Patchy indirect effects of predation: predators contribute to landscape heterogeneity and ecosystem function via localized pathways

Predators are widely recognized for their irreplaceable roles in influencing the abundance and traits of lower trophic levels. Predators also have irreplaceable roles in shaping community interactions and ecological processes via highly localized pathways (i.e. effects with well-defined and measurable spatio–temporal boundaries), irrespective of their influence on prey density or behavior. We synthesized empirical and theoretical research describing how predators – particularly medium- and large-sized carnivores – have indirect ecological effects confined to discrete landscape patches, processes we have termed ‘patchy indirect effects (PIEs) of predation'. Predators generate PIEs via three main localized pathways: generating and distributing prey carcasses, creating ecological hotspots by concentrating nutrients derived from prey, and killing ecosystem engineers that create patches. In each pathway, the indirect effects are limited to discrete areas with measurable spatial and temporal boundaries (i.e. patches). Our synthesis reveals the diverse and complex ways that predators indirectly affect other species via patches, ranging from mediating scavenger interactions to influencing parasite/disease transmission risk, and from altering ecosystem biogeochemistry to facilitating local biodiversity. We provide basic guidelines on how these effects can be quantified at the patch and landscape scales, and discuss how predator-mediated patches ultimately contribute to landscape heterogeneity and ecosystem functioning. Whereas density- and trait-mediated indirect effects of predation generally occur through population-scale changes, PIEs of predation occur through individual- and patch-level pathways. Our synthesis provides a more holistic view of the functional role of predation in ecosystems by addressing how predators create patchy landscapes via localized pathways, in addition to influencing the abundance and behavior of lower trophic levels.