Multiple effects of a Gracilaria vermiculophylla invasion on estuarine mudflat functioning and diversity

The invasive Japanese seaweed Gracilaria vermiculophylla has become established over the past several years in numerous European estuaries, from Portugal to Norway. In the Faou estuary (48.295°N-4.179°W, Brittany, France), it forms a dense population at the mud's surface. The effects of G. vermiculophylla on metabolism, diversity, and the food web were studied. Community gross primary production (GPP) and respiration (CR) during emersion, chlorophyll-a content, macrofaunal and meiofaunal diversity and abundance, and stable isotopes (δ13C and δ15N) of representative macrofaunal species and main food sources were measured at low tide in winter, spring, summer 2014, and winter 2015. Results show significant seasonal variation in GPP and CR. Moreover, GPP was significantly higher in areas where G. vermiculophylla was present than in the control area (bare mud). However, this high GPP appeared to be linked to the increase in biomass in primary producers, with their efficiency (primary productivity, i.e. assimilation number) remaining relatively stable compared with the control area. Significant variation in abundance of meiofauna and macrofauna was also detected and new epifaunal species were collected, mainly in Gracilaria-colonized areas. Isotopic food-web Bayesian mixing models strongly suggested that G. vermiculophylla plays a major role in the diet of some dominant species. Mechanisms interacting with the functioning and diversity of the mudflat are discussed. Finally, the invasive seaweed G. vermiculophylla affected the mudflat ecosystem in three ways: as a new primary producer (increase in metabolism), as a habitat-forming species (changes in diversity and abundance of macrofauna and meiofauna), and as a new abundant food source, likely through the detrital pathway.