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 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.     

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.     

Blue crab,<Emphasis Type="Italic">Callinectes sapidus</Emphasis>, response to the invasive common reed,<Emphasis Type="Italic">Phragmites australis</Emphasis>: Abundance,size, sex ratio,and molting frequency

Much effort has been directed recently at restoring marshes, by the removal of the invasive common reed,Phragmites australis, yet it is not clear how fish and invertebrates have responded either to the invasion ofPhragmites or to marsh restoration. The blue crab,Callinectes sapidus, uses marsh habitats during much of its benthic life. We investigated the response of blue crabs toPhragmites invasion and restoration efforts by comparing crab abundance (catch per unit effort), mean size and size frequency distribution, sex ratio, and molting of crabs in three physically similar areas differing in marsh vegetation;Spartina-dominated,Phragmites-dominated, and a treated area (Phragmites removed and now dominated bySpartina) in one marsh in the upper portion of Delaware Bay. Field sampling occurred monthly (April to November) from 1999 to 2001 using replicate daytime otter trawls in large marsh creeks. Crabs were categorized by carapace width into recruits (<30 mm), juveniles (30–115 mm), and adults (>115 mm). Juveniles dominated the system, representing 69.4% of all crabs. Similar monthly increases in mean size and molting patterns during the growing season (May–August) occurred inSpartina (natural and treated sites) andPhragmites sites suggesting that, subtidal habitats, used for molting, in these areas do not differ. More juveniles in the feeding molt stage (i.e., intermolt) than in other molt stages and more recruits predominantly in the feeding molt stage than adults were inSpartina, suggesting differences in the marsh surfaces used as feeding habitats withSpartina being preferred. Sex ratios of each life history stage were skewed towards males, but this was related to the low salinity of Alloway Creek, rather than marsh surface vegetation. Our results suggest that marsh surface vegetation influences the way blue crabs use marsh surface habitats, thus restoration efforts focusing on changing vegetation type may have a positive influence on blue crabs.     

Increased sediment accretion rates following invasion byPhragmites australis: The role of litter

Negative connotations of invasive plants worldwide have implicated them as the bearers of unfavorable ecosystem change. We contrasted 5-yr-old and 20-yr-oldPhragmites populations with pre-invasion areas occupied byTypha spp. andPanicum virgatum in an oligohaline tidal marsh of Chesapeake Bay. Peak live biomass was 3 times greater, while standing dead and litter was twice as great in the 20-yr-oldPhragmites. It is this abundance of concentrated litter on the marsh surface of maturePhragmites populations that we implicate as encouraging the trapping of organic and mineral matter. The rate of vertical accretion in 20-yr-oldPhragmites populations is 3–4 mm yr⁻¹ above the adjacent populations. By integrating the constant initial concentration and constant rate of supply models on individual²¹⁰Pb cores, we estimate thatPhragmites populations require a minimum of 7-yr post-colonization to enhance rates of accretion in this system. In ligh of the considerable loss of marsh habitat from relative sea-level rise, this finding contests the view that invasion creates strictly undesirable change at the ecosystem level.     

Utilization of <Emphasis Type="Italic">Spartina</Emphasis>- and <Emphasis Type="Italic">Phragmites</Emphasis>-Derived Dissolved Organic Matter by Bacteria and Ribbed Mussels (<Emphasis Type="Italic">Geukensia demissa</Emphasis>) from Delaware Bay Salt Marshes

