Reconstructing past biological invasions: niche shifts in response to invasive predators and competitors

Studying historic invasions can provide insight into the ongoing invasions that threaten global biodiversity. In this study, we reconsider the impacts of Littorina littorea and Carcinus maenas on the rocky intertidal community of the Gulf of Maine. Past research using invader-removal experiments demonstrated strong top-down effects of L. littorea on algal community structure; however, such removal experiments may overlook the long-term effects of niche shifts and local extinctions caused by invasive species. We considered how a niche-shift in the native littorine, Littorina saxatilis, may change the interpretation of L. littorea impacts. Using a factorial experiment crossing predator presence/absence with L. littorea presence/absence, we found that L. saxatilis is able to exert top-down control on ephemeral algae similar to that exerted by L.␣littorea and that both competition by L. littorea and predation by C. maenas have strong, negative impacts on L. saxatilis. We also found higher predation rates on protected shores and at lower tidal heights and preferential predation on L.␣saxatilis compared to L. littorea. While movement experiments demonstrate that behavioral response to tidal height is the proximate cause of L. saxatilis exclusion from the lower intertidal, our study suggests that the ultimate causes are the additive effects of competition from and predation by invasive species.