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Stukel, MR, Benitez-Nelson CR, Decima M, Taylor AG, Buchwald C, Landry MR.  2016.  The biological pump in the Costa Rica Dome: an open-ocean upwelling system with high new production and low export. Journal of Plankton Research. 38:348-365.   10.1093/plankt/fbv097   AbstractWebsite

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 Th-234: U-238 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/C-14-primary production) at 90-100 m ranging from 0.053 to 0.067. ThE-ratios (Th-234 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.

Freibott, A, Taylor AG, Selph KE, Liu HB, Zhang WC, Landry MR.  2016.  Biomass and composition of protistan grazers and heterotrophic bacteria in the Costa Rica Dome during summer 2010. Journal of Plankton Research. 38:230-243.   10.1093/plankt/fbv107   AbstractWebsite

We investigated biomass and composition of heterotrophic microbes in the Costa Rica Dome during June-July 2010 as part of a broader study of plankton trophic dynamics. Because picophytoplankton (, 2 mm) are known to dominate in this unique upwelling region, we hypothesized tight biomass relationships between size-determined predator-prey pairs (i.e. picoplankton-nano-grazers, nanoplankton-micro-grazers) within the microbial community. Integrated biomass of heterotrophic bacteria ranged from 180 to 487 mg C m(-2) and was significantly correlated with total autotrophic carbon. Heterotrophic protist (H-protist) biomass ranged more narrowly from 488 to 545 mg C m(-2), and was comprised of 60% dinoflagellates, 30% other flagellates and 11% ciliates. Nano-sized (<20 mu m) protists accounted for the majority (57%) of grazer biomass and were positively correlated with picoplankton, partially supporting our hypothesis, but nanoplankton and micro-grazers (>20 mu m) were not significantly correlated. The relative constancy of H-protist biomass among locations despite clear changes in integrated autotrophic biomass, Chl a, and primary production suggests that mesozooplankton may exert a tight top-down control on micro-grazers. Biomass-specific consumption rates of phytoplankton by protistan grazers suggest an instantaneous growth rate of 0.52 day(-1) for H-protists, similar to the growth rate of phytoplankton and consistent with a trophically balanced ecosystem dominated by piconanoplankton interactions.

Decima, M, Landry MR, Stukel MR, Lopez-Lopez L, Krause JW.  2016.  Mesozooplankton biomass and grazing in the Costa Rica Dome: amplifying variability through the plankton food web. Journal of Plankton Research. 38:317-330.   10.1093/plankt/fbv091   AbstractWebsite

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-mu m cells) with especially high concentrations of Synechococcus spp., we found high mesozooplankton biomass (similar to 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.

Taylor, AG, Landry MR, Freibott A, Selph KE, Gutierrez-Rodriguez A.  2016.  Patterns of microbial community biomass, composition and HPLC diagnostic pigments in the Costa Rica upwelling dome. Journal of Plankton Research. 38:183-198.   10.1093/plankt/fbv086   AbstractWebsite

We investigated biomass, size-structure, composition, depth distributions and spatial variability of the phytoplankton community in the Costa Rica Dome (CRD) in June-July 2010. Euphotic zone profiles were sampled daily during Lagrangian experiments in and out of the dome region, and the community was analyzed using a combination of digital epifluorescence microscopy, flow cytometry and HPLC pigments. The mean depth-integrated biomass of phytoplankton ranged 2-fold, from 1089 to 1858 mg C m(-2) (mean+/-SE = 1378+/-112 mg C m(-2)), among 4 water parcels tracked for 4 days. Corresponding mean (+/-SE) integrated values for total chlorophyll a (Chl a) and the ratio of autotrophic carbon to Chl a were 24.1+/-1.5 mg Chl a m(-2) and 57.5+/-3.4, respectively. Absolute and relative contributions of picophytoplankton (similar to 60%), Synechococcus (>33%) and Prochlorococcus (17%) to phytoplankton community biomass were highest in the central dome region, while >20 mu m phytoplankton accounted for <= 10%, and diatoms, <2%, of biomass in all areas. Nonetheless, autotrophic flagellates, dominated by dinoflagellates, exceeded biomass contributions of Synechococcus at all locations. Order-of-magnitude discrepancies in the relative contributions of diatoms (overestimated) and dinoflagellates (underestimated) based on diagnostic pigments relative to microscopy highlight potential significant biases associated with making community inferences from pigments.

Landry, MR, Selph KE, Decima M, Gutierrez-Rodriguez A, Stukel MR, Taylor AG, Pasulka AL.  2016.  Phytoplankton production and grazing balances in the Costa Rica Dome. Journal of Plankton Research. 38:366-379.   10.1093/plankt/fbv089   AbstractWebsite

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 C-14 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 similar to 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, KE, Landry MR, Taylor AG, Gutierrez-Rodriguez A, Stukel MR, Wokuluk J, Pasulka A.  2016.  Phytoplankton production and taxon-specific growth rates in the Costa Rica Dome. Journal of Plankton Research. 38:199-215.   10.1093/plankt/fbv063   AbstractWebsite

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 (C-14-incorporation) and group-specific growth and net growth rates (two-treatment sea-water 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 (similar to 0.5-0.9 day(-1)), but higher on the edge of the dome (similar to 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 (similar to 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.

Landry, MR, de Verneil A, Goes JI, Moffett JW.  2016.  Plankton dynamics and biogeochemical fluxes in the Costa Rica Dome: introduction to the CRD Flux and Zinc Experiments. Journal of Plankton Research. 38:167-182.   10.1093/plankt/fbv103   AbstractWebsite

The Costa Rica Dome (CRD) is an open-ocean upwelling system in the Eastern Tropical Pacific that overlies the ocean's largest oxygen minimum zone (OMZ). The region has unique characteristics, biomass dominance by picophytoplankton, suppressed diatoms, high biomass of higher consumers and presumptive trace metal limitation, but is poorly understood in terms of pelagic stock and process relationships, including productivity and production controls. Here, we describe the goals, project design, physical context and major findings of the Flux and Zinc Experiments cruise conducted in June-July 2010 to assess trophic flux relationships and elemental controls on phytoplankton in the CRD. Despite sampling during a year of suppressed summertime surface chlorophyll, cruise results show high productivity (similar to 1 g C m(-2) day(-1)), high new production relative to export, balanced production and grazing, disproportionate biomass-specific productivity of large phytoplankton and high zooplankton stocks. Zinc concentrations are low in surface waters relative to phosphorous and silicate in other regions, providing conditions conducive to picophytoplankton, like Synechococcus, with low Zn requirements. Experiments nonetheless highlight phytoplankton limitation or co-limitation by silicic acid, driven by a strong silica pump that is linked to low dissolution of biogenic silica in the cold shallow thermocline of the lower euphotic zone.