Long-term variation of macrobenthos in a mesohaline region of Chesapeake Bay

Macrobenthic species abundances and physical/chemical factors known of affect them were measured in a mesohaline region of the Chesapeake Bay from 1971 to 1982. Variation in species abundance due to station differences, seasonal patterns and year-to-year fluctuations in physical/chemical factors were quantified. The major source of variation in abundance of most species was that associated with seasonal recruitment cycles. Spatial variation in abundances of dominant species was small within habitats defined by sediment characteristics. However, abundance variation among sediment types was relatively large. All species showed significant year-to-year fluctuation in abundance, but no species had systematic long-term increases or decreases in abundance during the study period. Macrobenthos populations were persistent over the 11 years within abundance boundaries defined chiefly by fluctuations in salinity and dissolved oxygen concentration. Most species responded to salinity changes, especially those of an extreme nature, with predictable increases or decreases in abundance. Abundances of all species declined rapidly under conditions of low dissolved oxygen concentration (<2 ppm). Macrobenthos with planktonic developmental stages rapidly repopulated the region following periods of population decline. Repopulation by brooding species was slower.     

Long-Term Trends in Submersed Aquatic Vegetation (SAV) in Chesapeake Bay, USA, Related to Water Quality

Chesapeake Bay supports a diverse assemblage of marine and freshwater species of submersed aquatic vegetation (SAV) whose broad distributions are generally constrained by salinity. An annual aerial SAV monitoring program and a bi-monthly to monthly water quality monitoring program have been conducted throughout Chesapeake Bay since 1984. We performed an analysis of SAV abundance and up to 22 environmental variables potentially influencing SAV growth and abundance (1984–2006). Historically, SAV abundance has changed dramatically in Chesapeake Bay, and since 1984, when SAV abundance was at historic low levels, SAV has exhibited complex changes including long-term (decadal) increases and decreases, as well as some large, single-year changes. Chesapeake Bay SAV was grouped into three broad-scale community-types based on salinity regime, each with their own distinct group of species, and detailed analyses were conducted on these three community-types as well as on seven distinct case-study areas spanning the three salinity regimes. Different trends in SAV abundance were evident in the different salinity regimes. SAV abundance has (a) continually increased in the low-salinity region; (b) increased initially in the medium-salinity region, followed by fluctuating abundances; and (c) increased initially in the high-salinity region, followed by a subsequent decline. In all areas, consistent negative correlations between measures of SAV abundance and nitrogen loads or concentrations suggest that meadows are responsive to changes in inputs of nitrogen. For smaller case-study areas, different trends in SAV abundance were also noted including correlations to water clarity in high-salinity case-study areas, but nitrogen was highly correlated in all areas. Current maximum SAV coverage for almost all areas remain below restoration targets, indicating that SAV abundance and associated ecosystem services are currently limited by continued poor water quality, and specifically high nutrient concentrations, within Chesapeake Bay. The nutrient reductions noted in some tributaries, which were highly correlated to increases in SAV abundance, suggest management activities have already contributed to SAV increases in some areas, but the strong negative correlation throughout the Chesapeake Bay between nitrogen and SAV abundance also suggests that further nutrient reductions will be necessary for SAV to attain or exceed restoration targets throughout the bay.     

Effects of low dissolved oxygen events on the macrobenthos of the lower Chesapeake Bay

The effects of low dissolved oxygen or hypoxia (<2 mg l^?1) on macrobenthic infaunal community structure and composition in the lower Chesapeake Bay and its major tributaries, the Rappahannock, York, and James rivers are reported. Macrobenthic communities at hypoxia-affected stations were characterized by lower species diversity, lower biomass, a lower proportion of deep-dwelling biomass (deeper than 5 cm in the sediment), and changes in community composition. Higher dominance in density and biomass of opportunistic species (e.g., euryhaline annelids) and lower dominance of equilibrium species (e.g., long-lived bivalves and maldanid polychaetes) were observed at hypoxia-affected stations. Hypoxia-affected macrobenthic communities were found in the polyhaline deep western channel of the bay mainstem north of the Rappahannock River and in the mesohaline region of the lower Rappahannock River. No hypoxic effects on the infaunal macrobenthos were found in the York River, James River, or other deep-water channels of the lower Chesapeake Bay.     

