Intra-annual Variability in Primary Producer Groups and Nitrogen Dynamics in an Intermittently Closed Estuary Exposed to Mediterranean Climate

Nutrient dynamics in estuaries are temporally variable in response to changing physical–chemical conditions and biogeochemical processes involving primary producer groups such as phytoplankton, microphytobenthos, seagrass and macroalgae. In order to reveal intra-annual changes in the biomass of primary producer groups and associated changes in estuarine nutrient dynamics, we developed a box model, coupling water inflows and outflows and nitrogen dynamics in Wilson Inlet, a large, central-basin-dominated, intermittently closed estuary exposed to a Mediterranean climate in Western Australia. The model is calibrated and validated with monitoring data, aquatic plant biomass estimates and biogeochemical rate measurements. Macrophytes and their microalgal epiphytes appear to rapidly assimilate nutrients from the first flush from the catchment in winter, but this buffer capacity then ceases, and a phytoplankton ‘bloom’ develops in response to subsequent river runoff events in spring. Significant amounts of bioavailable nitrogen are exported to the ocean because phytoplankton predominance occurs while the sand bar is breached. Surface sediments play a key role for nitrogen dynamics: In late spring to autumn, high light availability at the sediment surface stimulates high primary production by microphytobenthos, leading to reduced benthic ammonium fluxes particularly in the deep basin. Microphytobenthos contributes about 60% of annual whole-system primary production. Despite high benthic primary production, nitrogen release from sediments is the biggest nitrogen source to the estuary.     

Biogeochemical Zones Within a Macrotidal,Dry-Tropical Fluvial-Marine Transition Area: A Dry-Season Perspective

The Fitzroy River delivers large amounts of nutrients and fine sediments to Keppel Bay (contiguous with the Great Barrier Reef Lagoon) during intermittent flow events. This study explores sources, forms and transformations of nutrients in Keppel Bay, and develops a functional process zonation that integrates seabed geochemistry and water column nutrient characteristics which are controlled by suspended sediment. The water column and seabed properties were investigated over two dry seasons, with supplementary core incubations taken to measure carbon decomposition rates and nutrient fluxes. Keppel Bay can be divided into three zones, the: zone of maximum resuspension (ZMR); coastal transitional zone (CTZ); and blue water zone (BWZ). Mineralisation of predominantly terrestrial organic matter occurs in the ZMR where nutrient uptake by phytoplankton is light limited. The CTZ and BWZ had higher light penetration and phytoplankton growth was likely limited by N and P, respectively. The identified zones conform to the bathymetry and hydrodynamic characteristics of the bay, allowing for the development of an integrated conceptual model accounting for the benthic and pelagic biogeochemical processes. Recognition of these different zones shows that considerable variation in benthic and water column properties is possible within a small system with the bathymetric and hydrodynamic characteristics of the fluidized bed reactor.     

Measured and modeled effects of benthic algae on eutrophication in Indian River-Rehoboth Bay, Delaware

A combination of field studies and mathematical modeling was used to examine the role of subtidal benthic algae in the eutrophication processes in two shallow estuarine systems. Field measurements indicated uptake by benthic algae retained ammonium and phosphate in the sediments when light at the sediment surface exceeded ≈150 μE m² s^?1. The measurements were used to calibrate a newly developed model of benthic algal activity. The benthic algal model was coupled with a hydrodynamic model, a eutrophication model, and a sediment diagenesis model. In the simulated ecosystem, benthic algae had a major influence in the intra-annual cycling of nitrogen and phosphorus. When nutrients were abundant in the water column (late winter and spring) they were transferred to the sediments through algal activity. Diagenesis released these nutrients to the water column in summer when nutrients were scarce. As a result of the nutrient transfer, annual primary production in the water column, in the presence of benthic algae, exceeded production in the absence of the algae.     

