Higher-order interaction between molluscs and sheep affecting seedling numbers in grassland

Abstract

Vertebrate and invertebrate herbivores are both important in mesotrophic grasslands and these two different classes of herbivore potentially interact in their effect upon plant populations. We used two field experiments to test for higher order interactions (HOIs) among sheep, slugs and seedlings, using the mechanistic definition that an HOI occurs when the presence of one species modifies the interaction between two others. In each experiment slug addition and slug-removal treatments were nested inside treatments that altered sheep grazing intensity and timing, and the emergence, of seedlings from experimentally sown seeds was monitored. In Experiment 1, seedling numbers of Cerastium fontanum were increased by intense summer grazing by sheep in both slug-addition and slugremoval treatment, but winter grazing by sheep only increased seedling emergence if slugs were removed. In Experiment 2, winter grazing by sheep significantly reduced total seedling emergence of four species sown ( Lotus corniculatus, Plantago lanceolata, Leucanthemum vulgare, Achillea millefolium), but the effect was only seen where slugs were removed. Though the experimental system is a relatively simple one with only four components (sheep, slugs, seedlings and the matrix vegetation), higher order interactions, a combination of direct and indirect effects and possible switching behaviour by slugs are all suggested by our results.