Use of detrital foods and assimilation of nitrogen by coastal detritivores

Two killifish common in east coast U.S.A. salt marshes,Cyprinodon variegatus Lacepede andFundulus heteroclitus Walbaum, differ in their ability to assimilate nitrogen from and grow on detritus.C. variegatus grew on a diet of detritus ofSpartina alterniflora Loisel, whileF. heteroclitus did not. In addition, when the fish were fed¹⁵N-labeledS. alterniflora detritus,¹⁵N:¹⁴N ratios inC. variegatus were higher than were ratios inF. heteroclitus. Therefore, even though both species ingest large amounts of detritus,C. variegatus makes more effective use of this portion of its diet. These dietary differences are corroborated by anatomical differences that suggest thatC. variegatus should make better use of detrital or plant tissues thanF. heteroclitus. In the label experiment, the degree of label in both fish was directly proportional to the degree of label in the food treatments. In previously published experiments designed to compare plant substrate with attached microbes as nitrogen sources for detritivores, %¹⁵N incorporated by a polychaete was also directly proportional to %¹⁵N in the detrital food. Therefore, it is difficult to distinguish between plant substrate and microbes as nitrogen sources for this detritivore.     

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.     

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.     

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.     

Tidal utilization of nekton in Delaware Bay restored and reference intertidal salt marsh creeks

Large-scale marsh restoration efforts were conducted to restore normal salt marsh structure and function to degraded marshes (i.e., former salt hay farms) in the mesohaline lower Delaware Bay. While nekton response has been previously evaluated for the marsh surface and subtidal creeks in these marshes, little effort has been focused on intertidal creeks. Nekton response in intertidal creeks was evaluated by sampling with seines to determine if restored (i.e., former salt hay farms restored in 1996) and reference (i.e., natural or relatively undisturbed) salt marshes were utilized by intertidal nekton in a similar manner. The overall nekton assemblage during June–October 2004–2005 was generally comprised of the same species in both the restored and reference marshes. Intertidal creek catches in both marsh types consisted primarily ofFundulus heteroclitus andMenidia menidia, with varying numbers of less abundant transient species present. Transient nekton were more abundant at restored marshes than reference marshes, but in insufficient numbers to cause differences in nekton assemblages. In both marsh types, low tide stages were characterized by resident nekton, dominated byF. heteroclitus, while high tide stages were characterized by a variable mix of transient and resident nekton. Assemblage level analyses indicated that intertidal creeks in restored and reference marshes were generally utilized in a similar manner by a similar nekton assemblage, so restoration efforts were deemed successful. This is in agreement with multiple comparative studies from the ame marshes examining fish, invertebrates, and vegetation in different marsh habitats.     

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.     

Salt-marsh tide pools as winter refuges for the mummichog,Fundulus heteroclitus,in New Jersey

Mummichog,Fundulus heteroclitus, were collected weekly from a southern New Jersey high-salinity salt marsh from October 1988 to June 1989 and from September 1989 to June 1990 to determine the overwintering habitat. Major habitat types sampled within the salt marsh were subtidal creek, intertidal creeks, and salt-marsh pools. Few individuals were collected in the intertidal creek or the subtidal creek from the end of October through the beginning of May for both years, when creek water temperatures were low. Both young-of-the-year and adults of both sexes were abundant in the salt-marsh pools (total lengths ranged from 29 mm to 125 mm) throughout the winter. In the spring, catch per unit effort (CPUE) within the tidal creek increased with increasing water temperature, while CPUE in marsh pools decreased with increases in estuarine water temperature. These collection patterns indicate that the majority ofF. heteroclitus may move from subtidal and intertidal creeks into salt-marsh pools in the late fall and leave in the spring. This seasonal movement could explain how fish survive winter environmental conditions because daily average water temperatures of salt-marsh pools were warmer than subtidal creek temperatures for much of the winter.     

