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.     

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.     

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.     

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.     

Comparison of Nekton Use for Cordgrass <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and Bulrush <Emphasis Type="Italic">Scirpus mariqueter</Emphasis> Marshes in the Yangtze River Estuary,China

To test whether invasive Spartina alterniflora marshes were functionally equivalent to native Scirpus mariqueter marshes, the present study used bottomless lift nets (20 m²) during 12 high-tide events from August to October 2008 to compare nekton densities and biomass between the two marsh types in the Dongtan wetland. A total of eight species of fish, two species of shrimp, and three species of crab were collected. So-iny mullet Chelon haematocheilus, keeled mullet Liza carinata, Asian freshwater goby Acanthogobius ommaturus, and ridge-tail prawn Exopalaemon carinicauda dominated samples from the two marsh types and accounted for over 90% of the total catch. There were significantly greater densities and biomass (p < 0.05) of total nekton (all species combined) and two mullets (C. haematocheilus and L. carinata) in S. alterniflora marshes than in S. mariqueter marshes in August 2008, while no significant differences (p > 0.05) between the two marsh types were observed for densities and biomass of any species or total nekton in September and October 2008. Non-metric multidimensional scaling ordination did not show clear separation of samples between the two marsh types (r = 0.071, p = 0.159). Furthermore, there were no habitat-specific differences (p > 0.05) in the size distributions of the three numerically dominant species (C. haematocheilus, L. carinata, and A. ommaturus). We concluded that S. alterniflora marshes were utilized by nekton in a fashion similar to their utilization of native S. mariqueter marshes under similar physical conditions.     

A comparison of sediment microbial communities associated with<Emphasis Type="Italic">Phragmites australis</Emphasis> and<Emphasis Type="Italic">Spartina alterniflora</Emphasis> in two brackish wetlands of New Jersey

The extensive spread ofPhragmites australis throughout brackish marshes on the East Coast of the United States is a major factor governing management and restoration decisions because it is assumed that biogeochemical functions are altered by the invasion. Microbial activity is important in providing wetland biogeochemical functions such as carbon and nitrogen cycling, but there is little known about sediment microbial communities inPhragmites marshes. Microbial populations associated with invasivePhragmites vegetation and with native salt marsh cordgrass,Spartina alterniflora, may differ in the relative abundance of microbial taxa (community structure) and in the ability of this biota to decompose organic substrates (community biogeochemical function). This study compares sediment microbial communities associated withPhragmites andSpartina vegetation in an undisturbed brackish marsh near Tuckerton, New Jersey (MUL), and in a brackish marsh in the anthropogenically affected Hackensack meadowlands (SMC). We use phospholipid fatty acid (PLFA) analysis and enzymataic activity to profile sediment microbial communities associated with both plants in each site. Sediment analyses include bulk density, total organic matter, and root biomass. PLFA profiles indicate that the microbial communities differ between sites with the undisturbed site exhibiting greater fatty acid richness (62 PLFA recovered from MUL versus 38 from SMC). Activity of the 5 enzymes analyzed (β-glucosidase, acid phosphatase, chitobiase, and 2 oxidases) was higher in the undisturbed site. Differences between vegetation species as measured by Principal Components Analysis were significantly greater at the undisturbed MUL site than at SMC, and patterns of enzyme activity and PLFAs did not correspond to patterns of root biomass. We suggest that in natural wetland sediments, macrophyte rhizosphere effects influence the community composition of sediment microbial populations. Physical and chemical site disturbances may impose limits on these rhizosphere effects, decreasing sediment microbial diversity and potentially, microbial biogeochemical functions.     

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.     

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.     

