<Emphasis Type="Italic">Spartina alterniflora</Emphasis> Biomass Allocation and Temperature: Implications for Salt Marsh Persistence with Sea-Level Rise

To predict the impacts of climate change, a better understanding is needed of the foundation species that build and maintain biogenic ecosystems. Spartina alterniflora Loisel (smooth cordgrass) is the dominant salt marsh-building plant along the US Atlantic coast. It maintains salt marsh elevation relative to sea level by the accumulation of aboveground biomass, which promotes sediment deposition and belowground biomass, which accretes as peat. Peat accumulation is particularly important in elevation maintenance at high latitudes where sediment supply tends to be limited. Latitudinal variation in S. alterniflora growth was quantified in eight salt marshes from Massachusetts to South Carolina. The hypothesis that allocation to aboveground and belowground biomass is phenotypically plastic was tested with transplant experiments among a subset of salt marshes along this gradient. Reciprocal transplants revealed that northern S. alterniflora decreased allocation to belowground biomass when grown in the south. Some northern plants also died when moved south, suggesting that northern S. alterniflora may be stressed by future warming. Southern plants that were moved north showed phenotypic plasticity in biomass allocation, but no mortality. Belowground biomass also decomposed more quickly in southern marshes. Our results suggest that warming will lead northern S. alterniflora to decrease belowground allocation and that belowground biomass will decompose more quickly, thus decreasing peat accumulation. Gradual temperature increases may allow for adaptation and acclimation, but our results suggest that warming will lower the ability of salt marshes to withstand sea-level rise.     

Stem Borer Frequency and Composition in Healthy <Emphasis Type="Italic">Spartina alterniflora</Emphasis> (smooth cordgrass) and Dieback Zones in a Southern Atlantic Coast Salt Marsh

Unprecedented Spartina alterniflora (smooth cordgrass) dieback has recently plagued Atlantic and Gulf coast US salt marshes. The hypothesized drivers of dieback vary geographically but include drought and herbivory. Stem-boring insect larvae may also contribute to dieback, but have thus far been overlooked. Here we describe stem borer frequency and composition among healthy S. alterniflora morphs (tall and short) and dieback zones in Dean Creek Marsh on Sapelo Island, GA. Overall stem borer frequency was highest in tall morph S. alterniflora and dieback zones (present in 46.7% and 39.5%, respectively). Healthy habitat types were characterized by six stem borer families with only two of those families observed in dieback zones. Taxa from these two families, previously reported by others to cause stem death, were found at their highest frequency in dieback zones. Although we cannot infer causation, our results raise the possibility of stem borers contributing to the formation of dieback zones, meriting further research.     

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.     

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.     

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.     

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.     

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 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.     

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.     

Factors contributing to the removal of a marine grass invader (<Emphasis Type="Italic">Spartina anglica</Emphasis>) and subsequent potential for habitat restoration

Our goal is to understand how removal regime and habitat type interact to influence removal success of a marine plant invader and the subsequent potential for restoration. In particular, we investigate the management program designed to eradicate the English cordgrass,Spartina anglica C. E. Hubbard, in marine intertidal habitats of Puget Sound, Washington, United States. Observational and manipulative experiments were used to measure the regrowth (vegetative growth), reinvasion (seedling recruitment), and restoration potential (return to native condition) of invaded habitats. Removal regime (consistent: yearly removal; interrupted: yearly removal with the last year missed) and habitat type (low salinity marsh, mudflat, cobble beach, and high salinity marsh sites) were considered. The response to removal regime was dramatic. Under consistent removal, cordgrass slowly declined but under interrupted removal, there was substantial regrowth of the invader. This pattern results from the resiliency of belowground biomass and the subsequent high aboveground productivity and seedling growth ofS. anglica. We also found that removal success depended on differences among sites that represent different habitat types. Cordgrass regrowth and reinvasion were substantially higher in the low salinity marsh sites where soils have lower salinity. We also found that at the low salinity marsh sites, some restoration of native plants and soil conditions was evident. At mudflat, cobble beach, and high salinity marsh sites, colonization of native vascular plants and algae not normally present, in the absence of the invasion, occurred. Whether these habitats will eventually revert back to the pre-invasion conditions over a longer period of time is unknown.     

