Habitat requirements for submerged aquatic vegetation in Chesapeake Bay: Water quality, light regime, and physical-chemical factors

We developed an algorithm for calculating habitat suitability for seagrasses and related submerged aquatic vegetation (SAV) at coastal sites where monitoring data are available for five water quality variables that govern light availability at the leaf surface. We developed independent estimates of the minimum light required for SAV survival both as a percentage of surface light passing though the water column to the depth of SAV growth (PLW _min) and as a percentage of light reaching reaching leaves through the epiphyte layer (PLL _min). Value were computed by applying, as inputs to this algorithm, statistically dervived values for water quality variables that correspond to thresholds for SAV presence in Chesapeake Bay. These estimates ofPLW _min andPLL _min compared well with the values established from a literature review. Calcultations account for tidal range, and total light attenuation is partitioned into water column and epiphyte contributions. Water column attenuation is further partitioned into effects of chlorophylla (chla), total suspended solids (TSS) and other substances. We used this algorithm to predict potential SAV presence throughout the Bay where calculated light available at plant leaves exceededPLL _min. Predictions closely matched results of aerial photographic monitoring surveys of SAV distribution. Correspondence between predictions and observations was particularly strong in the mesohaline and polythaline regions, which contain 75–80% of all potential SAV sites in this estuary. The method also allows for independent assessment of effects of physical and chemical factors other than light in limiting SAV growth and survival. Although this algorithm was developed with data from Chesapeake Bay, its general structure allows it to be calibrated and used as a quantitative tool for applying water quality data to define suitability of specific sites as habitats for SAV survival in diverse coastal environments worldwide.     

Ecosystem model of an estuarine submersed plant community: Calibration and simulation of eutrophication responses

As water quality in the Chesapeake Bay has declined over recent decades, formely healthy submersed plant communities have disappeared from littoral areas of the mesohaline estuary. A dynamic simulation model of shallow regions of bay tributaries (<1 m) was developed to investigate growth responses of submersed vascular plants to eutrophication and habitat degradation. Our objectives were to elucidate mechanisms responsible for the decline and to evaluate conditions required for plant restoration and survival. State varibles in the model are plant leaves, roots, phytoplankton, epiphytes, and detrital material. The model calculates biomass pools and biogeochemical rate processes over annual cycles with a time step of 6 h. Simulations were performed to investigate the influence of phytoplankton and epiphytes on the underwater light environment, how the balance of limiting resources (light and nutrients) controls growth and productivity of submersed plants, and conditions necessary, for the restoration of submersed vegetation. Model output for submersed plants was calibrated to baseline data from the mid 1970s (r²=0.86); simulations reproduced declines in plant biomass with increasing nutrient enrichment. Model experiments showed, that by increasing nutrient inputs 40% above levels observed in the 1960s, submersed plants disappeared within 1–2 yr due to enhanced growth of phytoplankton and epiphytes, which reduced light below required levels. Epiphytes were more important than were phytoplankton in attenuating light. The relationship between nutrient enrichment and plant loss rate was complex, as epiphyte density on leaf surfaces was not linearly related to nutrient levels. Relatively small nutrient increases could have a large effect on submersed plants because epiphyte density on leaves increased exponentially as leaf surface area decreased. Exchanges of organic carbon and nutrients between leaf and root compartments were seasonally variable and were critical for survival of submersed plants. The amount of root-rhizome material available for regrowth could control the outcome of nutrient reduction strategies. Consequently, model predictions of plant restoration success were highly dependent on initial conditions. The model is being used successfully as a research tool to interpret ecological relationships in the ongoing re-evaluation of management alternatives for submersed plant restoration.     

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.     

