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Multiple processes transport carbon into the deep ocean as part of the biological carbon pump, leading to long-term carbon sequestration. However, our ability to predict future changes in these processes is hampered by the absence of studies that have simultaneously quantified all carbon pump pathways. Here, we quantify carbon export and sequestration in the California Current Ecosystem resulting from (1) sinking particles, (2) active transport by diel vertical migration, and (3) the physical pump (subduction + vertical mixing of particles). We find that sinking particles are the most important and export 9.0 mmol C m −2 d −1 across 100-m depth while sequestering 3.9 Pg C. The physical pump exports more carbon from the shallow ocean than active transport (3.8 vs. 2.9 mmol C m −2 d −1 ), although active transport sequesters more carbon (1.0 vs. 0.8 Pg C) because of deeper remineralization depths. We discuss the implications of these results for understanding biological carbon pump responses to climate change. Biological carbon pump pathways combine to transport organic carbon into the deep ocean. This study shows that sinking particles sequester 4 Pg C, active transport sequesters 1 Pg C, and subduction sequesters 0.8 Pg C in the California Current Ecosystem.

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. Food-web and isotope-based trophic analyses suggest that blooms of salps (gelatinous grazers) can increase ecosystem efficiency and energy transfer to large organisms. These compensatory food-web dynamics may offset predicted trophic amplification.

Phytoplankton community structure is crucial to pelagic food webs and biogeochemical processes. Understanding size-based biomass distribution and carbon dynamics is essential for assessing their contributions to oceanic carbon cycling. This study quantifies plankton carbon (C) based size spectra, community composition, living to total particulate organic carbon (POC) and C:Chlorophyll a (C:Chl a ) ratios across biogeographical provinces in the Pacific sector of the Southern Ocean near the Subtropical Front (Chatham Rise, Aotearoa-New Zealand). We analyzed phytoplankton community composition using epifluorescence microscopy and flow cytometry, while quantifying size-fractionated Chl- a and POC to estimate normalized biomass, abundance size spectra, and C:Chl a ratios. On average, subtropical-influenced waters had lower macronutrients, higher total Chl a (1.1 ± 0.2 μg Chl a L -1 ) and were dominated by nanoplankton, which accounted for 45% of the total plankton community (35.2 ± 4.6 μg C L -1 ). In contrast, picoplankton dominated plankton communities within the subantarctic-influenced and accounted for 35% of the total plankton community (18.5 ± 0.9 μg C L -1 ) in these water with higher macronutrient concentrations and lower total Chl a concentrations (0.32 ± 0.06 μg Chl a L -1 ). Subantarctic-influenced regions had steeper (more negative) slopes for the normalized biomass size spectrum (average = -1.00) compared to subtropical-influenced waters (average = -0.78) indicating greater relative dominance of small taxa. The subantarctic-influenced region had ~2-fold higher surface average C:Chl a ratios compared to the subtropical-influenced region with picoplankton consistently having lower C:Chl a ratios, due to low Chl a values, than larger nano- or microplankton. Live plankton carbon contributed a median of 67% of total particulate organic carbon in the euphotic zone (non-living detritus comprises the remaining ~1/3), which is indicative of substantial primary production and rapid recycling by a strong microbial loop. Our study provides important insights into phytoplankton community structure, biomass distribution and their contribution to carbon sequestration in this region, highlighting the important roles of nanoplankton in subtropical productive waters and picoplankton in offshore subantarctic waters as well as a strong variation of C:Chl a across different phytoplankton size classes.

Sinking particles transport organic carbon produced in the surface ocean to the ocean interior, leading to net storage of atmospheric CO2 in the deep ocean. The rapid growth of in situ imaging technology has the potential to revolutionize our understanding of particle flux attenuation in the ocean; however, estimating particle flux from particle size and abundance (measured directly by in situ cameras) is challenging. Sinking rates are dependent on several factors, including particle excess density and porosity, which vary based on particle origin and type. Additionally, particle characteristics are transformed while sinking. We compare optically-measured particle size spectra profiles (Underwater Vision Profiler 5, UVP) with contemporaneous measurements of particle flux made using sediment traps and 234Th:238U disequilibrium on six process cruises from the California Current Ecosystem (CCE) LTER Program. These measurements allow us to assess the efficacy of using size-flux relationships to estimate fluxes from optical particle size measurements. We find that previously published parameterizations that estimate carbon flux from UVP profiles are a poor fit to direct flux measurements in the CCE. This discrepancy is found to result primarily from the important role of fecal pellets in particle flux. These pellets are primarily in a size range (i.e., 100 – 400 µm) that is not well-resolved as images by the UVP due to the resolution of the sensor. We develop a new, CCE-optimized algorithm for estimating carbon flux from UVP data in the southern California Current (Flux = ∑_(i=1)^x▒〖n_i A〖d_i〗^B ∆d_i 〗), with A = 13.45, B = 1.35, d = particle diameter (mm) and Flux in units of mg C m-2 d-1. We caution, however, that increased accuracy in flux estimates derived from optical instruments will require devices with greater resolution, the ability to differentiate fecal pellets from low porosity marine snow aggregates, and improved sampling of rapidly sinking fecal pellets. We also find that the particle size-flux relationships may be different within the euphotic zone than in the shallow twilight zone and hypothesize that the changing nature of sinking particles with depth must be considered when investigating the remineralization length scale of sinking particles in the ocean.

