Tidal hydrology and habitat utilization by resident nekton in<Emphasis Type="Italic">Pragmites</Emphasis> and Non-<Emphasis Type="Italic">Phragmites</Emphasis> Marshes

We compared nekton densities over a range of measured flooding conditions and locations withinPhragmites australis andSpartina alterniflora (salt marsh cordgrass) at the Charles Wheeler Salt Marsh, located on the lower Housatonic River estuary in southwestern Connecticut. Nekton were sampled on nine spring high tide events from May to October 2000 using bottomless lift nets positioned between 0–5 and 10–20 m from the creek edge. Flooding depth, duration, and frequency were measured from each vegetation type during each sampling month. Benthic macroinvertebrate density was also measured within each vegetation type in May, July, and September. Frequency of flooding was 52% lower and flooding depth and duration were also significantly reduced inP. australis relative toS. alterniflora. A total of 4,197 individuals representing 7 species, mostlyPalaemonetes pugio (dagger-blade grass shrimp) andFundulus heteroclitus (common mummichog), were captured.P. pugio densities were significantly greater inS. alterniflora as were benthic macroinvertebrate density and taxa richness during May, but not during June or October. Total fish density was not significantly different betweenP. australis andS. alterniflora and was independent of location on the marsh. Significantly more juvenileF. heteroclitus were collected withinS. alterniflora relative toP. australis in June and July, suggesting that recruitment of this species may be lower inP. australis habitat. Fish density generally did not vary predictably across the range of flooding depth and duration; there was a positive relationship between flooding depth and fish density inS. alterniflora. The measured reduction in flooding frequency (52%) withinP. australis at the Housatonic site would result in an average total monthly fish use, expressed as density, of 447 ind m⁻² forP. australis and 947 ind m⁻² forS. alterniflora. WhenP. australis expansion results in reduction of flooding frequency and duration, nekton community composition can change, access to the marsh surface is reduced twofold, and nursery habitat function may be impaired.     

Relationships of nitrogen loadings,residential development,and physical characteristics with plant structure in New England salt marshes

We examined the vascular plant species richness and the extent, density, and height ofSpartina species of ten Narragansett Bay, Rhode Island (United States) fringe salt marshes which had a wide range of residential land development and N-loadings associated with their watersheds. Significant inverse relationships of tallS. alterniflora with species richness and with the extent and density ofS. patens and shortS. alterniflora were observed. Extent and density ofS. patens and extent of shortS. alterniflora were positively and significantly related with plant species richness. Marsh elevation and area did not significantly correlate with plant structure. Flood tide height significantly and inversely correlated withS. patens, but did not significantly relate toS. alterniflora or plant species richness. Marsh width significantly and positively correlated with plant species richness andS. patens and inversely correlated with tallS. alterniflora. Significant inverse relationships were observed for N-load, % residential development, and slope withS. patens, shortS. alterniflora, and species richness, and significant positive relationships with tallS. alterniflora. The marsh slope and width were significantly correlated with N-load and residential development that made it difficult to determine to what extent anthropogenic stressors were contributing to the variation in the plant structure among the marshes. At five marhes with similar slopes, there were significant inverse relationships of N-load withS. patens (density and extent) and a positive relationship with tallS. alterniflora (extent). Although there were no significant relationships of slope with the plant metrics among the five sites, other physical factors, such as the flood tide height and marsh width, significantly correlated with the extent and density ofSpartina species. Significant relationships of N-load with plant structure (albeit confounded by the effect of the physical characteristics) support the hypothesis of competitive displacement of dominant marsh plants under elevated nitrogen. It is likely that the varying plant structure in New England marshes is a response to a combination of natural factors and multiple anthropogenic stressors (e.g., eutrophication and sea level rise).     