Response of submersed aquatic vegetation (SAV) in Lake Pontchartrain, Louisiana to the 1997–2001 El Niño Southern Oscillation shifts

Lake Pontchartrain is a large, shallow, low salinity estuary north of New Orleans, Louisiana. It is a water quality impaired system with restoration efforts in progress. One restoration goal is the reestablishment of historic submersed aquatic vegetation (SAV;Vallisneria americana Michx. andRuppia maritima L.), which has been in a state of decline since first studied in 1953. Annual SAV surveys and monthly water quality monitoring were conducted at four to five sites from 1996 through 2003 to evaluate trends and determine the causes of SAV change. We found a rapid increase in the distribution and abundance ofR. maritima in 1999 that persisted through 2002. An El Niño Southern Oscillation shift occurred between 1997 and 2001, which produced a drought in southern Louisiana as an ancillary effect of La Niña. This study was conducted to investigate causal links between the El Niño to La Niña climate phase shift and SAV change. We found that salinity and water clarity increased during La Niña. Increased water clarity produced a rapid increase in the euryhaline speciesR. maritima in deeper water and at historic sites where SAV had not been found since 1953. As salinity increased, the freshwater speciesV. americana andMyriophyllum spicatum L. declined, andNajas guadalupensis (Spreng.) Magnus andPotamogeton perfoliatus L. disappeared. In 2003, after the La Niña phase, salinity and water clarity decreased,R. maritima decreased, and the freshwater species increased, butP. perfoliatus was still absent. We found that salinity controlled SAV species composition, and water clarity controlled SAV colonization depth (Z_col=2.3/K_d). Our study demonstrated that climatic shifts cause cyclic changes in Lake Pontchartrain SAV and that restoration could be accomplished by improving water clarity. Due to the sensitivity of SAV to environmental change, similar responses to short-term and long-term climate changes should occur in other estuarine systems.     

The asiatic clam (Corbicula fluminea) invasion and system-level ecological change in the Potomac River Estuary near Washington,D.C.

The exotic freshwater clam speciesCorbicula fluminea (Asiatic clam) was first reported in the tidal freshwater Potomac estuary near Washington, D.C., in 1977, and was found in benthic surveys, conducted in 1978, 1982, 1984, 1986, and 1992. In 1981 a tripling of water clarity was reported in the region of the clam beds, followed in 1983 by reapperance of submerged aquatic vegetation (SAV) absent for 50 yr. Submerged aquatic vegetation (SAV) has been surveyed and mapped over the entire Potomac estuary region in almost every year from 1976 to 1993 by aerial photography, as part of the United States Environmental Protection Agency's Chesapeake Bay program. Fish surveys in 1986 found populations increased up to 7× in beds of SAV. Starting in 1984, the Washington, D.C. Christmas Bird Census reported significant increases in several aquatic bird populations both nonmigratory and migratory. An extensive benthic survey in September 1986 estimated a spring-summer population of 8.7×10⁶ kg Asiatic clams (wet weight including shell) in the 5-km region of the Potomac below Washington, D.C. This population was calculated as having the capacity to filter one-third to all of the water in this region of the estuary daily, depending on river flow. The 1986 clam population was smaller than that of 1984 and the 1992 population was 25% of that in 1986. Since 1986, SAV acreage has been decreasing in this area of the Potomac. Aquatic bird populations have declined. Yearly nuisance algae (Microcystis) blooms, which had been absent since 1983, reappeared in 1993. This paper presents evidence to support the theory the invasive Asiatic clam population in the 10 km below Washington, D.C., was responsible for SAV resurgence through filtration affecting turbidity. It suggests the clam populations triggered system-level changes in biota, including increase and decrease in local Potomac estuary populations (SAV, bird, fish, algae) over 10 yr, from 1983 to 1993. Major changes in the Asiatic clam population took place approximately 2 yr before parallel changes in SAV acreage were observed.     

