Distribution and nutrient status of haplotypes of the marsh grassPhragmites australis along the Rappahannock River in Virginia

We compared the distribution and nutrient status of native haplotype F ofPhragmites australis along the freshwater to mesohaline tidal marsh gradient of the Rappahannock River, Virginia, for comparison with the nonnative, invasive haploty M. Using GIS analysis of aerial photography and GPS-based ground truthing, we identified 55 separate clones of native haplotype F comprising a total of 3.68 ha (range 0.002–0.734 ha), all found in tidal wetlands where surface water salinity was 0 psu. We identified 219 separate clones of the invasive haplotype M covering 68.3 ha along the same stretch of river (range 0.004–11.86 ha), found in wetlands where salinity ranged from 0 to 11 psu. From 15 separate clones for each haplotype, average carbon content in leaves of the native was significantly higher than the invasive (43.90±0.08% versus 42.82±0.15%, F_1,28=20.938, p<0.01), and nitrogen content was significantly lower (2.22±0.03% versus 2.58±0.07%, F_1,28=11.972, p<0.01). The average C:N:P ratio for leaf tissue was 1100∶48∶1 for haplotype F and 1084∶56∶1 for haplotype M. Relative to the native, the invasive haplotype forms larger stands distributed throughout a broader estuarine reach and incorporates more nitrogen in leaf tissue. From a management standpoint, nativePhragmites protection should focus on deterring nonnative haplotype invasion through the minimization of both adjacent upland disturbance and nutrient enrichment in tidal freshwater marshes.     

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).     

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

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

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.     

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

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

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.     

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.     

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

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

Association Between <Emphasis Type="BoldItalic">Fusarium</Emphasis> spp. on <Emphasis Type="BoldItalic">Spartina alterniflora</Emphasis> and Dieback Sites in Connecticut and Massachusetts

Sudden vegetation dieback (SVD) is defined as the loss and lack of recovery of smooth cordgrass (Spartina alterniflora) in salt marshes. A new species of a moderately pathogenic fungus called Fusarium palustre is consistently found in SVD sites, but greenhouse tests revealed that it is not capable of causing mortality of healthy plants. Similarly, root-knot nematodes (Meloidogyne spartinae) are also found in SVD sites, but their incidence in marshes affected by SVD is not known. To understand more about the ecology of F. palustre and M. spartinae, salt marshes along Connecticut’s Long Island Sound and Massachusetts’ Cape Cod that exhibited SVD and those that did not, were visited during the summers of 2007, 2008, and 2009. Belowground and aboveground tissues of smooth cordgrass plants from 18 marshes were removed, washed, and assayed for Fusarium spp. to determine if patterns between the incidence of the different species of Fusarium, their virulence on S. alterniflora, root-knot nematodes (M. spartinae), and the health of the marsh could be revealed. There were significantly more colonies of Fusarium growing from plants in SVD sites (6.1%) than in healthy marshes where no SVD was present (<1.0%). The incidence of Fusarium spp. from plants at the perimeter of the SVD site was not statistically different from asymptomatic plants 10–20 m from the SVD edge. The majority of isolates could be assigned to one of two species, F. palustre or another slightly pathogenic group called Fusarium cf. incarnatum (88% in 2007, 62% in 2008, and 96% in 2009). The ratio of F. palustre to F. cf. incarnatum was 6.7, 2.7, or 2.1 for 2007, 2008, or 2009, respectively. Greenhouse tests on healthy S. alterniflora revealed that isolates of F. palustre were more virulent than F. cf. incarnatum, regardless of whether they were recovered from plants in healthy marshes or in SVD sites. Root-knot nematodes were found sporadically and could not be associated with SVD. Factorial greenhouse experiments did not demonstrate any interaction between F. palustre and M. spartinae providing no experimental evidence that combining Fusarium and root-knot nematodes could cause mortality. The presence of Fusarium on S. alterniflora in healthy marshes also suggests an endophytic relationship that may subsequently function in the breakdown of tissue when plants are compromised.     

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.     

