Strength of asymmetric competition between predators in food webs ruled by fluctuating prey: the case of foxes in tundra

In food webs heavily influenced by multi‐annual population fluctuations of key herbivores, predator species may differ in their functional and numerical responses as well as their competitive ability. Focusing on red and arctic fox in tundra with cyclic populations of rodents as key prey, we develop a model to predict how population dynamics of a dominant and versatile predator (red fox) impacted long‐term growth rate of a subdominant and less versatile predator (arctic fox). We compare three realistic scenarios of red fox performance: (1) a numerical response scenario where red fox acted as a resident rodent specialist exhibiting population cycles lagging one year after the rodent cycle, (2) an aggregative response scenario where red fox shifted between tundra and a nearby ecosystem (i.e. boreal forest) so as to track rodent peaks in tundra without delay, and (3) a constant subsidy scenario in which the red fox population was stabilized at the same mean density as in the other two scenarios. For all three scenarios it is assumed that the arctic fox responded numerically as a rodent specialist and that the mechanisms of competition is of a interference type for space, in which the arctic fox is excluded from the most resource rich patches in tundra. Arctic fox is impacted most by the constant subsidy scenario and least by the numerical response scenario. The differential effects of the scenarios stemmed from cyclic phase‐dependent sensitivity to competition mediated by changes in temporal mean and variance of available prey to the subdominant predator. A general implication from our result is that external resource subsidies (prey or habitats), monopolized by the dominant competitor, can significantly reduce the likelihood for co‐existence within the predator guild. In terms of conservation of vulnerable arctic fox populations this means that the likelihood of extinction increases with increasing amount of subsidies (e.g. carcasses of large herbivores or marine resources) in tundra and nearby forest areas, since it will act to both increase and stabilize populations of red fox.     

Diet of arctic foxes (Alopex lagopus) in Iceland

Arctic foxes, Alopex lagopus , live in low productivity arctic and northern tundra habitats, where they generally prey heavily on lemmings. In Iceland, however, no lemmings are present, and the foxes have a very varied diet, including plants such as seaweed and black crowberries, a wide range of birds and invertebrates, and carcasses of large mammals such as seals, reindeer, and sheep. Marked seasonal, geographical and inter-annual differences confirm arctic foxes in Iceland as opportunistic feeders. There are coastal and inland foxes: coastal foxes feed mainly on prey derived directly or indirectly from the ocean, particularly various seabirds and seals, while inland foxes feed largely on migrant birds, such as geese, waders and passerines in summer, and ptarmigan in winter. Despite their reputation for killing lambs, in this study, lamb carcasses were found at only 19.4% of 1125 fox dens, 44% of which had only one carcass. The distance to the nearest farm and the physical condition of lambs were major determinants of the number of carcasses found at a den. We discuss the implications of arctic foxes' diet for population dynamics and group formation, and for management practices.     

Predator–rodent–plant interactions along a coast–inland gradient in Fennoscandian tundra

Spatial variation in the strength of trophic cascades in arctic tundra has been related to flows of subsidies across ecosystem boundaries. Here, we ask whether the input of marine subsidies in tundra systems would cause spatial variation in the strength of rodent–plant interactions between coastal areas, where predators have access to marine‐derived resources, and non‐subsidized inland areas of northern Fennoscandia. We present a detailed evaluation of predator–rodent–vegetation interactions along a coast‐inland gradient, during the 2011 rodent outbreak and the two following decline years, by using direct assessments of rodent impacts and tracing of marine‐derived nutrients in the food web. Our results revealed that the main rodent predator during summer, the long‐tailed jaeger Stercorarius longicaudus, did not benefit from marine resources while breeding (relative dietary proportion in chicks’ diet = 0–3%). Contrary to this pattern, parasitic jaegers S. parasiticus, bred exclusively near the coast and preyed effectively on both marine resources (41% of chicks’ diet) and rodents (12%). Mammalian predators also showed a higher activity during winter near the coast. Despite overall higher predator numbers, no evidence was found for lower rodent population growth rates during the three monitoring summers and for weaker rodent grazing impacts in the coastal area. Instead, we documented pronounced damages caused by lemmings and voles on bryophytes and vascular plants, especially dwarf shrubs (e.g. Vaccinum myrtillus) all along the coast–inland gradient. Taken together, our results did not support the hypothesis that marine subsidies would trigger a trophic cascade in coastal tundra areas of northern Fennoscandia during a major rodent outbreak, probably due to a relatively low diversity of marine‐subsidized predators in the region. Comparative observational and experimental studies at large spatial scales in various arctic regions are absolutely necessary for a better understanding of factors causing regional variations in the functioning of arctic food webs.     

