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Abstract The eastern Indian Ocean is substantially under sampled with respect to the biological carbon pump – the suite of processes that transport the carbon fixed by phytoplankton into the deeper ocean. Using sediment traps and other ecosystem measurements, we quantified sinking organic matter flux and investigated the characteristics of sinking particles in waters overlying the Argo Abyssal Plain directly downstream of the Indonesian Throughflow off northwest Australia. Carbon export from the euphotic zone averaged 7.0 mmol C m^-2d^-1, which equated to an average export efficiency (export / net primary production) of 0.17. Sinking particle flux within the euphotic zone (beneath the mixed layer, but above the deep chlorophyll maximum) averaged slightly higher than flux at the base of the euphotic zone, suggesting that the deep euphotic zone was a depth stratum of net particle remineralization. Carbon flux attenuation continued into the twilight zone with a transfer efficiency (export at euphotic depth + 100m / export at euphotic depth) of 0.62 and an average Martin’sb-value of 1.1. Within the euphotic zone, fresh phytoplankton (chlorophyll associated with sinking particles, possibly contained within appendicularian houses) were an important component of sinking particles, but beneath the euphotic zone the fecal pellets of herbivorous zooplankton (phaeopigments) were more important. Changes in carbon and nitrogen isotopic composition with depth further reflected remineralization processes occurring as particles sank. We show similarities with biological carbon pump functioning in a similar semi-enclosed oligotrophic marginal sea, the Gulf of Mexico, including net remineralization across the deep chlorophyll maximum. Submitted to: Deep-sea Research II HighlightsDespite low productivity, export efficiency was 17% of primary productionFlux attenuation beneath the euphotic zone (EZ) was low for a tropical regionSinking particle flux from the upper to lower EZ exceeded export from lower EZThe deep EZ was a stratum of net particle remineralization (and net heterotrophy) 

Abstract The current conventional paradigm of ocean food web structure inserts one full level or more of microzooplankton heterotrophic consumption, a substantial energy drop, between phytoplankton and mesozooplankton. Using a dataset with contemporaneous measurements of primary production (PP), size-fractioned mesozooplankton biomass, and micro- and mesozooplankton grazing rates from 10 tropical to temperate ocean ecosystems, we examined whether the structural inefficiencies in this paradigm allow sufficient energy transfer to support active metabolism and growth of observed zooplankton standing stocks. Zooplankton carbon requirements (ZCR) were determined from allometric equations that account for ecosystem differences in temperature and size structure. ZCRs were relatively low (∼30% of PP or less) for both oligotrophic systems and bloom biomass accumulation in eutrophic coastal waters. Higher relative ZCRs (>30% PP) were associated with elevated mesozooplankton grazing scenarios (bloom declines, abundant salps), advective subsidies, and open-ocean upwelling systems. Microzooplankton generally dominated as grazers of PP but were equal or secondary to direct herbivory as nutritional support for mesozooplankton in five of eight regional studies. All systems were able to satisfy ZCR within the conventional food-web interpretation, but balanced open-ocean upwelling systems required the most efficient alignments of contributions from microzooplankton grazing, direct herbivory, and carnivory to do so. 

Abstract Gelatinous filter feeders (e.g., salps, doliolids, and pyrosomes) have high filtration rates and can feed at predator:prey size ratios exceeding 10,000:1, yet are seldom included in ecosystem or climate models. We investigated foodweb and trophic dynamics in the presence and absence of salp blooms using traditional productivity and grazing measurements combined with compound-specific isotopic analysis of amino acids estimation of trophic position during Lagrangian framework experiments in the Southern Ocean. Trophic positions of salps ranging 10–132 mm in size were 2.2 ± 0.3 (mean ± std) compared to 2.6 ± 0.4 for smaller (mostly crustacean) mesozooplankton. The mostly herbivorous salp trophic position was maintained despite biomass dominance of ~10-µm-sized primary producers. We show that potential energy flux to >10-cm organisms increases by approximately an order of magnitude when salps are abundant, even without substantial alteration to primary production. Comparison to a wider dataset from other marine regions shows that alterations to herbivore communities are a better predictor of ecosystem transfer efficiency than primary-producer dynamics. These results suggest that diverse consumer communities and intraguild predation complicate climate change predictions (e.g., trophic amplification) based on linear food chains. These compensatory foodweb dynamics should be included in models that forecast marine ecosystem responses to warming and reduced nutrient supply. 

