Pelagic tunicates (salps, pyrosomes) and fishes generate jelly-falls and/or fecal pellets that sink roughly 10 times faster than bulk oceanic detritus, but their impacts on biogeochemical cycles in the ocean interior are poorly understood. Using a coupled physical-biogeochemical model, we find that fast-sinking detritus decreased global net primary production and surface export, but increased deep sequestration and transfer efficiency in much of the extratropics and upwelling zones. Fast-sinking detritus generally decreased total suboxic and hypoxic volumes, reducing a “large oxygen minimum zone (OMZ)” bias common in global biogeochemical models. Newly aerobic regions at OMZ edges exhibited reduced transfer efficiencies in contrast with global tendencies. Reductions in water column denitrification resulting from improved OMZs improved simulated nitrate deficits relative to phosphate. The carbon flux to the benthos increased by 11% with fast-sinking detritus from fishes and pelagic tunicates, yet simulated benthic fluxes remained on the lower end of observation-based estimates.

Phytoplankton stoichiometry modulates the interaction between carbon, nitrogen and phosphorus cycles, yet most biogeochemical models represent phytoplankton C:N:P as constants. This simplification has been linked to Earth System Model (ESM) biases and potential misrepresentation of biogeochemical responses to climate change. Here we integrate key elements of the Adaptive Trait Optimization Model (ATOM) for phytoplankton stoichiometry with the Carbon, Ocean Biogeochemistry and Lower Trophics (COBALT) ocean biogeochemical model. Within a series of global ocean-ice-ecosystem retrospective simulations, ATOM-COBALT reproduced observations of particulate organic matter N:P, and compared to static N:P, exhibited reduced phytoplankton P-limitation, enhanced N-fixation, and increased low-latitude export, leading to improved consistency with observations. Two mechanisms together drove these patterns: the growth hypothesis and frugal P-utilization during scarcity. The addition of translation compensation- differential temperature dependencies of photosynthetic relative to biosynthetic processes- led to relatively modest strengthening of N:P variations and biogeochemical responses relative to growth-plus-frugality. Comparison of the multi-mechanism model herein against frugality-only models suggest that both can capture observed N:P patterns and produce qualitatively similar biogeochemical effects. There are, however, quantitative response differences and different responses across N:P mechanisms are expected under climate change- with the growth rate mechanism adding a distinct biogeochemical footprint in highly-productive low-latitude regions. These results suggest that variable phytoplankton N:P makes some biogeochemical processes resilient to environmental changes, and support using dynamic N:P formulations with the ocean biogeochemical component of next generation of ESMs.

Although zooplankton play a substantial role in the biological carbon pump and serve as a crucial link between primary producers and higher trophic level consumers, the skillful representation of zooplankton is not often a focus of ocean biogeochemical models. Systematic evaluations of zooplankton in models could improve their representation, but so far, ocean biogeochemical skill assessment of Earth system model (ESM) ensembles have not included zooplankton. Here we use a recently developed global, observationally-based map of mesozooplankton biomass to assess the skill of mesozooplankton in six CMIP6 ESMs. We also employ a biome-based assessment of the ability of these models to reproduce the observed relationship between mesozooplankton biomass and surface chlorophyll. The combined analysis found that most models were able to reasonably simulate the large regional variations in mesozooplankton biomass at the global scale. Additionally, three of the ESMs simulated a mesozooplankton-chlorophyll relationship within the observational bounds, which we used as an emergent constraint on future mesozooplankton projections. We highlight where differences in model structure and parameters may give rise to varied mesozooplankton distributions under historic and future conditions, and the resultant wide ensemble spread in projected changes in mesozooplankton biomass. Despite differences, the strength of the mesozooplankton-chlorophyll relationships across all models was related to the projected changes in mesozooplankton biomass globally and in regional biomes. These results suggest that improved observations of mesozooplankton and their relationship to chlorophyll will better constrain projections of climate change impacts on these important animals.

The Marine Biogeochemistry Library (MARBL) is a prognostic ocean biogeochemistry model that simulates marine ecosystem dynamics and the coupled cycles of carbon, nitrogen, phosphorus, iron, silicon, and oxygen. MARBL is a component of the Community Earth System Model (CESM); it supports flexible ecosystem configuration of multiple phytoplankton and zooplankton functional types; it is also portable, designed to interface with multiple ocean circulation models. Here, we present scientific documentation of MARBL, describe its configuration in CESM2 experiments included in the Coupled Model Intercomparison Project version 6 (CMIP6), and evaluate its performance against a number of observational datasets. The model simulates an air-sea CO2 flux and many aspects of the carbon cycle in good agreement with observations. However, the simulated integrated uptake of anthropogenic CO2 is weak, which we link to poor thermocline ventilation, a feature evident in simulated chlorofluorocarbon distributions. This also contributes to larger-than-observed oxygen minimum zones. Moreover, radiocarbon distributions show that the simulated circulation in the deep North Pacific is extremely sluggish, yielding extensive oxygen depletion and nutrient trapping at depth. Surface macronutrient biases are generally positive at low latitudes and negative at high latitudes. CESM2 simulates globally-integrated net primary production (NPP) of 48 Pg C yr-1 and particulate export flux at 100 m of 7.1 Pg C yr-1. The impacts of climate change include an increase in globally-integrated NPP, but substantial declines in the North Atlantic. Particulate export is projected to decline globally, attributable to decreasing export efficiency associated with changes in phytoplankton community composition.