Rough-Legged Buzzards,Arctic Foxes and Red Foxes in a Tundra Ecosystem without Rodents

Small rodents with multi-annual population cycles strongly influence the dynamics of food webs, and in particular predator-prey interactions, across most of the tundra biome. Rodents are however absent from some arctic islands, and studies on performance of arctic predators under such circumstances may be very instructive since rodent cycles have been predicted to collapse in a warming Arctic. Here we document for the first time how three normally rodent-dependent predator species—rough-legged buzzard, arctic fox and red fox – perform in a low-arctic ecosystem with no rodents. During six years (in 2006-2008 and 2011-2013) we studied diet and breeding performance of these predators in the rodent-free Kolguev Island in Arctic Russia. The rough-legged buzzards, previously known to be a small rodent specialist, have only during the last two decades become established on Kolguev Island. The buzzards successfully breed on the island at stable low density, but with high productivity based on goslings and willow ptarmigan as their main prey – altogether representing a novel ecological situation for this species. Breeding density of arctic fox varied from year to year, but with stable productivity based on mainly geese as prey. The density dynamic of the arctic fox appeared to be correlated with the date of spring arrival of the geese. Red foxes breed regularly on the island but in very low numbers that appear to have been unchanged over a long period – a situation that resemble what has been recently documented from Arctic America. Our study suggests that the three predators found breeding on Kolguev Island possess capacities for shifting to changing circumstances in low-arctic ecosystem as long as other small - medium sized terrestrial herbivores are present in good numbers.     

The fluctuating world of a tundra predator guild: bottom‐up constraints overrule top‐down species interactions in winter

Global warming is predicted to change ecosystem functioning and structure in Arctic ecosystems by strengthening top‐down species interactions, i.e. predation pressure on small herbivores and interference between predators. Yet, previous research is biased towards the summer season. Due to greater abiotic constraints, Arctic ecosystem characteristics might be more pronounced in winter. Here we test the hypothesis that top‐down species interactions prevail over bottom‐up effects in Scandinavian mountain tundra (Northern Sweden) where effects of climate warming have been observed and top‐down interactions are expected to strengthen. But we test this ‘a priori’ hypothesis in winter and throughout the 3–4 yr rodent cycle, which imposes additional pulsed resource constraints. We used snowtracking data recorded in 12 winters (2004–2015) to analyse the spatial patterns of a tundra predator guild (arctic fox Vulpes lagopus, red fox Vulpes vulpes, wolverine Gulo gulo) and small prey (ptarmigan, Lagopus spp). The a priori top‐down hypothesis was then tested through structural equation modelling, for each phase of the rodent cycle. There was weak support for this hypothesis, with top‐down effects only discerned on arctic fox (weakly, by wolverine) and ptarmigan (by arctic fox) at intermediate and high rodent availability respectively. Overall, bottom‐up constraints appeared more influential on the winter community structure. Cold specialist predators (arctic fox and wolverine) showed variable landscape associations, while the boreal predator (red fox) appeared strongly dependent on productive habitats and ptarmigan abundance. Thus, we suggest that the unpredictability of food resources determines the winter ecology of the cold specialist predators, while the boreal predator relies on resource‐rich habitats. The constraints imposed by winters and temporary resource lows should therefore counteract productivity‐driven ecosystem change and have a stabilising effect on community structure. Hence, the interplay between summer and winter conditions should determine the rate of Arctic ecosystem change in the context of global warming.     

Reproductive responses to spatial and temporal prey availability in a coastal Arctic fox population

1.?Input of external subsidies in the Arctic may have substantial effects on predator populations that otherwise would have been limited by low local primary productivity. 2.?We explore life-history traits, age-specific fecundity, litter sizes and survival, and the population dynamics of an Arctic fox (Vulpes lagopus) population to explore the influence of the spatial distribution and temporal availability of its main prey; including both resident and migrating (external) prey resources. 3.?This study reveals that highly predictable cross-boundary subsidies from the marine food web, acting through seasonal access to seabirds, sustain larger local Arctic fox populations. Arctic fox dens located close to the coast in Svalbard were found to have higher occupancy rates, as expected from both high availability and high temporal and spatial predictability of prey resources (temporally stable external subsidies). Whereas the occupancy rate of inland dens varied between years in relation to the abundance of reindeer carcasses (temporally varying resident prey). 4.?With regard to demography, juvenile Arctic foxes in Svalbard have lower survival rates and a high age of first reproduction compared with other populations. We suggest this may be caused by a lack of unoccupied dens and a saturated population.     

