Rise of Ruppia in Chesapeake Bay: Climate change-driven turnover of foundation species creates new threats and management opportunities.

Global change has converted many structurally complex and ecologically and economically valuable coastlines to bare substrate. In the structural habitats that remain, climate-tolerant and opportunistic species are increasing in response to environmental extremes and variability. The shifting of dominant foundation species identity with climate change poses a unique conservation challenge because species vary in their responses to environmental stressors and to management. Here, we combine 35 y of watershed modeling and biogeochemical water quality data with species comprehensive aerial surveys to describe causes and consequences of turnover in seagrass foundation species across 26,000 ha of habitat in the Chesapeake Bay. Repeated marine heatwaves have caused 54% retraction of the formerly dominant eelgrass (Zostera marina) since 1991, allowing 171% expansion of the temperature-tolerant widgeongrass (Ruppia maritima) that has likewise benefited from large-scale nutrient reductions. However, this phase shift in dominant seagrass identity now presents two significant shifts for management: Widgeongrass meadows are not only responsible for rapid, extensive recoveries but also for the largest crashes over the last four decades; and, while adapted to high temperatures, are much more susceptible than eelgrass to nutrient pulses driven by springtime runoff. Thus, by selecting for rapid post-disturbance recolonization but low resistance to punctuated freshwater flow disturbance, climate change could threaten the Chesapeake Bay seagrass' ability to provide consistent fishery habitat and sustain functioning over time. We demonstrate that understanding the dynamics of the next generation of foundation species is a critical management priority, because shifts from relatively stable habitat to high interannual variability can have far-reaching consequences across marine and terrestrial ecosystems.

Decoupling of Estuarine Hypoxia and Acidification as Revealed by Historical Water Quality Data.

Hypoxia and acidification are commonly coupled in eutrophic aquatic environments because aerobic respiration is usually dominant in bottom waters and can lower dissolved oxygen (DO) and pH simultaneously. However, the degree of coupling, which can be weakened by non-aerobic respiration and CaCO3 cycling, has not been adequately assessed. In this study, we applied a box model to 20 years of water quality monitoring data to explore the relationship between hypoxia and acidification along the mainstem of Chesapeake Bay. In the early summer, dissolved inorganic carbon (DIC) production in mid-bay bottom waters was dominated by aerobic respiration, contributing to DO and pH declines. In contrast, late-summer DIC production was higher than that expected from aerobic respiration, suggesting potential buffering processes, such as calcium carbonate dissolution, which would elevate pH in hypoxic waters. These findings are consistent with contrasting seasonal relationships between riverine nitrogen (N) loads and hypoxic and acidified volumes. The N loads were associated with increased hypoxic and acidified volumes in June, but only increased hypoxic volumes in August, when acidified volume declines instead. Our study reveals that the magnitude of this decoupling varies interannually with watershed nutrient inputs, which has implications for the management of co-stressors in estuarine systems.

Data synthesis for environmental management: A case study of Chesapeake Bay.

Synthesizing large, complex data sets to inform resource managers towards effective environmental stewardship is a universal challenge. In Chesapeake Bay, a well-studied and intensively monitored estuary in North America, the challenge of synthesizing data on water quality and land use as factors related to a key habitat, submerged aquatic vegetation, was tackled by a team of scientists and resource managers operating at multiple levels of governance (state, federal). The synthesis effort took place over a two-year period (2016-2018), and the results were communicated widely to a) scientists via peer review publications and conference presentations; b) resource managers via web materials and workshop presentations; and c) the public through newspaper articles, radio interviews, and podcasts. The synthesis effort was initiated by resource managers at the United States Environmental Protection Agencys' Chesapeake Bay Program and 16 scientist participants were recruited from a diversity of organizations. Multiple short, immersive workshops were conducted regularly to conceptualize the problem, followed by data analysis and interpretation that supported the preparation of the synthetic products that were communicated widely. Reflections on the process indicate that there are a variety of structural and functional requirements, as well as enabling conditions, that need to be considered to achieve successful outcomes from synthesis efforts.

