Deegan Linda A.

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Linda A.

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Turnover rates of nitrogen stable isotopes in the salt marsh mummichog, Fundulus heteroclitus, following a laboratory diet switch

( 2005-09-19) Logan, John ; Haas, Heather ; Deegan, Linda A. ; Gaines, Emily F.

Nitrogen stable isotopes are frequently used in ecological studies to estimate trophic position and determine movement patterns. Knowledge of tissue-specific turnover and nitrogen discrimination for the study organisms is important for accurate interpretation of isotopic data. We measured δ15 N turnover in liver and muscle tissue in juvenile mummichogs, Fundulus heteroclitus, following a laboratory diet switch. Liver tissue turned over significantly faster than muscle tissue suggesting the potential for a multiple tissue stable isotope approach to study movement and trophic position over different time scales; metabolism contributed significantly to isotopic turnover for both liver and muscle. Nitrogen diet-tissue discrimination was estimated at between 0.0 and 1.2‰ for liver and –1.0 and 0.2‰ for muscle. This is the first experiment to demonstrate a significant variation in δ15 N turnover between liver and muscle tissues in a fish species.
Effects of regular salt marsh haying on marsh plants, algae, invertebrates and birds at Plum Island Sound, Massachusetts

( 2008-10) Buchsbaum, Robert N. ; Deegan, Linda A. ; Horowitz, Julie ; Garritt, Robert H. ; Giblin, Anne E. ; Ludlam, John P. ; Shull, David H.

The haying of salt marshes, a traditional activity since colonial times in New England, still occurs in about 400 ha of marsh in the Plum Island Sound estuary in northeastern Massachusetts. We took advantage of this haying activity to investigate how the periodic large-scale removal of aboveground biomass affects a number of marsh processes. Hayed marshes were no different from adjacent reference marshes in plant species density (species per area) and end-of-year aboveground biomass, but did differ in vegetation composition. Spartina patens was more abundant in hayed marshes than S. alterniflora, and the reverse was true in reference marshes. The differences in relative covers of these plant species were not associated with any differences between hayed and reference marshes in the elevations of the marsh platform. Instead it suggested that S. patens was more tolerant of haying than S. alterniflora. S. patens had higher stem densities in hayed marshes than it did in reference marshes, suggesting that periodic cutting stimulated tillering of this species. Although we predicted that haying would stimulate benthic chlorophyll production by opening up the canopy, we found differences to be inconsistent, possibly due to the relatively rapid regrowth of S. patens and to grazing by invertebrates on the algae. The pulmonate snail, Melampus bidendatus was depleted in its δ13C content in the hayed marsh compared to the reference, suggesting a diet shift to benthic algae in hayed marshes. The stable isotope ratios of a number of other consumer species were not affected by haying activity. Migratory shorebirds cue in to recently hayed marshes and may contribute to short term declines in some invertebrate species, however the number of taxa per unit area of marsh surface invertebrates and their overall abundances were unaffected by haying over the long term. Haying had no impact on nutrient concentrations in creeks just downstream from hayed plots, but the sediments of hayed marshes were lower in total N and P compared to references. In sum, haying appeared to affect plant species composition but had only short-term affects on consumer organisms. This contrasts with many grassland ecosystems, where an intermediate level of disturbance, such as by grazing, increases species diversity and may stimulate productivity. From a management perspective, periodic mowing could be a way to maintain S. patens habitats and the suite of species with which they are associated.
Seasonal use of a New England estuary by foraging contingents of migratory striped bass

(American Fisheries Society, 2009-12-10) Pautzke, Sarah M. ; Mather, Martha E. ; Finn, John T. ; Deegan, Linda A. ; Muth, Robert M.

Using acoustic telemetry on migratory striped bass Morone saxatilis in Plum Island Estuary (PIE), Massachusetts, we found that striped bass (335–634 mm total length) tagged in the spring and summer of 2005 (n = 14) and 2006 (n = 46) stayed in the estuary for an average of 66.0 d in 2005 and 72.2 d in 2006. Striped bass spent the most time in two specific reaches: middle Plum Island Sound and lower Rowley River. In both years, three different use-groups of striped bass were observed in PIE. Short-term visitors (n = 24) stayed in the estuary only briefly (range = 5–20 d). Two groups of seasonal residents stayed for more than 30 d, either in the Rowley River (n = 14) or in Plum Island Sound (n = 22). Within PIE, the two seasonal-resident use-groups may be foraging contingents that learn how to feed efficiently in specific parts of the estuary. These distinct within-estuary use patterns could have different implications for striped bass condition and prey impact.
Long-term survival of adult Arctic grayling (Thymallus arcticus) in the Kuparuk River, Alaska

(National Research Council Canada, 2004-12-23) Buzby, Karen M. ; Deegan, Linda A.

