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In territorial species, observed density dependence is often manifest in lowered reproductive output at high population density where individuals have fewer resources or are forced to inhabit low-quality territories. The Arctic fox (Vulpes lagopus) in Iceland is territorial throughout the year and feeds mostly on birds, since lemmings are absent from the country. Thus, the population does not exhibit short-term population cycles that are evident in most of the species’ geographical range. The population has, however, gone through a major long-term fluctuation in population size. Because of the stability in hunting effort and reliable hunting records since 1958, the total number of adult foxes killed annually can be used as an index of population size (N t). An index of carrying capacity (K) from population growth data for five separate time blocks during 1958–2007 revealed considerable variation in K and allowed a novel definition of population density in terms of K, or N t/K. Correlation analysis suggested that the reproductive rate was largely determined by the proportion of territorial foxes in the population. Variation in litter size and cub mortality was, on the other hand, related to climatic variation. Thus, Arctic foxes in Iceland engage in typical contest competition but can adapt their territory sizes in response to both temporal and spatial variation in carrying capacity, resulting in surprisingly little variation in litter size.

Predation is an ecologically important process, and intra-guild interactions may substantially influence the ecological effects of predator species. Despite a rapid expansion in the use of mathematical graph theory to describe trophic relations, network approaches have rarely been used to study interactions within predator assemblages. Assemblages of diurnal raptors are subject to substantial intra and interspecific competition. Here we used the novel approach of applying analyses based on network topology to species-specific data on the stable isotopes 13C and 15N in feathers to evaluate patterns of relative resource utilization within a guild of diurnal raptors in northern Sweden. Our guild consisted of the golden eagle (Aquila chrysaetos), the gyrfalcon (Falco rusticolus), the peregrine falcon (Falco peregrinus) and the rough-legged buzzard (Buteo lagopus). We found a modular trophic interaction structure within the guild, but the interactions were less nested than expected by chance. These results suggest low redundancy and hence a strong ecological importance of individual species. Our data also suggested that species were less connected through intra-guild interactions than expected by chance. We interpret our results as a convergence on specific isotope niches, and that body size and different hunting behaviour may mediate competition within these niches. We finally highlight that generalist predators could be ecologically important by linking specialist predator species with disparate dietary niches.

Assessments of temporal variation in diets are important for our understanding of the ecology of many vertebrates. Ratios of naturally occurring stable isotopes in animal tissues are a combination of the source elements and tissue specific fractionation processes, and can thus reveal dietary information. We review three different approaches that have been used to resolve temporal diet variation through analysis of stable isotopes. The most straightforward approach is to compare samples from the same type of tissue that has been sampled over time. This approach is suited to address either long or short-term dietary variation, depending on sample regime and which tissue that is sampled. Second, one can compare tissues with different metabolic rates. Since the elements in a given tissue have been assimilating during time spans specific to its metabolic rate, tissues with different metabolic rates will reflect dietary records over different periods. Third, comparisons of sections from tissues with progressive growth, such as hair, feathers, claws and teeth, will reveal temporal variation since these tissues will retain isotopic values in a chronological order. These latter two approaches are mainly suited to address questions regarding intermediate and short-term dietary variation. Knowledge of tissue specific metabolic rates, which determine the molecular turnover for a specific tissue, is of central importance for all these comparisons. Estimates of isotopic fractionation between source and measured target are important if specific hypotheses regarding the source elements are addressed. Estimates of isotopic fractionation, or at least of differences in fractionation between tissues, are necessary if different tissues are compared. We urge for more laboratory experiments aimed at improving our understanding of differential assimilation of dietary components, isotopic fractionation and metabolic routing. We further encourage more studies on reptiles and amphibians, and generally more studies utilizing multiple tissues with different turnover rates.

Demographic bottlenecks can result in a loss of genetic variation due to the bottleneck effect and subsequent genetic drift. The arctic fox population in Scandinavia went through a severe demographic bottleneck in the early 20th century, and is today classified as critically endangered. In this study, we investigated the pre-bottleneck genetic variation in Scandinavia and compared it to modern samples from Scandinavia and North Russia. Variation in the mtDNA control region and five microsatellite loci was examined through ancient DNA analysis on museum specimens. The microsatellite data from the museum specimens was further used to simulate the expected effect of the bottleneck. The arctic foxes in Scandinavia have lost approximately 25% of the microsatellite alleles and four out of seven mtDNA haplotypes. The results also suggest that the genetic differentiation between North Russia and Scandinavia has doubled over the last 100 years. However, the level of heterozygosity was significantly higher than expected from the simulations. This highlights both the advantage of using museum specimens and the importance of generating specific predictions in conservation genetics.

