Antarctic krill (Euphausia superba) are swarming, oceanic crustaceans, up to two inches long, and best known as prey for whales and penguinsbut they have another important role. With their large size, high biomass and daily vertical migrations they transport and transform essential nutrients, stimulate primary productivity and influence the carbon sink. Antarctic krill are also fished by the Southern Ocean's largest fishery. Yet how krill fishing impacts nutrient fertilisation and the carbon sink in the Southern Ocean is poorly understood. Our synthesis shows fishery management should consider the influential biogeochemical role of both adult and larval Antarctic krill. O cean biogeochemical cycles are paramount in regulating atmospheric carbon dioxide (CO 2) levels and in governing the nutrients available for phytoplankton growth 1. As phytoplankton are essential in most marine food webs, biogeochemistry is also important in fuelling fishery production 2. The role of phytoplankton in atmospheric CO 2 drawdown and fish production has been the central focus of many biogeochemical studies (e.g., refs. 3,4). However, despite evidence of their potential importance, higher organisms (metazoa) such as zooplankton (e.g., copepods and salps), nekton (e.g., adult krill and fish), seabirds and mammals 5-12 , have received less attention concerning their roles in the global biogeochemical cycles. One of the main mechanisms by which metazoa can influence biogeochemical cycles is through the biological pump 1 (Fig. 1). The biological pump describes a suite of biological processes that ultimately sequester atmospheric CO 2 into the deep ocean on long timescales. During photosynthesis in the surface, ocean phytoplankton produce organic matter and a fraction (< 40 %) sinks to deeper waters 13. It is estimated that 5-12 Gt C is exported from the global surface ocean annually 14 , with herbivorous metazoa contributing to the biological pump by releasing fast-sinking faecal pellets, respiring carbon at depth originally assimilated in the surface ocean and by excreting nutrients near the surface promoting further phytoplankton

Fecal pellets (FP) are a key component of the biological carbon pump, as they can, under some circumstances, efficiently transfer carbon to depth. Like other forms of particulate organic carbon (POC), they can be remineralized in the ocean interior (particularly in the upper 200 m), or alternatively they can be preserved in the sediments. The controls on the attenuation of FP flux with depth are not fully understood, in particular, the relative contributions of zooplankton fragmentation and microbial/zooplankton respiration to FP loss. Collection of sinking particles using Marine Snow Catchers at three ecologically contrasting sites in the Scotia Sea, Antarctica, revealed large differences in POC flux composition (5-96% FP) and flux attenuation despite similar temperatures. To determine the importance of microbial respiration on FP loss in the upper mesopelagic, we made the first ever measurements of small scale oxygen gradients through the boundary layer at the interface of krill FP collected from the Scotia Sea. Estimated carbon-specific respiration rates of microbes within FP (0.010-0.065 d 21 ) were too low to account for the observed large decreases in FP flux over the upper 200 m. Therefore, the observed rapid declines in downward FP flux in the upper mesopelagic are more likely to be caused by zooplankton, through coprophagy, coprorhexy, and coprochaly. Microbial respiration is likely to be more important in regions of higher temperatures, and at times of the year, or in depths of the ocean, where zooplankton abundances are low and therefore grazing and fragmentation processes are reduced.

The efficiency of the ocean's biological carbon pump (BCP eff -here the product of particle export and transfer efficiencies) plays a key role in the air-sea partitioning of CO 2 . Despite its importance in the global carbon cycle, the biological processes that control BCP eff are poorly known. We investigate the potential role that zooplankton play in the biological carbon pump using both in situ observations and model output. Observed and modelled estimates of fast, slow, and total sinking fluxes are presented from three oceanic sites: the Atlantic sector of the Southern Ocean, the temperate North Atlantic, and the equatorial Pacific oxygen minimum zone (OMZ). We find that observed particle export efficiency is inversely related to primary production likely due to zooplankton grazing, in direct contrast to the model estimates. The model and observations show strongest agreement in remineralization coefficients and BCP eff at the OMZ site where zooplankton processing of particles in the mesopelagic zone is thought to be low. As the model has limited representation of zooplankton-mediated remineralization processes, we suggest that these results point to the importance of zooplankton in setting BCP eff , including particle grazing and fragmentation, and the effect of diel vertical migration. We suggest that improving parameterizations of zooplankton processes may increase the fidelity of biogeochemical model estimates of the biological carbon pump. Future changes in climate such as the expansion of OMZs may decrease the role of zooplankton in the biological carbon pump globally, hence increasing its efficiency.Published by Copernicus Publications on behalf of the European Geosciences Union.

The biological carbon pump drives a flux of particulate organic carbon (POC) through the ocean and affects atmospheric levels of carbon dioxide. Short term, episodic flux events are hard to capture with current observational techniques and may thus be underrepresented in POC flux estimates. We model the potential hidden flux of POC originating from Antarctic krill, whose swarming behaviour could result in a major conduit of carbon to depth through their rapid exploitation of phytoplankton blooms and bulk egestion of rapidly sinking faecal pellets (FPs). Our model results suggest a seasonal krill FP export flux of 0.039 GT C across the Southern Ocean marginal ice zone, corresponding to 17–61% (mean 35%) of current satellite-derived export estimates for this zone. The magnitude of our conservatively estimated flux highlights the important role of large, swarming macrozooplankton in POC export and, the need to incorporate such processes more mechanistically to improve model projections.

