Effects of herbaceous plant functional groups on the dynamics and structure of an alpine lichen heath: the results of a removal experiment

Alpine lichen heaths are polydominant, low-productive communities where lichens and, at some localities, the dwarf shrub Vaccinium vitis-idaea prevail. To analyse the role of herbaceous dominants in the structure of alpine heaths, we established a removal experiment. We tested (1) whether dominant graminoids and forbs differ in their effects on species richness, phytomass, and abundance of individual species; and (2) which remaining species are able to replace the removed dominants. Permanent plots were established in 1996 in the Teberda Reserve (the Northwestern Caucasus, Russia) at 2750 m a.s.l. with four treatments: (1) control, (2) forb removal, (3) graminoid removal, (4) graminoid and forb removal. Target species were clipped every year to avoid regrowth. We counted the shoot numbers of all vascular plants yearly from 1996 to 2009. In 2010, we harvested the aboveground phytomass. We used mixed-effects models to assess the responses of the remaining species and phytomass fractions to the removal. The biomass of the removed dominants was almost compensated for by the remaining species, but due to diffuse competition none of the subordinate species became a new dominant. Graminoid removal reduced community weighted mean leaf dry matter content and litter accumulation. Species number per plot completely recovered after graminoid, not forb removal. Shoot numbers of Antennaria dioica, Arenaria lychnidea, Campanula tridentata, Ranunculus oreophilus, and Trifolium polyphyllum showed positive responses to graminoid removal. Anemone speciosa shoot numbers decreased after graminoid removal. In alpine heath, graminoid dominants contribute more to community structure than forb dominants.