The Intertidal Burrowing Crab <Emphasis Type="Italic">Neohelice</Emphasis> (=<Emphasis Type="Italic">Chasmagnathus</Emphasis>) <Emphasis Type="Italic">granulata</Emphasis> Positively Affects Foraging of Rodents in South Western Atlantic Salt Marshes

The role of positive and indirect interactions is often crucial in communities with intense abiotic stress such as salt marshes. The burrowing crab, Neohelice (=Chasmagnathus) granulata, is the dominant benthic macroinvertebrate of southwest Atlantic marshes (southern Brazil to Northern Argentinean Patagonia), having strong direct and indirect effects on marsh soil and, in consequence, on marsh vegetation and primary consumers. In this work, we investigate if this crab indirectly modifies habitat use by the granivorous rodents, Akodon azarae and Oligoryzomys flavescens, by increasing nutrient availability and thus enhancing seed production by the marsh plant Spartina densiflora. The study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°32′ S). Rodent frequencies in S. densiflora were positively correlated with crab densities throughout the low and middle marsh. Additionally, the highest quality of S. densiflora and inflorescence density was recorded at the highest crab densities. Experimental manipulation of crab densities shows that N. granulata indirectly enhances the performance of S. densiflora (e.g., decreased fiber content and C/N ratios) and increases density of seeds. Moreover, N. granulata also facilitates S. densiflora seed availability to rodents by concentrating them in sediment mound at their burrows entrances. Experimental rodent exclusions showed that rodent species used S. densiflora seeds, a variable positively related to crab burrow density. Thus, our results show that N. granulata drives the granivorous rodent distribution and the intensity of seeds–rodent interaction trough facilitative and indirect interactions in marsh community.     

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.     

Growth and Movements of Mummichogs (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) Along Armored and Vegetated Estuarine Shorelines

Alteration of estuarine shorelines associated with increased urbanization can significantly impact biota and food webs. This study determined the impact of shoreline alteration on growth and movement of the estuarine fish Fundulus heteroclitus in a tributary of the Delaware Coastal Bays. Fundulus heteroclitus is abundant along the east coast of the USA, and is an important trophic link between marsh and subtidal estuary. The restricted home range of F. heteroclitus allowed discrete sampling, and fish growth comparisons, along 35–65-m long stretches of fringing Spartina alterniflora and Phragmites australis marsh, riprap, and bulkhead. Fundulus heteroclitus were tagged with decimal Coded Wire Tags. Of 725 tagged F. heteroclitus, 89 were recaptured 30–63 days later. Mean growth rate (0.06–0.15 mm day^?1 across all shoreline types) was greatest at riprap, lowest at Spartina and Phragmites, and intermediate at bulkhead, where growth was not significantly different from any other shoreline. This suggests that discernible environments exist along different shoreline types, even at the scale of tens of meters. No difference in movement distance was detected at different shoreline types; most individuals displayed a high degree of site fidelity. Forty-seven percent were recaptured within 5 m of their tagging location, although alongshore movements up to 475 m were recorded. Estimates of relative F. heteroclitus productivity, using relative density data from a concurrent study, were highest along Spartina and Phragmites, intermediate at riprap, and lowest at bulkhead. Therefore, despite greater growth rates along riprap than at vegetated shores, armoring reduces abundance sufficiently to negatively impact localized productivity of F. heteroclitus.     

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.     

Growth and production of the mummichog (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) in a restored salt marsh

