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Perceived predation risk varies in space and time. Foraging in this landscape of fear alters forager-resource interactions via cascading nonconsumptive effects. Estimating these indirect effects is difficult in natural systems. Here, we applied a novel measure to quantify the diversity at giving-up density that allows to test how spatial variation in perceived predation risk modifies the diversity of multispecies resources at local and regional spatial levels. Furthermore, we evaluated whether the nonconsumptive effects on resource species diversity can be explained by the preferences of foragers for specific functional traits and by the forager species richness. We exposed rodents of a natural community to artificial food patches, each containing an initial multispecies resource community of eight species (10 items each) mixed in sand. We sampled 35 landscapes, each containing seven patches in a spatial array, to disentangle effects at local (patch) and landscape levels. We used vegetation height as a proxy for perceived predation risk. After a period of three nights, we counted how many and which resource species were left in each patch to measure giving-up density and resource diversity at the local level (alpha diversity) and the regional level (gamma diversity and beta diversity). Furthermore, we used wildlife cameras to identify foragers and assess their species richness. With increasing vegetation height, i.e., decreasing perceived predation risk, giving-up density, and local alpha and regional gamma diversity decreased, and patches became less similar within a landscape (beta diversity increased). Foragers consumed more of the bigger and most caloric resources. The higher the forager species richness, the lower the giving-up density, and alpha and gamma diversity. Overall, spatial variation of perceived predation risk of foragers had measurable cascading effects on local and regional resource species biodiversity, independent of the forager species. Thus, nonconsumptive predation effects modify forager-resource interactions and might act as an equalizing mechanism for species coexistence.
Bioturbation contributes to soil formation and ecosystem functioning. With respect to the active transport of matter by voles, bioturbation may be considered as a very dynamic process among those shaping soil formation and biogeochemistry. The present study aimed at characterizing and quantifying the effects of bioturbation by voles on soil water relations and carbon and nitrogen stocks. Bioturbation effects were examined based on a field set up in a luvic arenosol comprising of eight 50 x 50 m enclosures with greatly different numbers of common vole (Microtus arvalis L., ca. 35-150 individuals ha(-1) mth(-1)). Eleven key soil variables were analyzed: bulk density, infiltration rate, saturated hydraulic conductivity, water holding capacity, contents of soil organic carbon (SOC) and total nitrogen (N), CO2 emission potential, C/N ratio, the stable isotopic signatures of C-13 and N-15, and pH. The highest vole densities were hypothesized to cause significant changes in some variables within 21 months. Results showed that land history had still a major influence, as eight key variables displayed an additional or sole influence of topography. However, the delta N-15 at depths of 10-20 and 20-30 cm decreased and increased with increasing vole numbers, respectively. Also the CO2 emission potential from soil collected at a depth of 15-30 cm decreased and the C/N ratio at 5-10 cm depth narrowed with increasing vole numbers. These variables indicated the first influence of voles on the respective mineralization processes in some soil layers. Tendencies of vole activity homogenizing SOC and N contents across layers were not significant. The results of the other seven key variables did not confirm significant effects of voles. Thus overall, we found mainly a first response of variables that are indicative for changes in biogeochemical dynamics but not yet of those representing changes in pools.