007. In 2007, all tundra habitats had a probability of <27 % of being grubbed whilst in 2012, there was a probability for being grubbed of more than 34 % in the least suitable habitat and around 59 % in the most suitable habitat. The increase in grubbing prevalence demonstrates a great need for monitoring the expanding pink-footed goose population and its impact on the tundra landscape.     

Where might the western Svalbard tundra be vulnerable to pink‐footed goose (Anser brachyrhynchus) population expansion? Clues from species distribution models

Pink-footed geese ( Anser brachyrhynchus ) breed in the Arctic, where their populations have doubled since the 1980s. There is concern that nesting geese disturb the fragile tundra and lead to a trophic cascade with strong top-down effects on vegetation and soil processes. A better understanding of the distribution of geese and factors that influence nest site selection is needed to highlight potential problem areas and assess the potential for further population expansion. To help infer the importance of environmental variables on nest site selection, we built generalized additive models using nest observations collected in 2003 and 2004 from the Sassendalen valley, Svalbard, along with a suite of geographical information system explanatory predictors. The fit of the models was very high (explaining over 72% of the deviance), and predictive power to independent samples indicated useful predictions that could discriminate between presences and absence of nests very well (area under the receiver operating characteristic curves exceeded 0.88). Significant predictors of nest site selection included elevation, slope, aspect, percentage of snow cover, percentage of foraging habitat cover, and a spatial autocovariate. Spatial predictions were applied to the broader Nordenskiöldsland region of Svalbard and highlighted the importance of previously unsurveyed locations for nesting.     

The dilemma of where to nest: influence of spring snow cover,food proximity and predator abundance on reproductive success of an arctic-breeding migratory herbivore is dependent on nesting habitat choice

Pink-footed geese Anser brachyrhynchus nest in two contrasting but commonly found habitats: steep cliffs and open tundra slopes. In Svalbard, we compared nest densities and nesting success in these two environments over ten breeding seasons to assess the impact of spring snow cover, food availability to nesting adults and arctic fox Vulpes lagopus (main terrestrial predator) abundance. In years with extensive spring snow cover, fewer geese at both colonies attempted to breed, possibly because snow cover limited pre-nesting feeding opportunities, leaving adults in poor breeding condition. Nesting success at the steep cliff colony was lower with extensive spring snow cover; such conditions force birds to commit to repeated and prolonged recess periods at far distant feeding areas, leaving nests open to predation. By contrast, nesting success at the open tundra slope was not affected by spring snow cover; even if birds were apparently in poor condition they could feed immediately adjacent to their nests and defend them from predators. Foxes were the main nest predator in the open tundra slopes but avian predators likely had a larger impact at the steep cliffs colony. Thus, the relative inaccessibility of the cliffs habitat may bring protection from foxes but also deprives geese from readily accessing feeding areas, with the best prospects for successful nesting in low spring snow cover years. Our findings indicate that spring snow cover, predator abundance and food proximity did not uniformly influence nesting success of this herbivore, and their effects were dependent on nesting habitat choice.     

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.     

Historical and ecological determinants of genetic structure in arctic canids

Wolves (Canis lupus) and arctic foxes (Alopex lagopus) are the only canid species found throughout the mainland tundra and arctic islands of North America. Contrasting evolutionary histories, and the contemporary ecology of each species, have combined to produce their divergent population genetic characteristics. Arctic foxes are more variable than wolves, and both island and mainland fox populations possess similarly high microsatellite variation. These differences result from larger effective population sizes in arctic foxes, and the fact that, unlike wolves, foxes were not isolated in discrete refugia during the Pleistocene. Despite the large physical distances and distinct ecotypes represented, a single, panmictic population of arctic foxes was found which spans the Svalbard Archipelago and the North American range of the species. This pattern likely reflects both the absence of historical population bottlenecks and current, high levels of gene flow following frequent long-distance foraging movements. In contrast, genetic structure in wolves correlates strongly to transitions in habitat type, and is probably determined by natal habitat-biased dispersal. Nonrandom dispersal may be cued by relative levels of vegetation cover between tundra and forest habitats, but especially by wolf prey specialization on ungulate species of familiar type and behaviour (sedentary or migratory). Results presented here suggest that, through its influence on sea ice, vegetation, prey dynamics and distribution, continued arctic climate change may have effects as dramatic as those of the Pleistocene on the genetic structure of arctic canid species.     

