A selection from Stockholm University publication database

• The fluctuating world of a tundra predator guild

  2019. Marianne Stoessel (et al.). Ecography 42 (3), 488-499

  Article

  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.

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• Homage to Hersteinsson and Macdonald

  2017. Bodil Elmhagen (et al.). Polar Research 36 (suppl. 1)

  Article

  Climate change can have a marked effect on the distribution and abundance of some species, as well as their interspecific interactions. In 1992, before ecological effects of anthropogenic climate change had developed into a topical research field, Hersteinsson and Macdonald published a seminal paper hypothesizing that the northern distribution limit of the red fox (Vulpes vulpes) is determined by food availability and ultimately climate, while the southern distribution limit of the Arctic fox (Vulpes lagopus) is determined by interspecific competition with the larger red fox. This hypothesis has inspired extensive research in several parts of the circumpolar distribution range of the Arctic fox. Over the past 25 years, it was shown that red foxes can exclude Arctic foxes from dens, space and food resources, and that red foxes kill and sometimes consume Arctic foxes. When the red fox increases to ecologically effective densities, it can cause Arctic fox decline, extirpation and range contraction, while conservation actions involving red fox culling can lead to Arctic fox recovery. Red fox advance in productive tundra, concurrent with Arctic fox retreat from this habitat, support the original hypothesis that climate warming will alter the geographical ranges of the species. However, recent studies show that anthropogenic subsidies also drive red fox advance, allowing red fox establishment north of its climate-imposed distribution limit. We conclude that synergies between anthropogenic subsidies and climate warming will speed up Arctic ecosystem change, allowing mobile species to establish and thrive in human-provided refugia, with potential spill-over effects in surrounding ecosystems.

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• The changing contribution of top-down and bottom-up limitation of mesopredators during 220 years of land use and climate change

  2017. Marianne Pasanen-Mortensen (et al.). Journal of Animal Ecology 86 (3), 566-576

  Article

  Apex predators may buffer bottom-up driven ecosystem change, as top-down suppression may dampen herbivore and mesopredator responses to increased resource availability. However, theory suggests that for this buffering capacity to be realized, the equilibrium abundance of apex predators must increase. This raises the question: will apex predators maintain herbivore/mesopredator limitation, if bottom-up change relaxes resource constraints? Here, we explore changes in mesopredator (red fox Vulpes vulpes) abundance over 220years in response to eradication and recovery of an apex predator (Eurasian lynx Lynx lynx), and changes in land use and climate which are linked to resource availability. A three-step approach was used. First, recent data from Finland and Sweden were modelled to estimate linear effects of lynx density, land use and winter temperature on fox density. Second, lynx density, land use and winter temperature was estimated in a 22650km(2) focal area in boreal and boreo-nemoral Sweden in the years 1830, 1920, 2010 and 2050. Third, the models and estimates were used to project historic and future fox densities in the focal area. Projected fox density was lowest in 1830 when lynx density was high, winters cold and the proportion of cropland low. Fox density peaked in 1920 due to lynx eradication, a mesopredator release boosted by favourable bottom-up changes - milder winters and cropland expansion. By 2010, lynx recolonization had reduced fox density, but it remained higher than in 1830, partly due to the bottom-up changes. Comparing 1830 to 2010, the contribution of top-down limitation decreased, while environment enrichment relaxed bottom-up limitation. Future scenarios indicated that by 2050, lynx density would have to increase by 79% to compensate for a projected climate-driven increase in fox density. We highlight that although top-down limitation in theory can buffer bottom-up change, this requires compensatory changes in apex predator abundance. Hence apex predator recolonization/recovery to historical levels would not be sufficient to compensate for widespread changes in climate and land use, which have relaxed the resource constraints for many herbivores and mesopredators. Variation in bottom-up conditions may also contribute to context dependence in apex predator effects.