The effect of nitrogen loading on a brackish estuarine faunal community: A stable isotope approach

Coastal ecosystems worldwide face increased nutrient enrichment from shoreline and watershed development and atmospheric pollution. We investigated the response of the faunal community of a small microtidal estuary dominated byRuppia maritima (widgeon grass) in Maine, United States, to increased nitrogen loading using an in situ mesocosm enrichment experiment. Community response was characterized by assessing quantitative shifts in macroinvertebrate community composition and identifying changes in food web structure using stable carbon and nitrogen isotope ratios of producers and consumers. The community was dominated by brackish water invertebrates including midge larvae, oligochaetes, damselfly larvae, amphipods, and ostracods. Experimental nutrient additions resulted in significantly lower densities of herbivorous chironomids and predatory damselflies and greater densities of deposit feeding oligochaetes. Grazing midge larvae (Chironomidae:Dicrotendipes, Cricotopus) consumed epiphytic algae under both natural and enriched conditions. Deposit feedingChironomus was dependent on allochthonous sources of detritus under natural conditions and exhibited a shift to autochthonous sources of detritus under enriched conditions. PredatoryEnallagma primarily consumed grazing chironomids under all but the highest loading conditions. Experimental nutrient loading resulted in an increase in generalist deposit feeders dependent on autochthonous sources of detritus.     

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.     

Life history of the spotfin killifish,fundulus luciae (pisces: cyprinodontidae), in Fox Creek Marsh,Virginia

Fundulus luciae was collected for a year at Fox Creek Marsh and, occasionally, at other salt marshes along the York River, Virginia. It occurred in high intertidal areas in brackish, sometimes oxygen deficient shallow ditches, mudholes and tidal rivulets located in stands ofSpartina alterniflora. Preserved specimens were examined to determine the reproductive season, fecundity, diet and associated metazoan parasites of the species. Developing ova were present from mid-April to mid-August and exhibited a broad size range; the number of large ova (>1.6 mm diameter) never exceeded 16 per female. Stomach contents consisted mainly of detritus, diatoms, ostracods, dipterans and copepods. All major metazoan parasite groups except Cestoda were represented onF. luciae; Monogenea were most numerous, with 57.2% incidence. Eggs and larvae were described. Preserved larvae ofF. luciae were distinguished from those ofF. heteroclitus by their dorsal pigmentation pattern. It was concluded thatF. luciae, a purportedly rare species, is not rare, but probably restricted to high intertidal salt marshes where it may have been previously overlooked.     

Gut contents of common mummichogs,Fundulus heteroclitus L., in a restored impounded marsh and in natural reference marshes

We examined the gut contents of mummichogs, Fundulus heteroclitus L., entering and leaving ditches in three marsh regions within the Barn Island Wildlife Management Area in Connecticut: a restored impounded valley marsh, a marsh below the impoundment dike (Headquarters Marsh), and an unimpounded valley marsh (Davis Marsh). On the Headquarters Marsh and at certain times on the other two marshes, fish entered the ditches on the flooding tide with relatively little food in their guts and left them on the following ebbing tide with considerably more food in their guts. Since the high tides did not flood the surface of the high marsh, it appears that the ditches are important foraging areas. Major components of the gut contents were detritus, algae, amphipods, tanaids, copepods, and insects. During the summer, fish in the restored impounded marsh consumed less food per unit body weight than did fish inhabiting the other marsh regions.     

Effects of Life History Strategy on Fish Distribution and Use of Estuarine Salt Marsh and Shallow-Water Flat Habitats

To assess the potential for habitat isolation effects on estuarine nekton, we used two species with different dispersal abilities and life history strategies, mummichog (Fundulus heteroclitus) and pinfish (Lagodon rhomboides) to examine: (1) distribution trends among estuarine shallow-water flat and various intertidal salt marsh habitats and (2) the influence of salt marsh habitat size and isolation. Collections were conducted using baited minnow traps set within nonisolated interior marshes (interior), nonisolated fringing marshes (nonisolated), isolated island marshes (isolated), and shallow-water flat habitats (flat) that were adjacent to isolated and nonisolated marshes. Size range of individuals collected included juvenile and adult F. heteroclitus (20–82-mm standard length) and L. rhomboides (22–151-mm standard length). During high tide, F. heteroclitus exclusively used marsh habitats, particularly high marsh, whereas L. rhomboides used marshes and flats. F. heteroclitus abundance followed an interior > nonisolated > isolated pattern. L. rhomboides abundance patterns were less consistent but followed a nonisolated > isolated > interior pattern. A size-dependent water depth relationship was observed for both species and suggests size class partitioning of marsh and flat habitats during high tide. Minimum water depth (~31 cm) restricted L. rhomboides populations in marshes, while maximum water depth (~69 cm) restricted F. heteroclitus population use of marshes and movement between marsh habitats. Disparities in F. heteroclitus young of year contribution between isolated compared to nonisolated and interior marsh types suggests isolated marshes acted as population sinks and were dependent on adult emigrants. Resident and transient salt marsh nekton species utilize estuarine habitats in different ways and these fundamental differences can translate into how estuarine landscape might affect nekton.     