Phragmites australis has been invading Spartina-alterniflora-dominated salt marshes throughout the mid-Atlantic. Although, Phragmites has high rates of primary production, it is not known whether this species supports lower trophic levels of a marsh food web in the same manner as Spartina. Using several related photochemical and biological assays, we compared patterns of organic matter flow of plant primary production through a key salt marsh metazoan, the ribbed mussel (Geukensia demissa), using a bacterial intermediate. Dissolved organic matter (DOM) was derived from plants collected from a Delaware Bay salt marsh and grown in the laboratory with ¹⁴C-CO₂. Bacterial utilization of plant-derived DOM measured as carbon mineralization revealed that both species provided bioavailable DOM to native salt marsh bacteria. Total carbon mineralization after 19 days was higher for Spartina treatments (36% ¹⁴CO₂ ± 3 SE) compared with Phragmites treatments (29% ±2 SE; Wilcoxon–Kruskal–Wallis rank sums test, P < 0.01). Pre-exposing DOM to natural sunlight only enhanced or decreased bioavailability of the DOM to the bacterioplankton during initial measurements (e.g., 7 days or less) but these differences were not significant over the course of the incubations. Mixtures of ¹⁴C-labeled bacterioplankton (and possibly organic flocs) from ¹⁴C-DOM treatments were cleared by G. demissa at similar rates between Spartina and Phragmites treatments. Moreover, ¹⁴C assimilation efficiencies for material ingested by mussels were high for both plant sources ranging from 74% to 90% and not significantly different between plant sources. Sunlight exposure did not affect the nutritional value of the bacterioplankton DOM assemblage for mussels. There are many possible trophic and habitat differences between Spartina- and Phragmites-dominated marshes that could affect G. demissa but the fate of vascular plant dissolved organic carbon in the DOM to bacterioplankton to mussel trophic pathway appears comparable between these marsh types.     

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.     

Effects of common reed (Phragmites australis) invasion on marsh surface macrofauna: Response of fishes and decapod crustaceans

The tidally inundated marsh surface is an importnat site for energy exchanges for many resident and transient species. In many areas along the East Coast of the U.S. the dominant vegetation,Spartina alterniflora, has been replaced by the common reed (Phragmites australis). This shift has caused concern about the impact ofPhragmites on marsh fauna but research in this area has been limited. During 1997 and 1998, we examined the effect ofPhragmites on fish and decapod crustacean use of the marsh surface in the brackish water reaches of the Mullica River, in southern New Jersey, U.S. Fish and decapod crustaceans were sampled with an array of shallow pit traps (rectangular glass dishes, 27.5×17.5×3.7 cm) and with flumes (1.3 m wide×10 m long of 3.2-mm mesh). Fish (2–60 mm TL) dominated pit trap collections withFundulus heteroclitus andFundulus luciae significantly more abundant atSpartina sites.Fundulus heteroclitus was also the dominant fish (15–275 mm TL) collected in flumes but collections with this gear, including a number of species not collected in pit traps, showed no distinct preferences for different marsh vegetation types. Decapod crustaceans (1–48 mm CW) collected in pit traps were generally less abundant than fishes withCallinectes sapidus andPalaemonetes spp. most abundant inSpartina, whileRhithropanopeus harrisii was most abundant inPhragmites. The same decapod crustacean species (2–186 mm CW) dominanted the flume collections and, similar to the pattern of fish collected by the flumes, there were no distinct habitat preferences for different marsh vegetation types. As a result of these observations, with different sampling techniques, it appears there is an overall negative effect ofPhragmites on larval and small juvenile fish but less or no effect on larger fish and decapods crustaceans.     

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.     

Comparison of fish and macroinvertebrate use of<Emphasis Type="Italic">Typha angustifolia,Phragmites australis</Emphasis>, and treated<Emphasis Type="Italic">Phragmites</Emphasis> marshes along the lower Connecticut River

Since 1965 large areas of lower Connecticut River tidelands have been converted from high diversity brackish meadow andTypha angustifolia marsh to near monocultures ofPhragmites australis. This study addresses the impact ofPhragmites invasion on fish and crustacean use of oligohaline high marsh. During spring tides from early June through early September 2000, fishes and crustaceans leaving flooded marsh along 3 km of the Lieutenant River, a lower Connecticut River tributary, were captured with Breder traps at 90 sites, equally distributed amongPhragmites, Typha, and treated (herbicide and mowing)Phragmites areas. Pit traps, 18 per vegetation type in 2000 and 30 each inPhragmites andTypha in 2001, caught larvae and juveniles at distances of up to 30 m into the marsh interior. There were no significant differences in fish species compositions or abundances among the vegetation types. Size distributions, size specific biomasses, and diets ofFundulus heteroclitus, the numerically dominant fish, were also similar. The shrimpPalaemonetes pugio was more abundant inPhragmites than in other types of vegetation, whereas the fiddler crabUca minax was least numerous inPhragmites. Mean numbers ofF. heteroclitus andP. pugio caught per site event were positively correlated with increasing site hydroperiod. Significantly moreF. heteroclitus were captured along the upper reach of the river where marsh elevations were lower than farther downstream. MoreF. heteroclitus and fewerP. pugio andU. minax were captured during the day than at night. A relatively small number of larval and juvenileFundulus sp. were captured in pit traps, but consistently fewer inPhragmites than inTypha, suggesting thatTypha and brackish meadow marshes may provide better nursery habitat. Vegetation was sampled along a 30 m transect at each trap site in 2000. Plant species diversity was greatest in treatedPhragmites areas and lowest inPhragmites sites.     