Nutrient uptake in a highly turbid estuary (the Humber, United Kingdom) and adjacent coastal waters

Surveys were conducted in April and June 1995 to quantify the uptake of dissolved nutrients in a highly turbid estuary (the Humber, United Kingdom) and to determine the factors controlling nutrient uptake rates. A combination of isotope labelling methods were used in conjunction with on-deck incubation techniques to estimate the uptake of dissolved nutrients (PO₄ ^3?, NH₄ ⁺, NO₃ ^?, and urea) in surface samples collected from coastal waters. Similarly, isotope labelling and laboratory incubgation techniques were employed to estimate dissolved nitrogen uptake (NH₄ ⁺, NO₃ ^?, and urea) in surface samples collected from the estuary mouth. Nutrient uptake rates were at the low end of ranges for coastal and estuarine environments reported in the literature. Concentrations of chlorophyll and the availability of photosynthetically active radiation were identified as potentially important factors controlling the uptake rates of nutrients. Uptake rates of dissolved nitrogen in the Humber mouth appeared to be related to the location of smapling sites. Depletion rates of dissolved nutrients in situ were estimated on the basis of integrated water column nutrient uptake rates and indicated assimilation of up to 16% of nutrients in the entire water column. Estimated depletion rates did not indicate preferential loss of any of the nutrient species investigated.     

Impacts of Climate-Related Drivers on the Benthic Nutrient Filter in a Shallow Photic Estuary

In shallow photic systems, the benthic filter, including microphytobenthos and denitrifiers, is important in preventing or reducing release of remineralized NH₄ ⁺ to the water column. Its effectiveness can be impacted by climate-related drivers, including temperature and storminess, which by increasing wind and freshwater delivery can resuspend sediment, reduce salinity and deliver nutrients, total suspended solids, and chromophoric dissolved organic matter (CDOM) to coastal systems. Increases in temperature and freshwater delivery may initiate a cascade of responses affecting benthic metabolism with impacts on sediment properties, which in turn regulate nitrogen cycling processes that either sequester (via microphytobenthos), remove (via denitrification), or increase sediment nitrogen (via remineralization, nitrogen fixation, and dissimilatory nitrate reduction to ammonium). We conducted a seasonal study at shallow stations to assess the effects of freshwater inflow, temperature, wind, light, and CDOM on sediment properties, benthic metabolism, nitrogen cycling processes, and the effectiveness of the benthic filter. We also conducted a depth study to constrain seasonally varying parameters such as temperature to better assess the effects of light availability and water depth on benthic processes. Based on relationships observed between climatic drivers and response variables, we predict a reduction in the effectiveness of the benthic filter over the long term with feedbacks that will increase effluxes of N to the water column with the potential to contribute to system eutrophication. This may push shallow systems past a tipping point where trophic status moves from net autotrophy toward net heterotrophy, with new baselines characterized by degraded water quality.     

Ammonium and phosphate dynamics in a Virginia salt marsh

Experimental chambers were used in a Virginia salt marsh to partition the tidal flux of dissolved nutrients occurring at the marsh surface and in the water column. On five dates from June to October 1989, six replicate chambers in the short Spartina alterniflora zone were monitored over complete tidal cycles. When reservoir water, used to simulate tidal flooding in the chambers, was initially low in dissolved nutrients, the marsh surface was a source of both ammonium and phosphate to the water column. Calculations of the physical processes of diffusion and advection could not account for total nutrient release from the marsh surface. We hypothesize the primary source of nutrients was organic matter mineralization in surface sediments, which released nutrients into the flooding water column. Assimilation (uptake) of phosphate measured in water-column incubation experiments was nearly equal to phosphate released from the marsh surface. Surface release of ammonium, however, was somewhat greater than water-column uptake. In this salt marsh, benthic production and release of ammonium and phosphate is comparable in magnitude to pelagic consumption, thereby yielding only a small “net” transfer of these nutrients to the estuary.     

Going West: Nutrient Limitation of Primary Production in the Northern Gulf of Mexico and the Importance of the Atchafalaya River