<Emphasis Type="Italic">Phragmites australis</Emphasis> (Common Reed) Invasion in the Rhode River Subestuary of the Chesapeake Bay: Disentangling the Effects of Foliar Nutrients,Genetic Diversity,Patch Size,and Seed Viability

The invasion and expansion of the introduced haplotype of Phragmites australis across North America is of growing concern. Previous studies in the Chesapeake Bay region found that Phragmites was more abundant, had higher foliar nitrogen, and produced more viable seeds in brackish wetland subestuaries with more anthropogenic development of the watershed. Here, we focus on a different scale and address issues related to the invasion of Phragmites within a single subestuary, the Rhode River. We evaluated patterns in seed viability, foliar nutrient concentrations, patch size, and genetic variation in ten Phragmites patches in wetlands that occur in the side of the subestuary that is surrounded by forest and 10 patches in wetlands that are in the side of the subestuary that has extensive anthropogenic development. Seed viability varied from 0–60% among the 20 patches but did not differ significantly between patches on the developed vs. forested sides of the Rhode River. Foliar nutrients also did not differ between patches on the two sides of the Rhode River. Seed viability, however, was negatively related to foliar nutrients. Most Phragmites patches consisted of >1 genotype. Larger patches had multiple genotypes, and patches with more genotypes produced more viable seeds. Our study indicates that the Rhode River subestuary behaves as one system with no differences in the measured Phragmites variables between the forested vs. developed sides of the watershed. Our findings also suggest a cyclical process by which Phragmites can spread: larger patches contain more genetic diversity, which increases the chances for cross-fertilization. The subsequent increased production of viable seeds can increase local levels of genetic diversity, which can further facilitate the spread of Phragmites by seed.     

Shifting nutrient limitation and eutrophication effects in marsh vegetation across estuarine salinity gradients

In light of widespread coastal eutrophication, identifying which nutrients limit vegetation and the community consequences when limitation is relaxed is critical to maintaining the health of estuarine marshes. Studies in temperate salt marshes have generally identified nitrogen (N) as the primary limiting nutrient for marsh vegetation, but the limiting nutrient in low salinity tidal marshes is unknown. I use a 3-yr nutrient addition experiment in mid elevation,Spartina patens dominated marshes that vary in salinity along two estuaries in southern Maine to examine variation in nutrient effects. Nutrient limitation shifted across estuarine salinity gradients; salt and brackish marsh vegetation was N limited, while oligohaline marsh vegetation was co-limited by N and phosphorus (P). Plant tissue analysis ofS. patens showed plants in the highest salinity marshes had the greatest percent N, despite N limitation, suggesting that N limitation in salt marshes is partially driven by a high demand for N to aid in salinity tolerance. Fertilization had little effect on species composition in monospecificS. patents stands of salt and brackish marshes, but N+P treatments in species-rich oligohaline marshes significantly altered community composition, favoring dominance by high aboveground producing plants. Eutrophication by both N and P has the potential to greatly reduce the characteristic high diversity of oligohaline marshes. Inputs of both nutrients in coastal watersheds must be managed to protect the diversity and functioning of the full range of estuarine marshes.     

The distribution and utility of sea‐level indicators in Eurasian sub‐Arctic salt marshes (White Sea,Russia)

In support of efforts to reconstruct relative sea level (RSL), we investigated the utility of foraminifera, diatoms and bulk‐sediment geochemistry (δ¹³C, C:N and parameters measured by Rock‐Eval pyrolysis) as sea‐level indicators in Eurasian sub‐Arctic salt marshes. At three salt marshes (<15 km apart) in Dvina Bay (White Sea, Russia), we collected surface sediment samples along transects from subtidal to Taiga forest environments. Foraminifera at all sites formed bipartite assemblages, where elevations below mean high higher water (MHHW) were dominated by Miliammina spp. and elevations between MHHW and the highest occurrence of foraminifera were dominated by Jadammina macrescens and Balticammina pseudomacrescens. Five high‐diversity groups of diatoms were identified and they displayed pronounced variability amongst the study sites. Bulk‐sediment geochemistry recognized two groups (clastic‐dominated environments below MHHW and organic‐rich environments above MHHW). As one group included subtidal elevations and the other included supratidal elevations, we conclude that the measured geochemical parameters are not stand‐alone sea‐level indicators. Core JT2012 captured a regressive sediment succession of clastic, tidal‐flat sediment overlain by salt‐marsh organic silt and freshwater peat. The salt‐marsh sediment accumulated at 2804±52 years before present and preserved foraminifera (Jadammina macrescens and Balticammina pseudomacrescens) with good analogy to modern assemblages indicating that RSL was +2.60±0.47 m at this time. Diatoms confirm that marine influence decreased through time, but the lack of analogy between modern and core assemblages limited their utility as sea‐level indicators. Geochemical parameters also indicate a reduction in marine influence through time. We conclude that RSL reconstructions derived from salt‐marsh sediment preserved beneath Eurasian sub‐Arctic peatlands can provide valuable insight into the spatio‐temporal evolution of the Fennoscandian and Eurasian ice sheets.     

