The importance of litter for interactions between terrestrial plants and invertebrates

Sammanfattning: According to the exploitation ecosystem hypothesis (EEH), terrestrial ecosystems are characterized by well defined trophic levels and strong trophic interactions with community level tropic cascades. In unproductive terrestrial habitats as tundra heaths, the energy shunt from litter and apparent competition between herbivores and detritivores are expected to be important for the structure and dynamics of the invertebrate community. The aim of this study was to test this hypothesis by investigating if plant litter accumulation was affecting the invertebrate community on a nutrient-poor tundra heath. The study was performed during one summer on the highland part of Joatka research area, in the north of Norway. The experimental area included 16 plots (100 m2 each), of which 12 had been littermanipulated. On four plots the amount of litter was increased by 100 %, on four by 200 % and on four by 400 %. Four plots were untreated and used as control plots. Invertebrates were collected by emergence traps (which cover an area of 1 m2), one trap on each plot and one pitfall trap inside each emergence trap. During the study period, traps were emptied and moved twice, resulting in three sampling periods. The invertebrates collected were counted and their length was measured, than all invertebrates were sorted into taxa and trophic guilds. During the study period, herbivore grazing damage was investigated on all 16 experimental plots, signs of herbivores on leaves of vascular plants in an area covering 3 m2 per plot were noted, for every leaf with signs of herbivory the percentage of leaf area removed was estimated. Plant biomass and plant species composition were estimated in all experimental plots by harvesting above-ground plant parts. In each plot, two squares were randomly chosen and all biomass in this square was collected. Plant biomass was sorted in to following groups: dwarf birch, billberry, Salix herbacea, Salix spp, graminoids, herbs, lichens, mosses and dwarf shrub. Before weighing the plant material, it was stored in paper bags at room temperature and then dried for 48 h at 40°C. In order to detect fertilisation effects, all bilberry shoots that had been produced during the actual summer were separately weighted when analyzing the plant biomass. The result showed that the invertebrate community in this area is dominated by carnivores while detritivores, parasitoids and herbivores are quite rare, this was in accordance with previous studies made in the area. Litter manipulation did not create any significant variation in the community structure, but there was a slight tendency that carnivore biomass increased and biomass of herbivores decreased when litter was added to the system. In contrary to this, gracing activity especially on dwarf willow (Salix herbacea) increased in plots were 100 % and 200 % more litter was added. There is a positive correlation between biomass of herbivores and detritivores but the reason for this seems unclear. No fertilisation effect was detected in litter manipulated plots. The structure and dynamics of the actual community could not be described by the food web theory EEH and energy shunt from litter and apparent competition between herbivores and detritivores. It seems to be several complicating factors to take into consideration when describing this community.