In many long-lived species such as Arctic grayling (Thymallus arcticus), population growth rate is most sensitive to changes in adult survival probabilities. Understanding the factors that regulate adult survival in this species should provide insight into the population dynamics of this and other long-lived Arctic species. Using the program MARK, we analyzed 17 years of mark–recapture data to estimate survival rates for Arctic grayling in the Kuparuk River, Alaska, from 1985 to 2000. Mean annual survival rates overall ranged from 0.39 to 1.0 and averaged 0.71 ± 0.05 for resident and 0.75 ± 0.05 for nonresident fish. Spending the summer in the more productive fertilized zone of the experimental reach had no influence on survival despite higher productivity on all trophic levels and consistently higher growth rates in Arctic grayling. None of the environmental (stream temperature, discharge, winter severity, and incidence of drought) or population parameters (growth, condition factor, and mean fish size) that we examined explained significant amounts of variance in survival rates. The lack of responsiveness of survival to annual environmental conditions was unexpected and suggests that multiyear factors or life history tactics that maintain survival at the expense of growth and fecundity likely determine survival.
Use of non-natal estuaries by migratory striped bass (Morone saxatilis) in summer

(U.S. Dept. of Commerce, NOAA NMFS, 2009-07) Mather, Martha E. ; Finn, John T. ; Ferry, Kristen H. ; Deegan, Linda A. ; Nelson, Gary A.

For most migratory fish, little is known about the location and size of foraging areas or how long individuals remain in foraging areas, even though these attributes may affect their growth, survival, and impact on local prey. We tested whether striped bass (Morone saxatilis Walbaum), found in Massachusetts in summer, were migratory, how long they stayed in non-natal estuaries, whether observed spatial patterns differed from random model predictions, whether fish returned to the same area across multiple years, and whether fishing effort could explain recapture patterns. Anchor tags were attached to striped bass that were caught and released in Massachusetts in 1999 and 2000, and recaptured between 1999 and 2007. In fall, tagged striped bass were caught south of where they were released in summer, confirming that fish were coastal migrants. In the first summer, 77% and 100% of the recaptured fish in the Great Marsh and along the Massachusetts coast, respectively, were caught in the same place where they were released. About two thirds of all fish recaptured near where they were released were caught 2–7 years after tagging. Our study shows that smaller (400–500 mm total length) striped bass migrate hundreds of kilometers along the Atlantic Ocean coast, cease their mobile lifestyle in summer when they use a relatively localized area for foraging (<20 km2), and return to these same foraging areas in subsequent years.
Stable isotope monitoring of benthic–planktonic coupling using salt marsh fish

(Inter-Research, 2008-10-13) Fry, Brian ; Cieri, Matthew ; Hughes, Jeff ; Tobias, Craig R. ; Deegan, Linda A. ; Peterson, Bruce J.

Salt marshes are important coastal ecosystems whose trophic function can be monitored with stable isotopes of abundant fish biosentinel species such as the mummichog Fundulus heteroclitus and the Atlantic silverside Menidia menidia. We compared movement patterns and feeding biology of these species in the summers of 1999 and 2000 in the Rowley River salt marsh estuary north of Boston, Massachusetts, USA. A 15N tracer addition experiment showed that fish of both species were more resident than transient, with mummichogs resident at scales of 1 km or less. Natural abundance stable isotope C, N, and S distributions showed that mummichogs feed more strongly in the benthic food web while silversides feed more in the planktonic food web, with % benthic feeding respectively averaging 58 ± 5 and 32 ± 3% (mean ± 95% confidence limit, CL). For both species, isotope results indicated considerable individual specialization in foraging behavior, likely related to use of channel habitat versus use of the marsh. Power analysis showed that measuring 3 composite samples each comprising 10 to 15 individual fish should provide relatively low errors of 0.5‰ (95% CL) or less around stable isotope averages. Use of such composite samples in monitoring programs will allow detection of significant temporal and spatial changes in benthic-planktonic coupling for salt marsh ecosystems, as recorded in average fish diets. Analyzing some individual fish also is recommended to obtain more detailed information on fish food sources, feeding specializations, and end-member isotope values used in estimating importance of benthic and planktonic food sources.
Plant nitrogen dynamics in fertilized and natural New England salt marshes : a paired 15N tracer study