Land Use and Salinity Drive Changes in SAV Abundance and Community Composition

Conserving and restoring submerged aquatic vegetation (SAV) are key management goals for estuaries worldwide because SAV integrates many aspects of water quality and provides a wide range of ecosystem services. Management strategies are typically focused on aggregated abundance of several SAV species, because species cannot be easily distinguished in remotely sensed data. Human land use and shoreline alteration have been shown to negatively impact SAV abundance, but the effects have varied with study, spatial scale, and location. The differences in reported effects may be partly due to the focus on abundance, which overlooks within-community and among-community dynamics that generate total SAV abundance. We analyzed long-term SAV aerial survey data (1984–2009) and ground observations of community composition (1984–2012) in subestuaries of Chesapeake Bay to integrate variations in abundance with differences in community composition. We identified five communities (mixed freshwater, milfoil-Zannichellia, mixed mesohaline, Zannichellia, and Ruppia-Zostera). Temporal variations in SAV abundance were more strongly related to community identity than to terrestrial stressors, and responses to stressors differed among communities and among species. In one fifth of the subestuaries, the community identity changed during the study, and the probability of such a change was positively related to the prevalence of riprapped shoreline in the subestuary. Mixed freshwater communities had the highest rates of recovery, and this may have been driven by Hydrilla verticillata, which was the single best predictor of SAV recovery rate. Additional species-specific and community-specific research will likely yield better understanding of the factors affecting community identity and SAV abundance, more accurate predictive models, and more effective management strategies.     

Structural components of eelgrass (Zostera marina) meadows in the lower Chesapeake Bay—Decapod crustacea

Otter trawl collections of eelgrass habitats in the lower Chesapeake Bay during 1976–1977 produced 14 species of decapod crustaceans. These collections were dominated by palaemonid shrimp (Palaemonetes spp.), blue crabs (Callinectes sapidus), and sand shrimp (Crangon septemspinosa), each of which exhibited unimodal seasonal abundance curves with large summer peaks. Decapod abundance was positively correlated with plant biomass throughout the year. Decapod densities on vegetated bottoms were greater than on unvegetated bottoms, and nighttime abundance on each bottom type was greater than corresponding daytime abundance. Total decapod abundances in Chesapeake Bay eelgrass meadows appear to be much greater than those reported in North Carolina eelgrass or Gulf of Mexico turtlegrass habitats.     

Relationship between Hypoxia and Macrobenthic Production in Chesapeake Bay

Human development has degraded Chesapeake Bay's health, resulting in an increase in the extent and severity of hypoxia (≤2 mg O₂ l^-1). The Bay's hypoxic zones have an adverse effect on both community structure and secondary production of macrobenthos. From 1996 to 2004, the effect of hypoxia on macrobenthic production was assessed in Chesapeake Bay and its three main tributaries (Potomac, Rappahannock, and York Rivers). Each year, in the summer (late July???early September), 25 random samples of the benthic macrofauna were collected from each system, and macrobenthic production in the polyhaline and mesohaline regions was estimated using Edgar's allometric equation. Fluctuations in macrobenthic production were significantly correlated with dissolved oxygen. Macrobenthic production was 90 % lower during hypoxia relative to normoxia. As a result, there was a biomass loss of ~7,320–13,200 metric tons C over an area of 7,720 km², which is estimated to equate to a 20 % to 35 % displacement of the Bay's macrobenthic productivity during the summer. While higher consumers may benefit from easy access to stressed prey in some areas, the large spatial and temporal extent of seasonal hypoxia limits higher trophic level transfer, via the inhibition of macrobenthic production. Such a massive loss of macrobenthic production would be detrimental to the overall health of the Bay, as it comes at a time when epibenthic and demersal predators have high-energy demands.     