To investigate controls on phytoplankton production along the Louisiana coastal shelf, we mapped salinity, nutrient concentrations (dissolved inorganic nitrogen (DIN) and phosphorus (P_i), silicate (Si)), nutrient ratios (DIN/P_i), alkaline phosphatase activity, chlorophyll and ¹⁴C primary productivity on fine spatial scales during cruises in March, May, and July 2004. Additionally, resource limitation assays were undertaken in a range of salinity and nutrient regimes reflecting gradients typical of this region. Of these, seven showed P_i limitation, five revealed nitrogen (N) limitation, three exhibited light (L) limitation, and one bioassay had no growth. We found the phytoplankton community to shift from being predominately N limited in the early spring (March) to P limited in late spring and summer (May and July). Light limitation of phytoplankton production was recorded in several bioassays in July in water samples collected after peak annual flows from both the Mississippi and Atchafalaya Rivers. We also found that organic phosphorus, as glucose-6-phosphate, alleviated P limitation while phosphono-acetic acid had no effect. Whereas DIN/P_i and DIN/Si ratios in the initial water samples were good predictors of the outcome of phytoplankton production in response to inorganic nutrients, alkaline phosphatase activity was the best predictor when examining organic forms of phosphorus. We measured the rates of integrated primary production (0.33?C7.01 g C m^?2 d^?1), finding the highest rates within the Mississippi River delta and across Atchafalaya Bay at intermediate salinities. The lowest rates were measured along the outer shelf at the highest salinities and lowest nutrient concentrations (<0.1 ??M DIN and Pi). The results of this study indicate that P_i limitation of phytoplankton delays the assimilation of riverine DIN in the summer as the plume spreads across the shelf, pushing primary production over a larger region. Findings from water samples, taken adjacent the Atchafalaya River discharge, highlighted the importance of this riverine system to the overall production along the Louisiana coast.     

Spatial and Temporal Variability of Benthic Oxygen Demand and Nutrient Regeneration in an Anthropogenically Impacted New England Estuary

Strong benthic–pelagic coupling is an important characteristic of shallow coastal marine ecosystems. Building upon a rich history of benthic metabolism data, we measured oxygen uptake and nutrient fluxes across the sediment–water interface along a gradient of water column primary production in Narragansett Bay, RI (USA). Despite the strong gradients seen in water column production, sediment oxygen demand (SOD) and benthic nutrient fluxes did not exhibit a clear spatial pattern. Some of our sites had been studied in the 1970s and 1980s and thus allowed historical comparison. At these sites, we found that SOD and benthic fluxes have not changed uniformly throughout Narragansett Bay. In the uppermost portion of the bay, the Providence River Estuary, we observed a significant decrease in dissolved inorganic phosphorus fluxes which we attribute to management interventions. At another upper bay site, we observed significant declines in SOD and dissolved inorganic nitrogen fluxes which may be linked to climate-induced decreases in water column primary production and shifts in bloom phenology. In the 1970s, benthic nutrient regeneration supplied 50% to over 200% of the N and P needed to support primary production by phytoplankton. Summer nutrient regeneration in the Providence River Estuary and Upper bay now may only supply some 5–30% of the N and 3–20% of the P phytoplankton demand.     

Benthic metabolism and nutrient cycling in Boston Harbor, Massachusetts

To gain insight into the importance of the benthos in carbon and nutrient budgets of Boston Harbor and surrounding bays, we measured sediment-water exchanges of oxygen, total carbon dioxide (DIC), nitrogen (ammonium, nitrate+nitrite, urea, N₂O), silicate, and phosphorus at several stations in different sedimentary environments just prior to and subsequent to cessation of sewage sludge disposal in the harbor. The ratio of the average annual DIC release to O₂ uptake at three primary stations ranged from 0.84 to 1.99. Annual average DIC:DIN flux ratios were consistently greater than predicted from the Redfield ratio, suggesting substantial losses of mineralized N. The pattern was less clear for P: some stations showed evidence that the sediments were a sink for P while others appeared to be a net source to the water column over the study period. In general, temporal and spatial patterns of respiration, nutrient fluxes, and flux ratios were not consistently related to measures of sediment oxidation-reduction status such as Eh or dissolved sulfide. Sediments from Boston Harbor metabolize a relatively high percentage (46%) of the organic matter inputs from phytoplankton production and allochthonous inputs when compared to most estuarine systems. Nutrient regeneration from the benthos is equivalent to 40% of the N, 29% of the P, and more than 60% of the Si demand of the phytoplankton. However, the role of the benthos in supporting primary production at the present time may be minor as nutrient inputs from sewage and other sources exceed benthic fluxes of N and P by 10-fold and Si by 4-fold. Our estimates of denitrification from DIC:DIN fluxes suggests that about 45% of the N mineralized in the sediments is denitrified, which accounts for about 17% of the N inputs from land.     

