Top-down and sideways: Herbivory and cross-ecosystem connectivity shape restoration success at the salt marsh-upland ecotone.

Wetland restoration provides remarkable opportunities to understand vegetation dynamics and to inform success of future projects through rigorous restoration experiments. Salt marsh restoration typically focuses on physical factors such as sediment dynamics and elevation. Despite many demonstrations of strong top-down effects on salt marshes, the potential for consumers to affect salt marsh restoration projects has rarely been quantified. Recently, major restoration projects at the Elkhorn Slough National Estuarine Research Reserve in central California, USA provided an opportunity to examine how herbivory influences restoration success. We quantified the strength of consumer effects by comparing caged to uncaged plantings, and compared effects among plant species and sites. We used camera traps to detect which herbivores were most common and how their abundance varied spatially. Beyond characterizing consumer effects, we also tested management strategies for reducing negative effects of herbivory at the restoration sites, including caging, mowing, and acoustic playbacks of predator sounds. We found extremely strong consumer effects at sites with extensive stands of exotic forbs upland of the high marsh; uncaged restoration plants suffered heavy herbivory and high mortality, while most caged plants survived. Brush rabbits (Sylvilagus bachmani) were by far the most frequent consumers of these high marsh plants. Our work thus provides the first evidence of mammal consumers affecting salt marsh restoration success. Mowing of tall exotic forb cover adjacent to the marsh at one restoration site greatly reduced consumption, and nearly all monitored plantings survived at a second restoration site where construction had temporarily eliminated upland cover. Playbacks of predator sounds did not significantly affect restoration plantings, but restoration efforts in marsh communities vulnerable to terrestrial herbivory may benefit from concurrent restoration of predator communities in the upland habitats surrounding the marsh. A landscape approach is thus critical for recognizing linkages between terrestrial and marine vegetation.