Eva Fuglei

Trophic Interaction Cycles in Tundra Ecosystems and the Impact of Climate Change

Abstract While population cycles are geographically widespread, it is on arctic tundra that such cycles appear to be most influential for the functioning of the whole ecosystem. We give an overview of tundra species that exhibit population cycles and describe what are currently believed to be the causal mechanisms. Population cycles most likely originate from trophic interactions within the plant-based tundra food web, where lemmings, either as prey for carnivores or as consumers of plants, play the key role. The predominance of trophic interaction cycles at northern latitudes is ultimately related to climate, and such cycles should therefore be vulnerable to climate change. Recent evidence indicates that changes have already taken place in the dynamics of some key herbivores and their predators, consistent with the expected impacts of climate change. There is a strong need for large-scale integrated monitoring and research efforts to further document such changes and their ecosystem consequences.

Climate events synchronize the dynamics of a resident vertebrate community in the high Arctic.

All Together Now Environmental drivers, such as extreme weather events, impact population dynamics and can synchronize such dynamics across populations within a species. Given that many species depend on similar resources, such events might also be expected to synchronize dynamics across species, but the complexity of multispecies communities makes it difficult to reveal potential drivers in common. Hansen et al. (p. 313) took advantage of the simplicity of the year-round community on the high-arctic island of Spitsbergen to test for the presence of synchrony. Population fluctuations were synchronized across the three herbivore species (Svalbard reindeer, Svalbard rock ptarmigan, and sibling vole) and the single resident predator, the arctic fox, was in lagged synchrony. The driver of these fluctuations appears to be extreme winter rain-on-snow events that reduce the availability of winter forage due to ice cover. Reindeer, ptarmigan, vole, and fox populations on Svalbard respond together to extreme weather. Recently accumulated evidence has documented a climate impact on the demography and dynamics of single species, yet the impact at the community level is poorly understood. Here, we show that in Svalbard in the high Arctic, extreme weather events synchronize population fluctuations across an entire community of resident vertebrate herbivores and cause lagged correlations with the secondary consumer, the arctic fox. This synchronization is mainly driven by heavy rain on snow that encapsulates the vegetation in ice and blocks winter forage availability for herbivores. Thus, indirect and bottom-up climate forcing drives the population dynamics across all overwintering vertebrates. Icing is predicted to become more frequent in the circumpolar Arctic and may therefore strongly affect terrestrial ecosystem characteristics.

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.

Seasonal trends in body mass, food intake and resting metabolic rate, and induction of metabolic depression in arctic foxes (Alopex lagopus) at Svalbard.

Abstract Post-absorptive resting metabolic rates (RMRs), body mass and ad libitum food intake were recorded on an annual cycle in captive arctic foxes (Alopex lagopus) at Svalbard. During the light season in May and in the dark period in November, RMR during starvation and subsequent re-feeding were also measured. In contrast to earlier findings, the present study indicated a seasonal trend in post-absorptive RMR (in W · kg−1 and W · kg−0.75). The values in the light summer were 15% and 11% higher than the values in the dark winter, suggesting a physiological adaptation aiding energy conservation during winter in arctic foxes. Body mass and ad libitum food intake varied inversely through the year. A significant reduction in RMR (in W and W · kg−0.75) with starvation (metabolic depression) was recorded both in May and November, indicating an adaptation to starvation in arctic foxes. The lack of metabolic depression during a period of starvation that was concomitant with extremely cold ambient temperatures in November 1994 indicates that metabolic responses to starvation may be masked by thermoregulatory needs. At very low ambient temperatures, arctic foxes may require increased heat production which cannot be achieved via below-average rates of metabolism.

Echinococcus multilocularis on Svalbard: introduction of an intermediate host has enabled the local life-cycle

The taeniid tapeworm Echinococcus multilocularis is here reported for the first time at the Svalbard Archipelago in the Norwegian Arctic. This new finding is interesting because the establishment of E. multilocularis is due to a recent anthropogenic introduction of its intermediate host--the sibling vole Microtus rossiaemeridionalis at Svalbard. The parasite itself has probably become naturally transferred to Svalbard due to migratory movements of its final host--the arctic fox Alopex lagopus between source areas for E. multilocularis in Siberia and Svalbard. We report macroscopically determined prevalence of E. multilocularis from a sample of 224 voles trapped in August in 1999 and 2000. The prevalence was among the highest ever recorded in intermediate hosts and was dependent on age and sex of the hosts approaching 100% in overwintered males. The high prevalence and the simplicity of the vole-arctic fox-E. multilocularis system at Svalbard makes it an eminent model system for further epidemiological studies.

Sea ice occurrence predicts genetic isolation in the Arctic fox

Unlike Oceanic islands, the islands of the Arctic Sea are not completely isolated from migration by terrestrial vertebrates. The pack ice connects many Arctic Sea islands to the mainland during winter months. The Arctic fox (Alopex lagopus), which has a circumpolar distribution, populates numerous islands in the Arctic Sea. In this study, we used genetic data from 20 different populations, spanning the entire distribution of the Arctic fox, to identify barriers to dispersal. Specifically, we considered geographical distance, occurrence of sea ice, winter temperature, ecotype, and the presence of red fox and polar bear as nonexclusive factors that influence the dispersal behaviour of individuals. Using distance‐based redundancy analysis and the BIOENV procedure, we showed that occurrence of sea ice is the key predictor and explained 40–60% of the genetic distance among populations. In addition, our analysis identified the Commander and Pribilof Islands Arctic populations as genetically unique suggesting they deserve special attention from a conservation perspective.