The mummichog,Fundulus heteroclitus, is one of the most important macrofaunal components of salt marsh surfaces and an important link to subtidal areas of the adjacent estuary along the east coast of the U.S. We estimated growth, population size, and production of the mummichog in a restored marsh in order to improve our understanding of the role of this resident fish and to evaluate the success of the restoration. The restored marsh, covering 234 ha, was a former salt hay farm located in the mesohaline portion of Delaware Bay that was restored to tidal influence in August 1996. We separated the mummichog population into two components based on life history stage and summer habitat use patterns. One component, consisting of adults and large young-of-the-year (YOY), exhibited tidal movements to and from the marsh surface and the subtidal creeks. These were examined with an intensive mark and recapture program using coded wire tags. Another component, consisting of small YOY, remained on the marsh surface throughout the tidal cycle. Throw traps were used to sample these small YOY. The mean annual population density of adults and large YOY for the entire marsh was approximately 1.2 fish m⁻² and mean monthly density peaked at 2.9 fish m⁻². The mean annual density of small YOY on the marsh surface was 15.1 fish m⁻² and mean monthly density peaked at 41.4 fish m⁻². Size and season influenced the growth rate of individual fish and instantaneous growth rates ranged from 0.03 to 2.26 mo⁻¹. Total annual mummichog production was estimated to be 8.37 g dw m⁻² yr⁻¹, with adults and large YOY contributing 28.4% (2.38 g dw m⁻² yr⁻¹) and small YOY on the marsh surface contributing 71.6% (5.99 g dw m⁻² yr⁻¹). The seasonal use and population densities were comparable to previous studies in natural marshes while growth and production of mummichog in this restored marsh appeared to be higher. Coupled with the results of other studies on the feeding, movement, and habitat use of this species in this restored marsh, the species has responded well to the restoration.     

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.     

<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.     

Ecosystem Engineering Effects of <Emphasis Type="Italic">Aster tripolium</Emphasis> and <Emphasis Type="Italic">Salicornia procumbens</Emphasis> Salt Marsh on Macrofaunal Community Structure

This paper examines how perennial Aster tripolium and annual Salicornia procumbens salt marshes alter the biomass, density, taxon diversity, and community structure of benthic macrofauna, and also examines the role of elevation, sediment grain size, plant cover, and marsh age. Core samples were collected on a fixed grid on an intertidal flat in the Westerschelde estuary (51.4° N, 4.1° E) over 5 years (2004–2008) of salt marsh development. In unvegetated areas, macrobenthic biomass, density, and taxon diversity were highest when elevation was highest, benthic diatoms were most abundant, and sediment median grain size was smallest. In contrast, in salt marsh areas, macrobenthic biomass and taxon diversity increased with median grain size, while the effects of elevation and diatom abundance on macrobenthic biomass, density, and diversity were not significant. In fine sediments, macrofaunal community structure in the salt marsh was particularly affected; common polychaetes such as Nereis diversicolor, Heteromastus filiformis, and Pygospio elegans had low abundance and oligochaetes had high abundance. Marsh age had a negative influence on the density of macrofauna, and A. tripolium stands had lower macrofaunal densities than the younger S. procumbens stands. There were no significant effects of marsh age, plant cover, and vegetation type on macrobenthic biomass, taxon diversity, and community structure. The results highlight that ecosystem engineering effects of salt marsh plants on macrofauna are conditional. Organic enrichment of the sediment and mechanical hindering of macrofaunal activity by plant roots are proposed as plausible mechanisms for the influence of the salt marsh plants on macrofauna.     

Effects of an Omnivorous Katydid,Salinity, and Nutrients on a Planthopper-<Emphasis Type="Italic">Spartina</Emphasis> Food Web

Top–down and bottom–up effects interact to structure communities, especially in salt marshes, which contain strong gradients in bottom–up drivers such as salinity and nutrients. How omnivorous consumers respond to variation in prey availability and plant quality is poorly understood. We used a mesocosm experiment to examine how salinity, nutrients, an omnivore (the katydid Orchelimum fidicinium) and an herbivore (the planthopper Prokelisia spp.) interacted to structure a simplified salt marsh food web based on the marsh grass Spartina alterniflora. Bottom–up effects were strong, with both salinity and nutrients decreasing leaf C/N and increasing Prokelisia abundance. Top–down effects on plants were also strong, with both the herbivore and the omnivore affecting S. alterniflora traits and growth, especially when nutrients or salt were added. In contrast, top–down control by Orchelimum of Prokelisia was independent of bottom–up conditions. Orchelimum grew best on a diet containing both Spartina and Prokelisia, and in contrast to a sympatric omnivorous crab, did not shift to an animal-based diet when prey were present, suggesting that it is constrained to consume a mixed diet. These results suggest that the trophic effects of omnivores depend on omnivore behavior, dietary constraints, and ability to suppress lower trophic levels, and that omnivorous katydids may play a previously unrecognized role in salt marsh food webs.     