Long-Term Trends of Nutrients and Phytoplankton in Chesapeake Bay

Climate effects on hydrology impart high variability to water-quality properties, including nutrient loadings, concentrations, and phytoplankton biomass as chlorophyll-a (chl-a), in estuarine and coastal ecosystems. Resolving long-term trends of these properties requires that we distinguish climate effects from secular changes reflecting anthropogenic eutrophication. Here, we test the hypothesis that strong climatic contrasts leading to irregular dry and wet periods contribute significantly to interannual variability of mean annual values of water-quality properties using in situ data for Chesapeake Bay. Climate effects are quantified using annual freshwater discharge from the Susquehanna River together with a synoptic climatology for the Chesapeake Bay region based on predominant sea-level pressure patterns. Time series of water-quality properties are analyzed using historical (1945–1983) and recent (1984–2012) data for the bay adjusted for climate effects on hydrology. Contemporary monitoring by the Chesapeake Bay Program (CBP) provides data for a period since mid-1984 that is significantly impacted by anthropogenic eutrophication, while historical data back to 1945 serve as historical context for a period prior to severe impairments. The generalized additive model (GAM) and the generalized additive mixed model (GAMM) are developed for nutrient loadings and concentrations (total nitrogen—TN, nitrate?+?nitrate—NO₂?+?NO₃) at the Susquehanna River and water-quality properties in the bay proper, including dissolved nutrients (NO₂?+?NO₃, orthophosphate—PO₄), chl-a, diffuse light attenuation coefficient (K _D (PAR)), and chl-a/TN. Each statistical model consists of a sum of nonlinear functions to generate flow-adjusted time series and compute long-term trends accounting for climate effects on hydrology. We present results identifying successive periods of (1) eutrophication ca. 1945–1980 characterized by approximately doubled TN and NO₂?+?NO₃ loadings, leading to increased chl-a and associated ecosystem impairments, and (2) modest decreases of TN and NO₂?+?NO₃ loadings from 1981 to 2012, signaling a partial reversal of nutrient over-enrichment. Comparison of our findings with long-term trends of water-quality properties for a variety of estuarine and coastal ecosystems around the world reveals that trends for Chesapeake Bay are weaker than for other systems subject to strenuous management efforts, suggesting that more aggressive actions than those undertaken to date will be required to counter anthropogenic eutrophication of this valuable resource.     

The Influence of Storms on Water Quality and Phytoplankton Dynamics in the Tidal James River

Due to the unpredictable nature of intense storms and logistical constraints of sampling during storms, little is known about their immediate and long-term impacts on water quality in adjacent aquatic ecosystems. By combining targeted experiments with routine monitoring, we evaluated immediate impacts of two successive storm events on water quality and phytoplankton community response in the tidal James River and compared these findings to a non-storm year. The James River is a subestuary of the Chesapeake Bay and sampling was conducted before, during, and after Hurricane Irene and Tropical Storm (TS) Lee in 2011 and during the same time period (late summer/early fall) in 2012 when there were no storms. We collected and compiled data on nutrient and chlorophyll a concentrations, phytoplankton abundance, nitrogen uptake, primary productivity rates, and surface salinity, temperature, and turbidity in the meso- and polyhaline segments of the James River. Hurricane Irene introduced significant amounts of freshwater over the entire James River and Chesapeake Bay watersheds, while rainfall from TS Lee fell primarily on the tidal fresh region of the James River and headwaters of the Chesapeake Bay. Dinoflagellates dominated the algal community in the meso- and polyhaline segments prior to the storms in 2011, and a mixed diatom community emerged after the storms. In the mesohaline river segment, cyanobacteria abundance increased after TS Lee when salinities were depressed, likely due to washout from the oligohaline and tidal fresh regions of the river. In 2012, dinoflagellates dominated the community in both segments of the river during late summer but diatoms were also abundant and their biomass fluctuated throughout the summer and fall. Cyanobacteria were not present in either segment. Overall, we observed that the high-intensity rainfall from Hurricane Irene combined with high flushing in the headwaters as a result of TS Lee likely reduced primary productivity and altered community composition in the mesohaline segment but not the more estuarine-influenced polyhaline segment. Understanding the influence of high freshwater flow with a short residence time associated with storms is key to the planning and management of estuarine restoration as such disturbances are projected to increase as a result of climate change.     