<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.     

Helophyte zonation in two regulated estuarine areas in the Netherlands: Vegetation analysis and relationships with hydrological factors

The relationship between hydrological conditions and riparian helophyte vegetation was studied in two freshwater estuaries that differed in tidal regulation in order to assess the effects of large-scale hydrological regulation on the fringe vegetation. Vegetation and environmental variables were sampled for 206 sites in the Rhine-Meuse estuary (146 sites) and the IJsselmeer region (60 sites) in the Netherlands. These samples were classified into 10 vegetation types, all of which were dominated by tall helophytes. The most common vegetation types were dominated byPhragmites australis andTypha angustifolia, which formed both monospecific stands and mixtures. Tall Cyperaceae dominated three vegetation types (dominated individually bySchoenoplectus lacustris, Bolboschoenus maritimus, andSchoenoplectus tabernaemontani).Acorus calamus. Principal components analysis of the species composition of vegetation fringing open-water areas and associated environmental data revealed complex gradients incorporating differences in water depth, water-level fluctuation, were exposure, and sedimentation and/or erosion. The composition of the helophyte belts varied among the areas as the result of the differing times at which regulation occurred. Based on historical data, hydrological regulation of the estuaries has resulted in deterioration of intertidalSchoenoplectus andBolboschoenus stands, due to erosion and predation.P. australis stands have been invaded by terrestrial plant species or have been replaced byT. angustifolia. A scheme is presented of helophyte vegetation development under the influence of changes in the hydrological regime.     

Indirect effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> in the habitat use of Argentina’s South West Atlantic salt marsh birds

The burrowing crab,Chasmagnathus granulatus, is the dominant benthic macroinvertebrate of southwest Atlantic salt marshes and tidal flats, having strong ecosystem engineering effects by direct and indirect effects on soil, vegetation, invertebrates, fishes, and birds. Vegetation structure is a main component for bird habitat selection, since greater habitat complexity generates higher niche diversity. This environmental complexity can be modified by species interactions or disturbance events (i.e., by crabs), in turn modifying the associated community. The bird species of salt marshes of the southwest Atlantic are highly dependent on these ecosystems. We assessed the effects of the burrowing crab on the structure of the cordgrass,Spartina densiflora, marshes, and how these changes affected the composition and diversity of birds. This study was conducted at the Mar Chiquita Coastal Lagoon, Argentina (37°40′S, 57°23′W).S. densiflora marshes were classified within three categories: marshes without burrowing crabs, marshes with low burrow densities, and marshes with high burrow densities. We made vegetation and bird surveys during spring beforeS. densiflora produce seeds and in summer when seeds are abundant. We also manipulated inflorescence (i.e., spike) density (a variable affected by crabs) to assess the effect on bird habitat use. The highest inflorescence density ofS. densiflora and highest bird diversity were recorded at low burrow densities. Cover and height ofS. densiflora were negatively correlated with burrow density. The number of bird species and the number of red-capped wren-spinetail,Spartonoica maluroides, were negatively related with cover and height ofS. densiflora. Experiments confirmed that granivorous species used areas with higher spike densities, a variable related to crab burrow density. Burrowing crabs directly and indirectly affect the cover, height, spike density, and morphologic characteristic of seeds ofS. densiflora marshes. These effects indirectly affect the bird community that uses these marshes, being negatively related toS. maluroides and total bird abundance, and positively related to bird diversity.     

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.     

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.