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.
Decima, M., Landry, M. R., Stukel, M. R., Lopez-Lopez, L., & Krause, J. W. (2016). Mesozooplankton biomass and grazing in the Costa Rica Dome: amplifying variability through the plankton food web.
J Plankton Res, 38(2), 317–330.
Abstract: We investigated standing stocks and grazing rates of mesozooplankton assemblages in the Costa Rica Dome (CRD), an open-ocean upwelling ecosystem in the eastern tropical Pacific. While phytoplankton biomass in the CRD is dominated by picophytoplankton (<2-microm cells) with especially high concentrations of Synechococcus spp., we found high mesozooplankton biomass ( approximately 5 g dry weight m-2) and grazing impact (12-50% integrated water column chlorophyll a), indicative of efficient food web transfer from primary producers to higher levels. In contrast to the relative uniformity in water-column chlorophyll a and mesozooplankton biomass, variability in herbivory was substantial, with lower rates in the central dome region and higher rates in areas offset from the dome center. While grazing rates were unrelated to total phytoplankton, correlations with cyanobacteria (negative) and biogenic SiO2 production (positive) suggest that partitioning of primary production among phytoplankton sizes contributes to the variability observed in mesozooplankton metrics. We propose that advection of upwelled waters away from the dome center is accompanied by changes in mesozooplankton composition and grazing rates, reflecting small changes within the primary producers. Small changes within the phytoplankton community resulting in large changes in the mesozooplankton suggest that the variability in lower trophic level dynamics was effectively amplified through the food web.
Kelly, T. B., Davison, P. C., Goericke, R., Landry, M. R., Ohman, M. D., & Stukel, M., R. (2019). The Importance of Mesozooplankton Diel Vertical Migration for Sustaining a Mesopelagic Food Web.
FRONTIERS IN MARINE SCIENCE, 6.
Abstract: We used extensive ecological and biogeochemical measurements obtained from quasi-Lagrangian experiments during two California Current Ecosystem Long-Term Ecosystem Research cruises to analyze carbon fluxes between the epipelagic and mesopelagic zones using a linear inverse ecosystem model (LIEM). Measurement constraints on the model include C-14 primary productivity, dilution-based microzooplankton grazing rates, gut pigment-based mesozooplankton grazing rates (on multiple zooplankton size classes), Th-234:U-238 disequilibrium and sediment trap measured carbon export, and metabolic requirements of micronekton, zooplankton, and bacteria. A likelihood approach (Markov Chain Monte Carlo) was used to estimate the resulting flow uncertainties from a sample of potential flux networks. Results highlight the importance of mesozooplankton active transport (i.e., diel vertical migration) in supplying the carbon demand of mesopelagic organisms and sequestering carbon dioxide from the atmosphere. In nine water parcels ranging from a coastal bloom to offshore oligotrophic conditions, mesozooplankton active transport accounted for 18-84% (median: 42%) of the total carbon transfer to the mesopelagic, with gravitational settling of POC (12-55%; median: 37%), and subduction (2-32%; median: 14%) providing the majority of the remainder. Vertically migrating zooplankton contributed to downward carbon flux through respiration and excretion at depth and via mortality losses to predatory zooplankton and mesopelagic fish (e.g., myctophids and gonostomatids). Sensitivity analyses showed that the results of the LIEM were robust to changes in nekton metabolic demand, rates of bacterial production, and mesozooplankton gross growth efficiency. This analysis suggests that prior estimates of zooplankton active transport based on conservative estimates of standard (rather than active) metabolism are likely too low.
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.
Krause, J. W., Stukel, M. R., Taylor, A. G., Taniguchi, D. A. A., De Verneil, A., & Landry, M. R. (2016). Net biogenic silica production and the contribution of diatoms to new production and organic matter export in the Costa Rica Dome ecosystem.
J Plankton Res, 38(2), 216–229.
Abstract: We determined the net rate of biogenic silica (bSiO2) production and estimated the diatom contribution to new production and organic matter export in the Costa Rica Dome during summer 2010. The shallow thermocline significantly reduces bSiO2 dissolution rates below the mixed layer, leading to significant enhancement of bSiO2 relative to organic matter (silicate-pump condition). This may explain why deep export of bSiO2 in this region is elevated by an order of magnitude relative to comparable systems. Diatom carbon, relative to autotrophic carbon, was low (<3%); however, the contribution of diatoms to new production averaged 3 and 13% using independent approaches. The 4-old discrepancy between methods may be explained by a low average C:Si ratio ( approximately 1.4) for the net produced diatom C relative to the net produced bSiO2. We speculate that this low production ratio is not the result of reduced C, but may arise from a significant contribution of non-diatom silicifying organisms to bSiO2 production. The contribution of diatoms to organic matter export was minor (5.7%). These results, and those of the broader project, suggest substantial food-web transformation of diatom organic matter in the euphotic zone, which creates enriched bSiO2 relative to organic matter within the exported material.
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.
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., 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.