Stem-boring insects of fragmented Calamagrostis habitats: herbivore–parasitoid community structure and the unpredictability of grass shoot abundance


MARK DUBBERT Agroecology, University of Göttingen, Waldweg 26, 37073 Göttingen, TEJA TSCHARNTKE Agroecology, University of Göttingen, Waldweg 26, 37073 Göttingen,


Abstract.1. Communities of stem-boring insects attacking the grass Calamagrostis epigeios (L.) Roth were analysed from twenty-five pure stands of C. epigeios differing in area and isolation. Insect communities comprised nine phytophagous and eighteen entomophagous species. Most abundant herbivores (attacking > 1% of all shoots) were Tetramesa eximia (Giraud) (Hym. Eurytomidae), Eurytoma sp. (Hym. Eurytomidae), Lasioptera calamagrostidis (Rübs.) (Dipt. Cecidomyiidae), and Eriopeltis sp. (Hom. Pseudococcidae). About 28% of the twenty-seven species were monophagous, 61% oligophagous (restricted to Poaceae), and 11% polyphagous. Although herbivores generally attacked thick shoots, species-specific differences in the attacked mean diameter were found.2. The main purpose of this study was to analyse the relative effects of area, isolation and unpredictability of available food resources on the abundance of herbivores, parasitoids and resulting tritrophic effects. Variability of shoot abundance varied greatly between years, and even more between habitats within the same year. On average, shoot density per habitat increased or decreased 2.4-fold between succeeding years.3. Variability of grass shoot abundance was not related to habitat characteristics such as area, isolation, biomass, shoot density, changes in area or shoot density between years, or shoot features like internode number or diameter, but was negatively influenced by shading of trees. Similarly, habitat area or isolation did not correlate with other habitat features.4. Grass shoot abundance was the most important predictor of species abundance and attack rate of herbivores. In contrast to expectations, area and isolation of habitats did not explain differences in species richness, and only influenced abundance of three species. In addition, even the experimental removal of all grass shoots (and thereby, of all stem-boring insects) from five habitats in winter did not negatively influence colonization the following year. These results suggest a surprisingly high mobility of this rather specialized community of stem-boring insects that appeared to superimpose on area and isolation effects. Unpredictability of grass shoot abundance was presumably the main selection pressure for the obviously high dispersal abilities of the Calamagrostis insects.5. Per cent parasitism was mainly influenced by the percentage of attacked shoots. Parasitism of T. eximia by its major parasitoid also correlated with habitat connectivity, emphasizing the hypothesis that a continuum of habitats should be particularly favourable for the third trophic level.