Temporal changes in zooplankton indicators highlight a bottom-up process in the Bay of Marseille (NW Mediterranean Sea)

Sixteen years (2005-2020) of zooplankton monitoring in the Bay of Marseille (N-W Mediterranean Sea) are analyzed in relation to physical, meteorological, climatic and biotic data. Samples were collected every two weeks by a vertical haul (0-55 m) of a 200 mu m plankton net. Different indices characterizing the mesozooplankton are compared: biomass dry weight of four size fractions between 200 and 2000 mu m; abundances of the whole of the mesozooplankton and of 13 main taxonomic groups defined from plankton imagery; seasonal onset timing of each zooplankton group; and two other types of indices: the first characterized diversity based on abundance data, and the second was derived from zooplankton size spectra shape. The clearest pattern in the environmental compartment was an overall decreasing trend in nutrients, shifts in phytoplankton metrics (i.e. size structure and particulate organic matter), and changes in winter conditions (i.e. increasing temperatures, precipitation and NAO). Interannual patterns in the mesozooplankton community were: (i) a decrease of total abundance (ii) a decrease in biomass for the four size fractions, with an earlier decrease for the 1000-2000 mu m size fraction (in 2008); (iii) a reduced dominance of copepods (calanoids and oithonoids) and a concomitant increase in abundance of other taxonomic groups (crustaceans, pteropods, chaetognaths, salps) which induced higher diversity; (iv) a first shift in size spectra towards smaller sizes in 2009, when the 1000-2000 mu m size fraction biomass decreased, and a second shift towards larger sizes in 2013 along with increased diversity; and (iv) a later onset in the phenology for some zooplankton variables and earlier onset for salps. Concomitant changes in the phytoplankton compartment, winter environmental conditions, zooplankton community structure (in size and diversity) and zooplankton phenology marked by a shift in 2013 suggest bottom-up control of the pelagic ecosystem.