Response of an arctic predator guild to collapsing lemming cycles

Alpine and arctic lemming populations appear to be highly sensitive to climate change, and when faced with warmer and shorter winters, their well-known high-amplitude population cycles may collapse. Being keystone species in tundra ecosystems, changed lemming dynamics may convey significant knock-on effects on trophically linked species. Here, we analyse long-term (1988–2010), community-wide monitoring data from two sites in high-arctic Greenland and document how a collapse in collared lemming cyclicity affects the population dynamics of the predator guild. Dramatic changes were observed in two highly specialized lemming predators: snowy owl and stoat. Following the lemming cycle collapse, snowy owl fledgling production declined by 98 per cent, and there was indication of a severe population decline of stoats at one site. The less specialized long-tailed skua and the generalist arctic fox were more loosely coupled to the lemming dynamics. Still, the lemming collapse had noticeable effects on their reproductive performance. Predator responses differed somewhat between sites in all species and could arise from site-specific differences in lemming dynamics, intra-guild interactions or subsidies from other resources. Nevertheless, population extinctions and community restructuring of this arctic endemic predator guild are likely if the lemming dynamics are maintained at the current non-cyclic, low-density state.

Congruent responses to weather variability in high arctic herbivores

Assessing the role of weather in the dynamics of wildlife populations is a pressing task in the face of rapid environmental change. Rodents and ruminants are abundant herbivore species in most Arctic ecosystems, many of which are experiencing particularly rapid climate change. Their different life-history characteristics, with the exception of their trophic position, suggest that they should show different responses to environmental variation. Here we show that the only mammalian herbivores on the Arctic islands of Svalbard, reindeer (Rangifer tarandus) and sibling voles (Microtus levis), exhibit strong synchrony in population parameters. This synchrony is due to rain-on-snow events that cause ground ice and demonstrates that climate impacts can be similarly integrated and expressed in species with highly contrasting life histories. The finding suggests that responses of wildlife populations to climate variability and change might be more consistent in Polar regions than elsewhere owing to the strength of the climate impact and the simplicity of the ecosystem.

Direct high-resolution genotyping of Toxoplasma gondii in arctic foxes (Vulpes lagopus) in the remote arctic Svalbard archipelago reveals widespread clonal Type II lineage

Characterization of Toxoplasma gondii genotypes in hosts living in remote, isolated regions is important for elucidating the population structure and transmission mode of this parasite. Herein, we report the results of direct genotyping of T. gondii in brain tissue of arctic foxes (Vulpes lagopus) from the remote, virtually cat-free, high arctic islands of Svalbard. DNA extracts from brains of 167 seropositive arctic foxes (including four cases of fatal toxoplasmosis) and 11 seronegative arctic foxes were genotyped at 10 loci (SAG1, SAG2, SAG3, BTUB, GRA6, L358, c22-8, c29-2, PK1, and Apico) using the polymerase chain reaction-restriction fragment length polymorphism method. Of the 167 samples from seropositive foxes (including toxoplasmosis cases), 31 were genotyped at all 10 loci and 24 were genotyped at four to nine loci. To ensure confidence in T. gondii strain genotyping, samples for which less than four loci were genotyped were not considered positive. None of the 11 samples from seronegative foxes was positive for the 10 markers. Of the 55 samples that genotyped positively, 46 were of the Type II strain, 7 were of the Type III strain, and 2 were of atypical T. gondii strains. Five representative samples of the three genotypes were sequenced at loci SAG2, SAG3, GRA6, PK1, and UPRT-1. The DNA sequences confirmed the genotyping results. This study shows that the archetype Type II T. gondii strain, which is most widely distributed in North America and Europe, also predominates in arctic foxes on the Svalbard archipelago. This suggests that the T. gondii at this location originate from continental Europe and that transmission may be mediated by migrating birds. This study highlights the significance of long-distance transport of T. gondii and demonstrates that high-resolution genotyping protocols are useful for direct genetic studies of T. gondii when isolation of live parasites is infeasible.

Intrapopulation Variability Shaping Isotope Discrimination and Turnover: Experimental Evidence in Arctic Foxes

Background Tissue-specific stable isotope signatures can provide insights into the trophic ecology of consumers and their roles in food webs. Two parameters are central for making valid inferences based on stable isotopes, isotopic discrimination (difference in isotopic ratio between consumer and its diet) and turnover time (renewal process of molecules in a given tissue usually measured when half of the tissue composition has changed). We investigated simultaneously the effects of age, sex, and diet types on the variation of discrimination and half-life in nitrogen and carbon stable isotopes (δ15N and δ13C, respectively) in five tissues (blood cells, plasma, muscle, liver, nail, and hair) of a top predator, the arctic fox Vulpes lagopus. Methodology/Principal Findings We fed 40 farmed foxes (equal numbers of adults and yearlings of both sexes) with diet capturing the range of resources used by their wild counterparts. We found that, for a single species, six tissues, and three diet types, the range of discrimination values can be almost as large as what is known at the scale of the whole mammalian or avian class. Discrimination varied depending on sex, age, tissue, and diet types, ranging from 0.3‰ to 5.3‰ (mean  = 2.6‰) for δ15N and from 0.2‰ to 2.9‰ (mean  = 0.9‰) for δ13C. We also found an impact of population structure on δ15N half-life in blood cells. Varying across individuals, δ15N half-life in plasma (6 to 10 days) was also shorter than for δ13C (14 to 22 days), though δ15N and δ13C half-lives are usually considered as equal. Conclusion/Significance Overall, our multi-factorial experiment revealed that at least six levels of isotopic variations could co-occur in the same population. Our experimental analysis provides a framework for quantifying multiple sources of variation in isotopic discrimination and half-life that needs to be taken into account when designing and analysing ecological field studies.