1. The number of breeding dens and litter sizes of arctic foxes Alopex lagopus were recorded and the diet of the foxes was analysed during a ship-based expedition to 17 sites along the Siberian north coast. At the same time the cyclic dynamics of coexisting lemming species were examined. 2. The diet of arctic foxes was dominated by the Siberian lemming Lemmus sibiricus (on one site the Norwegian lemming L. lemmus), followed by the collared lemming Dicrostonyx torquatus. 3. The examined Lemmus sibiricus populations were in different phases of the lemming cycle as determined by age profiles and population densities. 4. The numerical response of arctic foxes to varying densities of Lemmus had a time lag of 1 year, producing a pattern of limit cycles in lemming-arctic fox interactions. Arctic fox litter sizes showed no time lag, but a linear relation to Lemmus densities. We found no evidence for a numerical response to population density changes in Dicrostonyx. 5. The functional or dietary response of arctic foxes followed a type II curve for Lemmus, but a type III response curve for Dicrostonyx. 6. Arctic foxes act as resident specialist for Lemmus and may increase the amplitude and period of their population cycles. For Dicrostonyx, on the other hand, arctic foxes act as generalists which suggests a capacity to dampen oscillations.

The functional response is a key element of predator–prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.

In order to discuss dietary change over time by the use of stable isotopes, it is necessary to sort out the underlying processes in isotopic variation. Together with the dietary signal other processes have been investigated, namely metabolic processes, collagen turnover and physical growth. However, growth and collagen turnover time have so far been neglected in dietary reconstruction based on stable isotopes. An earlier study suggested that cave bears (Ursus spelaeus) probably gave birth to cubs during dormancy. We provide an estimate of the effect on stable isotopes of growth and metabolism and discuss collagen turnover in a population of cave bears. Based on a quantitative model, we hypothesized that bear cubs lactated their mothers during their first and second winters, but were fed solid food together with lactation during their first summer. This demonstrates the need to include physical growth, metabolism and collagen turnover in dietary reconstruction. Whereas the effects of diet and metabolism are due to fractionation, growth and collagen turnover are dilution processes.

The Arctic fox (Vulpes lagopus L.) population on the Kola Peninsula occupies an intermediate, and potentially connecting, position between foxes living on the Scandinavian Peninsula and populations further east in Russia, but very little is known about the status of this population. Here we summarize data from the literature, forgotten archival sources about research in the first half of the twentieth century, and the results of several independent expeditions undertaken over the past two decades. These materials include data on fur harvesting, incomplete monitoring data from official winter track counts of game animals, local knowledge, and our own observations. Our research revealed the extremely poor state of the Arctic fox population on the Kola Peninsula. According to our estimates, the current population is likely isolated and consists of no more than a few dozen adults. The fur return data, together with long-term data on small rodent abundance, suggest that irregular and fading out lemming cycles were a major driver of the Arctic fox population decline. The thorough research from the 1930s contrasts strongly with the lack of interest in studying and monitoring the population in recent decades, which is not even listed as a threatened species in the regional Red Data Book. In fact, the work performed here filled a more than a half-century gap in the study of the population and allowed us to determine the urgent need to resume research and immediately take active measures to protect and promote the recovery of the species in the region.

The Swedish Tundra Northwest Expedition of 1999 visited 17 sites throughout the Canadian Arctic. At 12 sites that were intensively sampled we estimated the standing crop of plants and the densities of herbivores and predators with an array of trapping, visual surveys, and faecal-pellet transects. We developed a trophic-balance model using ECOPATH to integrate these observations and determine the fate of primary and secondary production in these tundra ecosystems, which spanned an 8-fold range of standing crop of plants. We estimated that about 13% of net primary production was consumed by herbivores, while over 70% of small-herbivore production was estimated to flow to predators. Only 9% of large-herbivore production was consumed by predators. Organization of Canadian Arctic ecosystems appears to be more top-down than bottom-up. Net primary production does not seem to be herbivore-limited at any site. This is the first attempt to integrate trophic dynamics over the entire Canadian Arctic.