Nitrogen Uptake by Native and Invasive Temperate Coastal Macrophytes: Importance of Dissolved Organic Nitrogen

We investigated if the success of the invasive common reed Phragmites australis could be attributed to a competitive ability to use dissolved organic nitrogen (DON) when compared to the dominant macrophyte Spartina alterniflora in tidal wetlands. Short-term nutrient uptake experiments were performed in the laboratory on two genetic lineages of Phragmites (native and introduced to North America) and S. alterniflora. Our results provide the first evidence for direct assimilation of DON by temperate marsh plants and indicate that amino acids are assimilated intact by all plant types at similar rates. Both Phragmites lineages had significantly greater urea–N assimilation rates than S. alterniflora, and the affinity for dissolved inorganic nitrogen (DIN) species was the greatest in native Phragmites > introduced Phragmites > S. alterniflora. Field studies demonstrated uptake of both DON and DIN in similar proportion as those determined in the laboratory experiments. Based on these uptake rates, we estimate that DON has the potential to account for up to 47% of N demand for Phragmites plants, and up to 24% for S. alterniflora plants. Additionally, we suggest that differences in N uptake between native and introduced Phragmites lineages explain one mechanism for the success of the introduced type under increasingly eutrophic conditions.     

Effects of common reed (<Emphasis Type="Italic">Phragmites australis</Emphasis>) expansions on nitrogen dynamics of tidal marshes of the northeastern U.S.

Several recent studies indicate that the replacement of extant species withPhragmites australis can alter the size of nitrogen (N) pools and fluxes within tidal marshes. Some common effects ofP. australis expansion are increased standing stocks of N, greater differentiation of N concentrations between plant tissues (high N leaves and low N stems), and slower whole-plant decay rates than competing species (e.g.,Spartina, Typha spp.). Some of the greater differences between marsh types involveP. australis effects on extractable and porewater pools of dissolved inorganic nitrogen (DIN) and N mineralization rates. Brackish and salt marshes show higher concentrations of DIN in porewater beneathSpartina spp. relative toP. australis, but this is not observed in freshwater tidal marshes whenP. australis is compared withTypha spp. or mixed plant assemblages. With few studies of concurrent N fluxes, the net effect ofP. australis on marsh N budgets is difficult to quantify for single sites and even more so between sites. The magnitude and direction of impacts ofP. australis on N cycles appears to be system-specific, driven more by the system and species being invaded than byP. australis itself. WhereP. australis is found to affect N pools and fluxes, we suggest these alterations result from increased biomass (both aboveground and belowground) and increased allocation of that biomass to recalcitrant stems. Because N pools are commonly greater inP. australis than in most other communities (due to plant and litter uptake), one of the most critical questions remaining is “From where is the extra N inP. australis communities coming?” It is important to determine if the source of the new N is imported (e.g., anthropogenic) or internallyproduced (e.g., fixed, remineralized organic matter). In order to estimate net impacts ofP. australis on marsh N budgets, we suggest that further research be focused on the N source that supports high standing stocks of N inP. australis biomass (external input versus internal cycling) and the relative rates of N loss from different marshes (burial versus subsurface flow versus denitrification).     

Impact of common reed,<Emphasis Type="Italic">Phragmites australis</Emphasis>, on essential fish habitat: Influence on reproduction,embryological development,and larval abundance of mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>)

The invasion ofSpartina marshes by the common reed,Phragmites australis, along the east coast of the United States over the last several decades has been well documented, although we know little about the impact of this invasion on the fish fauna and the few published papers seem contradictory. During 1999–2000 (May–September) we evaluated the fish response to vegetation type (Phragmites australis veersusSpartina alterniflora) by monitoring several aspects of fish early life history (egg deposition, embryonic development, hatching success, and larval and juvenile abundance) in low salinity marshes in the Mullica River in southern New Jersey. The dominant fish species using the marsh surface,Fundulus heteroclitus (93% of total catch, n=996 individuals), reproduced in both vegetation types with eggs deposited in leaf axils near the base of the plant inSpartina and in broken stems ofPhragmites during both years. These eggs also undergo successful embryonic development to hatching in both vegetation types. Larval and juvenile (5–75 mm total length, but 95% < 34 mm TL) abundance of this species is much reduced onPhragmites-dominated (mean CUPE=0.02, n=7 ind) marsh surface relative toSpartina (mean CPUE=2.31). These findings, and similar results for fish abundance in 1997 and 1998, indicate that theSpartima marsh surface is likely essential fish habitat for this species because it provides habitat for larvae and small juveniles, whilePhragmites does not. ThePhragmites invasion in brackish marshes may be having deleterious effects on fish populations and possibly on predators that prey uponF. heteroclitus, and as a result, marsh secondary production.     

