Predictive theory on how plant diversity promotes herbivore suppression through movement patterns, host associations, and predation promises a potential alternative to pesticide-intensive monoculture crop production. We used meta-analysis on 552 experiments in 45 articles published over the last 10 years to test if plant diversification schemes reduce herbivores and/or increase the natural enemies of herbivores as predicted by associational resistance hypotheses, the enemies hypothesis, and attraction and repellency model applications in agriculture. We found extensive support for these models with intercropping schemes, inclusion of flowering plants, and use of plants that repel herbivores or attract them away from the crop. Overall, herbivore suppression, enemy enhancement, and crop damage suppression effects were significantly stronger on diversified crops than on crops with none or fewer associated plant species. However, a relatively small, but significantly negative, mean effect size for crop yield indicated that pest-suppressive diversification schemes interfered with production, in part because of reducing densities of the main crop by replacing it with intercrops or non-crop plants. This first use of meta-analysis to evaluate the effects of diversification schemes, a potentially more powerful tool than tallies of significant positive and negative outcomes (vote-counting), revealed stronger overall effects on all parameters measured compared to previous reviews. Our analysis of the same articles used in a recent review facilitates comparisons of vote-counting and meta-analysis, and shows that pronounced results of the meta-analysis are not well explained by a reduction in articles that met its stricter criteria. Rather, compared to outcome counts, effect sizes were rarely neutral (equal to zero), and a mean effect size value for mixed outcomes could be calculated. Problematic statistical properties of vote-counting were avoided with meta-analysis, thus providing a more precise test of the hypotheses. The unambiguous and encouraging results from this meta-analysis of previous research should motivate ecologists to conduct more mechanistic experiments to improve the odds of designing effective crop diversification schemes for improved pest regulation and enhanced crop yield.

SignificanceDecades of research have fostered the now-prevalent assumption that noncrop habitat facilitates better pest suppression by providing shelter and food resources to the predators and parasitoids of crop pests. Based on our analysis of the largest pest-control database of its kind, noncrop habitat surrounding farm fields does affect multiple dimensions of pest control, but the actual responses of pests and enemies are highly variable across geographies and cropping systems. Because noncrop habitat often does not enhance biological control, more information about local farming contexts is needed before habitat conservation can be recommended as a viable pest-suppression strategy. Consequently, when pest control does not benefit from noncrop vegetation, farms will need to be carefully comanaged for competing conservation and production objectives.

Human land use threatens global biodiversity and compromises multiple ecosystem functions critical to food production. Whether crop yield–related ecosystem services can be maintained by a few dominant species or rely on high richness remains unclear. Using a global database from 89 studies (with 1475 locations), we partition the relative importance of species richness, abundance, and dominance for pollination; biological pest control; and final yields in the context of ongoing land-use change. Pollinator and enemy richness directly supported ecosystem services in addition to and independent of abundance and dominance. Up to 50% of the negative effects of landscape simplification on ecosystem services was due to richness losses of service-providing organisms, with negative consequences for crop yields. Maintaining the biodiversity of ecosystem service providers is therefore vital to sustain the flow of key agroecosystem benefits to society.

Claims about the role of predator diversity in maintaining ecosystem function and providing ecosystem services such as pest control are controversial, but evaluative tests are beginning to accumulate. Empirical and experimental comparisons of species-rich versus species-poor assemblages of entomophagous arthropods and vertebrates range from strong suppression to facilitative release of herbivorous arthropod prey. Top-down control can be strengthened when natural enemies complement each other, dampened by negative interactions, balanced by both factors, and driven by single influential species. A meta-analytic synthesis shows a significant overall effect of enemy richness increasing top-down control of herbivores, which is consistent in agricultural studies conducted in tropical versus temperate zones, in studies using caged versus open-field designs, but not so in nonagricultural habitats. Synthetic analyses address theory and help set precautionary policy for conserving ecological services broadly, while characterizing uncertainty associated with herbivore response to changes in enemy diversity.