R. Scott Warren

Coastal eutrophication as a driver of salt marsh loss

Salt marshes are highly productive coastal wetlands that provide important ecosystem services such as storm protection for coastal cities, nutrient removal and carbon sequestration. Despite protective measures, however, worldwide losses of these ecosystems have accelerated in recent decades. Here we present data from a nine-year whole-ecosystem nutrient-enrichment experiment. Our study demonstrates that nutrient enrichment, a global problem for coastal ecosystems, can be a driver of salt marsh loss. We show that nutrient levels commonly associated with coastal eutrophication increased above-ground leaf biomass, decreased the dense, below-ground biomass of bank-stabilizing roots, and increased microbial decomposition of organic matter. Alterations in these key ecosystem properties reduced geomorphic stability, resulting in creek-bank collapse with significant areas of creek-bank marsh converted to unvegetated mud. This pattern of marsh loss parallels observations for anthropogenically nutrient-enriched marshes worldwide, with creek-edge and bay-edge marsh evolving into mudflats and wider creeks. Our work suggests that current nutrient loading rates to many coastal ecosystems have overwhelmed the capacity of marshes to remove nitrogen without deleterious effects. Projected increases in nitrogen flux to the coast, related to increased fertilizer use required to feed an expanding human population, may rapidly result in a coastal landscape with less marsh, which would reduce the capacity of coastal regions to provide important ecological and economic services.

Vegetation change on a northeast tidal marsh: interaction of sea-level rise and marsh accretion

Increasing rates of relative sea—level rise (RSL) have been linked to coastal wetland losses along the Gulf of Mexico and elsewhere. While such losses have yet to be reported for New England tidal marshes, rapidly rising RSL may still be affecting these systems. Studies of the Wequetequock—Pawcatuck tidal marshes over four decades have documented dramatic changes in vegetation that appear to be related primarily to differential rates of marsh accretion and sea—level rise. Other environmental factors such as sediment supply and anthropogenic modifications of the system may be involved as well. When initially studied in 1947—1948 the high marsh supported a Juncus gerardi—Spartina patens belting pattern typical of many New England salt marshes. On the most of the marsh complex the former Juncus belt has been replaced by forbs, primarily Triglochin maritima, while the former S. patens high marsh is now a complex of vegetation types–stunted Spartina alterniflora, Distichlis Spicata, forbs, and relic stands of S. patens. These changes are documented by vegetation sampling that closely followed the 1947—1948 methods and by peat core analysis. Marsh elevations were determined by leveling, and the mean surface elevation of areas where the vegetation has changed is significantly lower than that of areas still supporting the earlier pattern (4.6 vs. 13.9 cm above mean tide level). The differences in surface elevation reflect differences in accretion of marsh peat. Calculations based on sandy overwash layers deposited during historically recorded storms as well as on experimentally placed marker horizons of known age indicate that stable areas have been accreting at the rate of local sea—level rise, 2.0—2.5 mm/yr at least since 1938; changed areas have accreted at about one half that rate. Lower surface elevations result in greater frequency and duration of tidal flooding, and thus in increased peat saturation, salinity, and sulfide concentrations, and in decreased redox potential, as directly measured over the growing season at both changed and stable sites. It is proposed that these edaphic changes have combined to favor establishment of a wetter, more open vegetation type dominated by to distinctive communities–Stunted S. alterniflora and forbs. Changes documented on the Wequetequock—Pawcatuck system have been observed on the other Long Island Sound marshes and may serve as a model for the potential effects of seal—level rise on New England tidal salt marshes.

Salt marsh vegetation change in response to tidal restriction

Vegetation change in response to restriction of the normal tidal prism of six Connecticut salt marshes is documented. Tidal flow at the study sites was restricted with tide gates and associated causeways and dikes for purposes of flood protection, mosquito control, and/or salt hay farming. One study site has been under a regime of reduced tidal flow since colonial times, while the duration of restriction at the other sites ranges from less than ten years to several decades. The data indicate that with tidal restriction there is a substantial reduction in soil water salinity, lowering of the water table level, as well as a relative drop in the marsh surface elevation. These factors are considered to favor the establishment and spread ofPhragmites australis (common reed grass) and other less salt-tolerant species, with an attendant loss ofSpartina-dominated marsh. Based on detailed vegetation mapping of the study sites, a generalized scheme is presented to describe the sequence of vegetation change from typicalSpartina- toPhragmites-dominated marshes. The restoration of thesePhragmites systems is feasible following the reintroduction of tidal flow. At several sites dominated byPhragmites, tidal flow was reintroduced after two decades of continuous restriction, resulting in a marked reduction inPhragmites height and the reestablishment of typical salt marsh vegetation along creekbanks. It is suggested that large-scale restoration efforts be initiated in order that these degraded systems once again assume their roles within the salt marsh-estuarine ecosystem.