Does the common reed,Phragmites australis, affect essential fish habitat?

Since the early 1900sPhragmites australis has been replacing other vegetation in Atlantic and gulf coast marshes at a rate of about 1% to 6% of the marsh surface per year. Vast areas of coastal marsh are now characterized by dense monotypic stands of this species. By virtue of its ability to build up the marsh surface,P. australis affects the landscape, hydrology, and hydroperiod of the marsh as well as drainage density, and other geomorphic features. Smoothed microtopography results in more difficult access to the marsh by nekton, and possibly reduced exchange of organic materials between the marsh and adjacent estuary. The pattern of replacement byP. australis results in fragmentation of existing stands ofSpartina alterniflora and other extant macrophytes, thereby altering landscape ecology and the ability of the marsh to support biodiversity and the production of marsh fauna.     

Primary production and nutrient content in two salt marsh species,<Emphasis Type="Italic">Atriplex portulacoides</Emphasis> L. and<Emphasis Type="Italic">Limoniastrum monopetalum</Emphasis> L., in Southern Portugal

Seasonal variation patterns of aboveground and belowground biomass, net primary production, and nutrient accumulation were assessed inAtriplex portulacoides L. andLimoniastrum monopetalum (L.) Boiss. in Castro Marim salt marsh, Portugal. Sampling was conducted for five periods during 2001–2002 (autumn, winter, spring, summer, and autumn). This study indicates that both species have a clear seasonal variation pattern for both aboveground and belowground biomass. Mean live biomass was 2516 g m⁻² yr⁻¹ forL. monopetalum and 598 g m⁻² yr⁻¹ forA. portulacoides. Peak living biomass, in spring for both species, was three times greater in the former, 3502 g m⁻² yr⁻¹, than in the latter, 1077 g m⁻² yr⁻¹. For both the Smalley (Groenendijk 1984) and Weigert and Evans (1964) methods, productivity ofL. monopetalum (2917 and 3635 g m⁻² yr⁻¹, respectively) was greater than that ofA. portulacoides (1002 and 1615 g m⁻² yr⁻¹, respectively). Belowground biomass ofL. monopetalum was 1.7 times greater than that ofA. portulacoides. In spite of this, the root:shoot ratio forA. monopetalum to aerial components. Leaf area index was similar for both species, but specific leaf area ofA. portulacoides was twice that ofL. monopetalum. The greatest nutrient contents were found in leaves. Leaf nitrogen content was maximum in summer for both species (14.6 mg g⁻¹ forA. portulacoides and 15.5 mg g⁻¹ forL. monopetalum). Leaf phosphorus concentration was minimum in summer (1.1 mg g⁻¹ inA. portulacoides and 1.2 mg g⁻¹ inL. monopetalum). Leaf potassium contents inA. portulacoides were around three times greater than inL. monopetalum. Leaf calcium contents inL. monopetalum were three times greater than inA. portulacoides. There was a pronounced seasonal variation of calcium content in the former, while in the latter no clear variation was registered. Both species exhibited a decrease in magnesium leaf contents in the summer period. Mangamese content inL. monopetalum leaves was tenfold that inA. portulacoides. Seasonal patterns of nutrient contents inA. portulacoides andL. monopetalum suggest that availability of these elements was not a limiting factor to biomass production.     

