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In contrast to its productive coastal margins, the open-ocean Gulf of Mexico (GoM) is notable for highly stratified surface waters with extremely low nutrient and chlorophyll concentrations. Field campaigns in 2017 and 2018 identified low rates of turbulent mixing, which combined with oligotrophic nutrient conditions, give very low estimates for diffusive flux of nitrate into the euphotic zone (< 1 µmol N m⁻²d⁻¹). Estimates of local N₂-fixation are similarly low. In comparison, measured export rates of sinking particulate organic nitrogen (PON) from the euphotic zone are 2 – 3 orders of magnitude higher (i.e. 462 – 1144 µmol N m⁻²d⁻¹). We reconcile these disparate findings with regional scale dynamics inferred independently from remote-sensing products and a regional biogeochemical model and find that laterally-sourced organic matter is sufficient to support >90% of open-ocean nitrogen export in the GoM. Results show that lateral transport needs to be closely considered in studies of biogeochemical balances, particularly for basins enclosed by productive coasts.

 

Bluefin tuna spawn in restricted areas of subtropical oligotrophic seas. Here, we investigate the zooplankton prey and feeding selectivity of early larval stages of Atlantic bluefin tuna (ABT, Thunnus thynnus) in larval rearing habitat of the Gulf of Mexico. Larvae and zooplankton were collected during two multi-day Lagrangian experiments during peak spawning in May 2017 and 2018. Larvae were categorized by flexion stage and standard length. We identified, enumerated and sized zooplankton from larval gut contents and in the ambient community. Ciliates were quantitatively important (up to 9%) in carbon-based diets of early larvae. As larvae grew, diet composition and prey selection shifted from small copepod nauplii and calanoid copepodites to larger podonid cladocerans, which accounted for up to 70% of ingested carbon. Even when cladoceran abundances were <0.2 m−3, they comprised 23% of postflexion stage diet. Feeding behaviors of larvae at different development stages were more specialized, and prey selection narrowed to appendicularians and primarily cladocerans when these taxa were more abundant. Our findings suggest that ABT larvae have the capacity to switch from passive selection, regulated by physical factors, to active selection of presumably energetically optimal prey.

 

Multiple different equations have been used to quantify nutrient uptake rates from stable isotope tracer label incorporation experiments. Each of these equations implicitly assumes an underlying model for phytoplankton nutrient uptake behavior within the incubation bottle and/or pelagic environment. However, the applicability of different equations remains in question and uncertainty arising from subjective choices of which equation to use is never reported. In this study, I use two approaches to investigate the conditions under which different nutrient uptake equations should be used. First, I utilized a moderate‐complexity pelagic ecosystem model that explicitly models the δ¹⁵N values of all model compartments (NEMURO + ¹⁵N) to conduct simulated nitrate uptake and ammonium uptake incubations and quantify the accuracy of different nutrient uptake equations. Second, I used results of deckboard diel nutrient uptake experiments to quantify the biases of 24‐h incubations relative to six consecutive 4‐h incubations. Using both approaches, I found that equations that account for nutrient regeneration (i.e., isotope dilution) are more accurate than equations that do not, particularly when nutrient concentrations are low but uptake is relatively high. Furthermore, I find that if the goal is to estimate in situ uptake rates it is appropriate to use an in situ correction to standard equations. I also present complete equations for quantifying uncertainty in nutrient uptake experiments using each nutrient uptake equation and make all of these calculations available as Excel spreadsheets and Matlab scripts.

 

Abstract The availability of nitrogen (N) in ocean surface waters affects rates of photosynthesis and marine ecosystem structure. In spite of low dissolved inorganic N concentrations, export production in oligotrophic waters is comparable to more nutrient replete regions. Prior observations raise the possibility that di-nitrogen (N2) fixation supplies a significant fraction of N supporting export production in the Gulf of Mexico. In this study, geochemical tools were used to quantify the relative and absolute importance of both subsurface nitrate and N2 fixation as sources of new N fueling export production in the oligotrophic Gulf of Mexico in May 2017 and May 2018. Comparing the isotopic composition (“δ15N”) of nitrate with the δ15N of sinking particulate N collected during five sediment trap deployments each lasting two to four days indicates that N2 fixation is typically not detected and that the majority (≥80%) of export production is supported by subsurface nitrate. Moreover, no gradients in upper ocean dissolved organic N and suspended particulate N concentration and/or δ15N were found that would indicate significant N2 fixation fluxes accumulated in these pools, consistent with low Trichodesmium spp. abundance. Finally, comparing the δ15N of sinking particulate N captured within vs. below the euphotic zone indicates that during late spring regenerated N is low in δ15N compared to sinking N. 

Abstract Atlantic bluefin tuna (ABT) (Thunnus thynnus) travel long distances to spawn in oligotrophic regions of the Gulf of Mexico (GoM) which suggests these regions offer some unique benefit to offspring survival. To better understand how larval survival varies within the GoM a spatially explicit, Lagrangian, individual-based model was developed that simulates dispersal and mortality of ABT early life stages within realistic predator and prey fields during the spawning periods from 1993 to 2012. The model estimates that starvation is the largest cumulative source of mortality associated with an early critical period. However, elevated predation on older larvae is identified as the main factor limiting survival to late postflexion. As a result, first-feeding larvae have higher survival on the shelf where food is abundant, whereas older larvae have higher survival in the open ocean with fewer predators, making the shelf break an optimal spawning area. The modeling framework developed in this study explicitly simulates both physical and biological factors that impact larval survival and hence could be used to support ecosystem based management efforts for ABT under current and future climate conditions. 

