Comparison of marine copepod outfluxes: nature, rate, fate and role in the carbon and nitrogen cycles.

We compare the nature of copepod outfluxes of nonliving matter, the factors controlling their rate and their fate, and finally their role, particularly their relative importance in the carbon and nitrogen cycle. Copepods release dissolved matter through excretion and respiration and particulate matter through production of faecal pellets, carcasses, moults, and dead eggs. Excretion liberates several organic C, N, and P compounds and inorganic N and P compounds, with inorganic compounds constituting the larger part. The faecal pellets of copepods are covered by a peritrophic membrane and have a highly variable size and content. There is less information on the nature of other copepod particulate products. The weight-specific rates of posthatch mortality, respiration, excretion, and faecal pellet production have similar C or N levels and are higher than those of moulting and egg mortality. In general, most important factors controlling these rates are temperature, body mass, food concentration, food quality, and faunistic composition. Physical and biological factors govern the vertical fate of copepod products by affecting their sedimentation speed and concentration gradient. The physical factors are sinking speed, advection, stratification, turbulent diffusion, and molecular diffusion. They influence the sedimentation speed and degradation of the copepod products. The biological factors are production, biodegradation (by zooplankton, nekton, and microorganisms) and vertical migration of copepods (diel or seasonal). Physical degradation and biodegradation by zooplankton and nekton are faster than biodegradation by microorganisms. The most important copepod outfluxes are excretion and faecal pellet production. Excretion offers inorganic nutrients that can be directly used by primary producers. Faecal pellets have a more important role in the vertical transport of elements than the other particulate products. Most investigation has focused on carbon burial in the form of copepod faecal pellets, measured by sediment traps, and on the role of ammonia excretion in nutrient recycling. Full evaluation of the role of copepod products in the transport and recycling of elements and compounds requires a quantification of all copepod products and their different fates, particularly detritiphagy, remineralization, and integration as marine snow.