The influence of<Emphasis Type="Italic">Phragmites</Emphasis> (common reed) on the distribution,abundance, and potential prey of a resident marsh fish in the Hackensack Meadowlands,New Jersey

This study investigates the influence ofPhragmites australis (common reed) invasion on the habitat of the resident marsh fish,Fundulus heteroclitus (mummichog) in the Hackensack Meadowlands, New Jersey. These abundant fish play an important role in the transfer of energy from the marsh surface to adjacent subtidal waters and thus estuarine food webs. The objectives of this 2-yr study (1999 and 2000) were to compare the distribution and abundance of the eggs, larvae, juveniles, and adults of mummichog and their invertebrate prey inhabitingSpartina alterniflora-dominated marshes withPhragmites-dominated marshes, and to experimentally investigate the influence of marsh surface microtoprography on larval fish abundance withinPhragmites-dominated marshes. In 2000, we verified that egg deposition does occur inPhragmites-dominated marshes. In both years, the abundance of larvae and small juveniles (4–20 mm TL) inS. alterniflora was significantly greater than inPhragmites-dominated marshes, while larger juveniles and adults (>20 mm TL) were similarly abundant in both habitat types. The overall abundance of larvae and small juveniles was significantly greater in experimentalPhragmites plots in which microtopography was manipulated to resemble that ofSpartina marshes than inPhragmites control plots. Major groups of invertebrate taxa differed between marsh types with potential prey for larval fish being significantly more abundant inS. alterniflora marshes.Phragmites-dominated marshes may not provide the most suitable habitat for the early life-history stages of the mummichog. The low abundance of larvae and small juveniles inPhragmites marshes is likely due to inadequate larval habitat and perhaps decreased prey availability for these early life history stages.     

Restoration of an impounded salt marsh in New England

The restoration of a 20 ha tidal marsh, impounded for 32, yr, in Stonington, Connecticut was studied to document vegetation change 10 yr after the reintroduction of tidal flushing. These data were then compared to a 1976 survey of the same marsh when it was in its freshest state and dominanted byTypha angustifolia. Currently,T. angustifolia remains vigorous only along the upland borders and in the upper reaches of the valley marsh. Live coverage ofT. angustifolia has declined from 74% to 16% and surviving stands are mostly stunted and depauperate. Other brackish species have also been adversely effected, except forPhragmites australis which has increased. In contrast, the salt marsh speciesSpartina alterniflora has dramatically expanded, from <1% to 45% cover over the last decade. Locally, high marsh species have also become established, covering another 20% of the marsh.     

Aboveground Biomass Patterns of Dominant Spartina Species and Their Relationship with Selected Abiotic Variables in Argentinean SW Atlantic Marshes

Salt marsh zonation patterns generate different abiotic and biotic conditions that can accentuate species inherent differences in primary production and biomass. In South West Atlantic marshes, there are two Spartina species: Spartina alterniflora in the low intertidal and Spartina densiflora in the high intertidal. These two species are generally found in all marshes but with different dominance: In some marshes, the S. densiflora zone occupies higher extents, and in others, the S. alterniflora zone is the one that prevails. We found through field sampling that, in six studied marshes, there is greater S. densiflora live and total (i.e., dead+live) aboveground biomass (g m^?2) in the marshes dominated by S. densiflora than in the ones dominated by S. alterniflora. Spartina alterniflora had similar aboveground biomass in the six marshes, regardless of the dominance of each species. When comparing the two Spartina species within each marsh, S. densiflora had greater live and total biomass in the marshes it dominates. In the marshes dominated by S. alterniflora, both species had similar live and total biomass. In all marshes, there was greater dead S. densiflora biomass. A multivariate analysis using selected abiotic factors (i.e., salinity, latitude, and tidal amplitude) showed that S. alterniflora aboveground biomass patterns are mainly correlated with salinity, while S. densiflora live biomass is mainly correlated with salinity and latitude, dead biomass with salinity and tidal amplitude, and total biomass with salinity alone. We conclude that in S. densiflora dominated marshes, the main processes of that species zone (i.e., nutrient accumulation) will be accentuated because of its higher biomass. We also conclude that climatic conditions, in combination with specific Spartina biotic and ambient abiotic parameters, can affect marsh ecological functions.     