Carbon budget for a subtropical seagrass dominated coastal lagoon: How important are seagrasses to total ecosystem net primary production?

It has been assumed that because seagrasses dominate macrophyte biomass in many estuaries they also dominate primary production. We tested this assumption by developing three carbon budgets to examine the contribution of autotrophic components to the total ecosystem net primary production (TENPP) of Lower Laguna Madre, Texas. The first budget coupled average photosynthetic parameters with average daily irradiance to calculate daily production. The second budget used average photosynthetic parameters and hourly in situ irradiance to estimate productivity. The third budget integrated temperature-adjusted photosynthetic parameters (using Q₁₀=2) and hourly in situ irradiance to estimate productivity. For each budget TENPP was calculated by integrating production from each autotroph based on the producers’ areal distribution within the entire Lower Laguna Madre. All budgets indicated that macroalgae account for 33–42% of TENPP and seagrasses consistently accounted for about 33–38%. The contribution by phytoplankton was consistently about 15–20%, and the contribution from the benthic microalgae varied between 8% and 36% of TENPP, although this may have been underestimated due to our exclusion of the within bed microphytobenthos component. The water column over the seagrass beds was net heterotrophic and consequently was a carbon sink consuming between 5% and 22% of TENPP, TENPP ranged between 5.41×10¹⁰ and 2.53×10¹¹ g C yr⁻¹, depending on which budget was used. The simplest, most idealized budget predicted the highest TENPP, while the more realistic budgets predicted lower values. Annual production rates estimated using the third budget forHalodule urightii andThalassia testudinum compare well with field data. Macroalgae and microalgae contribute 50–60% of TENPP, and seagrass may be more important as three-dimensional habitat (i.e., structure) than as a source of organic carbon to the water column in Lower Laguna Madre.     

Inorganic phosphorus uptake in a carbonate-dominated seagrass ecosystem

Seasonal phosphate (Pi) uptake kinetics were determined using chambers encompassing the water column, sediment and the entire system (water column + sediment + seagrass/epiphyte) in Florida Bay (FB) during 2003–2006 and on the Little Bahama Bank (LBB) during a cruise June, 2004. Pi uptake was a linear function of concentration at low Pi levels (< 2 μmo11^-1). Applying the Pi system rate constant (S_p) from western (177 ±50 x 10^-6 m s^-1) and eastern (272 ±66 x 10^-6 m s^-1) bay sites, and using Pi measured during the study (0.02 to 0.177 μmol Pi 1^-1), we calculated a Pi uptake rate of 0.30 to 2.62 mmol Pi m^-2 d^-1 for western and 0.47 to 4.16 mmol Pi m^-2 d^-1 for eastern bay sites which includes phytoplankton uptake (0.455 m height). During non-bloom conditions, phytoplankton dominated Pi uptake in the east (46%) and both phytoplankton and the seagrass-epiphyte consortium in the west (32 and 52%, respectively), with a smaller contribution by the sediment (15–20%). On LBB interior sites, the water column always dominated (≽94%) Pi uptake with a higher S_p (573-881 x 10^-6 m s^-1) than FB. During cyanobacterial blooms in FB (chla 17 μg 1^-1), the water column dominated Pi uptake (100%) and S_p was the highest (>2,800 x 10^-6 m s^-1) measured. Phytoplankton accounted for 88% of this sequestered Pi with only 12% in the acid extractable fraction, likely as calcium bound and/or adsorbed P, and only 1% attributable to small heterotrophs. When chl α levels declined (2 μg I^-1) Pi uptake was still dominated by phytoplankton (77%), the acid extractable pool increased (18%) and the heterotrophic community became more important (22%). In carbonate-dominated seagrass systems, Pi is primarily taken up by the water column biota and is subsequently remineralized/hydrolyzed in the water column or settles to the benthos where it becomes available to benthic primary producers.     