The Effect of Submerged Aquatic Vegetation Expansion on a Declining Turbidity Trend in the Sacramento-San Joaquin River Delta

Submerged aquatic vegetation (SAV) has well-documented effects on water clarity. SAV beds can slow water movement and reduce bed shear stress, promoting sedimentation and reducing suspension. However, estuaries have multiple controls on turbidity that make it difficult to determine the effect of SAV on water clarity. In this study, we investigated the effect of primarily invasive SAV expansion on a concomitant decline in turbidity in the Sacramento-San Joaquin River Delta. The objective of this study was to separate the effects of decreasing sediment supply from the watershed from increasing SAV cover to determine the effect of SAV on the declining turbidity trend. SAV cover was determined by airborne hyperspectral remote sensing and turbidity data from long-term monitoring records. The turbidity trends were corrected for the declining sediment supply using suspended-sediment concentration data from a station immediately upstream of the Delta. We found a significant negative trend in turbidity from 1975 to 2008, and when we removed the sediment supply signal from the trend it was still significant and negative, indicating that a factor other than sediment supply was responsible for part of the turbidity decline. Turbidity monitoring stations with high rates of SAV expansion had steeper and more significant turbidity trends than those with low SAV cover. Our findings suggest that SAV is an important (but not sole) factor in the turbidity decline, and we estimate that 21–70 % of the total declining turbidity trend is due to SAV expansion.     

Alternation of factors limiting phytoplankton production in the Cape Fear River Estuary

Phytoplankton nutrient limitation experiments were performed from 1994 to 1996 at three stations in the Cape Fear River Estuary, a riverine system originating in the North Carolina piedmont. Nutrient addition bioassays were conducted by spiking triplicate cubitainers with various nutrient combinations and determining algal response by analyzing chlorophyll a production and ¹⁴C uptake daily for 3 d. Ambient chlorophyll a, nutrient concentration, and associated physical data were collected throughout the estuary as well. At a turbid, nutrient-rich oligohaline station, significant responses to nutrient additions were rare, with light the likely principal factor limiting phytoplankton production. During summer at a mesohaline station, phytoplankton community displayed significant nitrogen (N) limitation, while both phosphorus (P) and N were occasionally limiting in spring with some N+P co-limitation. Light was apparently limiting during fall and winter when the water was turid and nutrient-rich, as well as during other months of heavy rainfall and runoff. A polyhaline station in the lower estuary had clearer water and displayed significant responses to nutrient additions during all enrichment experiments. At this site N limitation occurred in summer and fall, and P limitation (with strong N+P co-limitation) occurred in winter and spring. The data suggest there are two patterns controlling phytoplankton productivity in the Cape Fear system: 1) a longitudinal pattern of decreasing light limitation and increasing nutrient sensitivity along the salinity gradient, and 2) a seasonal alternation of N limitation, light limitation, and P limitation in the middle-to-lower estuary. Statistical analyses indicated upper watershed precipitation events led to increased flow, turbidity, light attenuation, and nutrient loading, and decreased chlorophyll a and nutrient limitation potential in the estuary. Periods of low rainfall and river flow led to reduced estuarine turbidity, higher chlorophyll a, lower ambient nutrients, and more pronounced nutrient limitation.     

Decadal Changes in Water Quality and Net Productivity of a Shallow Danish Estuary Following Significant Nutrient Reductions

We utilized an extensive data set (1977–2013) from a water quality monitoring program to investigate the recovery of a Danish estuary following large reductions in total phosphorus (TP) and total nitrogen (TN) loading. Monthly rates of net transport and biogeochemical transformation of dissolved inorganic nitrogen (DIN) and phosphorus (DIP) were computed in two basins of the estuary using a box model approach, and oxygen-based rates of net ecosystem production (NEP) were determined. Since 1990, nutrient loading was reduced by 58 % for nitrogen and 80 % for phosphorus, causing significant decreases in DIN (60 %) and DIP (85 %) concentrations. Reductions in nutrient loadings and concentrations reduced annual chlorophyll levels by 50 % in the inner estuary and improved Secchi depth by approximately 1 m during the same period, particularly in the summer period. In the outer, deeper region of the estuary trends in water quality was less evident. Improvements in the inner estuary were strongly coupled to declines in DIN. Thresholds of DIN and DIP concentrations limiting phytoplankton growth indicated that both regions of the estuary were nitrogen limited. NEP rates indicated the development of more net autotrophic conditions over time that were likely associated with higher benthic primary production stimulated by improved light conditions. Box model computations revealed a modest reduction in summer net production of DIP over time, despite the persistence of elevated fluxes for several years after external loads were reduced. Since the mid-1990s, nutrient loading and transformation were stable while nutrient concentrations continued to decline and water quality improved in the inner estuary. The oligotrophication trajectory involved an initial fast transformation and modest retention of nutrients followed by a gradual decline in the rate of improvement towards a new stable condition.     

