ABSTRACT
1. Experimental studies across biomes demonstrate that herbivores can have significant effects on ecosystem functioning. Herbivore effects, however, can be highly variable with studies demonstrating positive, neutral or negative relationships between herbivore presence and different components of ecosystems. Mixed effects are especially likely in the soil, where herbivore effects are largely indirect mediated through effects on plants. 2. We conducted a long-term experiment to disentangle the effects of non-native moose in boreal forests on plant communities, nutrient cycling, soil composition and soil organism communities. 3. To explore the effect of moose on soils, we conduct separate analyses on the soil organic and mineral horizons. Our data come from 11 paired exclosure-control plots in eastern and central Newfoundland, Canada that provide insight into 22-25 years of moose herbivory. We fit piecewise structural equations models (SEM) to data for the organic and mineral soil horizons to test different pathways linking moose to above-ground and below-ground functioning. 4. The SEMs revealed that moose exclusion had direct positive impacts on adult tree count and an indirect negative impact on shrub percent cover mediated by adult tree count. We detected no significant impact of moose on soil microbial C:N ratio or net nitrogen mineralization in the organic or mineral soil horizon. Soil temperature and moisture, however, was more than twice as variable in the presence (i.e. control) than absence (i.e. exclosure) of moose. Overall, we observed clear impacts of moose on above-ground forest components with limited indirect effects below-ground. Even after 22-25 years of exclusion, we did not find any evidence of moose impacts on soil microbial C:N ratio and net nitrogen mineralization. 5. Our long-term study and mechanistic path analysis demonstrates that soils can be resilient to ungulate herbivore effects despite evidence of strong effects above-ground. Long-term studies and analyses such as this one are relatively rare yet critical for reconciling some of the context-dependency observed across studies of ungulates effects on ecosystem functions. Such studies may be particularly valuable in ecosystems with short growing seasons such as the boreal forest.
