Reduction of damage by the exotic invasive conifer seed bug Leptoglossus occidentalis in mixed pine-birch plantations: A case of biotic resistance

The number of invasive non-native pests is increasing rapidly in forests as a result of global change. It is therefore important to prevent their damage in order to preserve the integrity of forest ecosystems and the associated services. According to the biotic resistance hypothesis, species-rich communities are less likely to be invaded. The associational resistance hypothesis states that insect herbivores are more likely to colonise and exploit plants surrounded by conspecific neighbours than heterospecific, non-host plant species. More diverse forests would therefore be less damaged by non-native pests than tree monocultures. We tested these hypotheses by comparing the damage caused to seeds by the western conifer bug Leptoglossus occidentalis, , an invasive insect native to North America, in plots of pure maritime pine and mixed plots of maritime pine and birch. These plots were in two tree diversity experiments in Europe. Mixed pine plots differed in terms of pine density (1250 versus 625 pines/ha), proportion of pine and birch (25 %, 50 %, 75 %) and spatial mixing pattern (aggregated versus dispersed). We sampled 635 cones in 37 plots. Overall, the proportion of seeds damaged by the invasive bug was significantly lower in mixed plots than in pure pine plots at both experimental sites. There was no significant effect of pine density or relative proportion in the mixtures on seed damage. Aggregated pines in the mixed plots were significantly more damaged than dispersed pines, suggesting that bugs may have done less damage when they had more difficulty locating their host trees. This is consistent with the hypothesis of lower plant apparency. These results support the view that forest plantations of mixed species, which are known to be less vulnerable to attack by native insects, may also be more resistant to infestation by exotic insects. Row-wise, intimate mixtures of two species represent a promising option for designing tree plantations that are more resistant and easier to manage.