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• Paws without claws? Ecological effects of large carnivores in anthropogenic landscapes

  2016. D. P. J. Kuijper (et al.). Proceedings of the Royal Society of London. Biological Sciences 283 (1841)

  Article

  Large carnivores are frequently presented as saviours of biodiversity and ecosystem functioning through their creation of trophic cascades, an idea largely based on studies coming primarily out of relatively natural landscapes. However, in large parts of the world, particularly in Europe, large carnivores live in and are returning to strongly human-modified ecosystems. At present, we lack a coherent framework to predict the effects of large carnivores in these anthropogenic landscapes. We review how human actions influence the ecological roles of large carnivores by affecting their density or behaviour or those of mesopredators or prey species. We argue that the potential for density-mediated trophic cascades in anthropogenic landscapes is limited to unproductive areas where even low carnivore numbers may impact prey densities or to the limited parts of the landscape where carnivores are allowed to reach ecologically functional densities. The potential for behaviourally mediated trophic cascades may be larger and more widespread, because even low carnivore densities affect prey behaviour. We conclude that predator-prey interactions in anthropogenic landscapes will be highly context-dependent and human actions will often attenuate the ecological effects of large carnivores. We highlight the knowledge gaps and outline a new research avenue to study the role of carnivores in anthropogenic landscapes.

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• A boreal invasion in response to climate change?

  2015. Bodil Elmhagen (et al.). Ambio 44 (1), 39-50

  Article

  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.

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• Status and Ecological Effects of the World's Largest Carnivores

  2014. William J. Ripple (et al.). Science 343 (6167), 151-+

  Article

  Large carnivores face serious threats and are experiencing massive declines in their populations and geographic ranges around the world. We highlight how these threats have affected the conservation status and ecological functioning of the 31 largest mammalian carnivores on Earth. Consistent with theory, empirical studies increasingly show that large carnivores have substantial effects on the structure and function of diverse ecosystems. Significant cascading trophic interactions, mediated by their prey or sympatric mesopredators, arise when some of these carnivores are extirpated from or repatriated to ecosystems. Unexpected effects of trophic cascades on various taxa and processes include changes to bird, mammal, invertebrate, and herpetofauna abundance or richness; subsidies to scavengers; altered disease dynamics; carbon sequestration; modified stream morphology; and crop damage. Promoting tolerance and coexistence with large carnivores is a crucial societal challenge that will ultimately determine the fate of Earth's largest carnivores and all that depends upon them, including humans.

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• Where lynx prevail, foxes will fail - limitation of a mesopredator in Eurasia

  2013. Marianne Pasanen-Mortensen, Markku Pyykonen, Bodil Elmhagen. Global Ecology and Biogeography 22 (7), 868-877

  Article

  Aim Climate change and loss of apex predators can affect ecosystem structure and function through modified limitation processes. We investigated, on a continental scale, whether mesopredator abundance is limited from the top down by large predators, as predicted by the mesopredator release hypothesis, or by bottom-up factors. The mesopredator in focus is the red fox Vulpes vulpes, a key predator in many ecosystems due to its strong effects on prey abundance. Location Europe and northern Asia. Methods Data on red fox density were compiled from published papers and reports. For each site, we collated presence-absence data on large carnivores (Lynx lynx, Canis lupus, Canisaureus) and remote sensing data for factors potentially related to bottom-up limitation (winter severity, summer temperature, human density, primary productivity, tree cover). The data were analysed through structural equation modelling. Results The presence of lynx had a direct negative effect on red foxes, suppressing fox abundance. Also winter severity had a negative effect on red fox abundance, and in Eurasia as a whole this effect was partially mediated through lynx. Within the lynx distribution range, winter severity was the only bottom-up factor significantly affecting red fox abundance. Outside the lynx distribution range, primary productivity, summer temperature and human density had a positive effect on red fox abundance. Main conclusions Our results show that apex predators can limit mesopredator abundance on a continental scale, thus supporting the mesopredator release hypothesis. Winter severity also affected red fox abundance, partially due to an interaction between lynx and winter conditions. On the continental scale a complex network of processes operates with varying effects depending on mediation processes. Our results imply that apex predators can have an important effect on ecosystem structure by limiting mesopredator abundance, and we suggest that apex predators may dampen increases in mesopredator abundance driven by global warming.