An investigation of salt marsh dieback in Georgia using field transplants

In 2001 and 2002, Georgia salt marshes experienced a dieback event that, affected more than 800 ha throughout the coastal zone. The dieback event was unprecedented in the state and affected bothSpartina alterniflora andJuncus roemerianus. A transplant study was conducted from May to October 2003 to determine if healthy plants could survive in dieback areas. Transplants were carried out at two locations on the Georgia coast in areas ofS. alterniflora dieback along the banks of tidal creeks, an area ofS. alterniflora dieback in the mid marsh, and aJ. roemerianus dieback, area in the mid marsh. Transplant survival was nearly 100% and growth (measured as increases in the height of the 5 tallest stems and the number of stems per experimental pot) was observed in both healthy (control) and dieback areas.J. roemerianus grew more slowly thanS. alterniflora, with no, observed increase in stem height and an average 38% increase in stem density as compared to an average 57% increase in stem height and 137% increase in stem density inS. alterniflora. Differences in growth were inconsistent but in most cases no significant differences were observed between healthy and dieback areas. Soil characteristics measured over the course of the experiment were generally comparable between healthy and dieback areas (redox potential averaged 69±123 [SD] across all observations at all sites, pH averaged 6.7 ± 0.3 and salinity averaged 24.9±4.4), but porewater ammonium (NH₄) concentration was often higher in dieback areas (overall mean NH₄ concentration, was 138±127 μM in dieback areas versus 33±40 μM in healthy areas). These results suggest that the cause of dieback was no, longer present at the time of this study and that transplants are a possibility for restoring affected areas.     

Determination of food web support and trophic position of the mummichog,Fundulus heteroclitus, in New Jersey smooth cordgrass (Spartina alterniflora), common reed (Phragmites australis), and restored salt marshes

The invasion ofPhragmites australis into tidal marshes formerly dominated bySpartina alterniflora has resulted in considerable interest in the consequences of this invasion for the ecological functions of marsh habitat. We examined the provision of trophic support for a resident marsh fish,Fundulus heteroclitus, in marshes dominated byP. australis, byS. alterniflora, and in restored marshes, using multiple stable isotope analysis. We first evaluated our ability to distinguish among potential primary producers using the multiple stable isotope approach. Within a tidal creek system we found significant marsh and elevation effects on microalgal isotope values, and sufficient variability and overlap in primary producer isotope values to create some difficulty in identifying unique end members. The food webs supportingF. heteroclitus production were examined using dual isotope plots. At both sites, the δ¹³C values ofF. heteroclitus were clustered over values for benthic microalgae (BMI) and approximately midway between δ¹³C values ofSpartina andPhragmites. Based on comparisons of fish and primary producer δ¹³C, δ¹⁵N, and δ³⁴S values, and consideration ofF. heteroclitus feeding habits, we conclude that BMI were a significant component of the food web supportingF. heteroclitus in these brackish marshes, especially recently-hatched fish occupying pools on the marsh surface. A 2‰ difference in δ¹³C betweenFundulus occupying nearly adjacentSpartina andPhragmites marshes may be indicative of relatively less reliance on BMI and greater reliance onPhragmites production inPhragmites-dominated marshes, a conclusion consistent with the reduced BMI biomass found inPhragmites marshes. The mean δ¹³C value ofF. heteroclitus from restored marshes was intermediate between values of fish from naturally occurringSpartina marshes and areas invaded byPhragmites. We also examined the isotopic evidence for ontogenetic changes in the trophic position of larval and juvenileF. heteroclitus. We found significant positive relationships betweenF. heteroclitus δ¹⁵N values and total length, reflective of an increase in trophic position as fish grow.F. heteroclitus δ¹⁵N values indicate that these fish are feeding approximately two trophic levels above primary producers.     

