Phytoplankton reference communities for Chesapeake Bay and its tidal tributaries

Phytoplankton reference communities for Chesapeake Bay were quantified from least-impaired water quality conditions using commonly measured parameters and indicators derived from measured parameters. A binning approach was developed to classify water quality. Least-impaired conditions had relatively high water column transparency and low concentrations of dissolved inorganic nitrogen and orthophosphate. Reference communities in all seasons and salinity zones are characterized by consistently low values of chlorophylla and pheophytin coupled with relative stable proportions of the phytoplankton taxonomic groups and low biomasses of key bloom-forming species. Chlorophyll cell content was lower and less variable and average cell size and seasonal picophytoplankton biomass tended to be greater in the mesohaline and polyhaline reference communities as compared to the impaired communities. Biomass concentrations of the nano-micro phytoplankton size fractions (2–200 μm) in 12 of the 16 season-specific and salinity-specific reference communities were the same or higher than those in impaired habitat conditions, suggesting that nutrient reductions will not decrease the quantity of edible phytoplankton food available to large consumers. High (bloom) and low (bust) biomass events within the impaired phytoplankton communities showed strikingly different chlorophyll cell content and turnover rates. Freshwater flow had little effect on phytoplankton responses to water quality condition in most of the estuary. Improved water column transparency, or clarity, through the reduction of suspended sediments will be particularly important in attaining the reference communities. Significant nitrogen load reductions are also required.     

System-wide submerged aquatic vegetation model for Chesapeake Bay

A predictive model of submerged aquatic vegetation (SAV) biomass is coupled to a eutrophication model of Chesapeake Bay. Domain of the model includes the mainstem of the bay as well as tidal portions of major embayments and tributaries. Three SAV communities are modeled: ZOSTERA, RUPPIA, and FRESHWATER. The model successfully computes the spatial distribution and abundance of SAV for the period 1985–1994. Spatial distribution is primarily determined by computed light attenuation. Sensivitity analysis to reductions in nutrient and solids loads indicates nutrient controls will enhance abundance primarily in areas that presently support SAV. Restoration of SAV to areas in which it does not presently exist requires solids controls, alone or in combination with nutrient controls. For regions in which SAV populations exist at the refuge level or greater, improvements in SAV abundance are expected within 2 to 10 years of load reductions. For regions in which no refuge population exists, recovery time is unpredictable and will depend on propagule supply.     

Phytoplankton index of biotic integrity for Chesapeake Bay and its tidal tributaries

A Phytoplankton Index of Biotic Integrity (P-IBI) was developed from data collected during 18 yr 91985–2002) of the Chesapeake Bay Water Quality Monitoring Program. Dissolved inorganic nitrogen (DIN), orthophosphate (PO₄), and Secchi depth were used to characterize phytoplankton habitat conditions. Low DIN and PO₄ concentrations and high Secchi depths characterized least-impaire conditions. Thirty-eight phytoplankton metrics were tested for their ability to discriminate between impaired and least-impaired habitat conditions. Twelve discriminatory metrics were chosen, and different combinations of these twelve metrics were scored and used to create phytoplankton community indexes for spring and summer in the four salinity regimes in Chesapeake Bay. The scoring criteria for each metric were based on the distribution of the metric’s values in least-impaired conditions relative to the distribution in impaired conditions. An independent data set and jackknife validation procedure were used to examine P-IBI performance. The P-IBI correctly classified 70.0–84.4% of the impaired and least-impaired samples, grouped by season and salinity, in the calibration data set. The P-IBI is a management tool to assess phytoplankton community status relative to estuarine nutrient and light conditions.     

