Peter Hellström

Carnivore conservation in practice: replicated management actions on a large spatial scale

Summary 1. More than a quarter of the world’s carnivores are threatened, often due to multiple and complex causes. Considerable research efforts are devoted to resolving the mechanisms behind these threats in order to provide a basis for relevant conservation actions. However, even when the underlying mechanisms are known, specific actions aimed at direct support for carnivores are difficult to implement and evaluate at efficient spatial and temporal scales. 2. We report on a 30-year inventory of the critically endangered Fennoscandian arctic fox Vulpes lagopus L., including yearly surveys of 600 fox dens covering 21 000 km 2 . These surveys showed that the population was close to extinction in 2000, with 40–60 adult animals left. However, the population subsequently showed a fourfold increase in size. 3. During this time period, conservation actions through supplementary feeding and predator removal were implemented in several regions across Scandinavia, encompassing 79% of the area. To evaluate these actions, we examined the effect of supplemental winter feeding and red fox control applied at different intensities in 10 regions. A path analysis indicated that 47% of the explained variation in population productivity could be attributed to lemming abundance, whereas winter feeding had a 29% effect and red fox control a 20% effect. 4. This confirms that arctic foxes are highly dependent on lemming population fluctuations but also shows that red foxes severely impact the viability of arctic foxes. This study also highlights the importance of implementing conservation actions on extensive spatial and temporal scales, with geographically dispersed actions to scientifically evaluate the effects. We note that population recovery was only seen in regions with a high intensity of management actions. 5. Synthesis and applications. The present study demonstrates that carnivore population declines may be reversed through extensive actions that target specific threats. Fennoscandian arctic fox is still endangered, due to low population connectivity and expected climate impacts on the distribution and dynamics of lemmings and red foxes. Climate warming is expected to contribute to both more irregular lemming dynamics and red fox appearance in tundra areas; however, the effects of climate change can be mitigated through intensive management actions such as supplemental feeding and red fox control.

A boreal invasion in response to climate change? Range shifts and community effects in the borderland between forest and tundra

It has been hypothesized that climate warming will allow southern species to advance north and invade northern ecosystems. We review the changes in the Swedish mammal and bird community in boreal forest and alpine tundra since the nineteenth century, as well as suggested drivers of change. Observed changes include (1) range expansion and increased abundance in southern birds, ungulates, and carnivores; (2) range contraction and decline in northern birds and carnivores; and (3) abundance decline or periodically disrupted dynamics in cyclic populations of small and medium-sized mammals and birds. The first warm spell, 1930–1960, stands out as a period of substantial faunal change. However, in addition to climate warming, suggested drivers of change include land use and other anthropogenic factors. We hypothesize all these drivers interacted, primarily favoring southern generalists. Future research should aim to distinguish between effects of climate and land-use change in boreal and tundra ecosystems.

Changes in Vole and Lemming Fluctuations in Northern Sweden 1960–2008 Revealed by Fox Dynamics

Cyclic dynamics with extensive spatial synchrony has long been regarded as characteristic of key herbivores at high latitudes. This contrasts to recent reports of fading cycles in arvicoline rodents in boreal and alpine Fennoscandia. We investigate the spatiotemporal dynamics of boreal red fox and alpine arctic fox in Sweden as a proxy for the dynamics of their main prey, voles and Norwegian lemming, respectively. We analyse data from five decades, 1960–2008, with wavelets and autocorrelation approaches. Cyclic dynamics were identified with at least one method in all populations (arctic fox n = 3, red fox n = 6). The dynamics were synchronous between populations, or coupled with a 1-yr lag, in 8 of 13 pairwise comparisons. Importantly though, the dynamics were heterogeneous in space and time. All analytical approaches identified fading cycles in the three arctic fox populations and two northern red fox populations. At least one method identified similar patterns in three southern red fox populations. Red fox dynamics were cyclic in the 1970s primarily, while arctic fox dynamics was cyclic until the late 1980s or early 1990s. When cyclic, 4-yr cycles dominated in arctic fox and northern red fox, whilst 3–4-yr cycles was found in southern red foxes. Significant cyclic regimes reappeared in the 1990s or 2000s in two red fox populations and one arctic fox population. Cycles and regionally coupled dynamics appeared associated in northern arctic and red foxes. This study supports accumulating evidence which suggests that cyclic and synchronous patterns in the dynamics of lemmings and voles are nonstationary in space and time. Furthermore, the similar patterns of change in both fox species indicate that persistence of cycles is governed by similar mechanisms in lemmings and voles.

Functional responses of the rough-legged buzzard in a multi-prey system

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.

Effects of Trap Density and Duration on Vole Abundance Indices

This study aims to investigate if patterns of immigration by voles into removal plots on the third day of trapping are evident in the grey-sided vole, and if altering the number of traps at each station will result in increased precision of the vole abundance estimate. Traps were placed using the small quadrat method, with one, three, or five traps placed at each corner. Traps were checked twice a day for five days. Mixed-effect models were used to investigate the relationship between the number of traps and the length of time the traps were out on the abundance index. There was no difference between having three or five traps. Having one trap resulted in an inflated estimate. Five traps had the highest number of successful trapping events, reducing the number of zeros in the data set and leaving fewer individuals unaccounted. There was a peak in catches on the third day, driven by younger individuals and by males. These are suspected immigrants that are exploiting the territories left by individuals trapped in the first two days, suggesting this is not a closed system.

Network topology of stable isotope interactions in a sub-arctic raptor guild

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 analyzes 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.