Impact of Armored Shorelines on Shore-Zone Fish Density in a Mid-Atlantic,USA, Estuary: Modulation by Hypoxia and Temperature

Anthropogenic modifications of estuarine environments, including shoreline hardening and corresponding alteration of water quality, are accelerating worldwide as human population increases in coastal regions. Estuarine fish species inhabiting temperate ecosystems are adapted to extreme variations in environmental conditions including water temperature, salinity, and dissolved oxygen across seasonal, daily, and hourly time scales. The present research utilized quantitative sampling to examine the spatiotemporal distribution of shore-zone estuarine fish species in association with four unique shoreline types across a range of water temperature and dissolved oxygen conditions. Fish were collected from the intertidal and shallow subtidal region of four shoreline types, Spartina alterniflora marsh, Phragmites australis marsh, riprap, and bulkhead, in the summer and fall of 2009 and 2010. Analyses were performed to (1) compare mean fish density among shoreline types across all water conditions and (2) explore relationships of the complete fish assemblage, three functional species groupings, and two fish species (Fundulus heteroclitus and Menidia menidia) to unique shoreline/water conditions. Significantly greater mean fish densities were found along S. alterniflora shorelines than armored shorelines. Several metrics including fish density, species richness, and occurrence rates suggest S. alterniflora shorelines may serve as a form of refuge habitat during periods of low dissolved oxygen and high temperatures for various species, particularly littoral-demersal species including F. heteroclitus. Potential mechanisms that could contribute to a habitat providing refuge during adverse water quality conditions include tempering of the adverse condition (decreased temperatures, increased dissolved oxygen), predation protection, and increased foraging opportunities.     

Rapidly entrainable tidal vertical migrations in the salt marsh snailLittoraria irrorata

The salt marsh periwinkleLittoraria irrorata (Say) remains on the substratum during low tide but climbs above the water on stalks ofSpartina alterniflora Loisel during high tide. Rhythmic tidal migrations may allowL. irrorata to avoid predators such as blue crabsCallinectes sapidus Rathbun that forage when the marsh is inundated. These tidal rhythms may be driven by endogenous clocks or they may be easily entrained. Snails with flexible and entrainable climbing rhythms may be able to avoid predators in unpredictable environments (e.g., when water unexpectedly covers the substratum as in storm surges). We tested the behavioral response ofL. irrorata to different simulated tidal regimes in the laboratory, and the effect of remaining above mean high water (MHW) on snail survivorship in a smallS. alterniflora salt marsh. In laboratory mesocosms, vertical snail position was measured under constant water levels, simulated tidal cycles, and simulated tidal cycles 180° out of phase (reversed). Under constant water levels, snails ceased to migrate vertically after 1 d. When exposed to tidal and reversed tidal cycles, snails migrated in synchrony with the appropriate simulated rhythm.L. irrorata entrained quickly to differing tidal cycles and maintained their position above the water surfce when water levels were high. In a field experiment, snails were tethered toS. alteriflora plants near the substratum and above MHW in the marsh for 1 wk to assess survival. Survival of snails tethered above MHW was sigificantly greater than for snails tethered at the base of plants; no snails in control cages died. Rapid alteration of tidal vertical migrations may allowL. irrorata to avoid predators that forage when water inundates the marsh predictably or unexpectedly.     