Spatial and Interannual Variability in Winter Mortality of the Blue Crab (<Emphasis Type="Italic">Callinectes sapidus</Emphasis>) in the Chesapeake Bay

The blue crab, Callinectes sapidus, is an ecologically and economically valuable species in Chesapeake Bay. Field surveys and laboratory experiments indicate that blue crab mortality is significant during severe winters. We applied a temperature and salinity-dependent survival model to empirical temperature and salinity data to explore spatial and interannual patterns in overwintering mortality. Harmonic regression analysis and geostatistical techniques were used to create spatially explicit maps of estimated winter duration, average temperature, average salinity, and resulting crab survival probability for the winters of 1990–2004. Predicted survival was highest in the warmer, saline waters of the lower Bay and decreased with increasing latitude up bay. There was also significant interannual variation with survival being lowest after the severe winters of 1996 and 2003. We combine the survival probability maps with maps of blue crab abundance to show how winter mortality may reduce blue crab abundance prior to the start of the harvesting season.     

Spatial and temporal differences of Atlantic menhaden (Brevoortia tyrannus) recruitment across major drainages (1966–2004) of the Chesapeake Bay watershed

Atlantic menhaden (Brevoortia tyrannus) is well known for its commercial and ecological importance and has been historically declining in the Chesapeake Bay (Maryland), one of its principal nursery habitats along the eastern coast. Using data from the Striped Bass Seine Survey of the Maryland Department of Natural Resources (2003), we evaluated how the distribution of Atlantic menhaden has changed from 1966 to 2004 for 12 river drainages. We observed significant or marginally significant declines in 42% of the drainages, with drainages of the northern Bay showing the majority of those declines. Continued recruitment to several drainages of the Bay may partly explain why the adult spawning population is not declining. We determined if temporal changes in abundance were related to changes in salinity or water quality for five major drainages of the watershed. For one of these drainages, the Patuxent River, differences in productivity across sites largely explained differences in abundance. For the four remaining drainages, differences in recruitment could not be explained by productivity or salinity gradients. While reducing nitrogen loading and enhancing water clarity may improve Atlantic menhaden production, we suggest that the role of offshore processes on large-scale declines has been largely neglected and studies on larval ingression are necessary for further elucidation of spatial and temporal patterns of juvenile distribution in the Chesapeake Bay.     

A spatially-explicit approach for estimating carrying capacity: An application for the Atlantic menhaden (Brevoortia tyrannus) in Chesapeake Bay

A spatially-explicit methodology was developed for estimating system carrying capacities of fish stocks, and used to estimate the seasonal and spatial patterns of carrying capacity of Chesapeake Bay for Atlantic menhaden (Brevoortia tyrannus). We used a spatially-explicit three-dimensional (3-D) model that divided the heterogeneous habitat of Chesapeake Bay into over 4,000 cubes. Each cube represented a volume of water that was characterized by a specific set of environmental variables (phytoplankton biomass, temperature, and dissolved oxygen) driven by the 3-D water quality model. Foraging and bioenergetics models transformed the environmental variables into measures of potential growth rates of menhaden. Potential carrying capacity of menhaden was estimated as a function of phytoplankton production, menhaden consumption rate, and potential growth rate, combining phytoplankton production, thermal habitat, and menhaden physiology into one ecological value that is a measure of habitat quality from the perspective of the fish. Seasonal analysis of the Chesapeake Bay carrying capacity for Atlantic menhaden suggested two bottleneck periods: one in early June and a second during the fall. The bottleneck in carrying capacity was at about 10 billion age-0 fish. Annual recruitment of age-0 menhaden for the entire Atlantic coast of the U.S. ranged from 1.2–18.6 billion fish between 1955 and 1986. It appears that carrying capacity of, Chesapeake Bay does not limit the coastwide production of young menhaden. Any conditions such as nutrient reduction strategies, further eutrophication, or global climatic warming, that may influence the carrying capacity during the fall or early June periods, may ultimately alter coastwide abundance of menhaden through changes in Chesapeake Bay carrying capacity.     

