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.
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.
Stukel, M. R., Decima, M., & Kelly, T. B. (2018). A new approach for incorporating 15N isotopic data into linear inverse ecosystem models with Markov Chain Monte Carlo sampling.
PLoS One, 13(6), e0199123.
Abstract: Oceanographic field programs often use delta15N biogeochemical measurements and in situ rate measurements to investigate nitrogen cycling and planktonic ecosystem structure. However, integrative modeling approaches capable of synthesizing these distinct measurement types are lacking. We develop a novel approach for incorporating delta15N isotopic data into existing Markov Chain Monte Carlo (MCMC) random walk methods for solving linear inverse ecosystem models. We test the ability of this approach to recover food web indices (nitrate uptake, nitrogen fixation, zooplankton trophic level, and secondary production) derived from forward models simulating the planktonic ecosystems of the California Current and Amazon River Plume. We show that the MCMC with delta15N approach typically does a better job of recovering ecosystem structure than the standard MCMC or L2 minimum norm (L2MN) approaches, and also outperforms an L2MN with delta15N approach. Furthermore, we find that the MCMC with delta15N approach is robust to the removal of input equations and hence is well suited to typical pelagic ecosystem studies for which the system is usually vastly under-constrained. Our approach is easily extendable for use with delta13C isotopic measurements or variable carbon:nitrogen stoichiometry.
Stukel, M. R., Décima, M., Landry, M. R., & Selph, K. E. (2018). Nitrogen and isotope flows through the Costa Rica Dome upwelling ecosystem: The crucial mesozooplankton role in export flux.
Global Biogeochemical Cycles, 32(12), 1815–1832.
Abstract: The Costa Rica Dome (CRD) is an open-ocean upwelling ecosystem, with high biomasses of picophytoplankton (especially Synechococcus), mesozooplankton, and higher trophic levels. To elucidate the food web pathways supporting the trophic structure and carbon export in this unique ecosystem, we used Markov Chain Monte Carlo techniques to assimilate data from four independent realizations of δ15N and planktonic rate measurements from the CRD into steady state, multicompartment ecosystem box models (linear inverse models). Model results present well-constrained snapshots of ecosystem nitrogen and stable isotope fluxes. New production is supported by upwelled nitrate, not nitrogen fixation. Protistivory (rather than herbivory) was the most important feeding mode for mesozooplankton, which rely heavily on microzooplankton prey. Mesozooplankton play a central role in vertical nitrogen export, primarily through active transport of nitrogen consumed in the surface layer and excreted at depth, which comprised an average 36-46% of total export. Detritus or aggregate feeding is also an important mode of resource acquisition by mesozooplankton and regeneration of nutrients within the euphotic zone. As a consequence, the ratio of passively sinking particle export to phytoplankton production is very low in the CRD. Comparisons to similar models constrained with data from the nearby equatorial Pacific demonstrate that the dominant role of vertical migrators to the biological pump is a unique feature of the CRD. However, both regions show efficient nitrogen transfer from mesozooplankton to higher trophic levels (as expected for regions with large fish, cetacean, and seabird populations) despite the dominance of protists as major grazers of phytoplankton.
Stukel, M. R., Kahru, M., Benitez-Nelson, C. R., Décima, M., Goericke, R., Landry, M. R., et al. (2015). Using Lagrangian-based process studies to test satellite algorithms of vertical carbon flux in the eastern North Pacific Ocean.
J. Geophys. Res. Oceans, 120(11), 7208–7222.
Stukel, M. R., Ohman, M. D., Kelly, T. B., & Biard, T. (2019). The Roles of Suspension-Feeding and Flux-Feeding Zooplankton as Gatekeepers of Particle Flux Into the Mesopelagic Ocean in the Northeast Pacific.
Front. Mar. Sci., 6.
