Meta-analysis of tree diversity effects on the abundance, diversity and activity of herbivores' enemies

The natural enemies hypothesis predicts that the abundance and diversity of antagonists such as predators and parasitoids of herbivores increases with the diversity of plants, which can lead to more effective top-down control of insect herbivores. However, although the hypothesis has received large support in agricultural systems, fewer studies have been conducted in forest ecosystems and a comprehensive synthesis of previous research is still lacking. We conducted a meta-analysis of 65 publications comparing the diversity, abundance or activity of various groups of natural enemies (including birds, bats, spiders and insect parasitoids) in pure vs. mixed forest stands. We tested the effects of forest biome, natural enemy taxon and type of study (managed vs experimental forest). We found a significant positive effect of forest tree diversity on natural enemy abundance and diversity but not on their activity. The effect of tree diversity on natural enemies was stronger towards lower latitudes but was not contingent on the natural enemy taxon. Overall, our study contributes substantially toward a better understanding of the natural enemies hypothesis in forest systems and provides new insights about the mechanisms involved. Furthermore, we outline potential avenues for strengthening forest resistance to the growing threat of herbivorous insects.

Contribution of hardwood trees to budworm – parasitoid food web dynamics

A major pest of Atlantic forests is the spruce budworm caterpillar which outbreaks every 35 years and causes large scale tree mortality. Historically, budworm management has largely ignored other species in the food web. Broadening the focus could reduce budworm outbreaks while balancing the multiple demands on our forests. However, the food web surrounding budworm including other caterpillar species that are attacked by budworm parasitoids has been relatively undersampled and under-researched. Therefore, we tested two hypotheses: the alternating hardwood-softwood parasitoids hypothesis where parasitoids attack other caterpillars on hardwoods when budworm are rare and attack budworm on balsam fir or other softwoods when budworm are plentiful, and the mixed stands natural enemies hypothesis where stands with a mixture of softwood and hardwood trees harbour greater abundances and diversity of budworm parasitoids. We tested these hypotheses using stable isotope analysis of budworm parasitoids and through community analyses of parasitoids sampled along a hardwood gradient. We found indications that parasitoids do attack caterpillars on hardwoods and budworm on balsam fir, but found mixed results for the natural enemies hypothesis. Our study highlights the importance for budworm management of understanding the dynamics of the food web surrounding budworm.

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. Summary 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.

Habitat Management for Pest Management: Limitations and Prospects

Habitat management is an important strategy for pest control in integrated pest management (IPM). Various categories of habitat management such as trap cropping, intercropping, natural enemy refuges such as ‘beetle banks’, and floral resources for parasitoids and predators, have been used in applied insect ecology for many years. In a broader sense, two mechanisms, the ‘enemies hypothesis’ and the ‘resource concentration hypothesis’ have been identified as acting independently or combined in pest population dynamics. The ‘enemies hypothesis’ directly supports the conservation and enhancement of natural enemies, floral resources such as shelter, nectar, alternative food sources, and pollen (SNAP) to improve conservation biological control. The ‘resource concentration hypothesis’ emphasizes how the host selection behavior of herbivores in a diverse habitat can reduce pest colonization in crops. This review emphasizes the potential of these approaches, as well as possible dis-services, and includes limitations and considerations needed to boost the efficacy of these strategies worldwide.

Urban tree pests and natural enemies respond to habitat at different spatial scales

Trees provide many ecosystem services in our urban environments. However, city trees are often stressed by pests that are typically higher than those in nearby natural areas. Our research highlights a potential mismatch in scale between the habitat elements that affect the densities of pests and their natural enemies on city trees. We tested a well-known ecological concept, the enemies hypothesis, in the city, where relationships of pests and their enemies have not been thoroughly studied. To test our hypothesis that natural enemies and aphid predation services on urban trees increase with more local structural complexity around trees, we collected data on crape myrtle trees on NC State University’s campus from 2016 to 2017. We measured local structural complexity of vegetation around study trees, quantified impervious cover among other urban habitat elements, collected crape myrtle aphids (Tinocallis kahawaluokalani) and their natural enemies on trees, and performed predation experiments. We found that aphid abundance was positively correlated with more impervious cover within 100 m of crape myrtle trees. Alternatively, greater local structural complexity within the 10 × 10 m area around crape myrtles correlated with a higher abundance of natural enemies. Aphid predation was mostly predicted by local structural complexity and impervious cover within 20 m of crape myrtle trees. Together, these findings suggest that although the impervious nature of our cities may mean higher densities of some pests, local landscapes around trees can play an important role in maintaining natural enemies and predation services that help regulate pest populations.