The Influence of Tree Diversity on Natural Enemies-a Review of the Enemies Hypothesis in Forests

Purpose of Review Natural enemies are an important component for forest functioning. By consuming herbivores, they can be effective top-down regulators of potential pest species. Tree mixtures are generally expected to have larger predator and parasitoid populations compared to monocultures. This assumption is based on the enemies hypothesis, a classical ecological concept predicting a positive relationship between plant diversity (and complexity) and natural enemies, which, in turn, should increase top-down control in more diverse environments. However, the enemies hypothesis has mostly been tested and supported in relatively simple agricultural ecosystems. Until recently, research in forests was sparse. We summarize the upcoming knowledge-base for forests and identify forest characteristics likely shaping relationships between tree diversity, natural enemies (abundance, species richness, diversity), and top-down control. We further identify possible implications for mixed species forestry and key knowledge gaps. Recent Findings Tree diversity (almost exclusively quantified as tree species richness) does not consistently increase enemy abundance, diversity, or result in herbivore control. Tests of the enemies hypothesis are largely based on aboveground natural enemies (mainly generalists) and have highly variable outcomes across taxa and study systems, sometimes even finding a decrease in predator diversity with increasing tree diversity. Recurrent effects of tree species identity and composition indicate that a closer focus on tree functional and phylogenetic diversity might help to foster a mechanistic understanding of the specific circumstances under which tree diversity can promote top-down control. Our review suggests that the enemies hypothesis may not unambiguously apply to forests. With trees as structurally complex organisms, even low-diversity forests can maintain a high degree of habitat heterogeneity and may provide niches for many predator and parasitoid species, possibly blurring correlations between tree and natural enemy diversity. Several further factors, such as latitude, identity effects, intraguild predation, or functional and phylogenetic components of biodiversity, may confound the predictions of the enemies hypothesis. We identify topics needing more research to fully understand under which conditions tree diversity increases natural enemy diversity and top-down control-knowledge that will be crucial for forest management.