(Inter-Research, 2008-02-07) Drake, Deanne C. ; Peterson, Bruce J. ; Deegan, Linda A. ; Harris, Lora A. ; Miller, E. E. ; Warren, R. Scott

We examined the effects of increased nutrient availability on nitrogen (N) dynamics in dominant New England salt marsh plants (tall and stunted Spartina alterniflora and S. patens) using paired large-scale nutrient and 15NO3– tracer additions. This study is one component of a long-term, large-scale, salt marsh nutrient and trophic manipulation study (the Trophic Cascades and Interacting Control Processes in a Detritus-based Aquatic Ecosystem [TIDE] Project). We compared physiological variables of plants in fertilized (~17× ambient N and P in incoming tidal water) and reference marsh systems to quantify NO3– uptake and uptake efficiency, allocation of N to tissues, end-of-season N resorption, leaf litter quality and other potential responses to increased nutrient availability. Reference system plants sequestered ~24.5 g NO3-N ha–1 d–1 in aboveground pools during mid-summer, while fertilized plants sequestered ~140 g NO3-N ha–1 d–1. However, NO3– uptake efficiency (% of total incoming NO3-N sequestered aboveground) was higher in the reference system (16.8%) than in the fertilized system (2.6%), suggesting that our fertilization rate (~70 µM NO3– in incoming water) approaches or exceeds the uptake saturation point for this vegetation community. Leaf litter quality was clearly affected by N availability; N resorption efficiency was lower in all plants of the fertilized system; senesced leaves from the fertilized creek contained ~43% (tall S. alterniflora), 23% (stunted S. alterniflora) and 15% (S. patens) more N per unit biomass than reference creek leaves.
Susceptibility of salt marshes to nutrient enrichment and predator removal

( 2006-03-15) Deegan, Linda A. ; Bowen, Jennifer L. ; Drake, Deanne C. ; Fleeger, John W. ; Friedrichs, Carl T. ; Galvan, Kari A. ; Hobbie, John E. ; Hopkinson, Charles S. ; Johnson, J. Michael ; Johnson, David S. ; LeMay, Lynsey E. ; Miller, Erin ; Peterson, Bruce J. ; Picard, Christian ; Sheldon, Sallie ; Sutherland, Michael ; Vallino, Joseph J. ; Warren, R. Scott

