Brzezinski, M. A., Krause, J. W., Bundy, R. M., Barbeau, K. A., Franks, P., Goericke, R., et al. (2015). Enhanced silica ballasting from iron stress sustains carbon export in a frontal zone within the California Current. J. Geophys. Res. Oceans, 120(7), 4654–4669.
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Kelly, T. B., Goericke, R., Kahru, M., Song, H., & Stukel, M. R. (2018). CCE II: Spatial and interannual variability in export efficiency and the biological pump in an eastern boundary current upwelling system with substantial lateral advection. Deep Sea Research Part I: Oceanographic Research Papers, 140, 14–25.
Abstract: Estimating interannual variability in carbon export is a key goal of many marine biogeochemical studies. However, due to variations in export mechanisms between regions, generalized models used to estimate global patterns in export often fail when used for intra-regional analysis. We present here a region-specific model of export production for the California Current Ecosystem (CCE) parameterized using intensive Lagrangian process studies conducted during El Niño-Southern Oscillation (ENSO) warm and neutral phases by the CCE Long-Term Ecological Research (LTER) program. We find that, contrary to expectations from prominent global algorithms, export efficiency (e-ratio = export / primary productivity) is positively correlated with temperature and negatively correlated with net primary productivity (NPP). We attribute these results to the substantial horizontal advection found within the region, and verify this assumption by using a Lagrangian particle tracking model to estimate water mass age. We further suggest that sinking particles in the CCE are comprised of a recently-produced, rapidly-sinking component (likely mesozooplankton fecal pellets) and a longer-lived, slowly-sinking component that is likely advected long distances prior to export. We determine a new algorithm for estimating particle export in the CCE from NPP (Export = 0.08 · NPP + 72 mg C m-2 d-1). We apply this algorithm to a two-decade long time series of NPP in the CCE to estimate spatial and interannual variability across multiple ENSO phases. Reduced export during the warm anomaly of 2014-2015 and El Niño 2015-2016 resulted primarily from decreased export in the coastal upwelling region of the CCE; the oligotrophic offshore region exhibited comparatively low seasonal and interannual variability in flux. The model resolves intra-regional patterns of in situ export measurements, and provides a valuable contrast to global export models.
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Kelly, T. B., Goericke, R., Kahru, M., Song, H., & Stukel, M. R. (2018). CCE II: Spatial and interannual variability in export efficiency and the biological pump in an eastern boundary current upwelling system with substantial lateral advection. Deep Sea Research Part I: Oceanographic Research Papers, 140, 14–25.
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Kranz, S. A., Wang, S., Kelly, T. B., Stukel, M. R., Goericke, R., Landry, M. R., et al. (2020). Lagrangian Studies of Marine Production: A Multimethod Assessment of Productivity Relationships in the California Current Ecosystem Upwelling Region. J. Geophys. Res. Oceans, 125(6).
Abstract: A multimethod process‐oriented investigation of diverse productivity measures in the California Current Ecosystem (CCE) Long‐Term Ecological Research study region, a complex physical environment, is presented. Seven multiday deployments covering a transition region from high to low productivity were conducted over two field expeditions (spring 2016 and summer 2017). Employing a Lagrangian study design, water parcels were followed over several days, comparing 24‐h in situ measurements (14C and 15NO3 ‐uptake, dilution estimates of phytoplankton growth, and microzooplankton grazing) with high‐resolution productivity measurements by fast repetition rate fluorometry (FRRF) and equilibrium inlet mass spectrometry (EIMS), and integrated carbon export measuremnts using sediment traps. Results show the importance of accounting for temporal and fine spatial scale variability when estimating ecosystem production. FRRF and EIMS measurements resolved diel patterns in gross primary and net community production. Diel productivity changes agreed well with comparably more traditional measurements. While differences in productivity metrics calculated over different time intervals were considerable, as those methods rely on different base assumptions, the data can be used to explain ecosystem processes which would otherwise have gone unnoticed. The processes resolved from this method comparison further understanding of temporal and spatial coupling and decoupling of surface productivity and potential carbon burial in a gradient from coastal to offshore ecosystems.
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Morrow, R. M., Ohman, M. D., Goericke, R., Kelly, T. B., Stephens, B. M., & Stukel, M. R. (2018). CCE V: Primary production, mesozooplankton grazing, and the biological pump in the California Current Ecosystem: Variability and response to El Niño. Deep Sea Research Part I: Oceanographic Research Papers, 140, 52–62.