How ecological neighbourhoods influence the structure of the scavenger guild in low arctic tundra

Aim How the ecological neighbourhoods of coast and forest affect arctic tundra ecosystems is a pressing question as the circumpolar tundra belt is shrinking under global warming. Mobile facultative scavengers are likely to negatively impact tundra biodiversity as dominant competitors or predators, if they spill over into tundra. Here, we provide the first quantitative assessments of the structure of a scavenger guild in low arctic tundra with emphasis on how it changes along spatial gradients from neighbouring ecosystems (i.e. forest and coast) and with altitude (i.e. productivity gradients). We also assess the likelihood of interactions between guild members that may negatively impact vulnerable tundra species. Location North‐eastern part of Norway. Methods Extensive records of scavenger prevalence were obtained by deploying automatic digital cameras at experimental carcasses in tundra regions covering several thousand square kilometres and three winters in northern Norway. Main conclusions We found short‐range neighbourhood effects of forest and coast within the tundra scavenger guild. Species richness declined steeply with decreasing distance from the neighbouring ecosystems, in particular subarctic forest, and with increasing altitude. Bird species with strongholds in forest (golden eagle Aquila chrysaetos and hooded crow Corvus cornix) or along the coast (white‐tailed eagle Haliaeetus albicilla) were mostly responsible for short‐range neighbourhood effects on guild structure. However, the two most abundant guild members, the common raven Corvus corax and the red fox Vulpes vulpes, exhibited no spatial patterns within the range of neighbourhoods and altitudes examined. There was a clear diurnal segregation in the use of carcasses between birds and mammals reducing the likelihood of direct interactions between these two taxa. Presence of red fox appeared to exclude the arctic fox Vulpes lagopus, the only endemic tundra species within the guild, from carcasses.     

Seasonal pulses of migratory prey and annual variation in small mammal abundance affect abundance and reproduction by arctic foxes

We examined how large seasonal influxes of migratory prey influenced population dynamics of arctic foxes and how this varied with fluctuations in small mammal (lemming and vole) abundance—the main prey of arctic foxes throughout most of their range. Specifically, we compared how arctic fox abundance, breeding density and litter size varied inside and outside a large goose colony and in relation to annual variation in small mammal abundance. Information-theoretic model selection showed that (1) breeding density and fox abundance were 2–3 times higher inside the colony than they were outside the colony and (2) litter size, breeding density and annual variation in fox abundance in the colony tracked fluctuations in lemming abundance. The influence of lemming abundance on reproduction and abundance of arctic foxes outside the colony was inconclusive, largely because fox densities outside the colony were low, which made it difficult to detect such relationships. Lemming abundance was, thus, the main factor governing reproduction and abundance of arctic foxes in the colony, whereas seasonal influxes of geese and their eggs provided foxes with external subsidies that elevated breeding density and fox abundance above that which lemmings could support. This study highlights (1) the relative importance of migratory prey and other foods on the abundance and reproduction by local consumers and (2) how migratory animals function as vectors of nutrient transfer between distant ecosystems such as Arctic environments and wintering areas by geese thousands of kilometres to the south.     

The breeding productivity of dark-bellied brent geese and curlew sandpipers in relation to changes in the numbers of arctic foxes and lemmings on the Taimyr Peninsula, Siberia —

There were about three-year cycles in the populations of arctic foxes, and the breeding productivities of brent geese and curlew sandpipers on the Taimyr Peninsula, Russia, The populations of arctic foxes and lemmings changed in synchrony. The breeding productivities of the birds tended to be good when the arctic foxes were increasing in numbers and poor when the arctic foxes were decreasing. There was a negative relationship between arctic fox numbers (or occupied lairs) and the breeding productivity of brent geese in the following year. Although there was evidence of wide-spread synchrony In the lemming cycle across the Taimyr Peninsula, some localities showed differences, However, such sites would still have been influenced by the general pattern of fox abundance in the typical tundra zone of the Taimyr Peninsula, where most of the arctic foxes breed and from which extensive movements of foxes occur after a decline in lemming numbers. The results support a prey-switching hypothesis (also known as the alternative prey hypothesis) whereby arctic foxes, and other predators, feed largely on lemmings when these are abundant or increasing, but switch to birds when the lemming population is small or declining. The relationships between arctic foxes, lemmings and brent geese may be further influenced by snowny owls which create fox-exclusion zones around their nests, thus providing safe nesting areas for the geese.     

