Publications

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2018
Stukel, MR, Decima M, Landry MR, Selph KE.  2018.  Nitrogen and Isotope Flows Through the Costa Rica Dome Upwelling Ecosystem: The Crucial Mesozooplankton Role in Export Flux. Global Biogeochemical Cycles. 32:1815-1832.   10.1029/2018gb005968   AbstractWebsite

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 N-15 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. Plain Language Summary Most of the world's oceanic regions can be divided into (1) low-nutrient areas where small algae dominate and crustaceans, fish, and whales are scarce or (2) productive areas where large algae dominate, crustaceans and higher trophic levels are abundant, and substantial carbon is transported to depth as part of the biological pump. The Costa Rica Dome (CRD) is a unique natural laboratory for investigating the relationships between algae, zooplankton, and marine biogeochemistry because it is a productive region dominated by cyanobacteria (small algae) that nevertheless sustains large populations of crustaceans, fish, and whales. We used a novel data assimilation tool to constrain a food web model using at-sea rate measurements of plankton activity and nitrogen stable isotopes. We found that protists are an important intermediate trophic level linking cyanobacteria and mesozooplankton. Efficient recycling by the zooplankton community facilitates nitrogen transfer to fish, whales, and seabirds. In the CRD, vertically migrating zooplankton (which feed in the surface during the night but descend to depth during the day to escape predators) play a particularly important role in transporting nitrogen (and carbon dioxide) from the surface to the deep ocean, where it can be removed from the atmosphere.

2017
Steinberg, DK, Landry MR.  2017.  Zooplankton and the ocean carbon cycle. Annual Review of Marine Sciences, Vol 9. 9:413-444., Palo Alto: Annual Reviews   10.1146/annurev-marine-010814-015924   Abstract

Marine zooplankton comprise a phylogenetically and functionally diverse assemblage of protistan and metazoan consumers that occupy multiple trophic levels in pelagic food webs. Within this complex network, carbon flows via alternative zooplankton pathways drive temporal and spatial variability in production-grazing coupling, nutrient cycling, export, and transfer efficiency to higher trophic levels. We explore current knowledge of the processing of zooplankton food ingestion by absorption, egestion, respiration, excretion, and growth (production) processes. On a global scale, carbon fluxes are reasonably constrained by the grazing impact of microzooplankton and the respiratory requirements of mesozooplankton but are sensitive to uncertainties in trophic structure. The relative importance, combined magnitude, and efficiency of export mechanisms (mucous feeding webs, fecal pellets, molts, carcasses, and vertical migrations) likewise reflect regional variability in community structure. Climate change is expected to broadly alter carbon cycling by zooplankton and to have direct impacts on key species.

2015
Stukel, MR, Kahru M, Benitez-Nelson CR, Decima M, Goericke R, Landry MR, Ohman MD.  2015.  Using Lagrangian-based process studies to test satellite algorithms of vertical carbon flux in the eastern North Pacific Ocean. Journal of Geophysical Research-Oceans. 120:7208-7222.   10.1002/2015jc011264   AbstractWebsite

The biological carbon pump is responsible for the transport of similar to 5-20 Pg C yr(-1) from the surface into the deep ocean but its variability is poorly understood due to an incomplete mechanistic understanding of the complex underlying planktonic processes. In fact, algorithms designed to estimate carbon export from satellite products incorporate fundamentally different assumptions about the relationships between plankton biomass, productivity, and export efficiency. To test the alternate formulations of export efficiency in remote-sensing algorithms formulated by Dunne et al. (2005), Laws et al. (2011), Henson et al. (2011), and Siegel et al. (2014), we have compiled in situ measurements (temperature, chlorophyll, primary production, phytoplankton biomass and size structure, grazing rates, net chlorophyll change, and carbon export) made during Lagrangian process studies on seven cruises in the California Current Ecosystem and Costa Rica Dome. A food-web based approach formulated by Siegel et al. (2014) performs as well or better than other empirical formulations, while simultaneously providing reasonable estimates of protozoan and mesozooplankton grazing rates. By tuning the Siegel et al. (2014) algorithm to match in situ grazing rates more accurately, we also obtain better in situ carbon export measurements. Adequate representations of food-web relationships and grazing dynamics are therefore crucial to improving the accuracy of export predictions made from satellite-derived products. Nevertheless, considerable unexplained variance in export remains and must be explored before we can reliably use remote sensing products to assess the impact of climate change on biologically mediated carbon sequestration.