Herbivore impacts to the moss layer determine tundra ecosystem response to grazing and warming

Herbivory and climate are key environmental drivers, shaping ecosystems at high latitudes. Here, we focus on how these two drivers act in concert, influencing the high arctic tundra. We aim to investigate mechanisms through which herbivory by geese influences vegetation and soil processes in tundra ecosystems under ambient and warmed conditions. To achieve this, two grazing treatments, clipping plus faecal additions and moss removal, were implemented in conjunction with passive warming. Our key finding was that, in many cases, the tundra ecosystem response was determined by treatment impacts on the moss layer. Moss removal reduced the remaining moss layer depth by 30% and increased peak grass biomass by 27%. These impacts were probably due to observed higher soil temperatures and decomposition rates associated with moss removal. The positive impact of moss removal on grass biomass was even greater with warming, further supporting this conclusion. In contrast, moss removal reduced dwarf shrub biomass possibly resulting from increased exposure to desiccating winds. An intact moss layer buffered the soil to increased air temperature and as a result there was no response of vascular plant productivity to warming over the course of this study. In fact, moss removal impacts on soil temperature were nearly double those of warming, suggesting that the moss layer is a key component in controlling soil conditions. The moss layer also absorbed nutrients from faeces, promoting moss growth. We conclude that both herbivory and warming influence this high arctic ecosystem but that herbivory is the stronger driver of the two. Disturbance to the moss layer resulted in a shift towards a more grass-dominated system with less abundant mosses and shrubs, a trend that was further enhanced by warming. Thus herbivore impacts to the moss layer are key to understanding arctic ecosystem response to grazing and warming.     

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.     

Density-dependent switches in diet: a likely mechanism for negative feedbacks on goose population increase?

Goose grazing on arctic tundra vegetation has shown both positive and negative effects on subsequent foraging conditions. To understand the potential of a density-dependent feedback on herbivore population size, the relation between grazing pressure and future foraging conditions is essential. We studied the effect of increasing grazing pressure of barnacle geese (Branta leucopsis) on Spitsbergen. During the establishment of a breeding colony in the period 1992–2004, the proportion of graminoids decreased in the diet of wild geese, while the percentage of mosses increased. Grazing trials with captive geese in an unexploited area showed a similar shift in diet composition. High-quality food plants were depleted within years and over years. Intake rate declined too and as consequence, metabolisable energy intake rate (MEIR) decreased rapidly with increasing grazing pressure. During three successive years of experimental grazing, MEIR decreased at all levels of grazing pressure and declined below minimal energetic requirements when grazing exceeded natural levels of grazing pressure. This suggests that foraging conditions rapidly decline with increasing grazing pressure in these low-productive habitats. The potential for density-dependent feedbacks on local population increase is discussed.     

Using habitat selection theories to predict the spatiotemporal distribution of migratory birds during stopover – a case study of pink‐footed geese Anser brachyrhynchus

Understanding how animals select for habitat and foraging resources therein is a crucial component of basic and applied ecology. The selection process is typically influenced by a variety of environmental conditions including the spatial and temporal variation in the quantity and quality of food resources, predation or disturbance risks, and inter‐ and intraspecific competition. Indeed, some of the most commonly employed ecological theories used to describe how animals choose foraging sites are: nutrient intake maximisation, density‐dependent habitat selection, central‐place foraging, and predation risk effects. Even though these theories are not mutually exclusive, rarely are multiple theoretical models considered concomitantly to assess which theory, or combination thereof, best predicts observed changes in habitat selection over space and time. Here, we tested which of the above theories best‐predicted habitat selection of Svalbard‐breeding pink‐footed geese at their main spring migration stopover site in mid‐Norway by computing a series of resource selection functions (RSFs) and their predictive ability (k‐fold cross validation scores). At this stopover site geese fuel intensively as a preparation for breeding and further migration. We found that the predation risk model and a combination of the density‐dependent and central‐place foraging models best‐predicted habitat selection during stopover as geese selected for larger fields where predation risk is typically lower and selection for foraging sites changed as a function of both distance to the roost site (i.e. central‐place) and changes in local density. In contrast to many other studies, the nutritional value of the available food resources did not appear to be a major limiting factor as geese used different food resources proportional to their availability. Our study shows that in an agricultural landscape where nutritional value of food resources is homogeneously high and resource availability changes rapidly; foraging behaviour of geese is largely a tradeoff between fast refuelling and disturbance/predator avoidance.     

Changes in the structure and function of northern Alaskan ecosystems when considering variable leaf‐out times across groupings of species in a dynamic vegetation model

The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.     

