Allometric scaling of indirect effects: body size ratios predict non-consumptive effects in multi-predator systems.

Non-consumptive effects (NCES) frequently lead to non-independent effects of multiple predators. While such emergent predator effects are ubiquitous in natural communities, the strength of these effects varies among studies and systems, making it difficult to predict a priory how changes in predator diversity influence prey suppression. Thus, identifying general scaling rules which can explain this variation of non-independent effects is vital for modelling natural communities and how they respond to biodiversity loss. Body size is a key trait determining the nature and strength of ecological interactions. While great progress has been made using allometric relationships to predict the interaction strength of predator-prey pairs, it is unknown whether similar relationships explain variation in the strength of NCEs, and how they are related to consumptive effects. Here, we experimentally manipulate the relative size difference of multiple predators to determine whether NCEs follow general allometric scaling relationships in an aquatic multi-predator system. Results demonstrate that the presence and strength of NCEs can vary dramatically across predator combinations. However, this variation scaled predictably with the size ratio of predators, increasing the size difference among predators increased NCEs. This pattern was driven by a size-mediated shift in 'food web motif' from competition to intraguild predation and a positive correlation of NCEs and intraguild predation rate. Results indicate that models which assume that consumers have independent effects are particularly likely to make erroneous predictions when predators differ substantially in size, but simple allometric relationships of NCEs could be used to correct this bias.