The sustainability of coastal ecosystems in the face of widespread environmental change is an issue of pressing concern throughout the world (Emeis et al. 2001). Coastal ecosystems form a dynamic interface between terrestrial and oceanic systems and are one of the most productive ecosystems in the world. Coastal systems probably serve more human uses than any other ecosystem and they have always been valued for their rich bounty of fish and shellfish. Coastal areas are also the sites of the nation’s and the world’s most intense commercial activity and population growth; worldwide, approximately 75% of the human population now lives in coastal regions (Emeis et al. 2001). Over the past three decades nutrient enrichment of coastal and estuarine waters has become the premier issue for both scientists and managers (National Research Council 2000). Our understanding of coastal eutrophication has been developed principally through monitoring of estuaries, with a focus on pelagic or subtidal habitats (National Research Council 2000, Cloern 2001). Because estuarine systems are usually nitrogen limited, NO3- is the most common nutrient responsible for cultural nutrient enrichment (Cloern 2001). Increased nitrogen delivery to pelagic habitats of estuaries produces the classic response of ecosystems to stress (altered primary producers and nutrient cycles and loss of secondary producer species and production; Nixon 1995, Rapport and Whitford 1999, Deegan et al. 2002). Salt marsh ecosystems have been thought of as not susceptible to nitrogen over-loading because early studies found added nitrogen increased marsh grass production (primarily Spartina spp., cordgrass) and concluded that salt marshes can adsorb excess nutrients in plants and salt marsh plant-derived organic matter as peat (Verhoeven et al. 2006). Detritus from Spartina is important in food webs (Deegan et al. 2000) and in creating peat that forms the physical structure of the marsh platform (Freidrichs and Perry 2001). However, the accumulation of peat and inputs of sediments and loss of peat through decomposition and sediment through erosion may be altered under high nutrient regimes and threaten the long-term stability of marsh systems. Nitrogen addition may lead to either net gain or loss of the marsh depending on the balance between increased marsh plant production and increased decomposition. Absolute change in marsh surface elevation is determined by marsh plant species composition, production and allocation to above- and belowground biomass, microbial decomposition, sedimentation, erosion and compaction (Friedrichs and Perry 2001). Levine et al. (1998) suggested that competitive dynamics among plants might be affected by nutrient enrichment, potentially altering relative abundance patterns favoring species with less belowground storage and thus lowering rates of peat formation. When combined with the observation that nutrient additions may also stimulate microbial respiration and decomposition (Morris and Bradley 1999), the net effect on the salt marsh under conditions of chronic nitrogen loading is a critical unknown. Although most research treats nutrient enrichment as a stand-alone stress, it never occurs in isolation from other perturbations. The effect of nutrient loading on species composition (both plants and animals) and the resultant structure and function of wetlands has been largely ignored when considering their ability to adsorb nutrients (Verhoeven et al. 2006). Recent studies suggest the response of estuaries to stress may depend on animal species composition (Silliman et al. 2005). Animal species composition may alter the balance between marsh gain and loss as animals may increase or decrease primary production, decomposition or N recycling (Pennings and Bertness 2001). Failure to understand interactions between nutrient loading and change in species composition may lead to underestimating the impacts of these stresses. The 'bottom up or top down' theory originated from the observation that nutrient availability (bottom up)sets the quantity of primary productivity, while other studies have shown that species composition (top down), particularly of top consumers, has a marked and cascading effect on ecosystems, including controlling species composition and nutrient cycling (Matson and Price 1992, Pace et al. 1999). Most examples of trophic cascades are in aquatic ecosystems with fairly simple, algal grazing pelagic food webs (Strong 1992). The rarity of trophic cascades in terrestrial systems has been attributed to the importance of detrital food webs (Polis 1999). Detritus-based aquatic ecosystems, such as salt marshes, bogs, and swamps, have classically been considered bottom-up or physically controlled ecosystems. Recent experiments, however, suggest that salt marshes may exhibit top-down control at several trophic levels (Silliman and Zeiman. 2001, Silliman and Bertness 2002, Quiñones-Rivera and Fleeger 2005). One abundant, ubiquitous predator, a small (<10 cm total length) killifish (Fundulus heteroclitus, mummichog) has been suggested to control benthic algal through a trophic cascade because they prey on the invertebrates that graze on the benthic algae (Kneib 1997, Sarda et al. 1998). In late summer, killifish are capable of consuming 3-10 times the creek meiofauna production and meiofauna in the absence of predators appear capable of grazing over 60% of the microalgal community per day (Carman et al. 1997). Strong top-down control by grazers is considered a moderating influence on the negative effects of elevated nutrients on algae (Worm et al. 2000). Small-scale nutrient additions and predator community exclusion experiments have demonstrated bottom-up and top-down control of macroinfauna in mudflats associated with salt marsh creeks (Posey et al. 1999, Posey et al. 2002). Together, these observations suggest mummichogs are at the top of a trophic cascade that controls benthic algae (Sarda et al. 1998). Mummichogs are also omnivorous and ingest algae, bulk detritus and the attached microbial community (D’Avanzo and Valiela 1990). As a result, marsh decomposition rates may be limited by top-down controls through trophic pathways or by release from competition with algae for nutrients. Whole-ecosystem experiments have shown that responses to stress are often not predictable from studies of the individual components (Schindler 1998). Developing the information needed to predict the interacting impacts of nutrient loading