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.     

Foraging by Two Estuarine Fishes, <Emphasis Type="Italic">Fundulus heteroclitus</Emphasis> and <Emphasis Type="Italic">Fundulus majalis</Emphasis>, on Juvenile Asian Shore Crabs (<Emphasis Type="Italic">Hemigrapsus sanguineus</Emphasis>) in Western Long Island Sound

The Asian shore crab, Hemigrapsus sanguineus, is a recent and particularly successful introduction to the east coast of the USA. Little research has been done on the utilization of Asian shore crabs for food by native species, a potential form of biocontrol. Over a 4-year period, we examined the gut contents of cogeners, Fundulus heteroclitus and Fundulus majalis, collected from two embayments in western Long Island Sound for the presence of juvenile H. sanguineus. Frequency (percent) of occurrence of food items in the guts of both species varied over year and study site. Asian shore crabs were consumed more often by F. heteroclitus than by F. majalis, but predation pressure by both species was low. Only 13% of F. heteroclitus and 7.7% of F. majalis found with food in their guts had ingested Asian shore crab remains. Of those, 1/3 had consumed whole crabs; the rest had only autotomized appendages in their guts. The mean carapace width of juvenile Asian shore crabs ingested by F. heteroclitus was 3.59 ± 2.22 mm (N = 33). Results of our study on killifish predation support the hypothesis that H. sanguienus abundance is partly explained by reduced impact of native predators (i.e., the “enemy release hypothesis”). Predation pressure of other potential enemies on both planktonic and benthic stages of the Asian shore crab must be investigated, however, to understand the full impact of predation on H. sanguineus population dynamics.     

Determinants of expansion for<Emphasis Type="Italic">Phragmites australis</Emphasis>, common reed,in natural and impacted coastal marshes

The rapid spread ofPhragmites australis in the coastal marshes of the Northeastern United States has been dramatic and noteworthy in that this native species appears to have gained competitive advantage across a broad range of habitats, from tidal salt marshes to freshwater wetlands. Concomitant with the spread has been a variety of human activities associated with coastal development as well as the displacement of nativeP. australis with aggressive European genotypes. This paper reviews the impacts caused by pure stands ofP. australis on the structure and functions of tidal marshes. To assess the determinants ofP. australis expansion, the physiological tolerance and competitive abilities of this species were examined using a field experiment.P. australis was planted in open tubes paired withSpartina alterniflora, Spartina patens, Juncus gerardii, Lythrum salicaria, andTypha angustifolia in low, medium, and high elevations at mesohaline (14‰), intermediate (18‰), and salt (23‰) marsh locations. Assessment of the physiological tolerance ofP. australis to conditions in tidal brackish and salt marshes indicated this plant is well suited to colonize creek banks as well as upper marsh edges. The competitive ability ofP. australis indicated it was a robust competitor relative to typical salt marsh plants. These results were not surprising since they agreed with field observations by other researchers and fit within current competition models throught to structure plant distribution within tidal marshes. Aspects ofP. australis expansion indicate superior competitive abilities based on attributes that fall outside the typical salt marsh or plant competition models. The alignment of some attributes with human impacts to coastal marshes provides a partial explanation of how this plant competes so well. To curb the spread of this invasive genotype, careful attention needs to be paid to human activities that affect certain marsh functions. Current infestations in tidal marshes should serve as a sentinel to indicate where human actions are likely promoting the invasion (e.g., through hydrologic impacts) and improved management is needed to sustain native plant assemblages (e.g., prohibit filling along margins).