Multi-decade Responses of a Tidal Creek System to Nutrient Load Reductions: Mattawoman Creek,Maryland USA

We developed a synthesis using diverse monitoring and modeling data for Mattawoman Creek, Maryland, USA to examine responses of this tidal freshwater tributary of the Potomac River estuary to a sharp reduction in point-source nutrient loading rate. Oligotrophication of these systems is not well understood; questions concerning recovery pathways, threshold responses, and lag times remain to be clarified and eventually generalized for application to other systems. Prior to load reductions Mattawoman Creek was eutrophic with poor water clarity (Secchi depth <0.5 m), no submerged aquatic vegetation (SAV), and large algal stocks (50–100 μg L^?1 chlorophyll-a). A substantial modification to a wastewater treatment plant reduced annual average nitrogen (N) loads from 30 to 12 g N m^?2 year^?1 and phosphorus (P) loads from 3.7 to 1.6 g P m^?2 year^?1. Load reductions for both N and P were initiated in 1991 and completed by 1995. There was no trend in diffuse N and P loads between 1985 and 2010. Following nutrient load reduction, NO₂?+?NO₃ and chlorophyll-a decreased and Secchi depth and SAV coverage and density increased with initial response lag times of one, four, three, one, and one year, respectively. A preliminary N budget was developed and indicated the following: diffuse sources currently dominate N inputs, estimates of long-term burial and denitrification were not large enough to balance the budget, sediment recycling of NH₄ was the single largest term in the budget, SAV uptake of N from sediments and water provided a modest seasonal-scale N sink, and the creek system acted as an N sink for imported Potomac River nitrogen. Finally, using a comparative approach utilizing data from other shallow, low-salinity Chesapeake Bay ecosystems, strong relationships were found between N loading and algal biomass and between algal biomass and water clarity, two key water quality variables used as indices of restoration in Chesapeake Bay.     

Influences of Salinity and Light Availability on Abundance and Distribution of Tidal Freshwater and Oligohaline Submersed Aquatic Vegetation

Submersed aquatic vegetation (SAV) communities have undergone declines worldwide, exposing them to invasions from non-native species. Over the past decade, the invasive species Hydrilla verticillata has been documented in several tributaries of the lower Chesapeake Bay, Virginia. We used annual aerial mapping surveys from 1998 to 2007, integrated with spatial analyses of water quality data, to analyze the patterns and rates of change of a H. verticillata-dominated SAV community and relate them to varying salinity and light conditions. Periods of declining SAV coverage corresponded to periods where salinities exceeded 7 and early growing season (April to May) Secchi depths were <0.4 m. Increases were driven by the expansion of H. verticillata along with several other species into the upper estuary, where some areas experienced an 80% increase in cover. Field investigations revealed H. verticillata dominance to be limited to the upper estuary where total suspended solid concentrations during the early growing season were <15 mg l⁻¹ and salinity remained <3. The effect of poor early growing season water clarity on annual SAV growth highlights the importance of water quality during this critical life stage. Periods of low clarity combined with periodic salinity intrusions may limit the dominance of H. verticillata in these types of estuarine systems. This study shows the importance of the use of these types of biologically relevant episodic events to supplement seasonal habitat requirements and also provides evidence for the potential important role of invasive species in SAV community recovery.     

