Plant apparency, an overlooked driver of associational resistance to insect herbivory

1. Herbivore regulation is one of the services provided by plant diversity in terrestrial ecosystems. It has been suggested that tree diversity decreases insect herbivory in forests, but recent studies have reported opposite patterns, indicating that tree diversity can trigger associational resistance or susceptibility. The mechanisms underlying the tree diversity-resistance relationship thus remain a matter of debate. 2. We assessed insect herbivory on pedunculate oak saplings (Quercus robur) in a large-scale experiment in which we manipulated tree diversity and identity by mixing oaks, birch and pine species. 3. Tree diversity at the plot scale had no effect on damage due to leaf chewers, but abundance of leaf miners decreased with increasing tree diversity. The magnitude of this associational resistance increased with host dilution, consistent with the 'resource concentration hypothesis'. 4. At a smaller scale, we estimated tree apparency as the difference in total height between focal oak saplings and their nearest neighbouring trees. Levels of oak infestation with leaf miners decreased significantly with decreasing tree apparency. As the probability of having taller neighbours increased with tree diversity, notably due to the increase in the proportion of faster growing nonhost trees, such as birches and pines, tree apparency may be seen as a 'hidden', sampling effect of tree diversity. 5. Synthesis. These findings suggest that greater host dilution and lower tree apparency contribute to associational resistance in young trees. They also highlight the importance of taking plant size into account as a covariate, to avoid misleading interpretations about the biodiversity-resistance relationship.