An avian terrestrial predator of the Arctic relies on the marine ecosystem during winter

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea‐ice between December and April. Analysis of high‐resolution satellite images of sea‐ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea‐ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea‐ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.     

Population structure of two rabies hosts relative to the known distribution of rabies virus variants in Alaska

For pathogens that infect multiple species, the distinction between reservoir hosts and spillover hosts is often difficult. In Alaska, three variants of the arctic rabies virus exist with distinct spatial distributions. We tested the hypothesis that rabies virus variant distribution corresponds to the population structure of the primary rabies hosts in Alaska, arctic foxes (Vulpes lagopus) and red foxes (Vulpes vulpes) to possibly distinguish reservoir and spillover hosts. We used mitochondrial DNA (mtDNA) sequence and nine microsatellites to assess population structure in those two species. mtDNA structure did not correspond to rabies virus variant structure in either species. Microsatellite analyses gave varying results. Bayesian clustering found two groups of arctic foxes in the coastal tundra region, but for red foxes it identified tundra and boreal types. Spatial Bayesian clustering and spatial principal components analysis identified 3 and 4 groups of arctic foxes, respectively, closely matching the distribution of rabies virus variants in the state. Red foxes, conversely, showed eight clusters comprising two regions (boreal and tundra) with much admixture. These results run contrary to previous beliefs that arctic fox show no fine‐scale spatial population structure. While we cannot rule out that the red fox is part of the maintenance host community for rabies in Alaska, the distribution of virus variants appears to be driven primarily by the arctic fox. Therefore, we show that host population genetics can be utilized to distinguish between maintenance and spillover hosts when used in conjunction with other approaches.     

Natal den selection by sympatric arctic and red foxes on Herschel Island,Yukon, Canada

In the twentieth century, red fox (Vulpes vulpes) expanded into the Canadian Arctic, where it competes with arctic fox (Vulpes lagopus) for food and shelter. Red fox dominates in physical interactions with the smaller arctic fox, but little is known about competition between them on the tundra. On Hershel Island, north Yukon, where these foxes are sympatric, we focused on natal den choice, a critical aspect of habitat selection. We tested the hypothesis that red fox displaces arctic fox from dens in prey-rich habitats. We applied an approach based on model comparisons to analyse a 10-year data set and identify factors important to den selection. Red fox selected dens in habitats that were more prey-rich in spring. When red foxes reproduced, arctic fox selected dens with good springtime access, notably many burrows unblocked by ice and snow. These provided the best refuge early in the reproductive season. In the absence of red foxes, arctic foxes selected dens offering good shelter (i.e. large isolated dens). Proximity to prey-rich habitats was consistently less important than the physical aspects of dens for arctic fox. Our study shows for the first time that red foxes in the tundra select dens associated primarily with prey-rich areas, while sympatric arctic foxes do not. These results fit a model of red fox competitively interfering with arctic fox, the first detailed study of such competition in a true arctic setting.     

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.     

The arctic fox Alopex lagopus in Fennoscandia: a victim of human-induced changes in interspecific competition and predation?

After a marked decline at the beginning of the 1900s, the arctic fox Alopex lagopus population in Fennoscandia has remained at a very low level. We suggest that the main cause for the population crash was winter starvation caused by (1) over-hunting of reindeer Rangifer tarandus populations, and thus reduced carcass availability in the mountains, and (2) increased interspecific competition for these carcasses because of increased invasion of red foxes Vulpes vulpes from lower altitudes. The failure of arctic fox populations to recover, despite increasing reindeer populations in the mid 1900s, can be explained by a concurrent strong increase in red fox numbers. Analyses of countywide hunting statistics from Norway 1891–1920 suggest that there actually was an increase in red fox numbers in the period of arctic fox decline, and that the increase in reindeer populations from the 1920s to the 1950s was accompanied by a new increase in red fox numbers. We conclude that restoring arctic fox populations most likely will require a substantial and lasting reduction of red fox populations.     

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.     