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.     

Distribution and abundance of tidal marshes along the coast of maine

Planimetry studies of coastal geology maps prepared by the Maine Geological Survey show that there is more than an order of magnitude more tidal marsh area in the state of Maine than documented in previously published estimates. The highly convoluted coast of Maine, which is approximately 5,970 km long, contains almost 79 km² of salt marsh, far more than any other New England state, New York, or the Bay of Fundy region. Reasonable estimates for the per-unit primary productivity of salt marshes lead to projections of total marsh productivity on the order of 10¹⁰ g dry weight yr^?1 for the Maine coast and 10¹¹ g dry weight yr^?1 for the Gulf of Maine as a whole. Distribution of tidal marsh area is strongly controlled by coastal geomorphology, which varies considerably along the coast of Maine. The salt marsh area is concentrated in the southwestern coastal region of arcuate bays, where marshes have developed behind sandy beaches. A series of long islands and bedrock peninsulas in the south-central portion of the coast also provides sheltered areas where large marshes occur. Northeast of Penobscot Bay salt marshes become more numerous and smaller in average areal extent. A lack of protection from waves, along with limited sources of glacio-fluvial and glacio-marine sediments, restricts the occurrence of salt marshes in that region to the frignes of coves and tidal rivers.     

Sources and transformations of organic matter in surface soils and sediments from a tidal estuary (North Inlet, South Carolina, USA)

Surface soil and sediment samples collected along a forest-brackish marsh-salt marsh transect in a southeastern U.S. estuary were separated into three different fractions (sand, macro-organic matter, and humus) based on size and density. Elemental, stable carbon isotope, and lignin analyses of these samples reveal important contrasts in the quantity, composition, and sources of organic matter, between forest and marsh sites. Elevated nitrogen contents in humus samples suggest nitrogen incorporation during humification is most extensive in forest soils relative to the marsh sites. The lignin compositions of the macro-organic and humus samples reflect the predominant type of vegetation at each site. Lignin phenol ratios indicate that woody and nonwoody litter from, gymnosperm and angiosperms trees (pines and oaks) is the major source of vascular plant-derived organic matter in the forest site and that angiosperm, grasses (Juncus andSpartina) are the major sources of lignin at the marsh sites. The phenol distributions also reveal that oxidative degradation of lignin is most extensive in the forest and brackish marsh zones whereas little lignin decay occurs in the salt marsh samples. In forest soils, most organic matter originates from highly altered forest vegetation while at the brackish marsh site organic matter is a mixture of degradedJuncus materials and microbial/algal remains. Organic matter in the salt marsh appears to be composed of a more complex mixture of sources, including degradedSpartina detritus as well as algal and microbial inputs. Microbial methane oxidation appears to be an important process and a source of¹³C depleted organic carbon in subsurface sediments at this site.     

AreSpartina marshes a replaceable resource? A functional approach to evaluation of marsh creation efforts