Changes in plankton community structure and function in response to variable freshwater flow in two tributaries of the Chesapeake Bay

The biomass of phytoplankton, microzooplankton, copepods, and gelatinous zooplankton were measured in two tributaries of the Chesapeake Bay during the springs of consecutive dry (below average freshwater flow), wet (above average freshwater flow), and average freshwater flow years. The potential for copepod control of microzooplankton biomass in the dry and wet years was evaluated by comparing the estimated grazing rates of microzooplankton by the dominant copepod species (Acartia spp. andEurytemora affinis) to microzooplankton growth rates and by calculating the percent of daily microzooplanton standing stock removed through copepod grazing. There were significant increases in phytoplankton and copepod biomass, but not for microzooplankton biomass in the wet year as compared to the dry year. The ctenophoreMnemiopsis leidyi was present during the dry year but was absent during the sampling period of the wet and average freshwater flow years. Grazing pressure on microzooplankton was greatest in the wet year, withAcartia spp. andE. affinis ingesting 0.21–2.64 μg of microzooplankton C copepod⁻¹ d⁻¹ and removing up to 60% of the microzooplankton standing stock per day. In the dry year, these copepod species ingested 0.10–0.73 μg of microzooplankton C copepod⁻¹ d⁻¹ with a maximum daily removal of approximately 3% of the microzooplankton standing stock. Potential copepod grazing pressure was significantly less than microzooplankton growth in the dry year, but was equivalent to microzooplankton growth in the wet year, implying strong top-down control of the microzooplankton community in the wet year. These results suggest that increased grazing control of microzooplankton populations by more copepods in the wet year released top-down control of phytoplankton. Reduced microzooplankton grazing, in conjunction with increased nutrient availability, resulted in large increases in phytoplankton biomass in the wet year. Increased freshwater flow has the potential to influence trophic cascades and the partitioning of plankton production in estuarine systems.     

The response of fishes to submerged aquatic vegetation complexity in two ecoregions of the mid-Atlantic bight: Buzzards Bay and Chesapeake Bay

Estuarine seagrass ecosystems provide important habitat for fish and invertebrates and changes in these systems may alter their ability to support fish. The response of fish assemblages to alteration of eelgrass (Zostera marina) ecosystems in two ecoregions of the Mid-Atlantic Bight (Buzzards Bay and Chesapeake Bay) was evaluated by sampling historical eelgrass sites that currently span a broad range of stress and habitat quality. In two widely separated ecoregions with very different fish faunas, degradation and loss of submerged aquatic vegetation (SAV) habitat has lead to declines in fish standing stock and species richness. The abundance, biomass, and species richness of the fish assemblage were significantly higher at sites that have high levels of eelgrass habitat complexity (biomass >100 wet g m^?2; density <100 shotts m^?2) compared to sites that have reduced eelgrass (biomass <100 wet g m^?2; density <100 shoots m^?2) or that have completely lost eelgrass. Abundance, biomass, and species richness at reduced eelgrass complexity sites also were more variable than at high eelgrass complexity habitats. Low SAV complexity sites had higher proportions of pelagic species that are not dependent on benthic habitat structure for feeding or refuge. Most species had greater abundance and were found more frequently at sites that have eelgrass. The replacement of SAV habitats by benthic macroalgae, which occurred in Buzzards Bay but not Chesapeake Bay, did not provide an equivalent habitat to seagrass. Nutrient enrichment-related degradation of eelgrass habitat has diminished the overall capacity of estuaries to support fish populations.     

Inputs,transformations, and transport of nitrogen and phosphorus in Chesapeake Bay and selected tributaries

