Copepod feeding in the ocean: scaling patterns, composition of their diet and the bias of estimates due to microzooplankton grazing during incubations

Here, we report insights from the compilation and analysis of data on marine calanoid copepod feeding rates in the ocean. Our study shows that food availability and body weight are major factors shaping copepod feeding rates in the field, with a relatively minor role of temperature. Although the maximal feeding rates of copepods that are observed in the field agree with the well-known 3/4 of body size scaling rule for animals, copepod feeding in the oceans is typically limited and departs from this rule. Ciliates and dinoflagellates appear to be highly relevant in the composition of copepod diets, and this represents an indirect increase in the flux of primary production that is likely to reach the upper trophic levels; this contribution is higher in the less productive systems and may help to explain accounts of proportionally higher standing stocks of copepods supported per unit of primary producer biomass in oligotrophic environments. Contrary to common belief, diatoms emerge from our dataset as small contributors to the diet of copepods, except in some very productive ecosystems. We have also evaluated the bias in the estimation of copepod grazing rates due to within-bottle trophic cascade effects caused by the removal of microheterotrophs by copepods. This release of microzooplankton grazing pressure accounts for a relevant, but moderate, increase in copepod grazing estimates (ca. 20-30%); this bias has an effect on both the carbon flux budgets through copepods and on our view of their diet composition. However, caution is recommended against the indiscriminate use of corrections because they may turn out to be overestimates of the bias. We advise that both uncorrected and corrected grazing rates should be provided in future studies, as they probably correspond to the lower and upper boundaries of the true grazing rates.