Marsh creation has come into increasing use as a measure to mitigate loss of valuable wetlands. However, few programs have addressed the functional ecological equivalence of man-made marshes and their natural counterparts. This study addresses structural and functional interactions in a man-made and two natural marshes. This was done by integrating substrate characteristics and marsh utilization by organisms of two trophic levels. Sediment properties, infaunal community composition, andFundulus heteroclitus marsh utilization were compared for a man-madeSpartina salt marsh (between ages 1 to 3 yr) in Dills Creek, North Carolina, and adjacent natural marshes to the east and west. East natural marsh and planted marsh sediment grain-size distributions were more similar to each other than to the west natural marsh due to shared drainage systems, but sediment organic content of the planted marsh was much lower than in either natural marsh. This difference was reflected in macrofaunal composition. Natural marsh sediments were inhabited primarily by subsurface, deposit-feeding oligochaetes whereas planted marsh sediments were dominated by the tube-building, surface-deposit feeding polychaetesStreblospio benedicti andManayunkia aestuarina. Infaunal differences were mirrored inFundulus diets. Natural marsh diets contained more detritus and insects, because oligochaetes, though abundant, were relatively inaccessible. Polychaetes and algae were major constituents of the planted marshFundulus diet. Though naturalmarsh fish may acquire a potentially less nutritive, detritus-based diet relative to the higher animal protein diet of the planted marsh fish,Fundulus abundances were markedly lower in the planted marsh than in the natural marshes, indicating fewer fish were being supported. LowerSpartina stem densities in the planted marsh may have provided inadequate protection from predation or insufficient spawning sites for the fundulids. After three years, the planted marsh remained functionally distinct from the adjacent natural marshes. Mitigation success at Dills Creek could have been improved by increasing tidal flushing, thereby enhancing, access to marine organisms and by mulching withSpartina wrack to increase sediment organic-matter content and porosity. Results from this study indicate that salt marshes should not be treated as a replaceable resource in the short term. The extreme spatial and temporal variability inherent to salt marshes make it virtually impossible to exactly replace a marsh by planting one on another site.     

Feeding ecology of o-group sea bass,Dicentrarchus labrax, in salt marshes of Mont Saint Michel Bay (France)

0-group sea bass,Dicentrarchus labrax, colonize intertidal marsh creeks of Mont Saint Michel Bay, France, on spring tides (e.g., 43% of the tides) during flood and return to coastal waters during ebb. Most arrived with empty stomachs (33%), and feed actively during their short stay in the creeks (from 1 to 2 h) where they consumed on average a minimum of 8% of their body weight. During flood tide, diet was dominated by mysids,Neomysis integer, which feed on marsh detritus. During ebb, when young sea bass left tidal marsh creeks, the majority had full stomachs (more than 98%) and diet was dominated by the most abundant marsh (including vegetated tidal flats and associated marsh creeks) resident amphipod,Orchestia gammarellus. Temporal and tidal effects on diet composition were shown to be insignificant. Foraging in vegetated flats occurs very rarely since they are only flooded by about 5% of the tides. It was shown that primary and secondary production of intertidal salt marshes play a fundamental role in the feeding of 0-group sea bass. This suggests that the well known nursery function of estuarine systems, which is usually restricted to subtidal and intertidal flats, ought to be extended to the supratidal, vegetated marshes and mainly to intertidal marsh creeks.     

The Effects of Plant Invasion and Ecosystem Restoration on Energy Flow through Salt Marsh Food Webs

We measured the effects of a plant invasion (Phragmites australis) on resident fish (Fundulus heteroclitus) in New England salt marshes by assessing diet quality at the food web base and by quantifying the importance of primary producers to secondary production using a recently developed Bayesian mixing model (Stable Isotope Analysis in R, “SIAR”). Spartina alterniflora, the dominant native plant, exhibited significantly greater leaf toughness and higher C/N ratios relative to P. australis. Benthic microalgae and phytoplankton (as suspended particulate matter) exhibited the lowest C/N indicating higher diet quality. We conducted a sensitivity analysis in SIAR by modeling F. heteroclitus at three separate trophic levels (1.5, 2.0, and 2.5) using species-specific discrimination factors to determine basal resource contributions. Overall, the best-fitting models include those that assume F. heteroclitus resides approximately 2.0 trophic levels above primary producers. Using discrimination factors from a range of data sources reported in the literature, our analyses revealed that consumers rely less on benthic microalgae and phytoplankton in restricted marshes (7–23 % and 11–44 %, respectively) relative to reference marshes (5–34 % and 23–48 %, respectively), resulting in a shift in diet toward invasive plant consumption (0–27 %). This is likely due to increased P. australis cover and marsh surface shading leading to decreased microalgal biomass, combined with reduced flooding of the marsh surface that favors terrestrial invertebrate assemblages. Restoration decreased the quantity of P. australis in the food web (0–15 %) and increased the importance of microalgae (1–30 %), phytoplankton (19–48 %), and native plants (23–63 %), indicating a shift in ecological recovery toward reference conditions.     

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.     

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.