Microphytobenthos: The ecological role of the “secret garden” of unvegetated,shallow-water marine habitats. I. Distribution,abundance and primary production

The microphytobenthos consists of unicellular eukaryotic algae and cyanobacteria that grow within the upper several millimeters of illuminated sediments, typically appearing only as a subtle brownish or greenish shading. The surficial layer of the sediment is a zone of intense microbial and geochemical activity and of considerable physical reworking. In many shallow ecosystems, the biomass of benthic microalgae often exceeds that of the phytoplankton in the overlying waters. Direct comparison of the abundance of benthic and suspended microalgae is complicated by the means used to measure biomass and by the vertical and horizontal distribution of the microphytobenthos in the sediment. Where biomass has been estimated as chlorophyll a, there may be negligible to large (40%) error due to interference by degradation products, except where chlorophyll is measured by high-performance liquid chromatography. The vertical distribution of microphytobenthos, aside from mat-forming species, is determined by the opposing effects of their vertical migration, which tends to concentrate them near the surface, and physical mixing by overlying currents, which tends to cause an even vertical distribution through the mixed layer of sediment. Uncertainties in vertical distribution are compounded by frequently patchy horizontal distribution. Under-sampling on small (<1 m) scales can lead to errors in the estimate that are comparable to the ranges of seasonal and geographic variation. These uncertainties are compounded by biases in the techniques used to estimate production by the microphytobenthos. In most environments studied, biomass (as chlorophyll a) and light availability appear to be the principal determinants of benthic primary production. The effect of variable light intensities on integral production can be described by a functional response curve. When normalized to the chlorophyll content of the surficial sediment, the residual variation in the data described by the functional response curve is due to changes in the chlorophyll-specific response to irradiance. Production by the benthos is often a significant fraction of production in the water column and microphytobenthos may contribute directly to water column production when they are resuspended. Thus on both the basis of biomass and biogeochemical reactivity, benthic microalgae play significant roles in system productivity and trophic dynamics, as well as such habitat characteristics as sediment stability. *** DIRECT SUPPORT *** A01BY074 00003     

Nutrient cycling in the sub-tropical Brunswick estuary,Australia

A combination of mixing plots, one-dimensional salt balance modelling, nutrient loading budgets, and benthic flux measurements were used to assess nutrient cycling pathways in the enriched sub-tropical Brunswick estuary during different freshwater flows. A simple model accounting for freshwater residence times and nutrient availability was found to be a good predictor of phytoplankton biomass along the estuary, and suggested that biomass accumulation may become nutrient-limited during low flows and that recycling within the water column is important during blooms. Dissolved inorganic nitrogen (DIN) cycling budgets were constructed for the estuary during different freshwater flows accounting for all major inputs (catchment, sewage, and urban) to the estuary. Internal cycling due to phytoplankton uptake (based on measured biomass) and sediment-water fluxes (based on measured rates in each estuarine reach) was considered. Four different nutrient cycling states were identified during the study. In high flow, freshwater residence times are less than 1 d, internal cycling processes are bypassed and virtually all dissolved, and most particulate, nutrients are delivered to the continental shelf. During the growth phase of a phytoplankton bloom enhanced recycling occurs as residence times increase sufficiently to allow biomass accumulation. Remineralization of phytoplankton detritus during this phase can supply up to 50% of phytoplankton DIN demands. In post-bloom conditions, DIN uptake by phytoplankton decreases in the autumn wet season when biomass doubling times begin to exceed residence times. OM supply to the sediments diminishes and the benthos becomes nutrient-limited, resulting in DIN uptake by the sediments. As flows decrease further in the dry season, there is tight recycling and phytoplankton blooms, and uptake by the sediments can account for the entire DIN loading to the estuary resulting in complete removal of DIN from the water column. The ocean is a potentially important source of DIN to the estuary at this time. The results of the DIN cycling budgets compared favorably with mixing plots of DIN at each time. The results suggest that a combination of different approaches may be useful in developing a more comprehensive understanding of nutrient cycling behavior and the effects of nutrient enrichment in estuaries.     