Abstract: Predicting marine carbon sequestration in a changing climate requires mechanistic understanding of the processes controlling sinking particle flux under different climatic conditions. The recent occurrence of a warm anomaly (2014-2015) followed by an El Nino (2015-2016) in the southern sector of the California Current System presented an opportunity to analyze changes in the biological carbon pump in response to altered climate forcing. We compare primary production, mesozooplankton grazing, and carbon export from the euphotic zone during quasi-Lagrangian experiments conducted in contrasting conditions: two cruises during warm years – one during the warm anomaly in 2014 and one toward the end of El Nino 2016 – and three cruises during El Ninoneutral years. Results showed no substantial differences in the relationships between vertical carbon export and its presumed drivers (primary production, mesozooplankton grazing) between warm and neutral years. Mesozooplankton fecal pellet enumeration and phaeopigment measurements both showed that fecal pellets were the dominant contributor to export in productive upwelling regions. In more oligotrophic regions, fluxes were dominated by amorphous marine snow with negligible pigment content. We found no evidence for a significant shift in the relationship between mesozooplankton grazing rate and chlorophyll concentration. However, massspecific grazing rates were lower at low-to-moderate chlorophyll concentrations during warm years relative to neutral years. We also detected a significant difference in the relationship between phytoplankton primary production and photosynthetically active radiation between years: at similar irradiance and nutrient concentrations, productivity decreased during the warm events. Whether these changes resulted from species composition changes remains to be determined. Overall, our results suggest that the processes driving export remain similar during different climate conditions, but that species compositional changes or other structural changes require further attention.
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Morrow, R. M., Ohman, M. D., Goericke, R., Kelly, T. B., Stephens, B. M., & Stukel, M. R. (2018). CCE V: Primary production, mesozooplankton grazing, and the biological pump in the California Current Ecosystem: Variability and response to El Niño. Deep Sea Research Part I: Oceanographic Research Papers, .
Abstract: Predicting marine carbon sequestration in a changing climate requires mechanistic understanding of the processes controlling sinking particle flux under different climatic conditions. The recent occurrence of a warm anomaly (2014�2015) followed by an El Niño (2015�2016) in the southern sector of the California Current System presented an opportunity to analyze changes in the biological carbon pump in response to altered climate forcing. We compare primary production, mesozooplankton grazing, and carbon export from the euphotic zone during quasi-Lagrangian experiments conducted in contrasting conditions: two cruises during warm years – one during the warm anomaly in 2014 and one toward the end of El Niño 2016 � and three cruises during El Niño-neutral years. Results showed no substantial differences in the relationships between vertical carbon export and its presumed drivers (primary production, mesozooplankton grazing) between warm and neutral years. Mesozooplankton fecal pellet enumeration and phaeopigment measurements both showed that fecal pellets were the dominant contributor to export in productive upwelling regions. In more oligotrophic regions, fluxes were dominated by amorphous marine snow with negligible pigment content. We found no evidence for a significant shift in the relationship between mesozooplankton grazing rate and chlorophyll concentration. However, mass-specific grazing rates were lower at low-to-moderate chlorophyll concentrations during warm years relative to neutral years. We also detected a significant difference in the relationship between phytoplankton primary production and photosynthetically active radiation between years: at similar irradiance and nutrient concentrations, productivity decreased during the warm events. Whether these changes resulted from species composition changes remains to be determined. Overall, our results suggest that the processes driving export remain similar during different climate conditions, but that species compositional changes or other structural changes require further attention
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Morrow, R. M., Ohman, M. D., Goericke, R., Kelly, T. B., Stephens, B. M., & Stukel, M. R. (2018). CCE V: Primary production, mesozooplankton grazing, and the biological pump in the California Current Ecosystem: Variability and response to El Niño. Deep Sea Research Part I: Oceanographic Research Papers, 140, 52–62.
Abstract: Predicting marine carbon sequestration in a changing climate requires mechanistic understanding of the processes controlling sinking particle flux under different climatic conditions. The recent occurrence of a warm anomaly (2014-2015) followed by an El Nino (2015-2016) in the southern sector of the California Current System presented an opportunity to analyze changes in the biological carbon pump in response to altered climate forcing. We compare primary production, mesozooplankton grazing, and carbon export from the euphotic zone during quasi-Lagrangian experiments conducted in contrasting conditions: two cruises during warm years – one during the warm anomaly in 2014 and one toward the end of El Nino 2016 – and three cruises during El Ninoneutral years. Results showed no substantial differences in the relationships between vertical carbon export and its presumed drivers (primary production, mesozooplankton grazing) between warm and neutral years. Mesozooplankton fecal pellet enumeration and phaeopigment measurements both showed that fecal pellets were the dominant contributor to export in productive upwelling regions. In more oligotrophic regions, fluxes were dominated by amorphous marine snow with negligible pigment content. We found no evidence for a significant shift in the relationship between mesozooplankton grazing rate and chlorophyll concentration. However, massspecific grazing rates were lower at low-to-moderate chlorophyll concentrations during warm years relative to neutral years. We also detected a significant difference in the relationship between phytoplankton primary production and photosynthetically active radiation between years: at similar irradiance and nutrient concentrations, productivity decreased during the warm events. Whether these changes resulted from species composition changes remains to be determined. Overall, our results suggest that the processes driving export remain similar during different climate conditions, but that species compositional changes or other structural changes require further attention.
