Description: The northern Humboldt Current upwelling system (HCS) belongs to the most productive marine ecosystems, providing five to eight times higher fisheries landings per unit area than other coastal upwelling systems. To solve this “Peruvian puzzle”, to elucidate the pelagic food-web structure and to better understand trophic interactions in the HCS, a combined stable isotope and fatty acid trophic biomarker approach was adopted for key zooplankton taxa and higher trophic positions with an extensive spatial coverage from 8.5 to 16°S and a vertical range down to 1,000 m depth. A pronounced regional shift by up to ∼5‰ in the δ15N baseline of the food web occurred from North to South. Besides regional shifts, δ15N ratios of particulate organic matter (POM) also tended to increase with depth, with differences of up to 3.8‰ between surface waters and the oxygen minimum zone. In consequence, suspension-feeding zooplankton permanently residing at depth had up to ∼6‰ higher δ15N signals than surface-living species or diel vertical migrants. The comprehensive data set covered over 20 zooplankton taxa and indicated that three crustacean species usually are key in the zooplankton community, i.e., the copepods Calanus chilensis at the surface and Eucalanus inermis in the pronounced OMZ and the krill Euphausia mucronata, resulting in an overall low number of major trophic pathways toward anchovies. In addition, the semi-pelagic squat lobster Pleuroncodes monodon appears to play a key role in the benthic-pelagic coupling, as indicated by highest δ13C’ ratios of −14.7‰. If feeding on benthic resources and by diel vertical migration, they provide a unique pathway for returning carbon and energy from the seafloor to the epipelagic layer, increasing the food supply for pelagic fish. Overall, these mechanisms result in a very efficient food chain, channeling energy toward higher trophic positions and partially explaining the “Peruvian puzzle” of enormous fish production in the HCS.

Description: Zooplankton plays a notable role in ocean biogeochemical cycles. However, it is often simulated as one generic group and top closure term in ocean biogeochemical models. This study presents the description of three zooplankton functional types (zPFTs, micro-, meso- and macrozooplankton) in the ocean biogeochemical model FESOM-REcoM. In the presented model, microzooplankton is a fast-growing herbivore group, mesozooplankton is another major consumer of phytoplankton, and macrozooplankton is a slow-growing group with a low temperature optimum. Meso- and macrozooplankton produce fast-sinking fecal pellets. With three zPFTs, the annual mean zooplankton biomass increases threefold to 210 Tg C. The new food web structure leads to a 25% increase in net primary production and a 10% decrease in export production globally. Consequently, the export ratio decreases from 17% to 12% in the model. The description of three zPFTs reduces model mismatches with observed dissolved inorganic nitrogen and chlorophyll concentrations in the South Pacific and the Arctic Ocean, respectively. Representation of three zPFTs also strongly affects phytoplankton phenology: Fast nutrient recycling by zooplankton sustains higher chlorophyll concentrations in summer and autumn. Additional zooplankton grazing delays the start of the phytoplankton bloom by 3 weeks and controls the magnitude of the bloom peak in the Southern Ocean. As a result, the system switches from a light-controlled Sverdrup system to a dilution-controlled Behrenfeld system. Overall, the results suggest that representation of multiple zPFTs is important to capture underlying processes that may shape the response of ecosystems and ecosystem services to on-going and future environmental change in model projections.