Do herbivores cause habitat degradation or vegetation state transition? Evidence from the tundra

Range expansion and increasing densities of large herbivores are held responsible for large-scale habitat degradation in a wide range of natural and semi-natural ecosystems. Herbivore-driven ecosystem changes frequently represent predictable transitions from one vegetation state to another. Whether such predictable changes justify the value judgement 'habitat degradation' may be debatable as this strongly depends on individual perspective.
To further the debate on herbivore-driven habitat degradation, I apply the concept of alternative stable states to arctic tundra as a framework to capture predictable stepwise vegetation transitions in which the productivity and hence herbivore-carrying capacity increases with grazing pressure. Specifically, evidence is provided that large parts of the tundra biome can be in either of three relatively discrete vegetation states and that changes in reindeer/caribou density are responsible for sudden, predictable but often reversible state transitions. From this, it appears that the relatively rapidly emerging vegetation changes do not necessarily equate to habitat degradation, but in many cases reflect predictable vegetation change. Acknowledgement of the existence of predictable state transitions in tundra ecosystems may help to evaluate the observed radical vegetation changes occurring throughout the reindeer/caribou range.     

Climate-Driven Effects of Fire on Winter Habitat for Caribou in the Alaskan-Yukon Arctic

Climatic warming has direct implications for fire-dominated disturbance patterns in northern ecosystems. A transforming wildfire regime is altering plant composition and successional patterns, thus affecting the distribution and potentially the abundance of large herbivores. Caribou (Rangifer tarandus) are an important subsistence resource for communities throughout the north and a species that depends on terrestrial lichen in late-successional forests and tundra systems. Projected increases in area burned and reductions in stand ages may reduce lichen availability within caribou winter ranges. Sufficient reductions in lichen abundance could alter the capacity of these areas to support caribou populations. To assess the potential role of a changing fire regime on winter habitat for caribou, we used a simulation modeling platform, two global circulation models (GCMs), and a moderate emissions scenario to project annual fire characteristics and the resulting abundance of lichen-producing vegetation types (i.e., spruce forests and tundra >60 years old) across a modeling domain that encompassed the winter ranges of the Central Arctic and Porcupine caribou herds in the Alaskan-Yukon Arctic. Fires were less numerous and smaller in tundra compared to spruce habitats throughout the 90-year projection for both GCMs. Given the more likely climate trajectory, we projected that the Porcupine caribou herd, which winters primarily in the boreal forest, could be expected to experience a greater reduction in lichen-producing winter habitats (−21%) than the Central Arctic herd that wintered primarily in the arctic tundra (−11%). Our results suggest that caribou herds wintering in boreal forest will undergo fire-driven reductions in lichen-producing habitats that will, at a minimum, alter their distribution. Range shifts of caribou resulting from fire-driven changes to winter habitat may diminish access to caribou for rural communities that reside in fire-prone areas.     

Changes in the Spatial Configuration and Strength of Trophic Control Across a Productivity Gradient During a Massive Rodent Outbreak

Understanding the determinants of spatial and temporal differences in the relative strength of consumer–resource interactions is an important endeavour in ecology. Here, we explore the necessary conditions for temporal shifts in the relative strength of rodent–plant interactions in an area characterised by profound spatial differences in trophic control, with predator–prey interactions prevailing in productive habitats and rodent–plant interactions dominating unproductive habitats of the forest–tundra ecotone. We report data obtained during the exceptionally massive rodent outbreak of 2010–2012 in northernmost Fennoscandia, including an experimental manipulation of herbivore access to vegetation plots across a large-scale productivity gradient, multiple observational measures of plant–rodent interactions linked to rodent abundance data and a large-scale survey of breeding avian predators and mammalian predator activity. Unexpectedly, rodent grazing impacts documented during the rodent outbreak were uniformly strong across the landscape, regardless of habitat productivity. The runaway response in rodent populations was facilitated by a high population growth rate in the early phase of the outbreak due to the extended absence of predators in productive habitats, concomitant with an exceptionally long-lasting lemming outbreak in unproductive habitats. Our results showed that spatio-temporal variation in trophic control also occurs in ecosystems structured according to the exploitation ecosystems hypothesis and emphasises the importance of long-term studies to capture nonlinear and stochastic features that shape ecosystem functioning. In this context, the temporary release from top–down regulation in productive habitats caused strong grazing impacts that may be crucial for the resilience of tundra ecosystems under the threat of climate change-driven shrub encroachment.     

Tundra be dammed: Beaver colonization of the Arctic

Increasing air temperatures are changing the arctic tundra biome. Permafrost is thawing, snow duration is decreasing, shrub vegetation is proliferating, and boreal wildlife is encroaching. Here we present evidence of the recent range expansion of North American beaver (Castor canadensis) into the Arctic, and consider how this ecosystem engineer might reshape the landscape, biodiversity, and ecosystem processes. We developed a remote sensing approach that maps formation and disappearance of ponds associated with beaver activity. Since 1999, 56 new beaver pond complexes were identified, indicating that beavers are colonizing a predominantly tundra region (18,293 km²) of northwest Alaska. It is unclear how improved tundra stream habitat, population rebound following overtrapping for furs, or other factors are contributing to beaver range expansion. We discuss rates and likely routes of tundra beaver colonization, as well as effects on permafrost, stream ice regimes, and freshwater and riparian habitat. Beaver ponds and associated hydrologic changes are thawing permafrost. Pond formation increases winter water temperatures in the pond and downstream, likely creating new and more varied aquatic habitat, but specific biological implications are unknown. Beavers create dynamic wetlands and are agents of disturbance that may enhance ecosystem responses to warming in the Arctic.     