and species composition change requires experiments with realistic alterations carried out at scales of space and time that include the complexities of real ecosystems. Whole ecosystem manipulation experiments have been used effectively in other ecosystems (Bormann and Likens 1979, Carpenter et al. 1995), but they are rare in coastal research. Experiments in salt marshes have traditionally been less than a few m2. Our understanding of the response of salt marsh plants to nutrient enrichment is from small (<10 m2), plot-level additions where uniform levels of dry inorganic fertilizer (20 to > 1000 g N m-2 y-1) are sprinkled on the marsh surface at low tide. Dry fertilizer additions were usually made every two weeks or monthly and the duration of elevated nutrient levels after these additions was usually not determined. Tidal water is the primary vector for N delivery to coastal marshes, suggesting that dry fertilizer addition to the marsh surface may not be the best basis for determining if Spartina production responds to nutrient enrichment of tidal waters. Similarly, our understanding of top-down controls in salt marshes also relies on small (1 - 4 m2) exclusion experiments that use cages to isolate communities from top consumers. While the design of these cage experiments has improved, there are some remaining drawbacks. For example, it is impossible to selectively exclude single species using cages, and recruitment or size-selective movement into or out of the cages may obscure interpretations. In addition, while these small-scale experiments provide insight into controls on isolated ecosystem processes, they do not allow for interaction among different parts of the ecosystem which may buffer or alter the impacts and are not appropriate for determining the effects of populations of larger more motile animals on whole-ecosystems or the effects of ecosystem changes on populations. For example, interactions may be caused when a motile species alters its distribution among the habitats available to it because of an experimental treatment. Small-scale experiments generally do not allow such events to happen. Complex feedbacks among physical and biological processes can alter accumulation rates and affect marsh elevation relative to sea level rise making extrapolation of small plot level experiments to whole marsh ecosystems problematic. We are conducting an ecosystem-scale, multi-year field experiment including both nutrient and biotic manipulations to coastal salt marsh ecosystems. We are testing, for the first time at the ecosystem level, the hypothesis that nutrient enrichment and species composition change have interactive effects across multiple levels of biological organization and a range of biogeochemical processes. We altered whole salt marsh creek watersheds (~60,000 m2 of saltmarsh) by addition of nutrients (15x ambient) in flooding waters and by a 60% reduction of a key fish species, the mummichog. Small marsh creek watersheds provide an ideal experimental setting because they have the spatial complexity, species composition and processes characteristic of the larger salt marsh ecosystem, which are often hundreds of thousands of m2. Manipulating entire salt marsh creeksheds allowed us to examine effects on large motile animals and the interactive effects of motile species changes on ecosystem processes without cage artifacts. Because our manipulations were done on whole-marsh ecosystems, we are able to evaluate the integrated and interactive effects on all habitats (e.g., water column, tidal creeks and marsh) and on populations. These experiments are similar in many respects to the small watershed experiments carried out in forested catchments. Our nutrient enrichment is novel compared to past studies in two important ways. We added nutrients (N and P) directly to the flooding tidal creek waters to mimic the way in which anthropogenic nutrients reach marsh ecosystems. All previous experimental salt marsh nutrient enrichment studies used a dose-response design with spatially uniform dry fertilizer loading on small plots (<10 m2). Nutrients carried in water will interact and reach parts of the ecosystem differently than dry fertilizer. Our enrichment method also creates a spatial gradient of nutrient loading across the landscape that is proportional to the frequency and depth of inundation in the marsh. Spatial gradients in loading within an ecosystem are typical in real world situations in many terrestrial and aquatic ecosystems. Because of our enrichment method, at any location in the ecosystem, nutrient load will be a function of the nutrient concentration in the water, the frequency and depth of tidal flooding and the reduction of nutrients from the flooding waters by other parts of the ecosystem. Uniform loading misses important aspects of the spatial complexity of ecosystem exposure and response. This work is organized around two questions that are central to understanding the long-term fate of coastal marshes: 1. Does chronic nutrient enrichment via flooding water increase primary production more than it stimulates microbial decomposition? 2. Do top-down controls change the response of the salt marsh ecosystem to nutrient enrichment? Here we present findings on the first 2 years of these experiments including 1) water chemistry, 2) standing stocks and species composition of benthic microalgae, 3) microbial production, 4) species composition and ecophysiology of macrophytes, 5) invertebrates, and 6) nekton. Because even highly eutrophic waters result in nutrient loading that is an order of magnitude less than most plot level experiments, we expected little stimulation of salt marsh vascular plant growth. However, moderate levels of nutrient enrichment in the water column were expected to increase benthic algal biomass and to stimulate bacterial activity and detrital decomposition throughout the ecosystem because of direct uptake of nitrogen from the water column and availability of more high quality organic matter from increased algal production. We predicted nutrient enrichment would increase invertebrate production because of an increase of high quality microalgal and microbial production at the base of the food web. Finally, we predicted that fish reduction would reduce predation on benthic invertebrates resulting in increased abundance of benthic invertebrates that would graze down the benthic algae.
Top-down and bottom-up control of infauna varies across the saltmarsh landscape

