Habitat heterogeneity is thought to affect top‐down control of herbivorous insects and contribute to population stability by providing a more attractive microhabitat for natural enemies, potentially leading to reduced population fluctuations. Identifying the parameters that contribute to habitat heterogeneity promoting top‐down control of herbivorous insects by natural enemies could facilitate appropriate management decisions, resulting in a decreased risk of pest insect outbreaks because of a higher level of predation. In our study, we measured the top‐down pressure exerted by small mammals on the cocoons of a notorious pest insect in pine forests, the European pine sawfly (Neodiprion sertifer), which is known to be regulated by small mammal predation. The forest stands used differed in heterogeneity measured in terms of differences in tree diversity and density, understory vegetation height, presence/absence, and density of dead wood. We found higher predation in more dense spots within forest stands. Further, the effect of dead wood on sawfly cocoon predation depended on the pine proportion in forest stands. The addition of dead wood in a manipulation experiment had a slight positive effect on cocoon predation, while dead wood removal caused a clear decrease in predation rate, and the decrease was more pronounced when the proportion of pine increased. Our results show that habitat heterogeneity affects predation by generalist predators on herbivorous insects. This knowledge could be applied to reduce the risk of insect outbreaks by applying management methods that increase heterogeneity in perennial systems such as forests and orchards, thus decreasing the levels of insect damage.
1 Observed lower levels of herbivory in mixed compared with monoculture stands have been hypothesized to depend on top-down forces, through higher predation pressure by natural enemies or through bottom-up mechanisms through plant quality effects on herbivore performance. 2 In this study, we compared the performance measured as host plant induced mortality, cocoon weight, and predation mortality of the European pine sawfly Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae) in mixed and monoculture forest stands. 3 We did not observe a difference in host plant induced mortality, cocoon weight, or predation mortality between mixed and monoculture forest stands. We did find an effect of local conditions around each experimental tree on pine sawfly performance. For example, the nitrogen content of pine needles is negatively affected by the proportion of pine around the experimental tree, which in turn increases the survival of sawfly larvae. 4 The results suggest that local conditions around individual trees are more important for the performance of the European pine sawfly than stand type, i.e. mixed or monoculture plant stands. 5 We conclude that the ongoing trend for diversification within commercial forestry calls for more research where the effects of both bottom-up and top-down effects are studied at several spatial scales.
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