The remineralization depth of particulate organic carbon (POC) fluxes exported from the surface ocean exerts a major control over atmospheric CO₂ levels. According to a long‐held paradigm most of the POC exported to depth is associated with large particles. However, recent lines of evidence suggest that slow‐sinking POC (SSPOC) may be an important contributor to this flux. Here we assess the circumstances under which this occurs. Our study uses samples collected using the Marine Snow Catcher throughout the Atlantic Ocean, from high latitudes to midlatitudes. We find median SSPOC concentrations of 5.5 μg L−1, 13 times smaller than suspended POC concentrations and 75 times higher than median fast‐sinking POC (FSPOC) concentrations (0.07 μg L−1). Export fluxes of SSPOC generally exceed FSPOC flux, with the exception being during a spring bloom sampled in the Southern Ocean. In the Southern Ocean SSPOC fluxes often increase with depth relative to FSPOC flux, likely due to midwater fragmentation of FSPOC, a process which may contribute to shallow mineralization of POC and hence to reduced carbon storage. Biogeochemical models do not generally reproduce this behavior, meaning that they likely overestimate long‐term ocean carbon storage.

Abstract. Atmospheric levels of carbon dioxide are tightly linked to the depth at which sinking particulate organic carbon (POC) is remineralised in the ocean. Rapid attenuation of downward POC flux typically occurs in the upper mesopelagic (top few hundred metres of the water column), with much slower loss rates deeper in the ocean. Currently, we lack understanding of the processes that drive POC attenuation, resulting in large uncertainties in the mesopelagic carbon budget. Attempts to balance the POC supply to the mesopelagic with respiration by zooplankton and microbes rarely succeed. Where a balance has been found, depth-resolved estimates reveal large compensating imbalances in the upper and lower mesopelagic. In particular, it has been suggested that respiration by free-living microbes and zooplankton in the upper mesopelagic are too low to explain the observed flux attenuation of POC within this layer. We test the hypothesis that particle-associated microbes contribute significantly to community respiration in the mesopelagic, measuring particle-associated microbial respiration of POC in the northeast Atlantic through shipboard measurements on individual marine snow aggregates collected at depth (36–500 m). We find very low rates of both absolute and carbon-specific particle-associated microbial respiration (< 3 % d−1), suggesting that this term cannot solve imbalances in the upper mesopelagic POC budget. The relative importance of particle-associated microbial respiration increases with depth, accounting for up to 33 % of POC loss in the mid-mesopelagic (128–500 m). We suggest that POC attenuation in the upper mesopelagic (36–128 m) is driven by the transformation of large, fast-sinking particles to smaller, slow-sinking and suspended particles via processes such as zooplankton fragmentation and solubilisation, and that this shift to non-sinking POC may help to explain imbalances in the mesopelagic carbon budget.

) and attenuation to estimates of krill FP production based on previous measurements of krill density and literature FP egestion rates, and estimated net krill FP attenuation rates in the upper mesopelagic. Calculated attenuation rates are sensitive to krill densities in the overlying water column but suggest that krill FP could be transferred efficiently through the upper mesopelagic, and, in agreement with our MSC attenuation estimates, could make large contributions to bathypelagic POC fluxes. Our study contrasts with some others which suggest rapid FP attenuation, highlighting the need for further work to constrain attenuation rates and assess how important the contribution of Antarctic krill FP could be to the Southern Ocean biological carbon pump.

&lt;p&gt;&lt;strong&gt;Abstract.&lt;/strong&gt; The efficiency of the ocean&amp;#8217;s biological carbon pump (BCP&lt;i&gt;eff&lt;/i&gt; &amp;#8211; here the product of particle export and transfer efficiencies) plays a key role in the air-sea partitioning of CO&lt;sub&gt;2&lt;/sub&gt;. Despite its importance in the global carbon cycle, the biological processes that control BCP&lt;i&gt;eff&lt;/i&gt; are poorly known. We investigate the potential role that zooplankton play in the biological carbon pump using both &lt;i&gt;in situ&lt;/i&gt; observations and model output. Observed and modelled estimates of fast, slow and total sinking fluxes are presented from three oceanic sites: the Atlantic sector of the Southern Ocean, the temperate North Atlantic and the equatorial Pacific oxygen minimum zone (OMZ). We find that observed particle export efficiency is inversely related to primary production likely due to zooplankton grazing, in direct contrast to the model estimates. The model and observations show strongest agreement in remineralization coefficients and BCP&lt;i&gt;eff&lt;/i&gt; at the OMZ site where zooplankton processing of particles in the mesopelagic zone is thought to be low. As the model has limited representation of zooplankton-mediated remineralization processes, we suggest that these results point to the importance of zooplankton in setting BCP&lt;i&gt;eff&lt;/i&gt;, including particle grazing and fragmentation, and the effect of diel vertical migration. We suggest that improving parameterizations of zooplankton processes may increase the fidelity of biogeochemical model estimates of the biological carbon pump. Future changes in climate such as the expansion of OMZs may decrease the role of zooplankton in the biological carbon pump globally, hence increasing its efficiency.&lt;/p&gt;