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Stukel, M. R., Aluwihare, L. I., Barbeau, K. A., Chekalyuk, A. M., Goericke, R., Miller, A. J., et al. (2017). Mesoscale ocean fronts enhance carbon export due to gravitational sinking and subduction. Proc Natl Acad Sci U S A, 114(6), 1252–1257.
Abstract: Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg Cm-2d-1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional approximately 225 mg Cm-2d-1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.
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Stukel, M. R., Benitez-Nelson, C. R., Decima, M., Taylor, A. G., Buchwald, C., & Landry, M. R. (2016). The biological pump in the Costa Rica Dome: an open-ocean upwelling system with high new production and low export. J Plankton Res, 38(2), 348–365.
Abstract: The Costa Rica Dome is a picophytoplankton-dominated, open-ocean upwelling system in the Eastern Tropical Pacific that overlies the ocean's largest oxygen minimum zone. To investigate the efficiency of the biological pump in this unique area, we used shallow (90-150 m) drifting sediment traps and 234Th:238U deficiency measurements to determine export fluxes of carbon, nitrogen and phosphorus in sinking particles. Simultaneous measurements of nitrate uptake and shallow water nitrification allowed us to assess the equilibrium balance of new and export production over a monthly timescale. While f-ratios (new:total production) were reasonably high (0.36 +/- 0.12, mean +/- standard deviation), export efficiencies were considerably lower. Sediment traps suggested e-ratios (export/14C-primary production) at 90-100 m ranging from 0.053 to 0.067. ThE-ratios (234Th disequilibrium-derived export) ranged from 0.038 to 0.088. C:N and N:P stoichiometries of sinking material were both greater than canonical (Redfield) ratios or measured C:N of suspended particulates, and they increased with depth, suggesting that both nitrogen and phosphorus were preferentially remineralized from sinking particles. Our results are consistent with an ecosystem in which mesozooplankton play a major role in energy transfer to higher trophic levels but are relatively inefficient in mediating vertical carbon flux to depth, leading to an imbalance between new production and sinking flux.
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Stukel, M. R., Biard, T., Krause, J. W., & Ohman, M. D. (2018). Large Phaeodaria in the twilight zone: Their role in the carbon cycle. Association for the Sciences of Limnology and Oceanography, .
Abstract: Advances in in situ imaging allow enumeration of abundant populations of large Rhizarians that compose a substantial proportion of total mesozooplankton biovolume. Using a quasi-Lagrangian sampling scheme, we quantified the abundance, vertical distributions, and sinking‐related mortality of Aulosphaeridae, an abundant family of Phaeodaria in the California Current Ecosystem. Inter‐cruise variability was high, with average concentrations at the depth of maximum abundance ranging from < 10 to > 300 cells m−3, with seasonal and interannual variability associated with temperature‐preferences and regional shoaling of the 10°C isotherm. Vertical profiles showed that these organisms were consistently most abundant at 100�150 m depth. Average turnover times with respect to sinking were 4.7�10.9 d, equating to minimum in situ population growth rates of ~ 0.1�0.2 d−1. Using simultaneous measurements of sinking organic carbon, we find that these organisms could only meet their carbon demand if their carbon : volume ratio were ~ 1 μg C mm−3. This value is substantially lower than previously used in global estimates of rhizarian biomass, but is reasonable for organisms that use large siliceous tests to inflate their cross‐sectional area without a concomitant increase in biomass. We found that Aulosphaeridae alone can intercept > 20% of sinking particles produced in the euphotic zone before these particles reach a depth of 300 m. Our results suggest that the local (and likely global) carbon biomass of Aulosphaeridae, and probably the large Rhizaria overall, needs to be revised downwards, but that these organisms nevertheless play a major role in carbon flux attenuation in the twilight zone.
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