Spatial responses of two herbivore groups to a geometrid larva on mountain birch.

Direct or plant-mediated interactions between herbivores may modify their spatial distribution among and within plants. In this study, we examined the effect of a leaf-chewing geometrid, the autumnal moth (Epirrita autumnata), on two different herbivore groups, leaf rolling Deporaus betulae weevils and Eriocrania spp. leafminers, both feeding on mountain birch (Betula pubescens ssp. czerepanovii). The exact locations of herbivores within tree canopies were mapped during three successive summers. In the first 2 years, some trees were artificially colonized by eggs of the autumnal moth to induce both rapid and delayed resistance in the foliage. The natural infection levels of the pathogenic rust fungus (Melampsoridium betulinum), potentially involved in species interactions, were also recorded. At the level of the whole tree, the density of D. betulae leaf rolls was lower in trees infested by the autumnal moth in the same year. However, the feeding locations within trees were partly segregated: D. betulae favoured shadier branches, while E. autumnata preferred the sunny parts of the canopy. The autumnal moth did not affect current- or following-year density of leafminers at the tree or branch level. Trees infected by rust had fewer leafminers in the same summer than noninfected trees. There were no interaction effects between defoliation by the autumnal moth and rust infection, and no delayed effects on the abundance of other herbivores the following year. Taken together, these findings suggest that the autumnal moth has a negative, partially plant-mediated impact on D. betulae, and can reduce the extent of current-year defoliation caused by D. betulae. This may be beneficial for the mountain birch, since the greater part of D. betulae damage occurs around or after the end of the larval period of the autumnal moth, which may be a critical time for tree recovery after moth outbreaks.