Interactions between sheep, rodents, graminoids, and bryophytes in an oceanic alpine ecosystem of low productivity

Abstract

Large herbivores directly affect plant communities in alpine ecosystems. In addition, they may compete with, or facilitate foraging by, small herbivores and also cause strong indirect effects on plants. We used an exclosure experiment to examine short-term (5-y) effects of cessation of sheep grazing on rodent grazing and plant communities in an oceanic alpine environment of low productivity with a long history of heavy sheep grazing. Exclusion of sheep significantly impacted plant communities. Vascular plant height increased, but Deschampsia flexuosa was the only vascular plant species that increased in sheep exclosures. Changes in the frequency of graminoids, herbs, and dwarf shrubs in exclosed plots were not related to cessation of grazing, but 6 bryophyte species significantly increased or decreased in response to exclusion of sheep. The absence of large grazers thus brought about a change in the species composition in favour of successional bryophytes and the preferred fodder plant. Neither vascular plant nor bryophyte species richness, nor the total cover of bryophytes and lichens, were affected. Cessation of sheep grazing reduced the level of rodent grazing. Rodent grazing correlated with changes in plant communities that led to reduced height and cover of vascular plants, reduced cryptogam cover, and reduced frequencies of 3 bryophyte species. A strong correlation between sheep fodder value index and rodent grazing indirectly indicated additive herbivory. In addition, some of the rodent effects were compensatory; e.g., Nardus stricta, which is not grazed by sheep, was significantly reduced by rodents. Our study points to a more central role of facilitation in structuring herbivore assemblages in the short term, with direct implications for the joint effects of large and small herbivores on the cover and frequency of graminoids and bryophytes.