In this paper we assemble and analyze quantitative annual input-export budgets for total nitrogen (TN) and total phosphorus (TP) for Chesapeake Bay and three of its tributary estuaries (Potomac, Patuxent, and Choptank rivers). The budgets include estimates of TN and TP sources (point, diffuse, and atmospheric), internal losses (burial in sediments, fisheries yields, and denitrification), storages in the water column and sediments, internal cycling rates (zooplankton excretion and net sediment-water flux), and net downstream exchange. Annual terrestrial and atmospheric inputs (average of 1985 and 1986 data) of TN and TP ranged from 4.3 g TN m^?2 yr^?1 to 29.3 g TN m^?2 yr^?1 and 0.32 g TP m^?2 yr^?1 to 2.42 g TP m^?2 yr^?1, respectively. These rates of TN and TP input represent 6-fold to 8-fold and 13-fold to 24-fold increases in loads to these systems since the precolonial period. A recent 11-yr record for the Susquehanna River indicates that annual loads of TN and TP have varied by about 2-fold and 4-fold, respectively. TN inputs increased and TP inputs decreased during the 11-yr period. The relative importance of nutrient sources varied among these estuaries: point sources of nutrients delivered about half the annual TN and TP load to the Patuxent and nearly 60% of TP inputs to the Choptank; diffuse sources contributed 60–70% of the TN and TP inputs to the mainstream Chesapeake and Potomac River. The direct deposition of atmospheric wet-fall to the surface waters of these estuaries represented 12% or less of annual TN and TP loads except in the Choptank River (37% of TN and 20% of TP). We found direct, although damped, relationships between annual rates of nutrient input, water-column and sediment nutrient stocks, and nutrient losses via burial in sediments and denitrification. Our budgets indicate that the annual mass balance of TN and TP is maintained by a net landward exchange of TP and, with one exception (Choptank River), a net seaward transport of TN. The budgets for all systems revealed that inorganic nutrients entering these estuaries from terrestrial and atmospheric sources are rapidly converted to particulate and organic forms. Discrepancies between our budgets and others in the literature were resolved by the inclusion of sediments derived from shoreline erosion. The greatest potential for errors in our budgets can be attributed to the absence of or uncertainties in estimates of atmospheric dry-fall, contributions of nutrients via groundwater, and the sedimentation rates used to calculate nutrient burial rates.     

Analysis of the abundance of submersed aquatic vegetation communities in the Chesapeake Bay

A procedure was developed using aboveground field biomass measurements of Chesapeake Bay submersed aquatic vegetation (SAV), yearly species identification surveys, annual photographic mapping at 1∶24,000 scale, and geographic information system (GIS) analyses to determine the SAV community type, biomass, and area of each mapped SAV bed in the bay and its tidal tributaries for the period of 1985 through 1996. Using species identifications provided through over 10,000 SAV ground survey observations, the 17 most abundant SAV species found in the bay were clustered into four species associations: ZOSTERA, RUPPIA, POTAMOGETON, and FRESHWATER MIXED. Monthly aboveground biomass values were then assigned to each bed or bed section based upon monthly biomass models developed for each community. High salinity communities (ZOSTERA) were found to dominate total bay SAV aboveground biomass during winter, spring, and summer. Lower salinity communities (RUPPIA, POTAMOGETON, and FRESHWATER MIXED) dominated in the fall. In 1996, total bay SAV standing stock was nearly 22,800 metric tons at annual maximum biomass in July encompassing an area of approximately 25,670 hectares. Minimum biomass in December and January of that year was less than 5,000 metric tons. SAV annual maximum biomass increased baywide from lows of less than 15,000 metric tons in 1985 and 1986 to nearly 25,000 metric tons during the 1991 to 1993 period, while area increased from approximately 20,000 to nearly 30,000 hectares during that same period. Year-to-year comparisons of maximum annual community abundance from 1985 to 1996 indicated that regrowth of SAV in the Chesapeake Bay from 1985–1993 occurred principally in the ZOSTERA community, with 85% of the baywide increase in biomass and 71% of the increase in are a occurring in that community. Maximum biomass of FRESHWATER MIXED SAV beds also increased from a low of 3,200 metric tons in 1985 to a high of 6,650 metric tons in 1993, while maximum biomass of both RUPPIA and POTAMOGETON beds fluctuated between 2,450 and 4,600 metric tons and 60 and 600 metric tons, respectively, during that same period with net declines of 7% and 43%, respectively, between 1985 and 1996. During the July period of annual, baywide, maximum SAV biomass, SAV beds in the Chesapeake Bay typically averaged approximately 0.86 metric tons of aboveground dry mass per hectare of bed area.     