Salp grazing is important in shaping planktonic food-web structure. However, little is known about the size ranges of their prey in the field or how grazing impacts size structure. This study investigated the feeding habits of seven different species of salps, representing a variety of sizes and life stages across subtropical and subantarctic waters east of New Zealand. Scanning electron microscopy was used to examine the gut contents of 58 salps, which were then compared to water column plankton communities characterized via epifluorescence microscopy, FlowCam, and flow cytometry. While most of the gut contents resembled ambient waters, substantial differences were found amongst some co-occurring species, such as increased retention of submicron bacteria amongst smaller salps like Thalia democratica. We found that even for those salps capable of feeding on bacteria efficiently, nanoplankton and small microplankton still made up the majority of gut biomass. Larger microplankton were rarer in the guts than in the water column, potentially suggesting an upper size-threshold in addition to the lower size-threshold that has been the focus of most previous work. Salp carbon-weighted predator to prey size ratios were variable, with the majority falling between 1000:1 and 10,000:1 depending largely on the size of the salp. Taken together our results indicate that despite being able to feed on submicron particles, picoplankton make up at most 26.4% (mean = 6.4%) of salp gut carbon and are relatively unimportant to the energetics of most salps in this region compared to nanoplankton such as small dinoflagellates and diatoms.

This study was made possible by funding from the Ministry for Business, Innovation and Employment (MBIE) of New Zealand, NIWA Coast and Oceans Food Webs (COES) and Ocean Flows (COOF), and the Royal Society of New Zealand Marsden Fast-track award to MD, and by U.S. National Science Foundation awards OCE- 1756610 and 1756465 to MS and KES.

Model code is available as a zip file (Supplementary Dataset 6) that includes a Matlab live script, which can be used to generate all figures and summary data in the manuscript. It is also available on GitHub: https://github.com/mstukel/SalpEcosystemEfficiency and on Zenodo: https://doi.org/10.5281/zenodo.12700512.

Salp grazing is important in shaping planktonic food-web structure. However, little is known about the size ranges of their prey in the field or how grazing impacts size structure. This study investigated the feeding habits of both the solitary and aggregate life stages of 7 different species of salps, representing a variety of sizes across subtropical and subantarctic waters east of New Zealand. Scanning electron microscopy was used to examine the gut contents of 58 salps, which were then compared to water column plankton communities characterized via epifluorescence microscopy, FlowCam, and flow cytometry. While most of the gut contents resembled ambient waters, substantial differences were found amongst some co-occurring species, such as increased retention of submicron bacteria by Thalia democratica. We found that even for those salps capable of feeding on bacteria efficiently, nanoplankton and small microplankton still made up the majority of gut biomass. Larger microplankton were rarer in the guts than in the water column, suggesting an upper size-threshold in addition to the lower size-threshold that has been the focus of most previous work. Salp carbon-weighted predator to prey size ratios were variable, with the majority falling between 1,000:1 and 10,000:1 depending largely on the size of the salp. Taken together our results indicate that despite being able to feed on submicron particles, picoplankton make up at most 26.4% (mean = 6.4%) of salp gut carbon and are relatively unimportant to the energetics of most salps in this region compared to nanoplankton such as small dinoflagellates and diatoms. Competing Interest Statement The authors have declared no competing interest. Footnotes * Additional clarification on PPSR and SDPPSR added to Introduction and Methods. Updates made to most values and resulting figures. Other minor changes to reflect edits by coauthors prior to submission for publication. Supplemental files also updated.

Whereas recruitment success for many fisheries depends on coincident timing of larvae with abundance peaks of their prey, less can be more in the tropical/subtropical spawning areas of bluefin tunas if lower but steady food resources are offset by reduced larval vulnerability to pelagic predators. To understand larval habitat characteristics for Southern Bluefin Tuna (SBT), we quantified microbial community carbon flows based on growth and grazing rates from depth profiles of dilution incubations and carbon biomass assessments from microscopy and flow cytometry (FCM) during their peak spawning off NW Australia (Indian Ocean) in February 2022. Two Chla-based estimates of phytoplankton production gave differing offsets due to cycling or mixotrophy, exceeding 14C net community production on average (677 ± 98 versus 447 ± 43 mg C m−2 d−1). Productivity was higher than in the Gulf of Mexico spawning area for Atlantic Bluefin Tuna but less than similar studies of oceanic upwelling regions. Microzooplankton grazing averaged 482 ± 63 mg C m− 2 d−1 (71 ± 13% of production). Two measurement variables for Prochlorococcus gave average production and grazing rates of 282 ± 36 and 248 ± 32 mg C m−2 d−1 (86 ± 6% grazed). Prochlorococcus comprised almost half of production and grazing fluxes in the upper (0-25 m) euphotic zone where SBT larvae reside. Prochlorococcus declined and eukaryotic phytoplankton and heterotrophic bacteria increased in relative importance in the lower euphotic zone. These results describe relatively classic open-ocean oligotrophic conditions as the food web base for nutritional flows to SBT larvae.