External nutrient sources,internal nutrient pools,and phytoplankton production in Chesapeake Bay

External nutrient loadings, internal nutrient pools, and phytoplankton production were examined for three major subsystems of the Chesapeake Bay Estuary—the upper Mainstem, the Patuxent Estuary, and the 01 Potomac Estuary—during 1985–1989. The atomic nitrogen to phosphorus ratios (TN:TP) of total loads to the 01 Mainstem, Patuxent, and the Potomac were 51, 29 and 35, respectively. Most of these loads entered at the head of the estuaries from riverine sources and major wastewater treatment plants. Approximately 7–16% for the nitrogen load entered the head of each estuary as particulate matter in contrast to 48–69% for phosphorus. This difference is hypothesized to favor a greater loss of phosphorus than nitrogen through sedimentation and burial. This process could be important in driving estuarine nitrogen to phosphorus ratios above those of inputs. Water column TN: TP ratios in the tidal fresh, oligohaline, and mesohaline salinity zones of each estuary ranged from 56 to 82 in the Mainstem, 27 to 48 in the Patuxent, and 72 to 126 in the Potomac. A major storm event in the Potomac watershed was shown to greatly increase the particulate fraction of nitrogen and phosphorus and lower the TN:TP in the river-borne loads. The load during the month that contained this storm (November 1985) accounted for 11% of the nitrogen and 31% of the phosphorus that was delivered to the estuary by the Potomac River during the entire 60-month period examined here. Within the Mainstem estuary, salinity dilution plots revealed strong net sources of ammonium and phosphate in the oligohaline to upper mesohaline region, indicating that these areas were sites of considerable internal recycling of nutrients to surface waters. The sedimentation of particulate nutrient loads in the upper reaches of the estuary is probably a major source of these recycled nutrients. A net sink of nitrate was indicated during summer. A combination of inputs and these internal recycling processes caused dissolved inorganic N to P ratios to approach 16:1 in the mesohaline zone of the Mainstem during late summer; this ratio was much higher at other times and in the lower salinity zones. Phytoplankton biomass in the mesohaline Mainstem reached a peak in spring and was relatively constant throughout the other seasons. Productivity was highest in spring and summer, accounting for approximately 33% and 44%, respectively, of the total annual productivity in this region. In the Patuxent and Potomac, the TN:TP ratios of external loads documented here are 2–4 times higher than those observed over the previous two decades. These changes are attributed to point-source phosphorus controls and the likelihood that nitrogen-rich nonpoint source inputs, including contributions from the atmosphere, have increased. These higher N:P ratios relative to Redfield proportions (16:1) now suggest a greater overall potential for phosphorus-limitation rather than nitrogen-limitation of phytoplankton in the areas studied.     

Benthic and Pelagic Primary Production in Different Nutrient Regimes

Benthic flora can contribute significantly to gross primary production (GPP) of shallow coastal waters where light reaches the sea bottom. We quantified and compared benthic and pelagic GPP along nutrient gradients in time and space in the shallow estuary, Limfjorden, Denmark, based on monitoring data combined with historical information. Limfjorden experienced a shift from a pristine, benthic-dominated clear water regime with high total GPP in the early twentieth century to a eutrophic, plankton-dominated regime still with high total GPP in the 1980s when nutrient loadings peaked. Recent reductions in nutrient loadings reduced pelagic GPP, particularly in spring, but water clarity and benthic GPP did not increase correspondingly, so total GPP declined. The most nutrient-rich basins have remained plankton-dominated, with benthic vegetation constrained to shallow waters. The results support existing evidence that total GPP of shallow coastal areas does not increase systematically with eutrophication. Furthermore, the results suggest that total GPP may decline temporarily during oligotrophication as pelagic GPP declines, while feedback mechanisms delay or prevent restoration to a state with benthic dominance of GPP.     

