Predator effects on a detritus-based food web are primarily mediated by non-trophic interactions

Predator effects on ecosystems can extend far beyond their prey and are often not solely lethally transmitted. Change in prey traits in response to predation risk can have important repercussions on community assembly and key ecosystem processes (i.e. trait-mediated indirect effects). In addition, some predators themselves alter habitat structure or nutrient cycling through ecological engineering effects. Tracking these non-trophic pathways is thus an important, yet challenging task to gain a better grasp of the functional role of predators. Multiple lines of evidence suggest that, in detritus-based food webs, non-trophic interactions may prevail over purely trophic interactions in determining predator effects on plant litter decomposition. This hypothesis was tested in a headwater stream by modulating the density of a flatworm predator (Polycelis felina) in enclosures containing oak (Quercus robur) leaf litter exposed to natural colonization by small invertebrates and microbial decomposers. Causal path modelling was used to infer how predator effects propagated through the food web. Flatworms accelerated litter decomposition through positive effects on microbial decomposers. The biomass of prey and non-prey invertebrates was not negatively affected by flatworms, suggesting that net predator effect on litter decomposition was primarily determined by non-trophic interactions. Flatworms enhanced the deposition and retention of fine sediments on leaf surface, thereby improving leaf colonization by invertebrates - most of which having strong affinities with interstitial habitats. This predator-induced improvement of habitat availability was attributed to the sticky nature of the mucus that flatworms secrete in copious amount while foraging. Results of path analyses further indicated that this bottom-up ecological engineering effect was as powerful as the top-down effect on invertebrate prey. Our findings suggest that predators have the potential to affect substantially carbon flow and nutrient cycling in detritus-based ecosystems and that this impact cannot be fully appreciated without considering non-trophic effects.