This review provides an integrated synthesis with timelines and evaluations of ecological responses to eutrophi- cation in Chesapeake Bay, the largest estuary in the USA. Analyses of dated sediment cores reveal initial evidence of organic enrichment in ~200 yr old strata, while signs of increased phytoplankton and decreased water clarity first appeared ~100 yr ago. Severe, recurring deep-water hypoxia and loss of diverse submersed vascular plants were first evident in the 1950s and 1960s, respectively. The degradation of these benthic habitats has contributed to declines in benthic macro- infauna in deep mesohaline regions of the Bay and blue crabs in shallow polyhaline areas. In contrast, copepods, which are heavily consumed in pelagic food chains, are relatively un- affected by nutrient-induced changes in phytoplankton. Intense mortality associated with fisheries and disease have caused a dramatic decline in eastern oyster stocks and associated Bay water filtration, which may have exacerbated eutrophication effects on phytoplankton and water clarity. Extensive tidal marshes, which have served as effective nutrient buffers along the Bay margins, are now being lost with rising sea level. Although the Bay's overall fisheries production has probably not been affected by eutrophication, decreases in the relative contribution of demersal fish and in the efficiency with which primary production is transferred to harvest suggest funda- mental shifts in trophic and habitat structures. Bay ecosystem responses to changes in nutrient loading are complicated by non-linear feedback mechanisms, including particle trapping and binding by benthic plants that increase water clarity, and by oxygen effects on benthic nutrient recycling efficiency. Observations in Bay tributaries undergoing recent reductions in nutrient input indicate relatively rapid recovery of some ecosystem functions but lags in the response of others.

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Eutrophication of Chesapeake Bay: historical trends and ecological interactions

Contemporary climate change is characterized both by increasing mean temperature and increasing climate variability such as heat waves, storms, and floods. How populations and communities cope with such climatic extremes is a question central to contemporary ecology and biodiversity conservation. Previous work has shown that species diversity can affect ecosystem functioning and resilience. Here, we show that genotypic diversity can replace the role of species diversity in a species-poor coastal ecosystem, and it may buffer against extreme climatic events. In a manipulative field experiment, increasing the genotypic diversity of the cosmopolitan seagrass Zostera marina enhanced biomass production, plant density, and faunal abundance, despite near-lethal water temperatures due to extreme warming across Europe. Net biodiversity effects were explained by genotypic complementarity rather than by selection of particularly robust genotypes. Positive effects on invertebrate fauna suggest that genetic diversity has second-order effects reaching higher trophic levels. Our results highlight the importance of maintaining genetic as well as species diversity to enhance ecosystem resilience in a world of increasing uncertainty.

/pdf/ecosystem-recovery-after-climatic-extremes-enhanced-by-3xqgd4whtw.pdf

Ecosystem recovery after climatic extremes enhanced by genotypic diversity

Submerged vegetation respond to increased nutrient loading through a shift from slow-growing seagrasses and large macroalgae to fast-growing macroalgae, and the ultimate dominance of phytoplankton at high nutrient loadings. This shift reflects a change from nutrient to light limitation along the eutrophication gradient. Slow-growing seagrasses and large macroalgae are good competitors when nutrients are limiting because they have relatively low nutrient requirements, are able of efficient internal nutrient recycling, and can access the elevated nutrient pools in the sediment. Fast-growing macroalgae and phytoplankton are superior competitors when light is limiting because they are positioned closer to the water surface, and capture and use light more efficiently. The important ecosystem consequences of altered nutrient regimes derive from the shift in dominant vegetation types. Slow-growing seagrasses and large macroalgae are longevous, decompose slowly, and experience only moderate grazing losse...

Submerged aquatic vegetation in relation to different nutrient regimes

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

https://www.jstor.org/stable/info/1352224

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

Water movement in freshwater and marine environments affects submersed macrophytes, which also mediate water movement. The result of this complex interaction also affects sediment dynamics in and around submersed macrophyte beds. This review defines known relationships and identifies areas that need additional research on the complex interactions among submersed macrophytes, water movement, and sediment dynamics. Four areas are addressed: (1) the effects of water movement on macrophytes, (2) the effects of macrophyte stands on water movement, (3) the effects of macrophyte beds on sedimentation within vegetated areas, and (4) the relationship between sediment resuspension and macrophytes. Water movement has a significant effect on macrophyte growth, typically stimulating both abundance and diversity of macrophytes at low to moderate velocities, but reducing growth at higher velocities. In turn, macrophyte beds reduce current velocities both within and adjacent to the beds, resulting in increased sedimentation and reduced turbidity. Reduced turbidity increases light availability to macrophytes, increasing their growth. Additionally, macrophytes affect the distribution, composition and particle size of sediments in both freshwater and marine environments. Therefore, establishment and persistence of macrophytes in both marine and freshwater environments provide important ecosystem services, including: (1) improving water quality; and (2) stabilizing sediments, reducing sediment resuspension, erosion and turbidity.

The interaction between water movement, sediment dynamics and submersed macrophytes

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m−2 d−1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.

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Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

Seasonal pulses of turbidity and their relations to eelgrass (Zostera marina L.) survival in an estuary

Studies of seagrass meadows have shown that the production of algal epiphytes attached to seagrass blades approaches 20% of the seagrass production and that epiphytes are more important as food for associated fauna than are the more refractory seagrass blades. Since epiphytes may compete with seagrasses for light and water column nutrients, excessive epiphytic fouling could have serious consequences for seagrass growth. We summarize much of the literature on epiphytegrazer relationships in seagrass meadows within the context of seagrass growth and production. We also provide insights from mathematical modeling simulations of these relationships for a Chesapeake BayZostera marina meadow. Finally we focus on future research needs for more completely understanding the influences that epiphyte grazers have on seagrass production.

Epiphyte-grazer relationships in seagrass meadows: Consequences for seagrass growth and production

Seasonal variations in eelgrass (Zostera marina L.) responses to nutrient enrichment and reduced light availability in experimental ecosystems

Ten years (1985–1994) of data were analyzed to investigate general patterns of phytoplankton and nutrient dynamics, and to identify major factors controlling those dynamics in the York River Estuary, Virginia. Algal blooms were observed during winter-spring followed by smaller summer blooms. Peak phytoplankton biomass during the winter-spring blooms occurred in the mid reach of the mesohaline zone whereas peak phytoplankton biomass during the summer bloom occurred in the tidal fresh-mesohaline transition zone. River discharge appears to be the major factor controlling the location and timing of the winter-spring blooms and the relative degree of potential N and P limitation. Phytoplankton biomass in tidal fresh water regions was limited by high flushing rates. Water residence time was less than cell doubling time during high flow seasons. Positive correlations between PAR at 1 m depth and chlorophylla suggested light limitation of phytoplankton in the tidal fresh-mesohaline transition zone. Relationships of salinity difference between surface and bottom water with chlorophylla distribution suggested the importance of tidal mixing for phytoplankton dynamics in the mesohaline zone. Accumulation of phytoplankton biomass in the mesohaline zone was generally controlled by N with the nutrient supply provided by benthic or bottom water remineralization.

Richard L. Wetzel

Papers

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

Seasonal pulses of turbidity and their relations to eelgrass (Zostera marina L.) survival in an estuary

Epiphyte-grazer relationships in seagrass meadows: Consequences for seagrass growth and production

Seasonal variations in eelgrass (Zostera marina L.) responses to nutrient enrichment and reduced light availability in experimental ecosystems

Spatial and temporal characteristics of nutrient and phytoplankton dynamics in the York River Estuary, Virginia: Analyses of long-term data