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

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 220 years 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 22 650 km2 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.

Simulated moose (Alces alces L.) browsing increases accumulation of secondary metabolites in bilberry (Vaccinium myrtillus L.) along gradients of habitat productivity and solar radiation.

We have addressed the impact of moose (Alces alces L.) on accumulation of secondary metabolites, lignin, and nitrogen in bilberry (Vaccinium myrtillus L.) along gradients of habitat productivity and solar radiation. The study was conducted within a long-term research project on direct and indirect impacts of moose on the ecosystem. In the experiment, browsing, defecation, and urination corresponding to four different moose densities were simulated for eight years before bilberry tissue was collected and analyzed. Some quantitatively dominant flavonoids were affected by the simulated moose browsing and by habitat productivity and light. The content of flavonoids increased with increasing moose density and light, and decreased with increasing habitat productivity. The higher concentration of secondary metabolites in bilberry from nutrient-poor sites may have resulted from the increased photosynthesis relative to growth, which facilitated secondary metabolism. The higher concentration of secondary metabolites in plants subjected to simulated moose- herbivory might have been caused in part by loss of biomass. In addition, in areas with high biomass loss, i.e., high moose density, a more open canopy was created and more solar radiation could have induced secondary metabolism.

Depression of belowground respiration rates at simulated high moose population densities in boreal forests.

Large herbivores can affect the carbon cycle in boreal forests by changing productivity and plant species composition, which in turn could ultimately alter litter production, nutrient cycling, and the partitioning between aboveground and belowground allocation of carbon. Here we experimentally tested how moose (Alces alces) at different simulated population densities affected belowground respiration rates (estimated as CO2 flux) in young boreal forest stands situated along a site productivity gradient. At high simulated population density, moose browsing considerably depressed belowground respiration rates (24-56% below that of no-moose controls) except during June, where the difference only was 10%. Moose browsing depressed belowground respiration the most on low-productivity sites. Soil moisture and temperature did not affect respiration rates. Impact of moose on belowground respiration was closely linked to litter production and followed Michaelis-Menten dynamics. The main mechanism by which moose decrease belowground respiration rates is likely their effect on photosynthetic biomass (especially decreased productivity of deciduous trees) and total litter production. An increased productivity of deciduous trees along the site productivity gradient causes an unequal effect of moose along the same gradient. The rapid growth of deciduous trees may offer higher resilience against negative effects of moose browsing on litter production and photosynthate allocation to roots.

Fear or food - abundance of red fox in relation to occurrence of lynx and wolf.

Apex predators may affect mesopredators through intraguild predation and/or supply of carrion from their prey, causing a trade-off between avoidance and attractiveness. We used wildlife triangle snow-tracking data to investigate the abundance of red fox (Vulpes vulpes) in relation to lynx (Lynx lynx) and wolf (Canis lupus) occurrence as well as land composition and vole (Microtus spp.) density. Data from the Swedish wolf-monitoring system and VHF/GPS-collared wolves were used to study the effect of wolf pack size and time since wolf territory establishment on fox abundance. Bottom-up processes were more influential than top-down effects as the proportion of arable land was the key indicator of fox abundance at the landscape level. At this spatial scale, there was no effect of wolf abundance on fox abundance, whereas lynx abundance had a positive effect. In contrast, at the wolf territory level there was a negative effect of wolves on fox abundance when including detailed information of pack size and time since territory establishment, whereas there was no effect of lynx abundance. This study shows that different apex predator species may affect mesopredator abundance in different ways and that the results may be dependent on the spatiotemporal scale and resolution of the data.

Response of moose hunters to predation following wolf return in Sweden.

BACKGROUND: Predation and hunter harvest constitute the main mortality factors affecting the size and dynamics of many exploited populations. The re-colonization by wolves (Canis lupus) of the Scandinavian Peninsula may therefore substantially reduce hunter harvest of moose (Alces alces), the main prey of wolves. METHODOLOGY/PRINCIPAL FINDINGS: We examined possible effects of wolf presence on hunter harvest in areas where we had data before and after wolf establishment (n = 25), and in additional areas that had been continuously exposed to wolf predation during at least ten years (n = 43). There was a general reduction in the total number of moose harvested (n = 31,827) during the ten year study period in all areas irrespective of presence of wolves or not. However, the reduction in hunter harvest was stronger within wolf territories compared to control areas without wolves. The reduction in harvest was larger in small (500-800 km2) compared to large (1,200-1,800 km2) wolf territories. In areas with newly established wolf territories moose management appeared to be adaptive with regard to both managers (hunting quotas) and to hunters (actual harvest). In these areas an instant reduction in moose harvest over-compensated the estimated number of moose killed annually by wolves and the composition of the hunted animals changed towards a lower proportion of adult females. CONCLUSIONS/SIGNIFICANCE: We show that the re-colonization of wolves may result in an almost instant functional response by another large predator-humans-that reduced the potential for a direct numerical effect on the density of wolves' main prey, the moose. Because most of the worlds' habitat that will be available for future colonization by large predators are likely to be strongly influenced by humans, human behavioural responses may constitute a key trait that govern the impact of large predators on their prey.