Nutrient-Replete Benthic Microalgae as a Source of Dissolved Organic Carbon to Coastal Waters

Dissolved organic carbon (DOC) flux dynamics were examined in the context of other biogeochemical cycles in intertidal sediments inhabited by benthic microalgae. In August 2003, gross oxygenic photosynthetic (GOP) rates, oxygen penetration depths, and benthic flux rates were quantified at seven sites along the Duplin River, GA, USA. Sediments contained abundant benthic microalgal (BMA) biomass with a maximum chlorophyll a concentration of 201 mg chl a m^?2. Oxygen microelectrodes were used to determine GOP rates and O₂ penetration depth, which were tightly correlated with light intensity. Baseline and ¹⁵N-nitrate amended benthic flux core incubations were employed to quantify benthic fluxes and to investigate the impact of BMA on sediment water exchange under nitrogen (N)-limited and N-replete conditions. Unamended sediments exhibited tight coupling between GOP and respiration and served as a sink for water column dissolved inorganic nitrogen (DIN) and a source of silicate and dissolved inorganic carbon (DIC). The BMA response to the N addition indicated sequential nutrient limitation, with N limitation followed by silicate limitation. In diel (light–dark) incubations, biological assimilation accounted for 83% to 150% of the nitrate uptake, while denitrification (DNF) and dissimilatory nitrate reduction to ammonium (DNRA) accounted for <7%; in contrast, under dark conditions, DNF and DNRA accounted for >40% of the NO₃ ^? uptake. The N addition shifted the metabolic status of the sediments from a balance of autotrophy and heterotrophy to net autotrophy under diel conditions, and the sediments served as a sink for water column DIN, silicate, and DIC but became a source of DOC, suggesting that the increased BMA production was decoupled from sediment bacterial consumption of DOC.     

Benthic nutrient remineralization in a coastal lagoon ecosystem

In situ measurements of the exchange of ammonia, nitrate plus nitrite, phosphate, and dissolved organic phosphorus between sediments and the overlying water column were made in a shallow coastal lagoon on the ocean coast of Rhode Island, U.S.A. The release of ammonia from mud sediments in the dark (20–440 μmol per m² per h) averaged ten times higher than from a sandy tidal flat (0–60 μmol per m² per h), and while mud sediments also released nitrate and phosphate, sandy sediments took up these nutrients. Fluxes of nutrients from mud sediments, but not from sandy areas, markedly increased with temperature. Ammonia release rates for mud sediments in the light (0–350 μmol per m² per h) were lower than those in the dark and it is estimated that some 25% of the ammonia released to the water column on an annual basis may be intercepted by the benthic microfloral community. Estimates of the annual net exchange of nutrients across the sediment-water interface, weighted by sediment type for the lagoon as a whole, showed a release of 450 mmol per m² of ammonia, 5 mmol per m² of phosphate, 5 mmol per m² of dissolved organic phosphorus, and an uptake of 80 mmol per m² of nitrate. Although rates of ammonia and nitrate exchange were comparable to those described for the deeper heterotrophic bottom communities of nearby Narragansett Bay, rates of benthic phosphate release were significantly lower. On an annual basis the Bay benthos released approximately 20 times more inorganic phosphate per unit area than did the lagoon benthos. As a result., the N/P ratio for the flux from the sediments was 74∶1 in the lagoon, compared with 16∶1 in “average” marine plankton and 8∶1 for the benthic flux from Narragansett Bay. The lack of remineralized phosphate in the lagoon, is reflected in water, column phosphate concentrations (always <1 μm) and water column N/P ratios (annual N/P=27) and suggests that the lagoon may show phosphate limitation rather than the nitrogen limitation commonly associated with marine systems.     

Sediment-water oxygen and nutrient exchanges along the longitudinal axis of Chesapeake Bay: Seasonal patterns,controlling factors and ecological significance