Vegetation Patterns and Processes in New England Salt Marshes

Salt marshes are grass-dominated tidal wetlands fringing the land-water interface of many temperate regions. They are an integral part of the larger tidal marsh-estuarine ecosystems, which are among the most biologically productive in the world. There is considerable evidence that fin and shell fisheries depend on the biological activities occurring on these tidal wetlands (Cruz 1973, Odum and Skjei 1974, Teal 1962). Tidal marshes play an important role in nutrient cycling within estuaries (Welsh 1980, Woodwell and Whitney 1977). They can also serve in pollution filtration, sediment accretion, and erosion control (Grant and Patrick 1970, Sanders and Ellis 1961, Windon 1977). Their distinctive flora, as well as their rich spectrum of waterfowl and other wildlife, make them attractive to sportsmen and naturalists. Their economic value has been estimated at more than

Rates, Patterns, and Impacts of PHRAGMITES AUSTRALIS Expansion and Effects of Experimental PHRAGMITES Control on Vegetation, Macroinvertebrates, and Fish within Tidelands of the Lower Connecticut River

197,600 per hectare (

Estuaries of the Northeastern United States: Habitat and Land Use Signatures

80,000 per acre, Gosselink et al. 1974). Chapman (1960) has described nine different geographical coastal marsh regions throughout the world. The New England type marshes, extending from Maine to New Jersey, have developed a distinctive layer of fibrous peat along resistant rocky shores, where silt has been somewhat limited compared to other marsh areas. In New England, they tend to occur as relatively small individual systems in drowned valleys or behind barrier beaches. This paper deals primarily with the origin, vegetation patterns, and productivity of southern New England marshes and the role of mans impact on them,

Development of a tidal marsh in a New England river valley

Phragmites expansion rates (linear at 1–3% yr−1) and impacts of this expansion on high marsh macroinvertebrates, aboveground production, and litter decomposition fromPhragmites and other marsh graminoids were studied along a polyhaline to oligohaline gradient. These parameters, and fish use of creeks and high marsh, were also studied inPhragmites control sites (herbicide, mowing, and combined herbicide/mow treatments).Phragmites clones established without obvious site preferences on oligohaline marshes, expanding radially. At higher salinities,Phragmites preferentially colonized creekbank levees and disturbed upland borders, then expanded into the central marsh. Hydroperiods, but not salinities or water table, distinguishedPhragmites-dominated transects. Pooled samples ofPhragmites leaves, stems, and flowers decompose more slowly than other marsh angiosperms;Phragmites leaves alone decompose as or more rapidly than those of cattail. AbovegroundPhragmites production was 1,300 to 2,400 g m−2 (about 23% of this as leaves), versus 600–800 g m−2 for polyhaline to mesohaline meadow and 1,300 g m−2 for oligohaline cattail-sedge marsh. Macroinvertebrates appear largely unaffected byPhragmites expansion or control efforts; distribution and densities are unrelated to elevation or hydroperiod, but densities are positively related to litter cover. Dominant fish captured leaving flooded marsh wereFundulus heteroclitus andAnguilla rostrata; both preyed heavily on marsh macroinvertebrates.A. rostrata andMorone americana tended to be more common inPhragmites, but otherwise there were no major differences in use patterns betweenPhragmites and brackish meadow vegetation. SAV and macroalgal cover were markedly lower within aPhragmites-dominated creek versus one withSpartina-dominated banks. The same fish species assemblage was trapped in both plus a third within the herbicide/mow treatment. Fish biomass was greatest from theSpartina creek and lowest from thePhragmites creek, reflecting abundances ofF. heteroclitus. Mowing depressedPhragmites aboveground production and increased stem density, but was ineffective for control.Phragmites, Spartina patens, andJuncus gerardii frequencies after herbicide-only treatment were 0.53-0.21; total live cover was <8% with a heavy litter and dense standing dead stems. After two growing seasonsAgrostis stolonifera/S. patens/J. gerardii brackish meadow characterized most of the herbicide/mow treatment area;Phragmites frequency here was 0.53, contributing 3% cover. Both values more than doubled after four years; a single treatment is ineffective for long-termPhragmites control.