Additive Partitioning of Diversity Reveals No Scale-dependent Impacts of Large Ungulates on the Structure of Tundra Plant Communities

Large herbivores can change ecosystem functioning by impacting plant diversity. However, although such impacts are expected to be scale-dependent in ecosystems with wide-roaming ungulates, scaling issues rarely enter empirical assessments. We here test the hypothesis that the impact of increased reindeer abundance on plant diversity in alpine tundra is scale-dependent. Based on potentially high productivity of the focal habitat units and hence the possibility of positive grazer impacts on plant diversity we predicted higher α and β diversity at the habitat scale where reindeer densities are high. We also explored whether there were differences in diversity patterns at larger scales, including the scale of reindeer management districts. We estimated grazing disturbance as high versus low reindeer density in selected districts (a total extent of 7421 km²) of Northern Norway where reindeer-induced vegetation shifts are debated. We focus on dominance patterns because they can quantify the vegetation state and thus performed additive partitioning of Simpson diversity on multiple scales assessing also species’ contributions to diversity. Contrary to our predictions, we found only weak scale-dependent effects of reindeer grazing on plant diversity. Under high reindeer densities there was evidence for a landscape-scale homogenization of the vegetation, but the predicted α and β diversity increases at the habitat scale were not found. Consistently through all scales considered, four shrub species contributed the most to plant diversity. These results contradict the idea that reindeer at high stocking densities induce shifts in plant species dominance in productive habitats. We conclude that context-dependencies such as spatial scales of management units and habitat types need to be explicitly considered in evaluations of the impacts of large ungulates on plant diversity.     

Birds,nutrients, and climate change: mtDNA haplotype diversity of Arctic <Emphasis Type="Italic">Daphnia</Emphasis> on Svalbard revisited

Ecosystems in the high Arctic are in transition due to climate change and species shifts. On the Svalbard archipelago, the average annual temperature has increased by more than 2 °C over the past 30 years, and there has been a striking increase in breeding populations of geese. Birds serve as a dominant source of nutrients (via faeces) and may also serve as vectors of dispersal of many small aquatic organisms. We compared samples of species and haplotype composition of the dominant freshwater crustacean Daphnia spp., from 1992, and those resampled in 2014 to see if these major impacts on Arctic freshwater ecosystems may also have affected this key grazer over the past three decades. The study covers tundra ponds that vary in levels of nutrients, abundance, and diversity of birds. Comparison of genetic mitochondrial DNA sequences revealed little change in haplotype and nucleotide diversity between 1992 and 2014, but higher species and haplotype diversity were found in nutrient-rich ponds that hosted large migratory bird populations. This could either reflect that high nutrient levels allow for the maintenance of higher levels of genetic diversity (i.e. haplotypes, lineages), that birds serve as vectors for the dispersal of clones, or likely a combination of both mechanisms.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Impacts of disturbance on migratory waterfowl

It is well known that disturbance from human activities can cause temporary changes in behaviour and locally affect temporal and spatial distribution of migratory and wintering waterfowl. But it is also known that, to some extent, birds can compensate for disturbance by altering their behaviour or habituating to human activities. Comparatively little is known about how these reactions to disturbance may impact on the large-scale dispersion of waterfowl and, ultimately, on their population dynamics. To be able to answer these questions, a better theoretical framework, based on optimal foraging theory incorporating predation risk, and field experiments are required. Furthermore, we need to study the waterfowl throughout their winter ranges to interpret the overall impacts of disturbance. This paper examines two cases where the impacts of disturbance have been assessed from field experiments. In one study, disturbance effects of shooting were tested by setting up experimental reserves in two Danish coastal wetlands. Over a 5-year period, these became two of the most important staging areas for coastal waterfowl, and the national totals of key species were significantly increased. A national management plan which will establish more than 50 new shooting-free refuges on Danish coastal areas within the next 5 years is likely to boost waterfowl numbers even more. Such retention of birds at more northerly sites on the fiyway should result in a more efficient resource utilization and may positively affect the population dynamics where numbers are affected by winter resources. In a second study, the impacts of disturbance by farmers on spring fattening of Pink-footed Geese Anser brachyrhynchus were analysed. In undisturbed areas in northern Norway, abdominal profiles of the geese increased rapidly, whereas in disturbed sites they did not. Subsequently, geese that had used undisturbed sites reproduced better than geese from disturbed sites.