ABSTRACT: In agricultural landscapes, the amount and organization of crops and semi-natural habitats (SNH) have the potential to promote a bundle of ecosystem services due to their influence on ecological community at multiple spatio-temporal scales. SNH are relatively undisturbed and are often source of complementary resources and refuges, therefore supporting more diverse and abundant natural pest enemies. However, the nexus of SNH proportion and organization with pest suppression is not trivial. It is thus crucial to understand how the behavior of pest and natural enemy species, the underlying landscape structure, and their interaction, may influence conservation biological control (CBC). Here, we develop a generative stochastic landscape model to simulate realistic agricultural landscape compositions and configurations of fields and linear elements. Generated landscapes are used as spatial support over which we simulate a spatially explicit predator-prey dynamic model. We find that increased SNH presence boosts predator populations by sustaining high predator density that regulates and keeps pest density below the pesticide application threshold. However, predator presence over all the landscape helps to stabilize the pest population by keeping it under this threshold, which tends to increase pest density at the landscape scale. In addition, the joint effect of SNH presence and predator dispersal ability among hedge and field interface results in a stronger pest regulation, which also limits pest growth. Considering properties of both fields and linear elements, such as local structure and geometric features, provides deeper insights for pest regulation; for example, hedge presence at crop field boundaries clearly strengthens CBC. Our results highlight that the integration of species behaviors and traits with landscape structure at multiple scales is necessary to provide useful insights for CBC. Author summary In the agricultural context, the loss of semi-natural surfaces often results in high pest abundance requiring elevated pesticide loads. Habitat heterogeneity resulting from the agricultural intermixing of arable fields and semi-natural areas is key to allow organism fluxes across agro-ecological interfaces by influencing ecological processes. Semi-natural habitats (SNH) are often restricted to linear structures, such as hedgerows, but they play an important role by hosting a large number of species. However, the effect of hedgerows is controversial, as it could result in a positive, ineffective or negative effect for CBC. Usually, the impacts of landscape structure on pest population dynamics and resulting CBC are assessed through field experiments with a specific focus, which cannot be generalized, lack flexibility and are limited by the need to manipulate relatively large landscapes. Here, we tackle the challenge to investigate the controversial role of semi-natural habitats for CBC by presenting a simulation-based approach, which allows us to characterize the joint influence of landscape structure and species traits on CBC service. Our study corroborates that spatial heterogeneity, species traits and their interactions are fundamental for CBC. We show that hedge presence alone is not sufficient to lead to strong pest reduction, but hedge-based predators help to maintain the pest density under the pesticide threshold. Instead, SNH presence coupled with appropriate predator traits leads to stronger decrease of pest population. Moreover, we highlight an important scaling effect of SNH, which at the local scale has an even more important impact on CBC as local properties are considered.
