Classical ecological theory states that in tritrophic systems, predators indirectly facilitate plants by reducing herbivore densities through consumption, while more recent work has revealed that predators can generate the same positive effect on plants non-consumptively by inducing changes in herbivore traits (e.g. feeding rates). Based on observations in US salt marshes dominated by vast monocultures of cordgrass, we hypothesized that sit-and-wait substratedwelling predators (crabs) could actually strengthen per capita impacts of potent grazers (snails), by non-consumptively inducing a vertical habitat shift of snails to their predation refuge high on canopy leaves that are vulnerable to grazing. A two-month field experiment supported this hypothesis, revealing that predators non-consumptively increased the mean climbing height of snails on grasses, increased grazing damage per leaf and ultimately suppressed cordgrass biomass, relative to controls. While seemingly counterintuitive, our results can be explained by (i) the vulnerability of refuge resources to grazing, and (ii) universal traits that drove the vertical habitat shift-i.e. relative habitat domains of predator and prey, and the hunting mode of the top predator. These results underline the fact that not only should we continue to incorporate non-consumptive effects into our understanding of top-down predator impacts, but we should do so in a spatially explicit manner.

Coral reef restoration aims to help threatened coral ecosystems recover from recent severe declines. Here we address whether coral fragments should be out‐planted individually or in larger aggregations. Theory suggests alternative possible outcomes: whereas out‐plants within aggregations might suffer from heightened negative interactions with neighbors (e.g. competition for space), they may alternatively benefit from positive interactions with neighbors (e.g. buffering wave disturbances). On a degraded reef in the Caribbean (St. Croix, USVI), using out‐plants of the critically endangered staghorn coral Acropora cervicornis, we experimentally tested how aggregation density (1–20 out‐planted coral fragments spaced at approximately 5 cm) influenced initial coral growth (over 3 months). Coral growth declined as a function of aggregation size, and out‐plants within larger aggregations had fewer and shorter secondary branches on average, indicative of horizontal competition for space. Our results therefore suggest that wide spacing of individuals will maximize the initial growth of out‐planted branching corals.

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top-down control of grazers. Grazer populations often exhibit size-structure, but the nature of multiple predator effects on suppression of size-structured prey has seldom been explicitly considered. In a southeastern US salt-marsh, we used both field (additive design) and mesocosm (additive-substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator-avoidance behavior (i.e. a non-consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller-bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non-consumptive effects differed from consumptive effects, with only the large predator inducing predator-avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size-spectrum of prey trophic regulation and non-consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non-consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size-structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.