Contrasting patterns of spread in interacting invasive species: Membranipora membranacea and Codium fragile off Nova Scotia

In the Northwest Atlantic, overgrowth of the competitively dominant, native kelps by an invasive bryozoan Membranipora membranacea increases frond erosion, which has facilitated the establishment and spread of the invasive macroalga Codium fragile ssp fragile. To document the spread of both introduced species along the Atlantic coast of Nova Scotia from initial introduction points (the 'epicentre') southwest of Halifax, we conducted video-surveys of shallow rocky habitats along the southwestern shore of Nova Scotia (100 km linear distance, encompassing the range of M. membranacea) in 2000, and then along the entire Atlantic coast in 2007 (650 km). Membranipora membranacea was observed continuously throughout the surveyed ranges in 2000 and 2007, wherever kelps were present, suggesting natural dispersal via planktonic larvae. Codium fragile was observed along 95 km of the surveyed range in 2000 and along 445 km in 2007, with a relatively patchy distribution beyond the epicentre, suggesting a combination of natural and anthropogenic dispersal mechanisms. Rockweed-dominated (Fucus spp.) or mixed algal assemblages common outside the epicentre may alter the interaction between M. membranacea and C. fragile, since seaweeds other than kelp are not subject to defoliation by the bryozoan. Percent cover of kelp at the epicentre generally increased from 2000 to 2007, while that of C. fragile generally decreased. Codium fragile was the dominant canopy alga at 54% of sites in 2000 and at only 15% of sites in 2007. These findings indicate that, at near decadal timescales, C. fragile does not prevent re-colonization by native kelps.