The shift in marine resource management from a compartmentalized approach of dealing with resources on a species basis to an approach based on management of spatially defined ecosystems requires an ...accurate accounting of energy flow. The flow of energy from primary production through the food web will ultimately limit upper trophic-level fishery yields. In this work, we examine the relationship between yield and several metrics including net primary production, chlorophyll concentration, particle-export ratio, and the ratio of secondary to primary production. We also evaluate the relationship between yield and two additional rate measures that describe the export of energy from the pelagic food web, particle export flux and mesozooplankton productivity. We found primary production is a poor predictor of global fishery yields for a sample of 52 large marine ecosystems. However, chlorophyll concentration, particle-export ratio, and the ratio of secondary to primary production were positively associated with yields. The latter two measures provide greater mechanistic insight into factors controlling fishery production than chlorophyll concentration alone. Particle export flux and mesozooplankton productivity were also significantly related to yield on a global basis. Collectively, our analyses suggest that factors related to the export of energy from pelagic food webs are critical to defining patterns of fishery yields. Such trophic patterns are associated with temperature and latitude and hence greater yields are associated with colder, high latitude ecosystems.
Zooplankton fecal pellet flux is a highly variable component of the biological carbon pump. While fecal pellets can comprise 0 to nearly 100% of particulate organic carbon collected in sediment ...traps, mechanisms for this variability remain poorly understood. Fecal pellet carbon flux is a complex function of several variables. We present a model that incorporates individual-scale metabolic processes to determine fecal pellet production rate, the relationship between body size and fecal pellet size, the relationship between fecal pellet size and sinking rate, and a function representing the breakdown of particles in the water column. When applied to copepod communities sampled by the continuous plankton recorder in the Gulf of Maine over 25 years, a seasonal pattern of fecal pellet carbon flux emerges. The interannual flux time series produced by the model reflects known oceanographic perturbations and shows how organism-scale processes can be scaled up to explain ecosystem level variability. We conclude that fecal pellet carbon flux in the Gulf of Maine is driven by copepod community size structure and copepod abundance, and that the fraction of fecal pellet carbon that reaches depth is a function of copepod size, rather than abundance. Changes in the physical environment which alter the size composition of the copepod community lead to variability in fecal pellet carbon flux. Our results indicate that incorporating size composition into biogeochemical models can more accurately constrain zooplankton-mediated carbon flux.
Here we document Alexandrium fundyense cyst abundance and distribution patterns over nine years (1997 and 2004–2011) in the coastal waters of the Gulf of Maine (GOM) and identify linkages between ...those patterns and several metrics of the severity or magnitude of blooms occurring before and after each autumn cyst survey. We also explore the relative utility of two measures of cyst abundance and demonstrate that GOM cyst counts can be normalized to sediment volume, revealing meaningful patterns equivalent to those determined with dry weight normalization.
Cyst concentrations were highly variable spatially. Two distinct seedbeds (defined here as accumulation zones with>300cystscm−3) are evident, one in the Bay of Fundy (BOF) and one in mid-coast Maine. Overall, seedbed locations remained relatively constant through time, but their area varied 3–4 fold, and total cyst abundance more than 10 fold among years. A major expansion of the mid-coast Maine seedbed occurred in 2009 following an unusually intense A. fundyense bloom with visible red-water conditions, but that feature disappeared by late 2010. The regional system thus has only two seedbeds with the bathymetry, sediment characteristics, currents, biology, and environmental conditions necessary to persist for decades or longer. Strong positive correlations were confirmed between the abundance of cysts in both the 0–1 and the 0–3cm layers of sediments in autumn and geographic measures of the extent of the bloom that occurred the next year (i.e., cysts→blooms), such as the length of coastline closed due to shellfish toxicity or the southernmost latitude of shellfish closures. In general, these metrics of bloom geographic extent did not correlate with the number of cysts in sediments following the blooms (blooms→cysts). There are, however, significant positive correlations between 0–3cm cyst abundances and metrics of the preceding bloom that are indicative of bloom intensity or vegetative cell abundance (e.g., cumulative shellfish toxicity, duration of detectable toxicity in shellfish, and bloom termination date). These data suggest that it may be possible to use cyst abundance to empirically forecast the geographic extent of the forthcoming bloom and, conversely, to use other metrics from bloom and toxicity events to forecast the size of the subsequent cyst population as the inoculum for the next year's bloom. This is an important step towards understanding the excystment/encystment cycle in A. fundyense bloom dynamics while also augmenting our predictive capability for this HAB-forming species in the GOM.
