The effects of trophic structure and soil fertility on the assembly of plant communities: a microcosm experiment

Using closed, ventilated outdoor microcosms, plant communities were allowed to assemble from a pool of 48 herbaceous species comprising a wide range of plant functional types. The experiment involved factorial manipulation of soil fertility, invertebrate herbivores (slugs, grass aphids) and their predators (ground beetles, ladybirds). In the absence of herbivores the vegetation on soils of high, moderate and low fertility was dominated by plant species which under natural field conditions are restricted to fertile soils. At high fertility, the rate of competitive exclusion was rapid compared with communities persisting at lower fertility which remained species-rich with high densities of individuals. The effects of herbivory were profound and attributable mainly to the slug Deroceras reticulatum. At moderate and low soil fertility the main effect of herbivores was to change the ranking of plant species; palatable grasses (e.g. Poa annua, Poa trivialis and Lolium perenne) were suppressed and plant species of slower growth rate (e.g. Festuca rubra, Anthoxanthum odoratum, Festuca ovina) were promoted. More subtle impacts of herbivory altered the size structure of plant populations suggesting the preference, by slugs, for seedlings and small (suppressed) plants. Although ladybirds and their larvae persisted in low numbers in some microcosms, carnivory was more strongly in evidence (lower densities of D. reticulatum and reduced consumption of lettuce discs) in relation to the activities of the ground beetle Pterostichus melanarius. Only in the low fertility treatment, however, was this potential translated into effects on vegetation; for several plant species the frequency of large individuals increased in the presence of P. melanarius. We conclude that outdoor microcosms provide a valuable ecological tool and, in particular, allow investigation of (a) the role of herbivores in promoting slow-growing plant species in the vegetation of infertile habitats and (b) the protection of palatable plants against herbivores by carnivores.