Benthic foraminifera populate a diverse range of marine habitats. Their ability to use alternative electron acceptors—nitrate (NO3−) or oxygen (O2)—makes them important mediators of benthic nitrogen cycling. Nevertheless, the metabolic scaling of the two alternative respiration pathways and the environmental determinants of foraminiferal denitrification rates are yet unknown. We measured denitrification and O2 respiration rates for 10 benthic foraminifer species sampled in the Peruvian oxygen minimum zone (OMZ). Denitrification and O2 respiration rates significantly scale sublinearly with the cell volume. The scaling is lower for O2 respiration than for denitrification, indicating that NO3− metabolism during denitrification is more efficient than O2 metabolism during aerobic respiration in foraminifera from the Peruvian OMZ. The negative correlation of the O2 respiration rate with the surface/volume ratio is steeper than for the denitrification rate. This is likely explained by the presence of an intracellular NO3− storage in denitrifying foraminifera. Furthermore, we observe an increasing mean cell volume of the Peruvian foraminifera, under higher NO3− availability. This suggests that the cell size of denitrifying foraminifera is not limited by O2 but rather by NO3− availability. Based on our findings, we develop a mathematical formulation of foraminiferal cell volume as a predictor of respiration and denitrification rates, which can further constrain foraminiferal biogeochemical cycling in biogeochemical models. Our findings show that NO3− is the preferred electron acceptor in foraminifera from the OMZ, where the foraminiferal contribution to denitrification is governed by the ratio between NO3− and O2.

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Abstract. The intraseasonal evolution of physical and biogeochemical properties during a coastal trapped wave event off central Peru is analysed using data from an extensive shipboard observational programme conducted between April and June 2017, and remote sensing data. The poleward velocities in the Peru–Chile Undercurrent were highly variable and strongly intensified to above 0.5 m s−1 between the middle and end of May. This intensification was likely caused by a first-baroclinic-mode downwelling coastal trapped wave, excited by a westerly wind anomaly at the Equator and originating at about 95∘ W. Local winds along the South American coast did not impact the wave. Although there is general agreement between the observed cross-shore-depth velocity structure of the coastal trapped wave and the velocity structure of first vertical mode solution of a linear wave model, there are differences in the details of the two flow distributions. The enhanced poleward flow increased water mass advection from the equatorial current system to the study site. The resulting shorter alongshore transit times between the Equator and the coast off central Peru led to a strong increase in nitrate concentrations, less anoxic water, likely less fixed nitrogen loss to N2 and a decrease of the nitrogen deficit compared to the situation before the poleward flow intensification. This study highlights the role of changes in the alongshore advection due to coastal trapped waves for the nutrient budget and the cumulative strength of N cycling in the Peruvian oxygen minimum zone. Enhanced availability of nitrate may impact a range of pelagic and benthic elemental cycles, as it represents a major electron acceptor for organic carbon degradation during denitrification and is involved in sulfide oxidation in sediments.

The Peruvian Upwelling System is characterized by high primary productivity fuelled by the supply of nutrients in a highly dynamic boundary circulation. The intraseasonal evolution of the physical and biogeochemical properties is 10 analysed based on shipboard observations and remote sensing conducted between April and June 2017 off central Peru. The poleward transport in the subsurface Peru Chile Undercurrent was highly variable and strongly intensified between mid and end of May. This intensification was likely caused by a first baroclinic mode downwelling coastal trapped wave excited at the equator at about 95°W that propagated poleward along the South American coast. The intensified poleward flow shortens the time of water mass advection from the equatorial current system to the study site. The impact of the anomalous advection 15 is mostly noticed in the nitrogen cycle because during the shorter time needed for poleward advection less fixed nitrogen loss occurs within the waters. This causes a strong increase of nitrate concentrations and a decrease in the nitrogen deficit. These changes suggest that the advection caused by the coastal trapped wave supersedes the simultaneous effect of anomalous downwelling in terms of nutrient response.