This article compares measurements of particle shape parameters from three-dimensional (3D) X-ray micro-computed tomography (μCT) and two-dimensional (2D) dynamic image analysis (DIA) from the ...optical microscopy of a coastal bioclastic calcareous sand from Western Australia. This biogenic sand from a high energy environment consists largely of the shells and tests of marine organisms and their clasts. A significant difference was observed between the two imaging techniques for measurements of aspect ratio, convexity, and sphericity. Measured values of aspect ratio, sphericity, and convexity are larger in 2D than in 3D. Correlation analysis indicates that sphericity is correlated with convexity in both 2D and 3D. These results are attributed to inherent limitations of DIA when applied to platy sand grains and to the shape being, in part, dependent on the biology of the grain rather than a purely random clastic process, like typical siliceous sands. The statistical data has also been fitted to Johnson Bounded Distribution for the ease of future use. Overall, this research demonstrates the need for high-quality 3D microscopy when conducting a micromechanical analysis of biogenic calcareous sands.
The Gulf of Maine (GoME) is a shelf region especially vulnerable to ocean acidification (OA) due to natural conditions of low pH and aragonite saturation states (Ω‐Ar). This study is the first to ...assess the major oceanic processes controlling seasonal variability of the carbonate system and its linkages with pteropod abundance in Wilkinson Basin in the GoME. Two years of seasonal sampling cruises suggest that water‐column carbonate chemistry in the region undergoes a seasonal cycle, wherein the annual cycle of stratification‐overturn, primary production, respiration‐remineralization and mixing all play important roles, at distinct spatiotemporal scales. Surface production was tightly coupled with remineralization in the benthic nepheloid layer during high production seasons, which results in occasional aragonite undersaturation. From spring to summer, carbonate chemistry in the surface across Wilkinson Basin reflects a transition from a production‐respiration balanced system to a net autotropic system. Mean water‐column Ω‐Ar and abundance of large thecosomatous pteropods show some correlation, although patchiness and discrete cohort reproductive success likely also influence their abundance. Overall, photosynthesis‐respiration is the primary driving force controlling Ω‐Ar variability during the spring‐to‐summer transition as well as over the seasonal cycle. However, calcium carbonate (CaCO3) dissolution appears to occur near bottom in fall and winter when bottom water Ω‐Ar is generally low but slightly above 1. This is accompanied by a decrease in pteropod abundance that is consistent with previous CaCO3 flux trap measurements. The region might experience persistent subsurface aragonite undersaturation in 30–40 years under continued ocean acidification.
Key Points
Aragonite saturation is primarily driven by photosynthesis and respiration‐remineralization in the water column of the Gulf of Maine
Calcium carbonate dissolution appears to occur from fall to winter in the deep portion of the Gulf of Maine
Consistent undersaturation of subsurface aragonite saturation is likely to occur in 30–40 years under acidification in the Gulf of Maine
In oceanic subtropical gyres, primary producers are numerically dominated by small (1-5 µm diameter) pro- and eukaryotic cells that primarily utilize recycled nutrients produced by rapid grazing ...turnover in a highly efficient microbial loop. Continuous losses of nitrogen (N) to depth by sinking, either as single cells, aggregates or fecal pellets, are balanced by both nitrate inputs at the base of the euphotic zone and N2-fixation. This input of new N to balance export losses (the biological pump) is a fundamental aspect of N cycling and central to understanding carbon fluxes in the ocean. In the Pacific Ocean, detailed N budgets at the time-series station HOT require upward transport of nitrate from the nutricline (80-100 m) into the surface layer (∼0-40 m) to balance productivity and export needs. However, concentration gradients are negligible and cannot support the fluxes. Physical processes can inject nitrate into the base of the euphotic zone, but the mechanisms for transporting this nitrate into the surface layer across many 10s of m in highly stratified systems are unknown. In these seas, vertical migration by the very largest (10(2)-10(3) µm diameter) phytoplankton is common as a survival strategy to obtain N from sub-euphotic zone depths. This vertical migration is driven by buoyancy changes rather than by flagellated movement and can provide upward N transport as nitrate (mM concentrations) in the cells. However, the contribution of vertical migration to nitrate transport has been difficult to quantify over the required basin scales. In this study, we use towed optical systems and isotopic tracers to show that migrating diatom (Rhizosolenia) mats are widespread in the N. Pacific Ocean from 140°W to 175°E and together with other migrating phytoplankton (Ethmodiscus, Halosphaera, Pyrocystis, and solitary Rhizosolenia) can mediate time-averaged transport of N (235 µmol N m(-2) d(-1)) equivalent to eddy nitrate injections (242 µmol NO3 (-) m(-2) d(-1)). This upward biotic transport can close N budgets in the upper 250 m of the central Pacific Ocean and together with diazotrophy creates a surface zone where biological nutrient inputs rather than physical processes dominate the new N flux. In addition to these numerically rare large migrators, there is evidence in the literature of ascending behavior in small phytoplankton that could contribute to upward flux as well. Although passive downward movement has dominated models of phytoplankton flux, there is now sufficient evidence to require a rethinking of this paradigm. Quantifying these fluxes is a challenge for the future and requires a reexamination of individual phytoplankton sinking rates as well as methods for capturing and enumerating ascending phytoplankton in the sea.
