Landry, M. R., Selph, K. E., Decima, M., Gutierrez-Rodriguez, A., Stukel, M. R., Taylor, A. G., et al. (2016). Phytoplankton production and grazing balances in the Costa Rica Dome. J Plankton Res , 38 (2), 366–379.
Abstract: We investigated phytoplankton production rates and grazing fates in the Costa Rica Dome (CRD) during summer 2010 based on dilution depth profiles analyzed by flow cytometry and pigments and mesozooplankton grazing assessed by gut fluorescence. Three community production estimates, from 14C uptake (1025 +/- 113 mg C m-2 day-1) and from dilution experiments analyzed for total Chla (990 +/- 106 mg C m-2 day-1) and flow cytometry populations (862 +/- 71 mg C m-2 day-1), exceeded regional ship-based values by 2-3-fold. Picophytoplankton accounted for 56% of community biomass and 39% of production. Production profiles extended deeper for Prochlorococcus (PRO) and picoeukaryotes than for Synechococcus (SYN) and larger eukaryotes, but 93% of total production occurred above 40 m. Microzooplankton consumed all PRO and SYN growth and two-third of total production. Positive net growth of larger eukaryotes in the upper 40 m was balanced by independently measured consumption by mesozooplankton. Among larger eukaryotes, diatoms contributed approximately 3% to production. On the basis of this analysis, the CRD region is characterized by high production and grazing turnover, comparable with or higher than estimates for the eastern equatorial Pacific. The region nonetheless displays characteristics atypical of high productivity, such as picophytoplankton dominance and suppressed diatom roles.
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.
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
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.
Selph, K. E., Landry, M. R., Taylor, A. G., Gutierrez-Rodriguez, A., Stukel, M. R., Wokuluk, J., et al. (2016). Phytoplankton production and taxon-specific growth rates in the Costa Rica Dome. J Plankton Res , 38 (2), 199–215.
Abstract: During summer 2010, we investigated phytoplankton production and growth rates at 19 stations in the eastern tropical Pacific, where winds and strong opposing currents generate the Costa Rica Dome (CRD), an open-ocean upwelling feature. Primary production (14C-incorporation) and group-specific growth and net growth rates (two-treatment seawater dilution method) were estimated from samples incubated in situ at eight depths. Our cruise coincided with a mild El Nino event, and only weak upwelling was observed in the CRD. Nevertheless, the highest phytoplankton abundances were found near the dome center. However, mixed-layer growth rates were lowest in the dome center ( approximately 0.5-0.9 day-1), but higher on the edge of the dome ( approximately 0.9-1.0 day-1) and in adjacent coastal waters (0.9-1.3 day-1). We found good agreement between independent methods to estimate growth rates. Mixed-layer growth rates of Prochlorococcus and Synechococcus were largely balanced by mortality, whereas eukaryotic phytoplankton showed positive net growth ( approximately 0.5-0.6 day-1), that is, growth available to support larger (mesozooplankton) consumer biomass. These are the first group-specific phytoplankton rate estimates in this region, and they demonstrate that integrated primary production is high, exceeding 1 g C m-2 day-1 on average, even during a period of reduced upwelling.
Shropshire, T., Morey, S. L., Chassignet, E. P., Bozec, A., Coles, V. J., Landry, M. R., et al. (2019). Quantifying spatiotemporal variability in zooplankton dynamics in the Gulf of Mexico with a physical-biogeochemical model.
Abstract: Zooplankton play an important role in global biogeochemistry and their secondary production supports valuable fisheries of the world's oceans. Currently, zooplankton abundances cannot be estimated using remote sensing techniques. Hence, coupled physical-biogeochemical models (PBMs) provide an important tool for studying zooplankton on regional and global scales. However, evaluating the accuracy of zooplankton abundance estimates from PBMs has been a major challenge as a result of sparse observations. In this study, we configure a PBM for the Gulf of Mexico (GoM) from 1993�2012 and validate the model against an extensive combination of in situ biomass and rate measurements including total mesozooplankton biomass, size-fractionated mesozooplankton biomass and grazing rates, microzooplankton specific grazing rates, surface chlorophyll, deep chlorophyll maximum depth, phytoplankton specific growth rates, and net primary production. Spatial variability in mesozooplankton biomass climatology observed in a multi-decadal database for the northern GoM is well resolved by the model with a statistically significant (p < 0.01) correlation of 0.90. Mesozooplankton secondary production for the region averaged 66 + 8 mt C yr−1 equivalent to approximately 10 % of NPP and ranged from 51 to 82 mt C yr−1. In terms of diet, model results from the shelf regions suggest that herbivory is the dominant feeding mode for small mesozooplankton (< 1-mm) whereas larger mesozooplankton are primarily carnivorous. However, in open-ocean, oligotrophic regions, both groups of mesozooplankton have proportionally greater reliance on heterotrophic protists as a food source. This highlights the important role of microbial and protistan food webs in sustaining mesozooplankton biomass in the GoM which serves as the primary food source for early life stages of many commercially-important fish species, including tuna.
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.
Stukel, M. R., & Barbeau, K. A. (2020). Investigating the Nutrient Landscape in a Coastal Upwelling Region and Its Relationship to the Biological Carbon Pump. Geophys. Res. Lett. , 47 (6), e2020GL087351.
Abstract: We investigated nutrient patterns and their relationship to vertical carbon export using results from 38 Lagrangian experiments in the California Current Ecosystem. The dominant mode of variability reflected onshore-offshore nutrient gradients. A secondary mode of variability was correlated with silica excess and dissolved iron and likely reflects regional patterns of iron-limitation. The biological carbon pump was enhanced in high nutrient and Fe-stressed regions. Patterns in the nutrient landscape proved to be better predictors of the vertical flux of sinking particles than contemporaneous measurements of net primary production. Our results suggest an important role for Fe-stressed diatoms in vertical carbon flux. They also suggest that either preferential recycling of N or non-Redfieldian nutrient uptake by diatoms may lead to high PO:NO and Si(OH):NO ratios, following export of P- and Si-enriched organic matter. Increased export following Fe-stress may partially explain inverse relationships between net primary productivity and export efficiency.
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.
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.