A hydrodynamic model-based approach to assess sampling approaches for dissolved oxygen criteria in the Chesapeake Bay.

Technological advances in water quality measurement systems have provided the potential to expand high-frequency observations into coastal monitoring programs. However, with limited resources for monitoring budgets in natural waters that exhibit high temporal and spatial variability in water quality, there is a need to identify the locations and time periods where these new technologies can be deployed for maximum efficacy. To advance the capacity to make quantitative and objective decisions on the selection of monitoring locations and sampling frequency, we combined high-resolution numerical model simulations and multi-frequency water quality measurements to conduct a power analysis comparing alternative sampling designs in the assessment of water quality in the Chesapeake Bay. Specifically, we evaluated candidate monitoring networks that deployed both conventional long-term fixed station monitoring in deep channel areas and short-term continuous monitoring technologies in near-shore, shallow areas to assess 30-day dissolved oxygen criteria in two Bay tributaries. We conducted a cumulative frequency diagrams analysis to quantify the accuracy of each monitoring scheme in evaluating compliance with respect to the model. We used a Monte Carlo simulation to incorporate the spatial and temporal uncertainty of criteria failure. We found that additional long-term biweekly channel and short-term continuous shallow sampling efforts can lead to statistically unbiased and improved assessments at local spatial extents (less than 0.2 proportion of the assessed water body), especially when additional sampling is added at stations representing hypoxic water areas. Stations that represented seaward regions of the tributaries were more valuable in maintaining unbiased assessments of dissolved oxygen criteria attainment. This analysis highlights the importance of statistical evaluation of ongoing monitoring programs and suggests an approach to identify efficient deployments of monitoring resources and to improve assessment of other water quality metrics in estuarine ecosystems.

Advancing estuarine ecological forecasts: seasonal hypoxia in Chesapeake Bay.

Ecological forecasts are quantitative tools that can guide ecosystem management. The coemergence of extensive environmental monitoring and quantitative frameworks allows for widespread development and continued improvement of ecological forecasting systems. We use a relatively simple estuarine hypoxia model to demonstrate advances in addressing some of the most critical challenges and opportunities of contemporary ecological forecasting, including predictive accuracy, uncertainty characterization, and management relevance. We explore the impacts of different combinations of forecast metrics, drivers, and driver time windows on predictive performance. We also incorporate multiple sets of state-variable observations from different sources and separately quantify model prediction error and measurement uncertainty through a flexible Bayesian hierarchical framework. Results illustrate the benefits of (1) adopting forecast metrics and drivers that strike an optimal balance between predictability and relevance to management, (2) incorporating multiple data sources in the calibration data set to separate and propagate different sources of uncertainty, and (3) using the model in scenario mode to probabilistically evaluate the effects of alternative management decisions on future ecosystem state. In the Chesapeake Bay, the subject of this case study, we find that average summer or total annual hypoxia metrics are more predictable than monthly metrics and that measurement error represents an important source of uncertainty. Application of the model in scenario mode suggests that absent watershed management actions over the past decades, long-term average hypoxia would have increased by 7% compared to 1985. Conversely, the model projects that if management goals currently in place to restore the Bay are met, long-term average hypoxia would eventually decrease by 32% with respect to the mid-1980s.

Interannual variability of air-water CO2 flux in a large eutrophic estuary.

Air-water CO2 fluxes in estuarine environments are characterized by high interannual variability, in part due to hydrological variability that alters estuarine carbonate chemistry through multiple physical and biogeochemical processes. To understand the relative contributions of these varied controls on interannual air-water CO2 fluxes in the mainstem Chesapeake Bay, we implemented both hindcast and scenario simulations using a coupled physical-biogeochemical model. Significant spatiotemporal variability in bay-wide fluxes was found over a 10-year period (1996-2005), where the mainstem Bay was primarily a net CO2 sink, except in drought periods. Sensitivity scenario results suggested substantial effects of riverine nutrient and organic matter (OM) inputs to CO2 flux variations. The high correlations between riverine inputs and upper-Bay fluxes were due to elevated respiration under increased OM inputs. The interannual flux variations in the lower Bay was mostly regulated by the nutrient inputs. Both nutrient and OM inputs contributed to the flux variability in the mid Bay. It is found that the interannual CO2 flux of the mainstem was most sensitive to riverine nutrient inputs associated with the hydrological changes. For each hindcast simulation we computed the ratio of organic carbon turnover time to water residence time (λ), a proxy for CO2 efflux potential, and found that the wetter periods had a relatively lower λ. The variability of mainstem CO2 fluxes can be well represented using a generic function of λ. The model results showed that higher river flows would lead to enhanced CO2 sinks into a large eutrophic estuary by promoting net autotrophy.