In the Gulf of Maine area (GoMA), as elsewhere in the ocean, the organisms of greatest numerical abundance are microbes. Viruses in GoMA are largely cyanophages and bacteriophages, including ...podoviruses which lack tails. There is also evidence of Mimivirus and Chlorovirus in the metagenome. Bacteria in GoMA comprise the dominant SAR11 phylotype cluster, and other abundant phylotypes such as SAR86-like cluster, SAR116-like cluster, Roseobacter, Rhodospirillaceae, Acidomicrobidae, Flavobacteriales, Cytophaga, and unclassified Alphaproteobacteria and Gammaproteobacteria clusters. Bacterial epibionts of the dinoflagellate Alexandrium fundyense include Rhodobacteraceae, Flavobacteriaceae, Cytophaga spp., Sulfitobacter spp., Sphingomonas spp., and unclassified Bacteroidetes. Phototrophic prokaryotes in GoMA include cyanobacteria that contain chlorophyll (mainly Synechococcus), aerobic anoxygenic phototrophs that contain bacteriochlorophyll, and bacteria that contain proteorhodopsin. Eukaryotic microalgae in GoMA include Bacillariophyceae, Dinophyceae, Prymnesiophyceae, Prasinophyceae, Trebouxiophyceae, Cryptophyceae, Dictyochophyceae, Chrysophyceae, Eustigmatophyceae, Pelagophyceae, Synurophyceae, and Xanthophyceae. There are no records of Bolidophyceae, Aurearenophyceae, Raphidophyceae, and Synchromophyceae in GoMA. In total, there are records for 665 names and 229 genera of microalgae. Heterotrophic eukaryotic protists in GoMA include Dinophyceae, Alveolata, Apicomplexa, amoeboid organisms, Labrynthulida, and heterotrophic marine stramenopiles (MAST). Ciliates include Strombidium, Lohmaniella, Tontonia, Strobilidium, Strombidinopsis and the mixotrophs Laboea strobila and Myrionecta rubrum (ex Mesodinium rubra). An inventory of selected microbial groups in each of 14 physiographic regions in GoMA is made by combining information on the depth-dependent variation of cell density and the depth-dependent variation of water volume. Across the entire GoMA, an estimate for the minimum abundance of cell-based microbes is 1.7×10(25) organisms. By one account, this number of microbes implies a richness of 10(5) to 10(6) taxa in the entire water volume of GoMA. Morphological diversity in microplankton is well-described but the true extent of taxonomic diversity, especially in the femtoplankton, picoplankton and nanoplankton--whether autotrophic, heterotrophic, or mixotrophic, is unknown.
We report planktonic foraminiferal fluxes (accumulation rates) and oxygen isotopes ( delta 18O) from a nine-month sediment trap deployment, and delta 18O from three sediment cores in Jordan Basin, ...Gulf of Maine. The sediment trap was deployed at 150m, about halfway to the basin floor, and samples were collected every three weeks between August 2010 and May 2011. The planktonic foraminiferal fauna in the trap is dominated by Neogloboquadrina incompta that reached a maximum flux in the second half of October. Oxygen isotope ratios on that species indicate that on average during the collecting period it lived in the surface mixed layer, when compared to predicted values based on data from a nearby hydrographic buoy from the same period. New large diameter piston cores from Jordan Basin are 25 and 28m long. Marine hemipelagic sediments are 25m thick, and the sharp contact with underlying red deglacial sediments is bracketed by two radiocarbon dates on bivalves that indicate ice-free conditions began 16,900 calibrated years ago. Radiocarbon dating of foraminifera indicates that the basin floor sediments (270-290m) accumulated at >3m/kyr during the Holocene, whereas rates were about one tenth that on the basin slope (230m). In principle, Jordan Basin sediments have the potential to provide time series with interannual resolution. Our results indicate the Holocene is marked by 2 degree C variability in SST, and the coldest events of the 20th century, during the mid 1960s and mid 1920s, appear to be recorded in the uppermost 50cm of the seafloor.
In 2003, the Georges Bank stock of haddock (
Melanogrammus aeglefinus
) experienced the largest recruitment event recorded during its assessed history. Several hypotheses have been advanced to ...explain recruitment variability in this much-scrutinized stock, including variability in the retention of eggs and larvae on Georges Bank, the timing of haddock spawning, and variability in the spring bloom, which influences larval growth and survival. Although these processes may contribute to the formation of successful year classes, none of the factors associated with these previous hypotheses provides an adequate explanation of the 2003 recruitment event. We analyzed data on the dynamics of the fall phytoplankton bloom the year prior to spawning and show it to be highly correlated with subsequent recruitment. We suggest that the fall bloom affects recruitment through enhanced condition of adults and by increasing the quantity and quality of their reproductive output, which in turn leads to a higher probability of survival of their offspring. Although synoptic data on the fall bloom are limited and our analyses are correlative, our purpose is to stimulate a rigorous test of this promising "parental condition hypothesis".