Oligohaline benthic invertebrate communities at two Chesapeake Bay power plants

Benthic invertebrate populations at the Surry power plant on James River, Virginia and the C. P. Crane power plant on Saltpeter Creek, Maryland exhibited large spatial and temporal variations. At C. P. Crane, where the cooling water is pumped between two tidal creeks, populations in the receiving creek exhibited five response patterns: 1) mitigation of a winter dieoff (Rangia cuneata, a brackish water clam), 2) acceleration of growth or development (R. cuneata; Scolecolepides viridis, a polychaete;Leptocheirus plumulosus, an amphipod; Tubificidae; andCoelotanypus sp., a dipteran), 3) importantion of larvae from the source water creek (S. viridis andCoelotanypus sp.), 4) extension of creek-dwelling species into the adjacent river (Coelotanypus sp. and other dipterans), and 5) increased severity of late summer population depressions (S. viridis andL. plumulosus). At Surry, where the cooling was no confined creek system at the discharge, and effects were less pronounced. The community in the Surry discharge zone resembled the community at downriver stations more closely than at upriver reference stations. TheL. plumulosus population near the Surry discharge was depressed, while theMytilopsis leucophaeta population was enhanced. No major ecological damage was attributed to either power plant, due in part to the resilience of estuarine endemic populations, but the unique features exhibited by each of the two sites support the argument that oligohaline estuarine zones should not be designateda priori for unregulated industrial development.     

Distribution and abundance of submerged aquatic vegetation in Chesapeake Bay: An historical perspective

An historical summary of the distribution and abundance of submerged aquatic vegetation (SAV) in the Chesapeake Bay is presented. Evidence suggests that SAV has generally been common throughout the bay over the last several hundred years with several fluctuations in abundance. The decline ofZostera marina (eelgrass) in the 1930’s and the rapid expansion ofMyriophyllum spicatum (watermilfoil) in the late 1950’s and early 1960’s were two significant events involving a single species. Since 1965, however, there has been a significant reduction of all species in most sections of the bay. Declines were first observed in the Patuxent, Potomac and sections of other rivers in the Maryland portion of the Bay between 1965 and 1970. Dramatic reductions were observed over the entire length of the bay from 1970 to 1975. Particularly severe losses were observed at the head of the bay around Susquehanna Flats as well as in numerous rivers along Maryland’s eastern and western shores. Changes in the lower, Virginia portion of the bay occurred primarily in the western tributaries. Greatest losses of vegetation occurred in the years following Tropical Storm Agnes in 1972. Since 1975 little regrowth has been observed in the Chesapeake Bay. Other areas along the Atlantic Coast of the U.S. during the same period have experienced no similar widespread decline. It thus appears that the factors affecting the recent changes in distribution and abundance of submerged vegetation in the bay are regional in nature. Causes for this decline may be related to changes in water quality, primarily increased eutrophication and turbidity.     

A pollution history of Chesapeake Bay

Present day anthropogenic fluxes of some heavy metals to central Chesapeake Bay appear to be intermediate to those of the southern California coastal region and those of Narragansett Bay. The natural fluxes, however, are in general higher. On the bases of Pb-210 and Pu-239 + 240 geochronologies and of the time changes in interstitial water compositions, there is a mixing of the upper 30 or so centimeters of the sediments in the mid-Chesapeake Bay area through bioturbation by burrowing mollusks and polychaetes. Coal, coke and charcoal levels reach one percent or more by dry weight in the deposits, primarily as a consequence of coal mining operations.     