Abstract: Zooplankton are important consumers of sinking particles in the ocean's twilight zone. However, the impact of different taxa depends on their feeding mode. In contrast to typical suspension-feeding zooplankton, flux-feeding taxa preferentially consume rapidly sinking particles that would otherwise penetrate into the deep ocean. To quantify the potential impact of two flux-feeding zooplankton taxa [Aulosphaeridae (Rhizaria), and Limacina helicina (euthecosome pteropod)] and the total suspension-feeding zooplankton community, we measured depth-stratified abundances of these organisms during six cruises in the California Current Ecosystem. Using allometric-scaling relationships, we computed the percentage of carbon flux intercepted by flux feeders and suspension feeders. These estimates were compared to direct measurements of carbon flux attenuation (CFA) made using drifting sediment traps and U-238-Th-234 disequilibrium. We found that CFA in the shallow twilight zone typically ranged from 500 to 1000 m mol organic C flux remineralized per 10-m vertical depth bin. This equated to approximately 6-10% of carbon flux remineralized/10 m. The two flux-feeding taxa considered in this study could account for a substantial proportion of this flux near the base of the euphotic zone. The mean flux attenuation attributable to Aulosphaeridae was 0.69%/10 m (median = 0.21%/10 m, interquartile range = 0.04-0.81%) at their depth of maximum abundance (similar to 100 m), which would equate to similar to 10% of total flux attenuation in this depth range. The maximum flux attenuation attributable to Aulosphaeridae reached 4.2%/10 m when these protists were most abundant. L. helicina, meanwhile, could intercept 0.45-1.6% of carbon flux/10 m, which was slightly greater (on average) than the Aulosphaeridae. In contrast, suspension-feeding zooplankton in the mesopelagic (including copepods, euphausiids, appendicularians, and ostracods) had combined clearance rates of 2-81 L m(-3) day(-1) (mean of 19.6 L m(-3) day(-1)). This implies a substantial impact on slowly sinking particles, but a negligible impact on the presumably rapidly sinking fecal pellets that comprised the majority of the material collected in sediment traps. Our results highlight the need for a greater research focus on the many taxa that potentially act as flux feeders in the oceanic twilight zone.
Wang, S., Kranz, S. A., Kelly, T. B., Song, H., Stukel, M. R., & Cassar, N. (2020). Lagrangian Studies of Net Community Production: The Effect of Diel and Multiday Nonsteady State Factors and Vertical Fluxes on O
2/Ar in a Dynamic Upwelling Region. J. Geophys. Res. Biogeosci., 125(6), e2019JG005569.
Abstract: The ratio of dissolved oxygen to argon in seawater is frequently employed to estimate rates of net community production (NCP) in the oceanic mixed layer. The in situ O2/Ar‐based method accounts for many physical factors that influence oxygen concentrations, permitting isolation of the biological oxygen signal produced by the balance of photosynthesis and respiration. However, this technique traditionally relies upon several assumptions when calculating the mixed‐layer O2/Ar budget, most notably the absence of vertical fluxes of O2/Ar and the principle that the air‐sea gas exchange of biological oxygen closely approximates net productivity rates. Employing a Lagrangian study design and leveraging data outputs from a regional physical oceanographic model, we conducted in situ measurements of O2/Ar in the California Current Ecosystem in spring 2016 and summer 2017 to evaluate these assumptions within a �worst‐case� field environment. Quantifying vertical fluxes, incorporating nonsteady state changes in O2/Ar, and comparing NCP estimates evaluated over several day versus longer timescales, we find differences in NCP metrics calculated over different time intervals to be considerable, also observing significant potential effects from vertical fluxes, particularly advection. Additionally, we observe strong diel variability in O2/Ar and NCP rates at multiple stations. Our results reemphasize the importance of accounting for vertical fluxes when interpreting O2/Ar‐derived NCP data and the potentially large effect of nonsteady state conditions on NCP evaluated over shorter timescales. In addition, diel cycles in surface O2/Ar can also bias interpretation of NCP data based on local productivity and the time of day when measurements were made.