Influence of land use on macrobenthic communities in nearshore estuarine habitats

Macrobenthic community indices were examined for their ability to characterize the influence of shoreline alteration and watershed land use in nearshore estuarine environments of the Chesapeake Bay, U.S.A. Twenty-three watersheds were surveyed in 2002 and 2003 for nearshore macrobenthic assemblages, environmental parameters (i.e., dissolved oxygen, pH, total suspended solids, salinity, and sediment composition), shoreline condition, and land use. Two indices of macrobenthic biological integrity, benthic index of biological integrity in the nearshore (B-IBI_N) and abundance biomass comparison (W-value), were evaluated for associations with environmental and shoreline condition, and riparian and watershed land use. Comparisons between nearshore measures of the B-IBI with offshore values (>2 m; Chesapeake Bay benthic index of biological integrity [B-IBI_CB]) were conducted to assess the ability of the index to reflect land use patterns at near and far proximities to shore. Nearshore macrobenthic communities were represented by a total of 94 species (mean number of species =9.2 ± 0.4 sample⁻¹), and were dominated by the phyla Arthropoda, Annelida, and Mollusca. Temporal variability in environmental conditions and macrobenthic abundance and biomass may be attributable to the notable increase in precipitation in 2003 that led to nutrient influxes and algal blooms. For the biotic indices applied in the nearshore, the highest scores were associated with forested watersheds (W-value, B-IBI_N). Ecological thresholds were identified with nonparametric change-point analysis, which indicated a significant reduction in B-IBI_N and W-value scores when the amount of developed shoreline exceeded 10% and developed watershed exceeded 12%, respectively.     

Changes in submerged aquatic macrophyte populations at the head of Chesapeake Bay, 1958–1975

Submerged aquatic plant populations in the Susquehanna Flats of the Chesapeake Bay were followed for 18 years. An exotic species, eurasian water milfoil,Myriophyllum spicatum, increased dramatically from 1958 to 1962; at the same time the dominant native species declined. After 1962, milfoil populations declined and the native rooted aquatics gradually began to return to their former levels. In the late 1960's all species declined and in 1972 almost disappeared from the Susquehanna Flats. These fluctuations may have been related to several interrelated environmental factors in the Chesapeake Bay, including tropical storms, turbidity, salinity and disease. The utilization of the Susquehanna Flats by waterfowl appears to be related to the abundance and species composition of the submerged macrophytes present.     

Suspension-feeding bivalves and the fate of primary production: An estuarine model applied to Chesapeake Bay

A probabilistic mathematical model of bivalve suspension-feeding in estuaries is based on bivalve abundance, filtering capacities, and water mixing parameters. We applied the model to five regions of the upper Chesapeake Bay, ranging from shallow tidal fresh habitats to deep mesohaline habitats, for the years 1985 to 1987. Model results indicated that existing suspension-feeding bivalves could consume more than 50% of annual primary production in shallow freshwater and oligohaline reaches of the upper Chesapeake Bay and Potomac River. In deep mesohaline portions of the Chesapeake Bay and Potomac River, suspension-feeding bivalves could consume only 10% of primary production. Independent estimates of benthic carbon demand based on benthic production supported the model predictions. Hydrodynamics of large estuaries restrict the potential of benthic suspension-feeders to crop phytoplankton production because the width and depth of these estuaries limit transport of pelagic waters to the littoral flanks of the estuaries where benthic suspension-feeders can be abundant. Benthic suspension-feeders are dominant consumers in shallow segments of the Chesapeake Bay system, but are suppressed in deeper segments. The suppression is below that set by hydrodynamic limits, and may be due to periodic hypoxia or other factors. Our results suggest that the proposed use of suspension-feeding bivalves to improve water quality of large estuaries will be limited by the depth and width of the estuary, unless the bivalves are suspended in the water column by artificial means.     