Effects of watershed and estuarine characteristics on the abundance of submerged aquatic vegetation in Chesapeake Bay subestuaries

Watershed land use can affect submerged aquatic vegetation (SAV) by elevating nutrient and sediment loading to estuaries. We analyzed the effects of watershed use and estuarine characteristics on the spatial variation of SAV abundance among 101 shallow subestuaries of Chesapeake Bay during 1984–2003. Areas of these subestuaries range from 0.1 to 101 km², and their associated local watershed areas range from 6 to 1664 km². Watershed land cover ranges from 6% to 81% forest, 1% to 64% cropland, 2% to 38% grassland, and 0.3% to 89% developed land. Landscape analyses were applied to develop a number of subestuary metrics (such as subestuary area, mouth width, elongation ratio, fractal dimension of shoreline, and the ratio of local watershed area to subestuary area) and watershed metrics (such as watershed area). Using mapped data from aerial SAV surveys, we calculated SAV coverage for each subestuary in each year during 1984–2003 as a proportion of potential SAV habitat (the area < 2 m deep). The variation in SAV abundance among subestuaries was strongly linked with subestuary and watershed characteristics. A regression tree model indicated that 60% of the variance in SAV abundance could be explained by subestuary fractal dimension, mean tidal range, local watershed dominant land cover, watershed to subestuary area ratio, and mean wave height. Similar explanatory powers were found in wet and dry years, but different independent variables were used. Repeated measures ANOVA with multiple-mean comparison showed that SAV abundance declined with the dominant watershed land cover in the order: forested, mixed-undisturbed, or mixed-developed > mixed-agricultural > agricultural > developed. Change-point analyses indicated strong threshold responses of SAV abundance to point source total nitrogen and phosphorus inputs, the ratio of local watershed area to subestuary area, and septic system density in the local watershed.     

Seasonal oxygen depletion in Chesapeake Bay

The spring freshet increases density stratification in Chesapeake Bay and minimizes oxygen transfer from the surface to the deep layer so that waters below 10 m depth experiece oxygen depletion which may lead to anoxia during June to September. Respiration in the water of the deep layer is the major factor contributing to oxygen depletion. Benthic respiration seems secondary. Organic matter from the previous year which has settled into the deep layer during winter provides most of the oxygen demand but some new production in the surface layer may sink and thus supplement the organic matter accumulated in the deep layer.     

Modeling of wind-induced destratification in Chesapeake Bay

It has been observed that storms in early fall can result in top-to-bottom mixing of Chesapeake Bay. A three-dimensional, time-dependent circulation model is used to examine this destratification process for September 1983, when extensive current and hydrographic data were available. The model bay is forced at the surface by observed hourly winds, at the ocean boundary by observed hourly surface and bottom salinities and sea level fluctuations, and at the head by observed daily discharges for a 28-d period. A second-moment, turbulence-closure submodel, with no adjustments from previous applications to its requisite coefficients, is used to calculate the vertical turbulence mixing coefficients. Comparisons with data inside the model domain indicate relative errors of 7% to 14% for sea level, 7% to 35% for current, and 11% to 21% for salinity. The tidal portion of the spectrum is modeled better than the subtidal portion. The model is used to examine both the mechanisms of wind mixing and the temporal and spatial distribution of vertical mixing within the estuary. Wind-driven internal shear is shown to be a more effective mechanism of inducing destratification than turbulence generated at the surface. The model is also used to show that the vertical temperature inversion which occurs in the fall does not affect the timing of the destratification as much as its completeness. The distribution of mid-depth vertical mixing shows highly variable values in the mid-bay region, where wind-induced mixing is dominant. This suggests that the source of oxygen to mid-bay bottom waters is similarly variable. Vertical turbulence mixing coefficients of 10^?2 cm² s^?1 (background) to 10³ cm² s^?1 were needed to simulate the September period, indicating the need for time-variable mixing in models of dissolved and suspended estuarine constituents.     