Monitoring food web changes in tide-restored salt marshes: A carbon stable isotope approach

Primary producer (angiosperms, macroalgae, submerged aquatic vegetation), suspended particulate matter, andFundulus heteroclitus isotope values (δ¹³C, δ¹⁵N, δ³⁴S) were examined to assess their use as indicators for changes in food web support functions in tidally-restored salt marshes. Study sites, located throughout the southern New England region (USA), ranged fromSpartina alterniflora-dominated reference marshes, marshes under various regimes and histories of tide restoration, and a severely tide-restrictedPhragmites australis marsh.Fundulus δ¹³C values were greater for fish from referenceSpartina marshes than for fish from adjacent tide-restricted or tide-restored marshes where higher percent cover of C₃ plants, lower water column salinities, and more negative dissolved inorganic δ¹³C values were observed. The difference inFundulus δ¹³C values between a tide-restrictedPhragmites marsh and an adjacent referenceSpartina marsh was great compared to the difference between marshes at various stages of tide restoration and their respective reference marshes, suggesting that food web support functions are restored as the degree of tidal restriction is lessened. While a multiple isotopic approach can provide valuable information for determining specific food sources to consumers, this study demonstrates that monitoringFundulus δ¹³C values alone may be useful to evaluate the trajectory of ecological change for marshes undergoing tidal restoration.     

Spatial and temporal dynamics of epiphytic microalgae on the cordgrassSpartina alterniflora in North Inlet Estuary, South Carolina

Epiphytic microbial biomass (as chlorophylla) was measured monthly in North Inlet Estuary, South Carolina, for 16 months on spatially distinct stem sections (bottom and middle) of dead and livingSpartina alterniflora growth forms (tall, medium, and short) exposed at low tide. The highest biomass was located on the bottom section of tall plants, presumably due to their relatively longer contact with creek water and associated phytoplankton, and their closer proximity to marsh sediments with associated benthic microalgae, both recruitment sources for epiphytes. Dead plants left standing from the previous year’s growth cycle had higher epiphytic biomass than living plants, which occurred mostly in late spring through fall. Epiphytic biomass was highest in the winter (mean of 1.77 mg chla (m² marsh)⁻¹) and lowest in the summer (mean of 0.34 mg chla (m² marsh)⁻¹). Because phytoplankton andSpartina production are lowest in the winter, the results emphasize the relative importance of epiphytes to growth of herbivores in this season.     

<Emphasis Type="Italic">Phragmites australis</Emphasis> invasion and expansion in tidal wetlands: Interactions among salinity,sulfide, and hydrology

Through their physiological effects on ion, oxygen, and carbon balance, respectively, salinity, sulfide, and prolonged flooding combine to constrain the invasion and spread ofPhragmites in tidal wetlands. Initial sites of vigorous invasion by seed germination and growth from rhizome fragments appear limited to sections of marsh where salinity is <10‰, sulfide concentrations are less than 0.1 mM, and flooding frequency is less than 10%. In polyhaline tidal wetlands the invasion sites include the upland fringe and some high marsh creek banks. The zones of potential invasion tend to be larger in marshes occupying lower-salinity portions of estuaries and in marshes that have been altered hydrologically. Owing to clonal integration and a positive feedback loop of growth-induced modification of edaphic soil conditions, however, a greater total area of wetland is susceptible toPhragmites expansion away from sites of establishment. Mature clones have been reported growing in different marshes with salinity up to 45‰, sulfide concentration up to 1.75 mM, and flooding frequency up to 100%. ForPhragmites establishment and expansion in tidal marshes, windows of opportunity open with microtopographic enhancement of subsurface drainage patterns, marsh-wide depression of flooding and salinity regimes, and variation in sea level driven by global warming and lunar nodal cycles. To avoidPhragmites monocultures, tidal wetland creation, restoration, and management must be considered within the context of these different scales of plant-environment interaction.     

