Predatory beetles facilitate plant growth by driving earthworms to lower soil layers

Theory suggests that predators of soil-improving, plant-facilitating detritivores (e.g. earthworms) should suppress plant growth via a negative tri-trophic cascade, but the empirical evidence is still largely lacking. We tested this prediction in an alpine meadow on the Tibetan Plateau by manipulating predatory beetles (presence/absence) and quantifying (i) direct effects on the density andbehaviour of earthworms; and (ii) indirect effects on soil properties and above-ground plant biomass. In the absence of predators, earthworms improved soil properties, but did not significantly affect plant biomass. Surprisingly, the presence of predators strengthened the positive effect of earthworms on soil properties leading to the emergence of a positive indirect effect of predators on plant biomass. We attribute this counter-intuitive result to: (i) the inability of predators to suppress overall earthworm density; and (ii) the predator-induced earthworm habitat shift from the upper to lower soil layer that enhanced their soil-modifying, plant-facilitating, effects. Our results reveal that plant-level consequences of predators as transmitted through detritivores can hinge on behaviour-mediated indirect interactions that have the potential to overturn predictions based solely on trophic interactions. This work calls for a closer examination of the effects of predators in detritus food webs and the development of spatially explicit theory capable of predicting the occurrence and consequences of predator-induced detritivore behavioural shifts