Wave-forced resuspension of upper Chesapeake Bay muds

Moored instruments were used to make observations of near bottom currents, waves, temperature, salinity, and turbidity at shallow (3.5 m and 5.5 m depth) dredged sediment disposal sites in upper Chesapeake Bay during the winters of 1990 and 1991 to investigate time-varying characteristics of resuspension processes over extended periods. Resulting time series data show the variability of two components of the suspended sediment concentration field. Background suspended sediment concentrations varied inversely with salinity and in direct relation to Susquehanna River flow. Muddy bottom sediments were also resuspended locally by both tidal currents and wind-wave forcing, resulting in short-term increases and decreases in suspended concentration, with higher peak concentrations near the bottom. In both years, episodes of wave-forced resuspension dominated tidal resuspension on an individual event basis, exceeding most tidal resuspension peaks by a factor of 3 to 5. The winds that generated the waves responsible for the observed resuspension events were not optimal for wave generation, however. Application of a simple wind-wave model showed that much greater wave-forced resuspension than that observed might be generated under the proper conditions. The consolidated sediments investigated in 1990 were less susceptible to both tidal and wave-forced resuspension than the recently deposited sediments investigated in 1991. There was also some indication that wave-forced resuspension increased erodibility of the bottom sediments on a short-term basis. Wave-forced resuspension is implicated as an important part of sediment transport processes in much of Chesapeake Bay. Its role in deeper, narrower, and more tidally energetic estuaries is not as clear, and should be investigated on a case-by-case basis.     

Observations of a Chesapeake Bay tidal front

This paper presents combined conductivity-temperature-depth (CTD), thermistor chain, current meter, and acoustic backscatter observations of a tidal front observed in the Chesapeake Bay. The data were obtained from a moored platform as the front migrated past the platform. The thermistor chain and CTD data show an interface that slopes steeply down from the surface to an asymptotic depth of 6 m, marking the bottom of the light-water pool. The thermistor chain data show much higher activity levels within the light-water pool as compared to the dense-water pool. Current meter data taken at 3 m show a pronounced shear in the currents upon crossing the frontal boundary. The acoustic backscatter from a layer of copepods positioned on the interface shows episodic occurrences of overturning at the interface. This observation is borne out by the concurrent thermistor chain data, which also show the overturning events.     

Sources, input pathways, and distributions of Fe, Cu, and Zn in a Chesapeake Bay tidal freshwater marsh

Tidal freshwater marshes exist at the interface between watersheds and estuaries, and thus may serve as critical buffers protecting estuaries from anthropogenic metal pollution. Bi-weekly samples of newly deposited marsh sediments were collected and analyzed for Cu, Zn, and Fe concentrations over 21 months from July 1995 to March 1997 in five distinct habitats at the head of Bush River, Maryland. Bi-weekly anthropogenic metal enrichments ranged from 0.9–4.7. Anthropogenic excess metal loadings averaged over 1996 ranged from 6–306 and 25–1302 μg cm⁻² year⁻¹ between sites for Cu and Zn, respectively. Based on Fe-normalized trace metal signatures, Susquehanna River sediment does not significantly contribute to upper Bush River. Organic matter was found to dilute total metal concentrations, whereas past studies suggested organics enhance labile metal content. Analysis of metal input pathways shows that marsh metals are primarily imported from nearby subtidal accumulations of historic watershed material by tidal flushing. Received: 29 April 1999 / Accepted: 7 December 1999     