Satellite observations estimating the effects of river discharge and wind-driven upwelling on phytoplankton dynamics in the Chesapeake Bay.

Phytoplankton growth in estuaries is regulated by a complex combination of physical factors with freshwater discharge usually playing a dominating role controlling nutrient and light availability. The role of other factors, including upwelling-generating winds, is still unclear because most estuaries are too small for upwelling to emerge. In this study, we used remotely sensed proxies of phytoplankton biomass and concentration of suspended mineral particles to compare the effect of river discharge with the effect of upwelling events associated with persistent along-channel southerly winds in the Chesapeake Bay, a large estuary where upwelling and its effects on biogeochemical dynamics have been previously reported. The surface chlorophyll-a concentrations (Chl-a) were estimated from Visible Infrared Imaging Radiometer Suite (VIIRS) satellite data using the Generalized Stacked-Constraints Model (GSCM) corrected for seasonal effects by comparing remotely sensed and field-measured data. Light limitation of phytoplankton growth was assessed from the concentration of suspended mineral particles estimated from the remotely sensed backscattering at blue (443 nm) wavelength bbp (443). The nine-year time series (2012-2020) of Chl-a and bbp (443) confirmed that a primary factor regulating phytoplankton growth in this nearshore eutrophic area is discharge from the Susquehanna River, and presumably the nutrients it delivers, with a time lag up to four months. Persistent southerly wind events (2-3 days with wind speed >4 m/s) affected the water column stratification in the central part of the bay but did not result in significant increases in remotely sensed Chl-a. Analysis of model simulations of selected upwelling-favorable wind events revealed that strong southerly winds resulted in well-defined lateral (East-West) responses but were insufficient to deliver high-nutrient water to the surface layer to support phytoplankton bloom. We conclude that, in the Chesapeake Bay, which is a large, eutrophic estuary, wind-driven upwelling of deep water plays a limited role in driving phytoplankton growth under most conditions compared with river discharge. Integr Environ Assess Manag 2022;18:921-938. © 2022 SETAC.

Nutrient limitation of phytoplankton in three tributaries of Chesapeake Bay: Detecting responses following nutrient reductions.

Many coastal ecosystems suffer from eutrophication, algal blooms, and dead zones due to excessive anthropogenic inputs of nitrogen (N) and phosphorus (P). This has led to regional restoration efforts that focus on managing watershed loads of N and P. In Chesapeake Bay, the largest estuary in the United States, dual nutrient reductions of N and P have been pursued since the 1980s. However, it remains unclear whether nutrient limitation - an indicator of restriction of algal growth by supplies of N and P - has changed in the tributaries of Chesapeake Bay following decades of reduction efforts. Toward that end, we analyzed historical data from nutrient-addition bioassay experiments and data from the Chesapeake Bay long-term water-quality monitoring program for six stations in three tidal tributaries (i.e., Patuxent, Potomac, and Choptank Rivers). Classification and regression tree (CART) models were developed using concurrent collections of water-quality parameters for each bioassay monitoring location during 1990-2003, which satisfactorily predicted the bioassay-based measures of nutrient limitation (classification accuracy = 96%). Predictions from the CART models using water-quality monitoring data showed enhanced nutrient limitation over the period of 1985-2020 at four of the six stations, including the downstream station in each of these three tributaries. These results indicate detectable, long-term water-quality improvements in the tidal tributaries. Overall, this research provides a new analytical tool for detecting signs of ecosystem recovery following nutrient reductions. More broadly, the approach can be adapted to other waterbodies with long-term bioassays and water-quality data sets to detect ecosystem recovery.

