In situ camera observations reveal major role of zooplankton in modulating marine snow formation during an upwelling-induced plankton bloom

Particle aggregation and the consequent formation of marine snow alter important properties of biogenic particles (size, sinking rate, degradability), thus playing a key role in controlling the vertical flux of organic matter to the deep ocean. However, there are still large uncertainties about rates and mechanisms of particle aggregation, as well as the role of plankton community structure in modifying biomass transfer from small particles to large fast-sinking aggregates. Here we present data from a high-resolution underwater camera system that we used to observe particle size distributions and formation of marine snow (aggregates >0.5 mm) over the course of a 9-week in situ mesocosm experiment in the Eastern Subtropical North Atlantic. After an oligotrophic phase of almost 4 weeks, addition of nutrient-rich deep water (650 m) initiated the development of a pronounced diatom bloom and the subsequent formation of large marine snow aggregates in all 8 mesocosms. We observed a substantial time lag between the peaks of chlorophyll a and marine snow biovolume of 9–12 days, which is much longer than previously reported and indicates a marked temporal decoupling of phytoplankton growth and marine snow formation during our study. Despite this time lag, our observations revealed substantial transfer of biomass from small particle sizes (single phytoplankton cells and chains) to marine snow aggregates of up to 2.5 mm diameter (ESD), with most of the biovolume being contained in the 0.5–1 mm size range. Notably, the abundance and community composition of mesozooplankton had a substantial influence on the temporal development of particle size spectra and formation of marine snow aggregates: While higher copepod abundances were related to reduced aggregate formation and biomass transfer towards larger particle sizes, the presence of appendicularia and doliolids enhanced formation of large marine snow. Furthermore, we combined in situ particle size distributions with measurements of particle sinking velocity to compute instantaneous (potential) vertical mass flux. However, somewhat surprisingly, we did not find a coherent relationship between our computed flux and measured vertical mass flux (collected by sediment traps in 15 m depth). Although the onset of measured vertical flux roughly coincided with the emergence of marine snow, we found substantial variability in mass flux among mesocosms that was not related to marine snow numbers, and was instead presumably driven by zooplankton-mediated alteration of sinking biomass and export of small particles (fecal pellets). Altogether, our findings highlight the role of zooplankton community composition and feeding interactions on particle size spectra and formation of marine snow aggregates, with important implications for our understanding of particle aggregation and vertical flux of organic matter in the ocean