Abstract The highly stratified, oligotrophic regions of the oceans are predominantly nitrogen limited in the surface ocean and light limited at the deep chlorophyll maximum (DCM). Hence, determining light and nitrogen co-limitation patterns for diverse phytoplankton taxa is crucial to understanding marine primary production throughout the euphotic zone. During two cruises in the deep-water Gulf of Mexico, we measured primary productivity (H13CO3−), nitrate uptake (15NO3−) and ammonium uptake (15NH4+) throughout the water column. Primary productivity declined with depth from the mixed layer to the DCM, averaging 27.1 mmol C m−2 d−1. The fraction of growth supported by NO3− was consistently low, with upper euphotic zone values ranging from 0.01 to 0.14 and lower euphotic zone values ranging from 0.03 to 0.44. Nitrate uptake showed strong diel patterns (maximum during the day), whereas ammonium uptake exhibited no diel variability. To parameterize taxon-specific phytoplankton nutrient and light utilization, we used a data assimilation approach (Bayesian Markov Chain Monte Carlo) including primary productivity, nutrient uptake and taxon-specific growth rate measurements. Parameters derived from this analysis define distinct niches for five phytoplankton taxa (Prochlorococcus, Synechococcus, diatoms, dinoflagellates and prymnesiophytes) and may be useful for constraining biogeochemical models of oligotrophic open-ocean systems. 

Abstract We used linear inverse ecosystem modeling techniques to assimilate data from extensive Lagrangian field experiments into a mass-balance constrained food web for the Gulf of Mexico open-ocean ecosystem. This region is highly oligotrophic, yet Atlantic bluefin tuna (ABT) travel long distances from feeding grounds in the North Atlantic to spawn there. Our results show extensive nutrient regeneration fueling primary productivity (mostly by cyanobacteria and other picophytoplankton) in the upper euphotic zone. The food web is dominated by the microbial loop (>70% of net primary productivity is respired by heterotrophic bacteria and protists that feed on them). By contrast, herbivorous food web pathways from phytoplankton to metazoan zooplankton process <10% of the net primary production in the mixed layer. Nevertheless, ABT larvae feed preferentially on podonid cladocerans and other suspension-feeding zooplankton, which in turn derive much of their nutrition from nano- and micro-phytoplankton (mixotrophic flagellates, and to a lesser extent, diatoms). This allows ABT larvae to maintain a comparatively low trophic level (~4.2 for preflexion and postflexion larvae), which increases trophic transfer from phytoplankton to larval fish. 

Abstract Phytoplankton growth and microzooplankton grazing rates were measured in repeated profiles of dilution experiments incubated in situ on a drift array in order to assess microbial production and food web characteristics in the oligotrophic bluefin tuna spawning habitat of the Gulf of Mexico (May peak spawning seasons, 2017–2018). Grazing often exceeded growth with the processes more balanced overall in the surface mixed layer, but biomass accumulated in the mid-euphotic zone. Community production estimates (260–500 mg C m−2 day−1) were low compared to similar open-ocean studies in the Pacific Ocean. Prochlorococcus was a consistent major contributor (113–204 mg C m−2 day−1) to productivity, while diatoms and dinoflagellates (2–10 and 4–13 mg C m−2 day−1, respectively) were consistently low. Prymnesiophytes, the most dynamic component (34–134 mg C m−2 day−1), co-dominated in 2017 experiments. Unexpected imbalances in grazing relative to production were observed for all picoplankton populations (Prochlorococcus, Synechococcus and heterotrophic bacteria), suggesting a trophic cascade in the absence of mesozooplankton predation on large microzooplankton. Study sites with abundant larval tuna had the shallowest deep chlorophyll maxima and significant net positive phytoplankton growth below the mixed layer. 

Abstract Biomass and composition of the phytoplankton community were investigated in the deep-water Gulf of Mexico (GoM) at the edges of Loop Current anticyclonic eddies during May 2017 and May 2018. Using flow cytometry, high-performance liquid chromatography pigments and microscopy, we found euphotic zone integrated chlorophyll a of ~10 mg m−2 and autotrophic carbon ranging from 463 to 1268 mg m−2, dominated by picoplankton (<2 μm cells). Phytoplankton assemblages were similar to the mean composition at the Bermuda Atlantic Time-series Study site, but differed from the Hawaii Ocean Times-series site. GoM phytoplankton biomass was ~2-fold higher at the deep chlorophyll maximum (DCM) relative to the mixed layer (ML). Prochlorococcus and prymnesiophytes were the dominant taxa throughout the euphotic zone; however, other eukaryotic taxa had significant biomass in the DCM. Shallower DCMs were correlated with more prymnesiophytes and prasinophytes (Type 3) and reduced Prochlorococcus. These trends in ML and DCM taxonomic composition likely reflect relative nutrient supply—with ML populations relying on remineralized ammonium as a nitrogen source, and the taxonomically diverse DCM populations using more nitrate. These spatially separated phytoplankton communities represent different pathways for primary production, with a dominance of picoplankton in the ML and more nano- and microplankton at the DCM.