Fish biomass size spectra in Chesapeake Bay

Biomass size spectra of pelagic fish were modeled to describe community structure, estimate potential fish production, and delineate trophic relationships in Chesapeake Bay. Spectra were constructed from midwater trawl collections each year in April, June–August, and October 1995–2000. The size spectra were bimodal: the first spectral dome corresponded to small zooplanktivorous fish, primarily bay anchovyAnchoa mitchilli; the second dome consisted of larger fish from several feeding guilds that are supported by multiple prey-predator linkages. Annual production estimates of pelagic fish, derived from a mean production to biomass ratio, varied nearly three-fold, ranging from 162 × 109 kcal (125 × 103 tons) in 1996 to 457 × 109 kcal (352 × 103 tons) in 2000. Seasonally, the biomass level and mean individual sizes of fish in the first dome increased from April to October, while the biomass level of the second dome was relatively stable. Regionally, biomass levels in the second dome were higher than biomasses in the first dome for the upper and lower Bay, but were minimal in the middle Bay where seasonal and episodic hypoxia occurs. To test a benthic-pelagic coupling hypothesis that could explain the higher biomass in the second domes for the lower and upper Bay, a cyclic size-spectrum model was fit that included only species in the zooplanktivorous-piscivorous fish guilds. The mean, normalized slope equaled ?1, indicating that zooplanktivorous fish may support piscivore production, but that a benthic-pelagic linkage is required to fully support fish production in the second dome. Interannual variability in slopes and intercepts of modeled size spectra was related to salinity, recruitment level of bay anchovy, and the primary axis of a correspondence analysis (salinity effect) on fish community structure. The spectral slope and intercept of normalized spectra were lowest in 1996, a near-record wet year. Results suggest that fish size spectra can be developed as useful indicators of ecosystem state and response to perturbations, especially if prey-predator relationships are explicitly represented.     

Wind Modulation of Dissolved Oxygen in Chesapeake Bay

A numerical circulation model with a simplified dissolved oxygen module is used to examine the importance of wind-driven ventilation of hypoxic waters in Chesapeake Bay. The model demonstrates that the interaction between wind-driven lateral circulation and enhanced vertical mixing over shoal regions is the dominant mechanism for providing oxygen to hypoxic sub-pycnocline waters. The effectiveness of this mechanism is strongly influenced by the direction of the wind forcing. Winds from the south are most effective at supplying oxygen to hypoxic regions, and winds from the west are shown to be least effective. Simple numerical simulations demonstrate that the volume of hypoxia in the bay is nearly 2.5 times bigger when the mean wind is from the southwest as compared to the southeast. These results provide support for a recent analysis that suggests much of the long-term variability of hypoxia in Chesapeake Bay can be explained by variations in the summertime wind direction.     

Scales of nutrient-limited phytoplankton productivity in Chesapeake Bay

The scales on which phytoplankton biomass vary in response to variable nutrient inputs depend on the nutrient status of the plankton community and on the capacity of consumers to respond to increases in phytoplankton productivity. Overenrichment and associated declines in water quality occur when phytoplankton growth rate becomes nutrient-saturated, the production and consumption of phytoplankton biomass become uncoupled in time and space, and phytoplankton biomass becomes high and varies on scales longer than phytoplankton generation times. In Chesapeake Bay, phytoplankton growth rates appear to be limited by dissolved inorganic phosphorus (DIP) during spring when biomass reaches its annual maximum and by dissolved inorganic nitrogen (DIN) during summer when phytoplankton growth rates are highest. However, despite high inputs of DIN and dissolved silicate (DSi) relative to DIP (molar ratios of N∶P and Si∶P>100), seasonal accumulations of phytoplankton biomass within the salt-intruded-reach of the bay appear to be limited by riverine DIN supply while the magnitude of the spring diatom bloom is governed by DSi supply. Seasonal imbalances between biomass production and consumption lead to massive accumulations of phytoplankton biomass (often>1,000 mg Chl-a m^?2) during spring, to spring-summer oxygen depletion (summer bottom water <20% saturation), and to exceptionally high levels of annual phytoplankton production (>400 g m^?2 yr^?1). Nitrogen-dependent seasonal accumulations of phytoplankton biomass and annual production occur as a consequence of differences in the rates and pathways of nitrogen and phosphorus cycling within the bay and underscore the importance of controlling nitrogen inputs to the mesohaline and lower reaches of the bay.