Changing restoration rules: Exotic bivalves interact with residence time and depth to control phytoplankton productivity

Non-native species are a prevalent ecosystem stressor that can interact with other stressors to confound resource management and restoration. We examine how interactions between physical habitat attributes and a particular category of non-native species (invasive bivalves) influence primary production in aquatic ecosystems. Using mathematical models, we show how intuitive relationships between phytoplankton productivity and controllable physical factors (water depth, hydraulic transport time) that hold in the absence of bivalves can be complicated - and even reversed - by rapid bivalve grazing. In lightlimited environments without bivalves, shallow, hydrodynamically slow'' habitats should generally have greater phytoplankton biomass and productivity than deeper, faster'' habitats. But shallower, slower environments can be less productive than deeper, faster ones if benthic grazing is strong. Moreover, shallower and slower waters exhibit a particularly broad range of possible productivity outcomes that can depend on whether bivalves are present. Since it is difficult to predict the response of non-native bivalves to habitat restoration, outcomes for new shallow, slow environments can be highly uncertain. Habitat depth and transport time should therefore not be used as indicators of phytoplankton biomass and production where bivalve colonization is possible. This study provides for ecosystem management a particular example of a broad lesson: abiotic ecosystem stressors should be managed with explicit consideration of interactions with other major (including biotic) stressors. We discuss the applicability and management implications of our models and results for a range of aquatic system types, with a case study focused on the Sacramento-San Joaquin Delta (California, USA). Simple mathematical models like those used here can illuminate interactions between ecosystem stressors and provide process-based guidance for resource managers as they develop strategies to augment valued populations, restore habitats, and manipulate ecosystem functions.