Rewilding with large herbivores: Positive direct and delayed effects of carrion on plant and arthropod communities.

Carrion of large animals is an extremely nutrient rich, ephemeral resource that is essential for many species, but is scarce in the anthropogenic Western-European landscape due to legislative restrictions. Rewilding, a novel conservation strategy that aims at restoring natural processes with minimal human intervention, is increasing in popularity and could lead to increased carrion availability in the landscape. It is therefore important to understand the effects of carrion on biodiversity. We investigated the direct and delayed (five months) effects of red deer (Cervus elaphus) carcasses on plants and arthropods in the Oostvaardersplassen, the Netherlands, one of the oldest rewilding sites in Europe. Specifically, we tested whether carrion has a positive direct effect on the abundances and diversity of various arthropod functional groups, as well as a delayed effect on the vegetation and arthropods through the increased nutrient availability. During the active decomposition stage in spring, we, not surprisingly, observed higher abundances of carrion associated species (scavengers and their specialized predators) at the carrion sites than at control sites without carrion, but no higher abundances of predators or detritivores. In late summer, after near-complete decomposition, plant biomass was five times higher, and nutritional plant quality (C:N ratio) was higher at the carrion sites than at the control sites. Arthropod abundance and diversity were also manifold higher, owing to higher numbers of herbivorous and predatory species. Regression analysis showed that abundances of herbivores and detritivores were positively related to plant biomass, and predator abundances were positively related to abundances of herbivores and detritivores, suggesting bottom-up effects propagating through the food chain. Our results show that even in a naturally nutrient-rich ecosystem like the Oostvaardersplassen, carrion can have strong positive effects on local plant biomass and nutritional quality and arthropod abundances, lasting the whole growing season. We found evidence that these effects were first directly caused by the presence of carrion, and later by the enhanced nutrient availability in the soil. This highlights the importance of the indirect pathways by which carrion can structure arthropod communities.