Nutrient limitation of phytoplankton in Chesapeake Bay: Development of an empirical approach for water-quality management.

Understanding the temporal and spatial roles of nutrient limitation on phytoplankton growth is necessary for developing successful management strategies. Chesapeake Bay has well-documented seasonal and spatial variations in nutrient limitation, but it remains unknown whether these patterns of nutrient limitation have changed in response to nutrient management efforts. We analyzed historical data from nutrient bioassay experiments (1992-2002) and data from long-term, fixed-site water-quality monitoring program (1990-2017) to develop empirical approaches for predicting nutrient limitation in the surface waters of the mainstem Bay. Results from classification and regression trees (CART) matched the seasonal and spatial patterns of bioassay-based nutrient limitation in the 1992-2002 period much better than two simpler, non-statistical approaches. An ensemble approach of three selected CART models satisfactorily reproduced the bioassay-based results (classification rate = 99%). This empirical approach can be used to characterize nutrient limitation from long-term water-quality monitoring data on much broader geographic and temporal scales than would be feasible using bioassays, providing a new tool for informing water-quality management. Results from our application of the approach to 21 tidal monitoring stations for the period of 2007-2017 showed modest changes in nutrient limitation patterns, with expanded areas of nitrogen-limitation and contracted areas of nutrient saturation (i.e., not limited by nitrogen or phosphorus). These changes imply that long-term reductions in nitrogen load have led to expanded areas with nutrient-limited phytoplankton growth in the Bay, reflecting long-term water-quality improvements in the context of nutrient enrichment. However, nutrient limitation patterns remain unchanged in the majority of the mainstem, suggesting that nutrient loads should be further reduced to achieve a less nutrient-saturated ecosystem.

Natural and Anthropogenic Drivers of Acidification in Large Estuaries.

Oceanic uptake of anthropogenic carbon dioxide (CO2) from the atmosphere has changed ocean biogeochemistry and threatened the health of organisms through a process known as ocean acidification (OA). Such large-scale changes affect ecosystem functions and can have impacts on societal uses, fisheries resources, and economies. In many large estuaries, anthropogenic CO2-induced acidification is enhanced by strong stratification, long water residence times, eutrophication, and a weak acid-base buffer capacity. In this article, we review how a variety of processes influence aquatic acid-base properties in estuarine waters, including coastal upwelling, river-ocean mixing, air-water gas exchange, biological production and subsequent aerobic and anaerobic respiration, calcium carbonate (CaCO3) dissolution, and benthic inputs. We emphasize the spatial and temporal dynamics of partial pressure of CO2 (pCO2), pH, and calcium carbonate mineral saturation states. Examples from three large estuaries-Chesapeake Bay, the Salish Sea, and Prince William Sound-are used to illustrate how natural and anthropogenic processes and climate change may manifest differently across estuaries, as well as the biological implications of OA on coastal calcifiers.

Discerning effects of warming, sea level rise and nutrient management on long-term hypoxia trends in Chesapeake Bay.

Analyses of dissolved oxygen concentration in Chesapeake Bay over the past three decades suggested seasonally-dependent changes in hypoxic volume and an earlier end of hypoxic conditions. While these studies hypothesized and evaluated multiple potential driving mechanisms, quantitative evidence for the relative effects of various drivers has yet to be presented. In this study, a coupled physical-biogeochemical model was used to conduct hindcast simulations between 1985 and 2016. Additional numerical experiments, in which the long-term trends in external drivers were removed, were analyzed to discern the separate effects of temperature increase, sea level rise and nutrient reduction. After the removal of seasonal and interannual variations, dissolved oxygen concentration in all regions of the estuary showed a statistically significant declining trend: ~0.1 mg/L per decade. Most of this decline occurred during winter and spring while May-August hypoxic volumes showed no changes and September hypoxic volume showed a slight decrease (~0.9 km3). Our simulations show that warming was the dominant driver of the long-term oxygen decline, overwhelming the effects of sea level rise and modest oxygen increases associated with nutrient reduction. There was no statistically significant trend in the initiation of hypoxia in spring, where the potential delay associated with nutrient reduction was offset by warming-induced oxygen declines, and both nutrient reduction and warming contributed to an earlier disintegration of hypoxia in the fall. These results suggest that recent warming has prevented oxygen improvements in Chesapeake Bay expected from nutrient input reductions and support the expectation that continued warming will serve to counter future nutrient management actions.