Identification of important primary producers in a Chesapeake Bay tidal creek system using stable isotopes of carbon and sulfur

The use of multiple stable isotopes in the study of trophic relationships in temperate estuaries has usually been limited to euhaline systems, in which phytoplankton, benthic microalgae, andSpartina alterniflora are major sources of organic matter for consumers. Within large estuaries such as Chesapeake Bay, however, many species of consumers are found in the upper mesohaline to oligohaline portions. These lower salinity wetlands have a greater abundance of macrophytes that use C3 photosynthesis to fix carbon, in addition toS. alterniflora, which fixes carbon via the C4 photosynthetic pathway. In a broad survey of the biota and sediments of a brackish tidal creek tributary to Chesapeake Bay, combined δ¹³C and δ³⁴S measurements disclosed a balanced contribution to secondary production from phytoplankton, C3 macrophytes,Spartina sp., and benthic microalgae. Surface sediment δ¹³C suggested that the organic matter from C3 plants was derived both from allochthonous sources (terrestrial runoff) and from autochthonous production (marsh macrophytes). Unlike most estuarine systems studied to date, which are dominated by algae (phytoplankton and benthic microalgae) and C4 macrophytes, C3 plants are of greater importance in the diets of consumers in this low-salinity creek system.     

An estuarine benthic index of biotic integrity for the Mid-Atlantic region of the United States. II. Index development

A benthic index of biotic integrity was developed for use in estuaries of the mid-Atlantic region of the United States (Delaware Bay estuary through Albemarle-Pamlico Sound). The index was developed for the Mid-Atlantic Integrated Assessment Program (MAIA) of the U.S. Environmental Protection Agency using procedures similar to those applied previously in Chesapeake Bay and southeastern estuaries, and was based on sampling in July through early October. Data from seven federal and state sampling programs were used to categorize sites as degraded or non-degraded based on dissolved oxygen, sediment contaminant, and sediment toxicity criteria. Various metrics of benthic community structure and function that distinguished between degraded and reference (non-degraded) sites were selected for each of five major habitat types defined by classification analysis of assemblages. Each metric was scored according to thresholds established on the distribution of values at reference sites, so that sites with low scoring metrics would be expected to show signs of degradation. For each habitat, metrics that correctly classified at least 50% of the degraded sites in the calibration data set were selected whenever possible to derive the index. The final index integrated the average score of the combination of metrics that performed best according to several criteria. Selected metrics included measures of productivity (abundance), diversity (number of taxa, Shannon-Wiener diversity, percent dominance), species composition and life history (percent abundance of pollution-indicative taxa, percent abundance of pollution-sensitive taxa, percent abundance of Bivalvia, Tanypodinae-Chironomidae abundance ratio), and trophic composition (percent abundance of deep-deposit feeders). The index correctly classified 82% of all sites in an independent data set. Classification efficiencies of sites were higher in the mesohaline and polyhaline habitats (81–92%) than in the oligohaline (71%) and the tidal freshwater (61%). Although application of the index to low salinity habitats should be done with caution, the MAIA index appeared to be quite reliable with a high likelihood of correctly identifying both degraded and non-degraded conditions. The index is expected to be of great utility in regional assessments as a tool for evaluating the integrity of benthic assemblages and tracking their condition over time.     

Seasonal and interannual dynamics of free-living bacterioplankton and microbially labile organic carbon along the salinity gradient of the Potomac River

Samples were collected at stations located in the mesohaline, oligohaline, and tidal fresh regions of the Potomac River, Maryland, between April 1998 and December 1999 to evaluate the seasonal distribution of bacterioplankton and microbially labile organic carbon (MLOC) in relation to hydrodynamic parameters (dissolved oxygen, salinity, and temperature). Bacterioplankton abundance (BA) averaged 13 × 10⁶ cells ml⁻¹ at all stations, a value that is higher than the average observed in many other temperate estuaries around the world, and were almost exclusively free-living. During the summer of 1998, BA often exceeded 30 × 10⁶ cells ml⁻¹ in the mesohaline region during periods of anoxia in subpycnocline waters. Dissolved MLOC typically accounted for 40% of total MLOC and on some occasions during summer and autumn accounted for 80%. A significant positive relationship between dissolved MLOC and BA was evident in the mesohaline Potomac River, the region where anoxia occurs each summer, but the regressions of particulate MLOC and chla on BA were not significant at this location. In the mesohaline Potomac River, BA regressed negatively and significantly on dissolved oxygen (r²=0.50, p<0.001). BA may be an important indicator of ecosystem health in this and other eutrophied estuaries, because of the relationships between BA, dissolved MLOC, and dissolved oxygen in the salinity stratified Potomac River and because free-living BA was elevated along the length of the river.     