Population dynamics of tundra voles in relation to configuration of willow thickets in southern arctic tundra

The areal extent and configuration of thickets of willow shrubs are currently changing in the Arctic both as an effect of global warming and changed browsing pressure of reindeer. These changes have been predicted to impact the distribution and abundance of wildlife species relying on willow thickets as habitat. We assessed the relation between variables quantifying willow thicket configuration and population dynamics of tundra voles (Microtus oeconomus) in three riparian regions in Finnmark, northern Norway, which were subject to intense browsing by semi-domesticated reindeer. The tundra vole, which exhibits 5-year population cycles in Finnmark, is the dominant small rodent species in riparian landscape elements in southern arctic tundra. In the course of a 4-year trapping study, tundra vole populations went through the cyclic phases of increase, peak and crash, however, with distinct differences between the three regions in the population dynamics. Within regions, the occupancy pattern during the increase phase was positively related to willow thicket configuration (in particular edge density and willow height) only in the region attaining the highest abundance and occupancy. However, local abundance was not clearly related to habitat features within any regions. The lack of consistency in the response of tundra vole populations to willow thicket configuration, as well as the positive relation between the degree of thicket shredding and tundra vole habitat occupancy in one of the regions, indicates that tundra voles will not be much affected by climate or browsing induced changes in the shrubbiness of the tundra in the future.     

Red fox takeover of arctic fox breeding den: an observation from Yamal Peninsula, Russia

Here, we report from the first direct observation of a red fox (Vulpes vulpes) intrusion on an arctic fox (Vulpes lagopus) breeding den from the southern Arctic tundra of Yamal Peninsula, Russia in 2007. At the same time, as a current range retraction of the original inhabitant of the circumpolar tundra zone the arctic fox is going on, the red fox is expanding their range from the south into arctic habitats. Thus, within large parts of the northern tundra areas the two species are sympatric which gives opportunities for direct interactions including interference competition. However, direct first-hand observations of such interactions are rare, especially in the Russian Arctic. In the present study, we observed one red fox taking over an arctic fox breeding den which resulted in den abandonment by the arctic fox. On July 19, eight arctic fox pups were observed on the den before the red fox was observed on the same den July 22. The pups were never seen at the den or elsewhere after the red fox was observed on the den for as long as we stayed in the area (until August 10). Our observation supports the view that direct interference with red fox on breeding dens may contribute to the range retraction of arctic foxes from the southern limits of the Arctic tundra in Russia.     

Inheritance of litter size at birth in farmed arctic foxes (Alopex lagopus, Canidae, Carnivora)

Natural populations of the arctic fox (Alopex lagopus, Canidae, Carnivora) differ drastically in their reproductive strategy. Coastal foxes, which depend on stable food resources, produce litters of moderate size. Inland foxes feed on small rodents, whose populations are characterized by cycling fluctuation. In the years with low food supply, inland fox populations have a very low rate of reproduction. In the years with high food supply, they undergo a population explosion. To gain insight into the genetic basis of the reproductive strategy of this species, we performed complex segregation analysis of the litter size in the extended pedigree of the farmed arctic foxes involving 20,665 interrelated animals. Complex segregation analysis was performed using a mixed model assuming that the trait was under control of a major gene and a large number of additive genetic and random factors. To check the significance of any major gene effect, we used Elston-Stewart transmission probability test. Our analysis demonstrated that the inheritance of this trait can be described within the frameworks of a major gene model with recessive control of low litter size. This model was also supported by the pattern of its familial segregation and by comparison of the distributions observed in the population and that expected under our model. We suggest that a system of balanced polymorphism for litter size in the farmed population might have been established in natural populations of arctic foxes as a result of adaptation to the drastic fluctuations in prey availability.     

Distinct patterns in alpine vegetation around dens of the Arctic fox

The arctic fox Alopex lagopus excavates its dens in gravely ridges and hillocks, and creates a local environment quite distinct from the surrounding tundra or heath landscape. In northern Sweden, the vegetation of 18 dens of the arctic fox was investigated, as well as reference areas off the dens but in geologically and topographically similar locations. The species composition showed considerable differences between den and reference areas, with grasses and forbs occurring more abundantly on the dens, and evergreen dwarf-shrubs occurring more in reference areas. The effect of the foxes' activities is thought to be either through mechanical soil disturbance, or through nutrient enrichment via scats, urine, and carcasses. This was expected to result in differences in plant traits with key functional roles in resource acquisition and regeneration, when comparing dens with reference areas. We hypothesised that the community mean of specific leaf area (SLA) would differ if nutrient enrichment was the more important effect, and that seed weight, inversely proportional to seed number per ramet and hence dispersal ability, would differ if soil disturbance was the more important effect. Specific leaf area showed a significant difference, indicating nutrient enrichment to be the most important effect of the arctic fox on the vegetation on its dens. Arctic foxes act as ecosystems engineers on a small scale, maintaining niches for relatively short-lived nutrient demanding species on their dens in spite of the dominance of long-lived ericaceous dwarf-shrubs in the landscape matrix. Thus, foxes contribute to the maintenance of species richness on the landscape level.     

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.