Community-level effects of co-occurring native and exotic ecosystem engineers

1. Non-indigenous ecosystem engineers can substantially affect native biodiversity by transforming the physical structure of habitats. In the Great Lakes–St. Lawrence River system, introduced dreissenid mussels (Dreissena polymorpha and D. bugensis) and the native benthic macroalga Cladophora act as ecosystem engineers by increasing substratum complexity and providing interstitial habitat for benthic macroinvertebrates. 2. We manipulated the topography and perimeter-to-area ratio of patches of dreissenid mussels in a series of colonisation experiments conducted at two sites in the St. Lawrence River. Experimental substrata were variably colonised by Cladophora, prompting us to examine (i) how the topography of Dreissena patches affects benthic macroinvertebrate diversity and (ii) the extent to which the effects of Dreissena are altered by the presence of another habitat-modifying organism (Cladophora). 3. The results of our first experiment suggested that a patchy distribution of dreissenid mussels is an important driver of benthic diversity at small spatial scales. The results of our second and third experiments suggested that a native habitat engineer, Cladophora, modifies the impact of Dreissena on benthic macroinvertebrate communities. 4. While macroalgal blooms have been linked to the large-scale impacts of Dreissena on light and nutrient availability, Dreissena shells inhibited Cladophora growth at our experimental scale. These findings demonstrate that the interactions between habitatmodifying species can complicate efforts to predict the community-level effects of an invasion.