Optical Changes in a Eutrophic Estuary During Reduced Nutrient Loadings

Loss of water clarity is one of the consequences of coastal eutrophication. Efforts have therefore been made to reduce external nutrient loadings of coastal waters. This paper documents improvements to water clarity between 1985 and 2008–2009 at four stations in the microtidal estuary Roskilde Fjord and find significant relationships to freshwater nutrient loadings. The paper then investigates to which extent changes in phytoplankton biomass (chlorophyll a (Chl a)), non-algal particulate organic matter (POM*), and residual attenuation in the water (K _b), respectively, can account for this optical improvement. Vertical light attenuation (K _d) declined, on average, by 34 %, accompanying a 71 % reduction of Chl a and an 80 % reduction of POM*. Residual attenuation declined by 26 % over the period in accordance with a measured 34 % decline of dissolved organic nitrogen. Analysis of simultaneous changes in light attenuation and Secchi depth also suggested a reduction of the scatter-to-absorption ratio over time. Considering the stronger reductions of particle concentrations than dissolved organic matter, the contribution of residual attenuation to vertical attenuation increased from 54 to 74 % in 1985 to 78 to 85 % in 2008–2009. Overall, efforts to reduce nutrient loading and improve water clarity appeared to have had a larger impact on POM* than on Chl a and colored dissolved organic matter concentrations in the estuary, which can account for the decrease in the scatter-to-absorption ratio. These optical changes lead to larger improvements of Secchi depth than of vertical light attenuation. The consequence of this is an overestimation (0.45–1.48 m) of the predicted increase of potential seagrass depth limits when based on Secchi depth rather than K _d.     

Magnitude and Patterns of Change in Submerged Aquatic Vegetation of the Tidal Freshwater Hudson River

Three aerial photography inventories were used to examine change in submerged aquatic vegetation (SAV) in the tidal freshwater Hudson River over the interval 1997 to 2007. Overall, there was about a 30 % net decline in SAV coverage although there were also many individual areas of expansion. The invasive water chestnut (Trapa natans) did not change appreciably in net cover over the interval, and there was replacement of SAV by water chestnut along with slightly fewer cases of SAV replacing the exotic. A fine-scale (100 m by 100 m quadrats) analysis showed that about 30 % of quadrats that supported vegetation changed by more than 10 % in plant cover and overall SAV was quite dynamic. SAV in the Hudson is limited by light which is in turn controlled by suspended sediment. SAV was rarely found at depths >1 m below low water, and interannual differences in clarity affected the ability of SAV beds to maintain locally supersaturated levels of dissolved oxygen. We found that location within the River channel (proximity to shore) influenced the magnitude and variability in change in SAV between census periods. The physical nature of the adjacent shoreline also affected the magnitude of change with greater declines in cover in areas next to hard-engineered shore types. SAV in the Hudson is highly dynamic, apparently quite resilient, and the control of light by suspended sediment rather than phytoplankton growth offers a contrast to eutrophication-influenced changes in other estuaries. Management and protection of SAV habitat must recognize the highly variable nature of plant cover and that absence in any particular year does not preclude future appearance of submerged plants at that location.     

Reversal of eutrophication following sewage treatment upgrades in the New River Estuary, North Carolina

The New River Estuary consists of a series of broad shallow lagoons draining a catchment area of 1,436 km², located in Onslow County, North Carolina. During the 1980s and 1990s it was considered one of the most eutrophic estuaries in the southeastern United States and sustained dense phytoplankton blooms, bottom water anoxia and hypoxia, toxic outbreaks of the dinoflagellatePfiesteria, and fish kills. High nutrient loading, especially of phosphorus (P), from municipal and military sewage treatment plants was the principal cause leading to the eutrophic conditions. Nutrient addition bioassay experiments showed that additions of nitrogen (N) but not P consistently yielded significant increases in phytoplankton production relative to controls. During 1998 the City of Jacksonville and the U.S. Marine Corps Base at Camp Lejeune completely upgraded their sewage treatment systems and achieved large improvements in nutrient removal, reducing point source inputs of N and P to the estuary by approximately 57% and 71%, respectively. The sewage treatment plant upgrades led to significant estuarine decreases in ammonium, orthophosphate, chlorophylla, and turbidity concentrations, and subsequent increases in bottom water dissolved oxygen (DO) and light penetration. The large reduction in phytoplankton biomass led to a large reduction in labile phytoplankton carbon, likely an important source of biochemical oxygen demand in this estuary. The upper estuary stations experienced increases in average bottom water DO of 0.9 to 1.4 mg l⁻¹, representing an improvement in benthic habitat for shellfish and other organisms. The reductions in light attenuation and turbidity should also improve the habitat conditions for growth of submersed aquatic vegetation, an important habitat for fish and shellfish.     