Evolution of tidal creek networks in a high sedimentation environment: A 5-year experiment at Tijuana Estuary, California

In a large (8 ha) salt marsh restoration site, we tested the effects of excavating tidal creeks patterned after reference systems. Our purposes were to enhance understanding of tidal creek networks and to test the need to excavate creeks during salt marsh restoration. We compared geomorphic changes in areas with and without creek networks (n = 3; each area 1.3 ha) and monitored creek cross-sectional areas, creek lengths, vertical accretion, and marsh surface elevations for 5 yr that included multiple sedimentation events. We hypothesized that cells with creeks would develop different marsh surface and creek network characteristics (i.e., surface elevation change, sedimentation rate, creek cross-sectional area, length, and drainage density). Marsh surface vertical accretion averaged 1.3 cm yr⁻¹ with large storm inputs, providing the opportunity to assess the response of the drainage network to extreme sedimentation rates. The constructed creeks initially filled due to high accretion rates but stabilized at cross-sectional areas matching, or on a trajectory toward, equilibrium values predicted by regional regression equations. Sedimentation on the marsh surface was greatest in low elevation areas and was not directly influenced by creeks. Time required for cross-sectional area stabilization ranged from 0 to > 5 yr, depending on creek order. First-order constructed creeks lengthened rapidly (mean rate of 1.3 m yr⁻¹) in areas of low elevation and low vegetation cover. New (volunteer) creeks formed rapidly in cells without creeks in areas with low elevation, low vegetation cover, and high elevation gradient (mean rate of 6.2 m yr⁻¹). After 5 yr, volunteer creeks were, at most, one-fourth the area of constructed creeks and had not yet reached the upper marsh plain. In just 4 yr, the site’s drainage density expanded from 0.018 to reference levels of 0.022 m m⁻². Pools also formed on the marsh plain due to sediment resuspension associated with wind-driven waves. We conclude that excavated creeks jump-started the development of drainage density and creek and channel dimensions, and that the tidal prism became similar to those of the reference site in 4–5 yr.     

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.     

Can Plant Competition and Diversity Reduce the Growth and Survival of Exotic <Emphasis Type="Italic">Phragmites australis</Emphasis> Invading a Tidal Marsh?

The rapid proliferation of Phragmites australis in North America has challenged resource managers to curb its expansion and reduce the loss of functional tidal marsh. We investigated whether native plant competition could reduce the ability of Phragmites to invade a tidal marsh, and if plant diversity (species richness, evenness, and composition) altered the competitive outcome. Immature Phragmites shoots and four native halophytes were transplanted to small but dense field plots (~1,200 shoots m⁻²) comprising three community structure types (Phragmites alone, Phragmites + 1 native species, and Phragmites + 4 native species). Interspecific competition significantly reduced Phragmites aboveground biomass, shoot length production, density, and survival by approximately 60%. Additionally, plots planted with greater native diversity contained Phragmites with the lowest growth and survival, potentially indicating diversity-enhanced resource competition. Competition consistently reduced the growth of Phragmites even under favorable conditions: lack of strong tidal flooding stresses as well as elevated nutrient pools.     