SUSCEPTIBILITY OF SALT MARSHES TO NUTRIENT ENRICHMENT AND PREDATOR REMOVAL

Geographic signatures are physical, chemical, biotic, and human-induced characteristics or processes that help define similar or unique features of estuaries along latitudinal or geographic gradients. Geomorphologically, estuaries of the northeastern U.S., from the Hudson River estuary and northward along the Gulf of Maine shoreline, are highly diverse because of a complex bedrock geology and glacial history. Back-barrier estuaries and lagoons occur within the northeast region, but the domiant type is the drowned-river valley, often with rocky shores. Tidal range and mean depth of northeast estuaries are generally greater when compared to estuaries of the more southern U.S. Atlantic coast and Gulf of Mexico. Because of small estuarine drainage basins, low riverine flows, a bedrock substrate, and dense forest cover, sediment loads in northeast estuaries are generally quite low and water clarity is high. Tidal marshes, seagrass meadows, intertidal mudflats, and rocky shores represent major habitat types that fringe northeast estuaries, supporting commercially-important fauna, forage nekton and benthos, and coastal bird communities, while also serving as links between deeper estuarine waters and habitats through detritus-based pathways. Regarding land use and water quality trends, portions of the northeast have a history of over a century of intense urbanization as reflected in increased total nitrogen and total phosphorus loadings to estuaries, with wastewater treatment facilities and atmospheric deposition being major sources. Agricultural inputs are relatively minor throughout the northeast, with relative importance increasing for coastal plain estuaries. Identifying geographic signatures provides an objective means for comparing the structure, function, and processes of estuaries along latitudinal gradients.

Restoration of an impounded salt marsh in New England

A model for the geomorphic and vegetation development of a river valley tidal marsh in southern New England (Connecticut) is based on both the species composition of roots and rhizomes and on the mineralogic sediments preserved in peat. The maximum depth of salt marsh peat is 3.8 m and in the deepest areas this can overlie up to 1.9 m of fresh to brackish water peat. Based on a radiocarbon date of 3670±140 yr before the present (B.P.) for basal peat at a depth of 4.0 m, vertical accretion rates have averaged ca. 1.1 mm yr−1. Salt marsh formation began in response to rising sea level 3800–4000 yr B.P., as brackish marshes, dominated by bulrush (Scirpus sp.), replaced freshwater wetlands along stream and river channels. Gradually salt marsh vegetation developed over submerging brackish marshes, adjacent uplands, and accreting tidal flats. By 3000 yr B.P. the lower estuary was tidal, with sufficient salinity for salt marsh to dominate most wetlands. Spikegrass (Distichlis spicata) was an important early colonizer in salt marsh formation and its role in marsh development has not been documented previously. Blackgrass (Juncus gerardi), currently a typical upper border species, appears in the peat record relatively recently, perhaps within the last few centuries. In contrast, reed (Phragmites australis) has been present for at least 3500 yr. The dominance of reed along the upper border today, however, appears to be a relatively recent phenomenon.

Tidal energy subsidy and standing crop production of Spartina alterniflora

Salt marsh ecosystems have been considered not susceptible to nitrogen overloading because early studies suggested that salt marshes adsorbed excess nutrients in plant growth. However, the possible effect of nutrient loading on species composition, and the combined effects of nutrients and altered species composition on structure and function, was largely ignored. Failure to understand interactions between nutrient loading and species composition may lead to severe underestimates of the impacts of stresses. We altered whole salt marsh ecosystems (;60 000 m 2 /treatment) by addition of nutrients in flooding waters and by reduction of a key predatory fish, the mummichog. We added nutrients (N and P; 15-fold increase over ambient conditions) directly to the flooding tide to mimic the way anthropogenic nutrients are delivered to marsh ecosystems. Despite the high concentrations (70 mmol N/L) achieved in the water column, our annual N loadings (15-60 g Nm � 2 � yr � 1 ) were an order of magnitude less than most plot-level fertilization experiments, yet we detected responses at several trophic levels. Preliminary calculations suggest that 30-40% of the added N was removed by the marsh during each tidal cycle. Creek bank Spartina alterniflora and high marsh S. patens production increased, but not stunted high marsh S. alterniflora. Microbial production increased in the fertilized creek bank S. alterniflora habitat where benthic microalgae also increased. We found top-down control of benthic microalgae by killifish, but only under nutrient addition and in the opposite direction (increase) than that predicted by a fish-invertebrate-microalgae trophic cascade. Surprisingly, infauna declined in abundance during the first season of fertilization and with fish removal. Our results demonstrate ecological effects of both nutrient addition and mummichog reduction at the whole-system level, including evidence for synergistic interactions.