Long-Term Trends of Water Quality and Biotic Metrics in Chesapeake Bay: 1986 to 2008

We analyzed trends in a 23-year period of water quality and biotic data for Chesapeake Bay. Indicators were used to detect trends of improving and worsening environmental health in 15 regions and 70 segments of the bay and to assess the estuarine ecosystem’s responses to reduced nutrient loading from point (i.e., sewage treatment facilities) and non-point (e.g., agricultural and urban land use) sources. Despite extensive restoration efforts, ecological health-related water quality (chlorophyll-a, dissolved oxygen, Secchi depth) and biotic (phytoplankton and benthic indices) metrics evaluated herein have generally shown little improvement (submerged aquatic vegetation was an exception), and water clarity and chlorophyll-a have considerably worsened since 1986. Nutrient and sediment inputs from higher-than-average annual flows after 1992 combined with those from highly developed Coastal Plain areas and compromised ecosystem resiliency are important factors responsible for worsening chlorophyll-a and Secchi depth trends in mesohaline and polyhaline zones from 1986 to 2008.     

Scientific Bases for Numerical Chlorophyll Criteria in Chesapeake Bay

In coastal ecosystems with long flushing times (weeks to months) relative to phytoplankton growth rates (hours to days), chlorophyll a (chl-a) integrates nutrient loading, making it a pivotal indicator with broad implications for ecosystem function and water-quality management. However, numerical chl-a criteria that capture the linkage between chl-a and ecosystem impairments associated with eutrophication (e.g., hypoxia, water clarity and loss of submerged aquatic vegetation, toxic algal blooms) have seldom been developed despite the vulnerability of these ecosystems to anthropogenic nutrient loading. Increases in fertilizer use, animal wastes, and population growth in the Chesapeake Bay watershed since World War II have led to increases in nutrient loading and chl-a. We describe the development of numerical chl-a criteria based on long-term research and monitoring of the bay. Baseline chl-a concentrations were derived using statistical models for historical data from the 1960s and 1970s, including terms to account for the effects of climate variability. This approach produced numerical chl-a criteria presented as geometric means and 90th percentile thresholds to be used as goals and compliance limits, respectively. We present scientific bases for these criteria that consider specific ecosystem impairments linked to increased chl-a, including low dissolved oxygen (DO), reduced water clarity, and toxic algal blooms. These multiple lines of evidence support numerical chl-a criteria consisting of seasonal mean chl-a across salinity zones ranging from 1.4 to 15 mg m^?3 as restoration goals and corresponding thresholds ranging from 4.3 to 45 mg m^?3 as compliance limits. Attainment of these goals and limits for chl-a is a precondition for attaining desired levels of DO, water clarity, and toxic phytoplankton prior to rapid human expansion in the watershed and associated increases of nutrient loading.     

Calculating optical water quality targets to restore and protect submersed aquatic vegetation: Overcoming problems in partitioning the diffuse attenuation coefficient for photosynthetically active radiation