Sediment-water oxygen and nutrient (NH₄ ⁺, NO₃ ^?+NO₂ ^?, DON, PO₄ ^3?, and DSi) fluxes were measured in three distinct regions of Chesapeake Bay at monthly intervals during 1 yr and for portions of several additional years. Examination of these data revealed strong spatial and temporal patterns. Most fluxes were greatest in the central bay (station MB), moderate in the high salinity lower bay (station SB) and reduced in the oligohaline upper bay (station NB). Sediment oxygen consumption (SOC) rates generally increased with increasing temperature until bottom water concentrations of dissolved oxygen (DO) fell below 2.5 mg l^?1, apparently limiting SOC rates. Fluxes of NH₄ ⁺ were elevated at temperatures >15°C and, when coupled with low bottom water DO concentrations (<5 mg l^?1), very large releases (>500 μmol N m^?2 h^?1) were observed. Nitrate + nitrite (NO₃ ^?+NO₂ ^?) exchanges were directed into sediments in areas where bottom water NO₃ ^?+NO₂ ^? concentrations were high (>18 μM N); sediment efflux of NO₃ ^?+NO₂ ^? occurred only in areas where bottom water NO₃ ^?+NO₂ ^? concentrations were relatively low (<11 μM N) and bottom waters well oxygenated. Phosphate fluxes were small except in areas of hypoxic and anoxic bottom waters; in those cases releases were high (50–150 μmol P m^?2 h^?1) but of short duration (2 mo). Dissolved silicate (DSi) fluxes were directed out of the sediments at all stations and appeared to be proportional to primary production in overlying waters. Dissolved organic nitrogen (DON) was released from the sediments at stations NB and SB and taken up by the sediments at station MB in summer months; DON fluxes were either small or noninterpretable during cooler months of the year. It appears that the amount and quality of organic matter reaching the sediments is of primary importance in determining the spatial variability and interannual differences in sediment nutrient fluxes along the axis of the bay. Surficial sediment chlorophyll-a, used as an indicator of labile sediment organic matter, was highly correlated with NH₄ ^?, PO₄ ^3?, and DSi fluxes but only after a temporal lag of about 1 mo was added between deposition events and sediment nutrient releases. Sediment O:N flux ratios indicated that substantial sediment nitrification-denitrification probably occurred at all sites during winter-spring but not summer-fall; N:P flux ratios were high in spring but much less than expected during summer, particularly at hypoxic and anoxic sites. Finally, a comparison of seasonal N and P demand by phytoplankton with sediment nutrient releases indicated that the sediments provide a substantial fraction of nutrients required by phytoplankton in summer, but not winter, especially in the mid bay region.     

Spatial and temporal patterns in sediment and water column nutrients in a eutrophic Southern California estuary

Quarterly field sampling was conducted to characterize variations in water column and sediment nutrients in a eutrophic southern California estuary with a history of frequent macroalgal blooms. Water column and sediment nutrient measures demonstrated that Upper Newport Bay (UNB) is a highly enriched estuary. High nitrate (NO₃ ⁻) loads from the river entered the estuary at all sampling times with a rainy season (winter) maximum estimated at 2,419 mol h⁻¹. This resulted in water NO₃ ⁻ concentration in the estuary near the river mouth at least one order of magnitude above all other sampling locations during every seasons; maximum mean water NO₃ ⁻ concentration was 800 μM during springer 1997. Phosphorus (P)-loading was high year round (5.7–90.4 mol h⁻¹) with no seasonal pattern. Sediment nitrogen (N)-content showed a seasonal pattern with a spring maximum declining through fall. sediment and water nutrients, as well as percent cover of three dominant macroalgae, varied between the main channel and tidal creeks. During all seasons, water column NO₃ ⁻ concentrations were higher in the main channel than in tidal creeks while tidal creeks had higher levels of sediment total Kjeldhal nitrogen (TKN) and P. During each of the four sampling periods, percent cover ofEntermorpha intestinalis andCeramium spp. was higher in tidal creeks than in the main channel, while percent cover ofUlva expansa was always higher in the main channel. Decreases in sediment N in both creek and channel habitats were concurrent with increases in macroalgal cover, possibly reflecting use of stored sediment TKN by macroalgae. Our data suggest a shift in primary nutrient sources for macroalgae in UNB from riverine input during winter and spring to recycling from sediments duirng summer and fall.     

Insights into estuarine benthic dissolved organic carbon (DOC) dynamics using δC-DOC values, phospholipid fatty acids and dissolved organic nutrient fluxes