<Emphasis Type="Italic">Rhizedra lutosa</Emphasis>, a natural enemy of<Emphasis Type="Italic">Phragmites australis</Emphasis> in North America

Rhizedra lutosa (Hübner) (Lepidoptera: Noctuidae), native to Europe, was first found in North America in 1988 in southern New Jersey. The insect is reported to cause serious damage toPhragmites australis in the Netherlands, but it now appears to be relatively unimportant throughout central Europe. We are investigating this insect as part of an effort to characterize existing natural enemies ofP. australis in North America toward a goal of biological control of this invasive plant. We trapped adults ofR. lutosa with blacklights in Rhode Island, Massachusetts, Connecticut, New York, Delaware, and Maryland; we also found it in Ohio and Pennsylvania.R. lutosa larvae feed in stems and rhizomes ofP. australis growing in dry sites. Based uponR. lutosa damage surveys andP. australis biomass measurements, we have determined that this insect does cause a reduction of plant growth in some sites, however, because of low moth densities, this effect is small. Despite spreading rapidly,R. lutosa densities do not appear to be increasing at sites we have investigated, and this insect does not currently appear to have much of an impact uponP. australis in North America.     

Nitrogen and phosphorus distribution and utilization bySpartina alterniflora in a Louisiana gulf coast marsh

Nitrogen and phosphorus content ofSpartina alterniflora Loisel and soil nitrogen were measured along a transect perpendicular to a stream in a Louisiana salt marsh in order to provide information on differences between the so-called streamside and inland regions. Total plant nitrogen and phosphorus levels in June and September tended to be greater at streamside than inland sites. Total soil nitrogen on a dry soil weight basis increased with distance inland from a natural stream toward an interdistributary basin in the marsh. Soil extractable ammonium-nitrogen levels measured in June were very low in vegetated streamside and inland areas, but they were much higher in inland areas devoid of plants. Nitrogen and phosphorus utilization byS. alterniflora was also investigated at an inland location in the salt marsh. Labelled ammonium-nitrogen and phosphate-phosphorus were added in May at a rate of 200 kg/ha to the soil of replicated plots. Added nitrogen significantly increased total above-ground plant biomass and plant height by 28 and 25%, respectively, 4 months after application. The ratio of belowground macro-organic matter to total aboveground biomass was decreased from 5.7 to 4.7 by the additional nitrogen. Added phosphorus did not significantly affect plant height and biomass. The use of¹⁵N-depleted nitrogen tracers showed that about half of the nitrogen in the aboveground portion ofS. alterniflora from 1 to 4 months after the nitrogen addition was derived from the added ammonium-nitrogen. After 4 months, 28 and 29% of the added labelled nitrogen was recovered in the aboverground and belowground biomass ofS. alterniflora, respectively. Recovery of added nitrogen was overestimated with a non-tracer method based on the difference in total nitrogen uptake between nitrogen-amended plots and untreated plots. Soil organic nitrogen comprised the majority of the nitrogen in the salt marsh. Nitrogen in the standing crop biomass ofS. alterniflora represented only about 2% of the total nitrogen in the plantsoil system of an inland marsh to a 20 cm soil depth.     

Responses of resident marsh fishes to stages ofPhragmites australis invasion in three mid Atlantic estuaries

Modification of brackish marshes by nonindigenousPhragmites australis has occurred across a broad geographical area in eastern North America. Among its effects on marsh processes,Phragmites may be increasingly unfavorable to marsh surface fishes as its invasion progresses within an estuary. We assessed the effect of thePhragmites invasion on resident marsh surface fishes by examining the population response ofFundulus heteroclitus (mummichog, 5–48 mm TL) andF. luciae (spotfin killifish, 5–41 mm TL) to four distinct invasion stages in three estuaries of the U.S. mid Atlantic region (New Jersey, Delaware, and Maryland). We documented precipitous declines in mean catch per unit effort ofF. heteroclitus in pit traps from natural marsh (51.6), through initial (33.8), early (12.3), and late invasion stages (2.4) across all sites. A similar pattern was documented forF. luciae, with mean catch per unit effort in pit traps declining from natural marsh (48.9), through initial (39.1), early (9.3), and late invasion stages (2.7). Population structure of both species also changed somewhat across invasion stages such that we collected a narrower size range of individuals of both species from late invasion stages. Patterns suggest that as thePhragmites invasion progresses, there is a decline in habitat function for larval and juvenileF. heteroclitus and an increased risk of extirpation ofF. luciae from brackish marshes along the east coast of the U.S.     

