Behaviorally-mediated trophic cascade attenuated by prey use of risky places at safe times.

The mere threat of predation may incite behavioral changes in prey that lead to community-wide impacts on productivity, biodiversity, and nutrient cycling. The paucity of experimental manipulations, however, has contributed to controversy over the strength of this pathway in wide-ranging vertebrate systems. We investigated whether simulated gray wolf (Canis lupus) presence can induce behaviorally-mediated trophic cascades, specifically, whether the 'fear' of wolf olfactory cues alone can change deer foraging behavior in ways that affect plants and soils. Wolves were recently removed from the Cedar Creek Ecosystem Science Reserve (Minnesota, USA), such that consumptively mediated predator effects were negligible. At 32 experimental plots, we crossed two nested treatments: wolf urine application and herbivore exclosures. We deployed camera traps to quantify how white-tailed deer (Odocoileus virginianus) adjusted their spatiotemporal habitat use, foraging, and vigilance in response to wolf cues and how these behavioral changes affected plant productivity, plant communities, and soil nutrients. Weekly applications of wolf urine significantly altered deer behavior, but deer responses did not cascade to affect plant or soil properties. Deer substantially reduced crepuscular activity at wolf-simulated sites compared to control locations. As wolves in this area predominantly hunted during mornings and evenings, this response potentially allows deer to maximize landscape use by accessing dangerous areas when temporal threat is low. Our experiment suggests that prey may be sensitive to 'dynamic' predation risk that is structured across both space and time and, consequentially, prey use of risky areas during safe times may attenuate behaviorally-mediated trophic cascades at the predator-prey interface.