( 2007-12) Fleeger, John W. ; Johnson, David S. ; Galvan, Kari A. ; Deegan, Linda A.

Responses of infaunal saltmarsh benthic invertebrates to whole-ecosystem fertilization and predator removal were quantified in Plum Island Estuary, Massachusetts, USA. Throughout a growing season, we enriched an experimental creek on each flooding tide to 70 mM NO3 - and 4 mM PO4 -3 (a 10 x increase in loading above background), and we reduced Fundulus heteroclitus density by 60% in a branch of the fertilized and a reference creek. Macroinfauna and meiofauna were sampled in creek (mudflat and creek wall), marsh edge (tall form Spartina alterniflora) and marsh platform (Spartina patens and stunted S. alterniflora) habitats before and after treatments were begun; responses were tested with BACI-design statistics. Treatment effects were most common in the mid-range of the inundation gradient. Most fertilization effects were on creek wall where ostracod abundance increased, indices of copepod reproduction increased and copepod and annelid communities were altered. These taxa may use epiphytes (that respond rapidly to fertilization) of filamentous algae as a food source. Killifish reduction effects on meiobenthic copepod abundance were detected at the marsh edge and suggest predator limitation. Fish reduction effects on annelids did not suggest top-down regulation in any habitat; however, fish reduction may have stimulated an increased predation rate on annelids by grass shrimp. Interactions between fertilization and fish reduction occurred under S. patens canopy where indirect predator reduction effects on annelids were indicated. No effects were observed in mudflat or stunted S. alterniflora habitats. Although the responses of infauna to fertilization and predator removal were largely independent and of similar mild intensity, our data suggests that the effects of ecological stressors vary across the marsh landscape.
Salt marsh ecosystem biogeochemical responses to nutrient enrichment : a paired 15N tracer study

(Ecological Society of America, 2009-09) Drake, Deanne C. ; Peterson, Bruce J. ; Galvan, Kari A. ; Deegan, Linda A. ; Hopkinson, Charles S. ; Johnson, J. Michael ; Koop-Jakobsen, Ketil ; LeMay, Lynsey E. ; Picard, Christian

We compared processing and fate of dissolved NO3− in two New England salt marsh ecosystems, one receiving natural flood tide concentrations of 1–4 μmol NO3−/L and the other receiving experimentally fertilized flood tides containing 70–100 μmol NO3−/L. We conducted simultaneous 15NO3− (isotope) tracer additions from 23 to 28 July 2005 in the reference (8.4 ha) and fertilized (12.4 ha) systems to compare N dynamics and fate. Two full tidal cycles were intensively studied during the paired tracer additions. Resulting mass balances showed that essentially 100% (0.48–0.61 mol NO3-N·ha−1·h−1) of incoming NO3− was assimilated, dissimilated, sorbed, or sedimented (processed) within a few hours in the reference system when NO3− concentrations were 1.3–1.8 μmol/L. In contrast, only 50–60% of incoming NO3− was processed in the fertilized system when NO3− concentrations were 84–96 μmol/L; the remainder was exported in ebb tidewater. Gross NO3− processing was 40 times higher in the fertilized system at 19.34–24.67 mol NO3-N·ha−1·h−1. Dissimilatory nitrate reduction to ammonium was evident in both systems during the first 48 h of the tracer additions but <1% of incoming 15NO3− was exported as 15NH4+. Nitrification rates calculated by 15NO3− dilution were 6.05 and 4.46 mol·ha−1·h−1 in the fertilized system but could not be accurately calculated in the reference system due to rapid (<4 h) NO3− turnover. Over the five-day paired tracer addition, sediments sequestered a small fraction of incoming NO3−, although the efficiency of sequestration was 3.8% in the reference system and 0.7% in the fertilized system. Gross sediment N sequestration rates were similar at 13.5 and 12.6 mol·ha−1·d−1, respectively. Macrophyte NO3− uptake efficiency, based on tracer incorporation in aboveground tissues, was considerably higher in the reference system (16.8%) than the fertilized system (2.6%), although bulk uptake of NO3− by plants was lower in the reference system (1.75 mol NO3−·ha−1·d−1) than the fertilized system (10 mol NO3−·ha−1·d−1). Nitrogen processing efficiency decreased with NO3− load in all pools, suggesting that the nutrient processing capacity of the marsh ecosystem was exceeded in the fertilized marsh.

Deegan Linda A.

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