Valuation of continuous and intermittent<Emphasis Type="Italic">Phragmites</Emphasis> control

Quantifying the relative value ofPhragmites australis andSpartina alterniflora habitat is important to evaluate the benefits and risks of different attempts to addressPhragmites expansion on the U.S. eastern seaboard. Two contrasting approaches commonly used to restore tidal marsh habitats invaded byPhragmites communities involve sprayingPhragmites with herbicide only when its coverage of a particular marsh area is near or close to 100%. Alternatively, after the first application, herbicide is annually applied on any surviving patches ofPhragmites present in a mosaic of other marsh vegetation. A model is introduced to evaluate the relative habitat value of these control regimes, here termed the Intermittent and Continuous. Compared to the Intermittent approach, the area of herbicide application in the Continuous approach is higher in the first 6 yr, but lower the reafter. The cumulative gain in habitat quality after 20 yr in either approach is sensitive to the presumed relative values ofPhragmites versusSpartina habitat, and may even be negative if they are nearly equal. Annual applications of herbicide to patches ofPhragmites appears to generate more habitat value and with less herbicide than occasional applications whenPhragmites cover is at is maximum.     

Life history strategies of estuarine nekton: The role of marsh macrophytes,benthic microalgae,and phytoplankton in the trophic spectrum

The stable isotope signatures of marine transient and resident nekton were used to investigate trophic linkages between primary producers, marsh macrophytes, phytoplankton, benthic microalgae, and consumers within the Delaware Bay. A whole estuary approach was used to compare the flux of nutrients from primary producers to juvenile weakfish (Cynoscion regalis), bay anchovy (Anchoa mitchilli), and white perch (Morone americana) in open waters of the lower and upper Bay and adjacent salt marshes dominated by eitherSpartina alterniflora orPhragmites australis. Our results suggest that trophic linkages vary significantly along the salinity gradient, reflecting the transition fromSpartina toPhragmites-dominated marshes, and secondarily, in a marsh to open water (offshore) direction at a given salinity. Superimposed on this pattern was a gradient in the proximate use of organic matter that depended on life history traits of each species ranging from pelagic to benthic in the order bay anchovy > weakfish > white perch.     

Essential Fish Habitat and Wetland Restoration Success: A Tier III Approach to the Biochemical Condition of Common Mummichog Fundulus heteroclitus in Common Reed Phragmites australis- and Smooth Cordgrass Spartina alterniflora-Dominated Salt Marshes

A tier III, essential fish habitat analysis was used to evaluate the biochemical condition of common mummichog Fundulus heteroclitus residing in two isolated tidal salt marshes, one a relatively undisturbed polyhaline site dominated by Spartina alterniflora and the other a meso-oligohaline site dominated by an invasive variety of Phragmites australis. Stable isotopes signatures of C, N, and S in whole tissue samples of F. heteroclitus were used to compare the trophic spectrum for this species in each marsh as a function of the dominant macrophytes present with additional contributions from phytoplankton and benthic microalgae. Allometry of wet mass and its components, water mass, lean protein mass and lipid mass in individual fish exhibited hyperallometric patterns; and average lipid mass fell within the range reported for most fundulids, including F. heteroclitus. Significant differences were also detected in the allocation of lipid classes to energy reserves in the form of triacylglycerols (TAG) and free fatty acids. These reserves, especially TAG, are critical for reproduction, migration, and overwintering survival in many taxa and were significantly lower in fish collected in the P. australis-dominated marsh. Relative to the relatively undisturbed Spartina-dominated site, we tentatively conclude that the P. australis-invaded marsh was an inferior habitat for F. heteroclitus.