Net primary production and decomposition of salt marshes of the Ebre delta (Catalonia,Spain)

Net primary production was measured in three characteristic salt marshes of the Ebre delta: anArthrocnemum macrostachyum salt marsh,A. macrostachyum-Sarcocornia fruticosa mixed salt marsh andS. fruticosa salt marsh. Above-ground and belowground biomass were harvested every 3 mo for 1 yr. Surface litter was also collected from each plot. Aboveground biomass was estimated from an indirect non-destructive method, based on the relationship between standing biomass and height of the vegetation. Decomposition of aboveground and belowground components was studied by the disappearance of plant material from litter bags in theS. fruticosa plot. Net primary production (aboveground and belowground) was calculated using the Smalley method. Standing biomass, litter, and primary production increased as soil salinity decreased. The annual average total aboveground plus belowground biomass was 872 g m⁻² in theA. macrostachyum marsh, 1,198 g m⁻² in theA. macrostachyum-S. fruticosa mixed marsh, and 3,766 g m⁻² in theS. fruticosa biomass (aboveground plus belowground) was 226, 445, and 1,094 g m⁻², respectively. Total aboveground plus below-ground net primary production was 240, 1,172, and 1,531 g m⁻² yr⁻¹. There was an exponential loss of weight during decomposition. Woody stems and roots, the most recalcitrant material, had 70% and 83% of the original material remaining after one year. Only 20–22% of leafy stem weight remained after one year. When results from the Mediterranean are compared to other salt marshes dominated by shrubbyChenopodiaceae in Mediterranean-type climates, a number of similarities emerge. There are similar zonation patterns, with elevation and maximum aboveground biomass and primary production occurring in the middle marsh. This is probably because of stress produced by waterlogging in the low marsh and by hypersalinity in the upper marsh.     

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.     

Connectivity Among Salt Marsh Subhabitats: Residency and Movements of the Mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>)

We examined connectivity among marsh subhabitats to determine the structural limits and important components of a polyhaline salt marsh by studying the patterns of abundance, residency, and movement of a numerically and ecologically dominant nektonic fish (mummichog, Fundulus heteroclitus). We captured, tagged (n = 14,040 individuals, 30–110 mm), and recaptured from Feb 2001 to Jul 2002, although most recaptures (75–95% by tagging location) occurred within 150 days. Seasonal residency and movements were common among most subhabitats based on catch per unit effort and recapture per unit effort. Thus, these (marsh pools, intertidal and subtidal creeks, and marsh surface) should be considered natural subhabitats within New England type salt marshes. Further, all these subhabitat types should be included in studies of salt marsh nekton and marsh restoration and creation activities.     

Mechanisms of marsh habitat alteration due to<Emphasis Type="Italic">Phragmites</Emphasis>: Response of young-of-the-year mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) to treatment for<Emphasis Type="Italic">Phragmites</Emphasis> removal

In recent decades, marshes naturally dominated bySpartina spp. have been replaced byPhragmites australis throughout the northeastern United States. We suggest that early in this invasion there was little effect on the fish fauna. As the invasion proceeds, the marsh surface habitat became more altered (i.e., elevated, flattened, reduced water-filled depressions, and reduced standing water), which resulted in a reduction of feeding, reproduction, and nursery function for fishes, especiallyFundulus spp. These potential changes in marsh habitat and function have resulted in numerous attempts to removePhragmites and restoreSpartina spp. To evaluate the response of marsh surface fishes toPhragmites treatment, we examined fish use in the brackish water reaches of Alloway Creek in the Delaware Bay estuary. ReferencePhragmites habitats were compared with referenceSpartina alterniflora-dominated habitats and sites treated (1996–1998) to removePhragmites to restore former vegetation (i.e., restored, now comprised of 100%Spartina). Fish were sampled with an array (n=9 at each site) of shallow pit traps (rectangular glass dishes, 27.5×17.5×3.7 cm). Small individuals (mean=17.5, 5–45 mm TL) dominated all pit trap collections. Fish abundance was highest at the restored (catch per unit effort [CPUE]=2.16) andSpartina (CPUE=0.81) sites with significantly lower values atPhragmites (CPUE=0.05) habitats. Samples were dominated by young-of-the-year mummichog,Fundulus heteroclitus (98% of total fish, n=631). The only other fish species collected was spotfin killifish,Fundulus luciae (2% of total catch, n=14), which was only present in restored andSpartina habitats. These observations suggest that the restored marsh is providing habitat (water-filled depressions on the marsh surface) for young-of-the-yearFundulus spp. These marshes are responding favorably to the restoration based on the much greater abundance of fish in restored versusPhragmites habitats and the overall similarity between restored andSpartina habitats.