Biogeochemical Controls on Coastal Hypoxia.

Aquatic environments experiencing low-oxygen conditions have been described as hypoxic, suboxic, or anoxic zones; oxygen minimum zones; and, in the popular media, the misnomer "dead zones." This review aims to elucidate important aspects underlying oxygen depletion in diverse coastal systems and provides a synthesis of general relationships between hypoxia and its controlling factors. After presenting a generic overview of the first-order processes, we review system-specific characteristics for selected estuaries where adjacent human settlements contribute to high nutrient loads, river-dominated shelves that receive large inputs of fresh water and anthropogenic nutrients, and upwelling regions where a supply of nutrient-rich, low-oxygen waters generates oxygen minimum zones without direct anthropogenic influence. We propose a nondimensional number that relates the hypoxia timescale and water residence time to guide the cross-system comparison. Our analysis reveals the basic principles underlying hypoxia generation in coastal systems and provides a framework for discussing future changes.

Detection of the effects of stormwater control measure in streams using a Bayesian BACI power analysis.

The unpredictable timing and magnitude of precipitation events and the spatiotemporal variability of constituent concentrations are major complications to effective monitoring of watershed nutrient and sediment loads. Furthermore, detecting small changes in constituent loads in response to implementation of Stormwater control measures (SCMs) against natural variability is a challenge. Nevertheless, regulatory frameworks that direct reductions of pollutants to streams frequently depend on the ability to quantify changes in loads after management interventions. The before-after-control impact (BACI) sampling design is often used to assess the effects of an environmental change made at a known point in time. However, this approach may be complicated to apply to nutrient and sediment loads in streams as the relative impact of SCMs on nutrient concentration conditional on the long term variability of discharges has not been evaluated. Multi-scale monitoring studies that provide estimates of the natural temporal and spatial variability of discharge and concentrations could provide useful information in designing a BACI study. Here we use data from the Baltimore Long Term Ecological Research (LTER) sites and urban restoration sites to develop multiple statistical measures of the effectiveness of a given monitoring scheme in revealing the hypothesized restoration effects in terms of hydrology and nutrient loads. Stratified sampling over baseflow and stormflow and the use of multiple control streams were useful tools to detect long term cumulative reductions in concentrations due to SCMs. Moderate reductions in concentration (20%), however, were not detectable with the design options considered. We emphasize that appropriate pre-planning of monitoring schemes and sampling frequency is essential to determine if the effects on constituent loads resulting from a given watershed restoration activity are measurable.

Progress and challenges in coupled hydrodynamic-ecological estuarine modeling.

Numerical modeling has emerged over the last several decades as a widely accepted tool for investigations in environmental sciences. In estuarine research, hydrodynamic and ecological models have moved along parallel tracks with regard to complexity, refinement, computational power, and incorporation of uncertainty. Coupled hydrodynamic-ecological models have been used to assess ecosystem processes and interactions, simulate future scenarios, and evaluate remedial actions in response to eutrophication, habitat loss, and freshwater diversion. The need to couple hydrodynamic and ecological models to address research and management questions is clear, because dynamic feedbacks between biotic and physical processes are critical interactions within ecosystems. In this review we present historical and modern perspectives on estuarine hydrodynamic and ecological modeling, consider model limitations, and address aspects of model linkage, skill assessment, and complexity. We discuss the balance between spatial and temporal resolution and present examples using different spatiotemporal scales. Finally, we recommend future lines of inquiry, approaches to balance complexity and uncertainty, and model transparency and utility. It is idealistic to think we can pursue a "theory of everything" for estuarine models, but recent advances suggest that models for both scientific investigations and management applications will continue to improve in terms of realism, precision, and accuracy.