Coffee berry borer control, but not coffee yield, is mediated by non-additive interaction between birds and ants across different cultivation systems

Coffee is one of the most traded tropical crops, cultivated in some of the most biodiverse regions on the planet. Coffee production can be seriously reduced by the coffee berry borer (CBB), a specialized beetle that feeds on the endosperm of coffee berries. Given the CBB's economic relevance, coffee-producing countries have developed extensive Integrated Pest Management programs. Nonetheless, most of these programs fail to incorporate CBB control by natural enemies such as birds and ants. While the effects of birds and ants on CBB suppression have been demonstrated to be positive when studied in isolation, their interactive effects have been little studied. To better understand the effects of the trophic interaction between birds and ants on CBB control, we conducted a full-factorial block experiment excluding birds and ants from coffee shrubs. We distributed 49 experimental blocks across three different coffee systems in a Colombian landscape: sun-exposed coffee, coffee-plantain intercropping, and shade coffee. We found birds to be key control agents of CBB: in the presence of ants, bird exclusion increased CBB infestation by 36%. However, in the absence of ants, birds had little effect on CBB infestation, demonstrating that the effects of birds and ants were non-additive. This suggests that birds control CBB through a trophic cascade mediated by ants. We also found that the effects of exclusions were modified by the cultivation system, with the shade coffee system consistently reducing CBB infestation. Our experiment demonstrates that crop diversification is an effective measure for integrating local enemies into IPM strategies. Nonetheless, we acknowledge that trophic interactions are highly complex and exhibit a context-dependency that can result in the suppression of biological pest control. Therefore, we recommend conducting future analysis on evaluating the effects of predator´s community composition to encourage the development of IPM programs that leverage biodiversity in agroecosystems.