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.     

Dispersion and recruitment of crab larvae in the Chesapeake Bay plume: Physical and biological controls

As part of the Microbial Exchanges and Coupling in Coastal Atlantic Systems (MECCAS) Project, crab larvae were collected in the shelf waters off Chesapeake Bay in June and August 1985 and April 1986. We conducted hydrographic (temperature, salinity, nutrients) and biological (chlorophyll, copepods) mapping in conjunction with Eulerian and Lagrangian time studies of the vertical distribution of crab larvae in the Chesapeake Bay plume. These abundance estimates are used with current meter records and drifter trajectories to infer mechanisms of larval crab dispersion to the shelf waters and recruitment back into Chesapeake Bay. The highest numbers of crab larvae were usually associated with the Chesapeake Bay plume, suggesting that it was the dominant source of crab larvae to shelf waters. Patches of crab larvae also were found in the higher salinity shelf waters, and possibly were remnants of previous plume discharge events. The distribution of crab larvae in the shelf waters changed on 1–2 d time scales as a consequence of both variations in the discharge rate of the Chesapeake Bay plume and local wind-driven currents. Downwelling-favorable winds (NW) intensified the coastal jet and confined the plume and crab larvae along the coast. In April during a downwelling event (when northwesterly winds predominated), crab zoeae were transported southward along the coast at speeds that at times exceeded 168 km d⁻¹. During June and August the upwelling-favorable winds (S, SW) opposed the anticyclonic turn of the plume and, via Ekman circulation, forced the plume and crab larvae to spread seaward. Plume velocities during these conditions generally were less than 48 km d⁻¹. The recruitment of crab larvae to Chesapeake Bay is facilitated in late summer by the dominance of southerly winds, which can reverse the southward flow of shelf waters. Periodic downwelling-favorable winds can result in surface waters and crab larvae moving toward the entrance of Chesapeake Bay. Approximately 27% of the larval crabs spend at least part of the day in bottom waters, which have a residual drift toward the bay mouth. There appears to be a variety of physical transport mechanisms that can enhance the recruitment of crab larvae into Chesapeake Bay.     

Factors influencing spatial and annual variability in eelgrass (Zostera marina L.) meadows in Willapa Bay, Washington, and Coos Bay, Oregon, estuaries

Environmental factors that influence annual variability and spatial differences (within and between estuaries) in eelgrass meadows (Zostera marine L.) were examined within Willapa Bay, Washington, and Coos Bay, Oregon, over a period of 4 years (1998–2001). A suite of eelgrass metrics were recorded annually at field sites that spanned the estuarine gradient from the marine-dominated to mesohaline region of each estuary. Plant density (shoots m^?2) of eelgrass was positively correlated with summer estuarine salinity and inversely correlated with water temperature gradients in the estuaries. Eelgrass density, biomass, and the incidence of flowering plants all increased substantially in Willapa Bay, and less so in Coos Bay, over the duration of the study. Warmer winters and cooler summers associated with the transition from El Niño to La Niña ocean conditions during the study period corresponded with this increase in eelgrass abundance and flowering. Large-scale changes in climate and nearshore ocean conditions may exert a strong regional influence on eelgrass abundance that can vary annually by as much as 700% in Willapa Bay. Lower levels of annual variability observed in Coos Bay may be due to the stronger and more direct influence of the nearshore Pacific Ocean on the Coos Bay study sites. The results suggest profound effects of climate variation on the abundance and flowering of eelgrass in Pacific Northwest coastal estuaries.