Benthic dissolved organic carbon (DOC) flux rates and changes in DOC isotope ratios, along with nutrient fluxes, phospholipid fatty acids concentration and carbon isotope ratios were measured in productive estuarine sediments over a diel cycle to determine the mechanisms driving benthic-pelagic coupling of DOC. There was uptake of DOC during the dark and efflux during the light at all sites. DOC uptake rates were related to benthic respiration (dark O₂ uptake) and effluxes were coupled to the trophic status (ratio of production to respiration) of the sediments. Highest uptake and efflux rates were observed at two high nutrient concentration sites. The DOC:DON ratio of water column dissolved organic matter (DOM) decreased during the dark and increased during the light indicating preferential uptake and release of carbon rich dissolved organic matter. The calculated carbon isotope ratio of the DOC taken up by the benthos was significantly more depleted than the bulk water column DOC pool, suggesting preferential uptake of selected components of the water column DOC pool. Generally the isotope ratio of the DOC released during the light was more enriched than that taken up during the dark, which suggests that the benthos has the potential to significantly alter the estuarine DOC pool. Uptake and efflux were coupled to respiration and algal grazing/mineralization, therefore increased nutrient loading may shift the composition of the estuarine DOC pool through changes in the magnitude of benthic DOC fluxes. A combination of biological (diel shifts in DOC production and consumption) and abiotic processes (flocculation) appear to be driving the observed benthic DOC dynamics at the study sites. This study was the first to measure carbon isotopic changes in the water column DOC pool due to benthic processes, and shows that the benthos can alter the estuarine DOC pool through diel differences in DOC uptake and efflux.     

Spatial and temporal patterns of factors influencing phytoplankton in a salt wedge estuary, the Swan River, Western Australia

During 1995 the phytoplankton in the Swan River were intensively sampled to assess biomass and species composition. Continuous measurements of fluorescence, salinity, and temperature were made weekly during 40 km sampling trips along the estuary and used to map the seasonal progression of the algal biomass. Weekly measurements of primary production were made and used to model net primary production from the vertical distribution of biomass, irradiance, and phytoplankton species composition. Potential nutrient limitation was assessed with “all but one” nutrient bioassays. The results indicate a complex mixture of potentially limiting factors, which vary in time and space. Although the data sequence is short, it suggests a annual succession pattern of diatoms, chlorophytes, diatoms, and finally dinoflagellates and cryptophytes in late summer-autumn. Peak seasonal biomass was observed during January to April. Mean annual chlorophylla biomass was greatest in upstream stations (5–9), where estimates of net primary production rates averaged 1.55 g C m^?2 d^?1 and gross primary production was 800–1000 g C m^?2 yr^?1. Potential nutrient limitation was most severe from November to May, although not during January 1995. Based on bioassay results, during the period of greatest potential for nutrient limitation, nitrogen was 15 to 30 times more limiting to biomass development than phosphate. Runoff due to consistent rainfall during winter eventually breaks down stratification and flushes the estuary with low-salinity, nutrient-rich water, producing, a light-limited, nutrient-rich aquatic ecosystem. Timing and magnitude of physical forcing events, mainly rainfall, appear critical in determining the susceptibility of this ecosystem to summer and autumn algal blooms.     

Spatial and Temporal Variations in <Emphasis Type="Italic">Pyrodinium bahamense</Emphasis> Cyst Concentrations in the Sediments of Bioluminescent Mangrove Lagoon,St. Croix,USVI

Mangrove Lagoon, located on the island of St. Croix, US Virgin Islands (USVI), is one of few actively bioluminescent lagoons in a location experiencing significant anthropogenic impacts. The bioluminescence is due to an abundance of the dinoflagellate Pyrodinium bahamense in the water column. We recovered surface sediments and sediment cores from Mangrove Lagoon to analyze the spatial distribution and temporal variability of P. bahamense cysts in this system. Surface sediment P. bahamense cyst concentrations ranged from 0 to 466 cysts g^?1 dry sediment, with higher abundances associated with elevated surface water nutrient concentrations and a mixed terrestrial–marine organic matter source regime. In combination with available bioassay data, we hypothesize that phytoplankton utilize nutrients rapidly and subsequent decay of organic matter makes nutrients available for dinoflagellates at the sediment–water interface in the eastern and northern quadrants of the lagoon. However, the nutrients are rapidly exhausted during counterclockwise lagoon circulation resulting in the decline of primary productivity and dinoflagellate abundance in the western quadrants. Downcore profiles suggest that P. bahamense blooms have been occurring for decades, declining in recent years. No cysts were present in sediments predating dredging activities of the 1960s that created Mangrove Lagoon. Recent reductions in cyst abundance may be the result of limited primary productivity caused by restricted water exchange with Salt River Bay due to shallowing of a sill at the mouth of the lagoon. This research highlights the need for more comprehensive geochemical and fossil analyses to better understand long-term ecological variability and inform conservation efforts of these unique habitats.