Abstract Photosynthesis in the surface ocean and subsequent export of a fraction of this fixed carbon leads to carbon dioxide sequestration in the deep ocean. Ecological relationships among plankton functional groups and theoretical relationships between particle size and sinking rate suggest that carbon export from the euphotic zone is more efficient when communities are dominated by large organisms. However, this hypothesis has never been tested against measured size spectra spanning the >5 orders of magnitude found in plankton communities. Using data from five ocean regions (California Current Ecosystem, North Pacific subtropical gyre, Costa Rica Dome, Gulf of Mexico, and Southern Ocean subtropical front), we quantified carbon‐based plankton size spectra from heterotrophic bacteria to metazoan zooplankton (size class cutoffs varied slightly between regions) and their relationship to net primary production and sinking particle flux. Slopes of the normalized biomass size spectra (NBSS) varied from −1.6 to −1.2 (median slope of −1.4 equates to large 1–10 mm organisms having a biomass equal to only 7.6% of the biomass in small 1–10 μm organisms). Net primary production was positively correlated with the NBSS slope, with a particularly strong relationship in the microbial portion of the size spectra. While organic carbon export co‐varied with NBSS slope, we found only weak evidence that export efficiency is related to plankton community size spectra. Multi‐variate statistical analysis suggested that properties of the NBSS added no explanatory power over chlorophyll, primary production, and temperature. Rather, the results suggest that both plankton size spectra and carbon export increase with increasing system productivity. 

Abstract The Southern Ocean contributes substantially to the global biological carbon pump (BCP). Salps in the Southern Ocean, in particular Salpa thompsoni , are important grazers that produce large, fast-sinking fecal pellets. Here, we quantify the salp bloom impacts on microbial dynamics and the BCP, by contrasting locations differing in salp bloom presence/absence. Salp blooms coincide with phytoplankton dominated by diatoms or prymnesiophytes, depending on water mass characteristics. Their grazing is comparable to microzooplankton during their early bloom, resulting in a decrease of ~1/3 of primary production, and negative phytoplankton rates of change are associated with all salp locations. Particle export in salp waters is always higher, ranging 2- to 8- fold (average 5-fold), compared to non-salp locations, exporting up to 46% of primary production out of the euphotic zone. BCP efficiency increases from 5 to 28% in salp areas, which is among the highest recorded in the global ocean. 

Abstract. The ability to constrain the mechanisms that transport organiccarbon into the deep ocean is complicated by the multiple physical,chemical, and ecological processes that intersect to create, transform, andtransport particles in the ocean. In this paper we develop andparameterize a data-assimilative model of the multiple pathways of thebiological carbon pump (NEMUROBCP). The mechanistic model is designedto represent sinking particle flux, active transport by vertically migratingzooplankton, and passive transport by subduction and vertical mixing, whilealso explicitly representing multiple biological and chemical propertiesmeasured directly in the field (including nutrients, phytoplankton andzooplankton taxa, carbon dioxide and oxygen, nitrogen isotopes, and234Thorium). Using 30 different data types (including standing stockand rate measurements related to nutrients, phytoplankton, zooplankton, andnon-living organic matter) from Lagrangian experiments conducted on 11cruises from four ocean regions, we conduct an objective statisticalparameterization of the model and generate 1 million different potentialparameter sets that are used for ensemble model simulations. The modelsimulates in situ parameters that were assimilated (net primary productionand gravitational particle flux) and parameters that were withheld(234Thorium and nitrogen isotopes) with reasonable accuracy. Modelresults show that gravitational flux of sinking particles and verticalmixing of organic matter from the euphotic zone are more importantbiological pump pathways than active transport by vertically migratingzooplankton. However, these processes are regionally variable, with sinkingparticles most important in oligotrophic areas of the Gulf of Mexico andCalifornia Current, sinking particles and vertical mixing roughly equivalentin productive coastal upwelling regions and the subtropical front in theSouthern Ocean, and active transport an important contributor in the easterntropical Pacific. We further find that mortality at depth is an importantcomponent of active transport when mesozooplankton biomass is high, but itis negligible in regions with low mesozooplankton biomass. Our results alsohighlight the high degree of uncertainty, particularly amongstmesozooplankton functional groups, that is derived from uncertainty in modelparameters. Indeed, variability in BCP pathways between simulations for aspecific location using different parameter sets (all with approximatelyequal misfit relative to observations) is comparable to variability in BCPpathways between regions. We discuss the implications of these results forother data-assimilation approaches and for studies that rely on non-ensemblemodel outputs. 

Abstract Marine snow, formed through the aggregation of phytoplankton and other organic matter, can be consumed by various types of zooplankton, affecting both planktonic trophic dynamics and the export of carbon to depth. This study focuses on how two factors—phytoplankton growth phase and species—affect copepod feeding on marine snow. To do this, we conducted a series of grazing experiments using gut pigment and stable isotope methods to quantify the ingestion of the copepod, Calanus pacificus, on both marine snow aggregates and individual phytoplankton. Results demonstrate that marine snow can represent a substantial food source for copepods, comparable to rates on individual phytoplankton. Moreover, we found that both the overall ingestion and the relative ingestion of aggregates vs. individual phytoplankton depended on phytoplankton growth phase for experiments conducted with the diatom Thalassiosira weissflogii. Although copepods consumed aggregates composed of Skeletonema marinoi at similar rates as those composed of T. weissflogii, no effect of growth phase was observed for S. marinoi. These findings suggest that marine snow can be an important source of nutrition for copepods, but that its role in planktonic food webs may differ depending on the phytoplankton community composition and the stage of phytoplankton blooms.