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• Changes in vole and lemming fluctuations in northern Sweden 1960-2008 revealed by fox dynamics

  2011. Bodil Elmhagen (et al.). Annales Zoologici Fennici 48 (3), 167-179

  Article

  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 spatio-temporal 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.

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• Top predators, mesopredators and their prey

  2010. Bodil Elmhagen (et al.). Journal of Animal Ecology 79 (4), 785-794

  Article

  1. The Mesopredator Release Hypothesis (MRH) suggests that top predator suppression of mesopredators is a key ecosystem function with cascading impacts on herbivore prey, but it remains to be shown that this top-down cascade impacts the large-scale structure of ecosystems.

  2. The Exploitation Ecosystems Hypothesis (EEH) predicts that regional ecosystem structures are determined by top-down exploitation and bottom-up productivity. In contrast to MRH, EEH assumes that interference among predators has a negligible impact on the structure of ecosystems with three trophic levels.

  3. We use the recolonisation of a top predator in a three-level boreal ecosystem as a natural experiment to test if large-scale biomass distributions and population trends support MRH. Inspired by EEH, we also test if top-down interference and bottom-up productivity impact regional ecosystem structures.

  4. We use data from the Finnish Wildlife Triangle Scheme which has monitored top predator (lynx Lynx lynx), mesopredator (red fox Vulpes vulpes) and prey (mountain hare Lepus timidus) abundance for 17 years in a 200 000 km² study area which covers a distinct productivity gradient.

  5. Fox biomass was lower than expected from productivity where lynx biomass was high, while hare biomass was lower than expected from productivity where fox biomass was high. Hence, where interference controlled fox abundance, lynx had an indirect positive impact on hare abundance as predicted by MRH. The rates of change indicated that lynx expansion gradually suppressed fox biomass.

  6. Lynx status caused shifts between ecosystem structures. In the “interference ecosystem”, lynx and hare biomass increased with productivity whilst fox biomass did not. In the “mesopredator release ecosystem”, fox biomass increased with productivity but hare biomass did not. Thus, biomass controlled top-down did not respond to changes in productivity. This fulfils a critical prediction of EEH.

  7. We conclude that the cascade involving top predators, mesopredators and their prey can determine large-scale biomass distribution patterns and regional ecosystem structures. Hence, interference within trophic levels has to be taken into account to understand how terrestrial ecosystem structures are shaped.

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• Trophic control of mesopredators in terrestrial ecosystems: top-down or bottom-up?

  2007. Bodil Elmhagen, Steve Rushton. Ecology Letters 10 (3), 197-206

  Article

  It has been argued that widespread extinctions of top predators have changed terrestrial ecosystem structures through mesopredator release, where increased abundances of medium-sized predators have detrimental effects on prey communities. This top-down concept has received much attention within conservation biology, but few studies have demonstrated the phenomenon. The concept has been criticized since alternative explanations involving bottom-up impacts from bioclimatic effects on ecosystem productivity and from anthropogenic habitat change are rarely considered. We analyse the response of a mesopredator (the red fox) to declines in top predators (wolf and Eurasian lynx) and agricultural expansion over 90 years in Sweden, taking bioclimatic effects into account. We show a top-down mesopredator release effect, but ecosystem productivity determined its strength. The impacts of agricultural activity were mediated by their effects on top predator populations. Thus, both top-down and bottom-up processes need to be understood for effective preservation of biodiversity in anthropogenically transformed ecosystems.

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Show all publications by Bodil Sofi Elmhagen at Stockholm University