Submersed aquatic vegetation (SAV) is an important component of shallow water estuarine systems that has declined drastically in recent decades. SAV has particularly high light requirements, and losses of SAV have, in many cases, been attributed to increased light attenuation in the water column, frequently due to coastal eutrophication. The desire to restore these valuable habitats to their historical levels has created the need for a simple but accurate management tool for translating light requirements into water quality targets capable of supporting SAV communities. A procedure for calculating water quality targets for concentrations of chlorophyll and total suspended solids (TSS) is derived, based on representing the diffuse attenuation coefficient for photosynthetically active radiation, K_d(PAR), as a linear function of contributions due to water plus colored dissolved organic matter (CDOM), chlorophyll, and TSS. It is assumed that K_d(PAR) conforms to the Lambert-Beer law. Target concentrations are determined as the intersection of a line representing intended reduction of TSS and chlorophyll by management actions, with another line describing the dependence of TSS on chlorophyll at a constant value of K_d(PAR). The validity of applying the Lambert-Beer law to K_d(PAR) in estuarine waters was tested by comparing the performance of a linear model of K_d(PAR) with data simulated using a more realistic model of light attenuation. The linear regression model tended to underestimate K_d(PAR) at high light attenuation, resulting in erroneous predictions of target concentrations at shallow restoration depths. The errors result more from the wide spectral bandwidth of PAR, than from irrecoverable nonlinearities in the diffuse attenuation coefficient per se. In spite of the failure of the Lambert-Beer law applied to K_d(PAR), the variation of TSS with chlorophyll at constant K_d(PAR) determined by the more mechanistic attenuation model was, nevertheless, highly linear. Use of the management tool based on intersecting lines is still possible, but coefficients in the line describing the dependence of TSS on chlorophyll at constant K_d(PAR) must be determined empirically by application of an optical model suitably calibrated for the region of interest. An example application of the procedure to data from the Rhode River, Maryland, indicates that approximately 15% reduction in both TSS and chlorophyll concentrations, or 50% reduction in chlorophyll alone, will be needed to restore conditions for growth of SAV to levels that existed in the late 1960s.     

Investigations of the availability and survival of submersed aquatic vegetation propagules in the tidal Potomac River

The establishment of submersed aquatic vegetation (SAV) at unvegetated sites in the freshwater tidal Potomac River was limited primarily by factors other than propagule availability. For two years, traps were used to quantify the amount of plant material reaching three unvegetated sites over the growing season. The calculated flux values provided a gross estimate of the flux of propagules that could potentially survive if other site factors were suitable. The mean flux ofHydrilla verticillata and all other species (≥0.01 gdw m^?2 d^?1) appeared sufficient to favor the establishment of vegetation, particularly considering the high viability (70–100%) of whole plants and fragments under controlled conditions. However, median water clarity values (i.e., for light attenuation, Secchi depth, total suspended solids, and chlorophylla) were below SAV restoration goals at all unvegetated sites. Additionally, sediments from unvegetated sites showed a potential for nitrogen limitation of the growth ofH. verticillata. Our findings support the hypothesis that in the tidal Potomac River, water clarity and nutrient (especially nitrogen) levels in sediment are key to plant community establishment.     

Nutrient Budgets and Management Actions in the Patuxent River Estuary,Maryland

Multi-year nitrogen (N) and phosphorus (P) budgets were developed for the Patuxent River estuary, a seasonally stratified and moderately eutrophic tributary of Chesapeake Bay. Major inputs (point, diffuse, septic, and direct atmospheric) were measured for 13 years during which, large reductions in P and then lesser reductions in N-loading occurred due to wastewater treatment plant improvements. Internal nutrient losses (denitrification and long-term burial of particulate N and P) were measured in tidal marshes and sub-tidal sediments throughout the estuary as were nutrient storage in the water column, sediments, and biota. Nutrient transport between the oligohaline and mesohaline zones and between the Patuxent and Chesapeake Bay was estimated using a salt and water balance model. Several major nutrient recycling terms were directly and indirectly evaluated and compared to new N and P inputs on seasonal and annual time-scales. Major findings included: (1) average terrestrial and atmospheric inputs of N and P were very close to the sum of internal losses plus export, suggesting that dominant processes are captured in these budgets; (2) both N and P export were a small fraction (13% and 28%, respectively) of inputs, about half of that expected for N based on water residence times, and almost all exported N and P were in organic forms; (3) the tidal marsh-oligohaline estuary, which by area comprised ~27% of the full estuarine system, removed about 46% and 74% of total annual upland N and P inputs, respectively; (4) recycled N and P were much larger sources of inorganic nutrients than new inputs during warm seasons and were similar in magnitude even during cold seasons; (5) there was clear evidence that major estuarine processes responded rapidly to inter-annual nutrient input variations; (6) historical nutrient input data and nutrient budget data from drought periods indicated that diffuse nutrient sources were dominant and that N loads need to be reduced by about 50% to restore water quality conditions to pre-eutrophic levels.