Birds and bats enhance cacao yield despite suppressing arthropod mesopredation

Bird- and bat-mediated biocontrol benefits the productivity of tropical commodity crops such as cacao, but the ecological interactions driving these ecosystem services remain poorly understood. Whereas birds and bats prey on herbivorous arthropods, they may also prey on arthropod mesopredators such as ants, with poorly understood consequences for pest biocontrol. We used a full-factorial experiment excluding birds, bats, and ants to assess their effects on (a) the abundance of multiple arthropod groups; (b) predation pressure on arthropods evaluated through artificial sentinel caterpillars; and (c) cacao yield over 1 year in shaded agroforestry systems of native cacao varieties in Peru. Birds and bats increased cacao yield by 118%, which translates in smallholder benefits of ca. US $959 ha^-1  year^-1 . Birds and bats decreased predation by ants and other arthropods, but contributed to the control of phytophagous taxa such as aphids and mealybugs. By contrast, ant presence increased the abundance of these sap-sucking insects, with negative impacts for cacao yield. Notably, high abundances of the dominant ant Nylanderia sp., known to attend sap-sucking insects, were associated with lower cacao yield along a distance gradient from the closest forest edge. According to these results, arthropod predation by birds and bats, rather than mesopredation by arthropods, was most responsible for increases in cacao yield. Moving forward, detailed research about their trophic interactions will be necessary to identify the cause of such benefits. Retaining and restoring the large benefits of birds and bats as well as minimizing disservices by other taxa in cacao agroforests can benefit from management schemes that prioritize preservation of shade trees and adjacent forests within agroforestry landscapes.