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.     

Short- and Long-Term Vegetative Propagation of Two <Emphasis Type="Italic">Spartina</Emphasis> Species on a Salt Marsh in Southern Brazil

Spartina alterniflora and Spartina densiflora are native salt marsh plants from the Atlantic coast; their habitats in Patos Lagoon estuary (southern Brazil) are characterized by a microtidal regime (<0.5 m) and, during El Niño events, high estuarine water levels and prolonged flooding due to elevated freshwater discharge from a 200,000-km² watershed. During and between El Niño events, the vegetative propagation of these two Spartina species in the largest estuary of southern Brazil (Patos Lagoon) was evaluated by monitoring transplanted plants for 10 years (short-term study) and interpreting aerial photos of natural stands for 56 years (long-term study). During the short-term study, S. alterniflora quickly occupied mud flats (up to 208 cm year^?1) by elongation of rhizomes, whereas S. densiflora showed a modest lateral spread (up to 13 cm year^?1) and generated dense circular-shaped stands. However, moderate and strong El Niño events can promote excessive flooding and positive anomalies in the estuarine water level that reduce the lateral spread and competitive ability of S. densiflora. During the long-term study, natural stands of S. alterniflora and S. densiflora had steady lateral spread rates of 152 and 5.2 cm year^?1, respectively, over mud flats. In the microtidal marshes of the southwest Atlantic, the continuous long-term lateral expansion of both Spartina species embodies periods of intense flooding stress (moderate and strong El Niños), when there is a decrease of vegetative propagation and less stressful low water periods of fast spread over mud flats (non-El Niño periods and weak intensity El Niños).     

Predation by the black-clawed mud crab,<Emphasis Type="Italic">Panopeus herbstii</Emphasis>, in Mid-Atlantic salt marshes: Further evidence for top-down control of marsh grass production

Although top-down control of plant growth has been shown in a variety of marine systems, it is widely thought to be unimportant in salt marshes. Recent caging experiments in Virginia and Georgia have challenged this notion and shown that the dominant marsh grazer (the periwinkle,Littoraria irrorata) not only suppresses plant growth, but can denude marsh substrate at high densities. In these same marshes, our field observations suggest that the black-clawed mud crab,Panopeus herbstii, is an abundant and potentially important top-down determinant of periwinkle density. No studies have quantitatively examinedPanopeus distribution or trophic interactions in marsh systems, and its potential impacts on community structure remained unexplored. We investigated distribution and feeding habits ofPanopeus in eight salt marshes along the Mid-Atlantic seashore (Delaware-North Carolina). We found that mud crabs were abundant in tall (4–82 ind m^?2), intermediate (0–15 ind m^?2), and short-form (0–5 ind m^?2)Spartina alterniflora zones in all marshes and that crab densities were negatively correlated with tidal height and positively correlated with bivalve density. Excavation of crab lairs r?utinely produced shells of plant-grazing snails (up to 36 lair^?1) and bivalves. Lab experiments confirmed that mud crabs readily consume these abundant marsh molluscs. To experimentally examine potential community effects of observed predation patterns, we manipulated crab and periwinkle densities in a 1-mo field experiment. Results showed thatPanopeus can suppress gastropod abundance and that predation rates increase with increasing snail density. In turn, crab suppression of snail density reduces grazing intensity on salt marsh cordgrass, suggesting presence of a trophic cascade. These results indicate that this previously under-appreciated consumer is an important and indirect determinant of community structure and contribute to a growing body of evidence challenging the long-standing notion that consumers play a minor role in regulating marsh plant growth.     

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.     

<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.     

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

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

Effects of the Herbicide Diuron on Cordgrass (<Emphasis Type="Italic">Spartina foliosa</Emphasis>) Reflectance and Photosynthetic Parameters

Early indicators of salt marsh plant stress are needed to detect stress before it is manifested as changes in biomass and coverage. We explored a variety of leaf-level spectral reflectance and fluorescence variables as indicators of stress in response to the herbicide diuron. Diuron, a Photosystem II inhibitor, is heavily used in areas adjacent to estuaries, but its ecological effects are just beginning to be recognized. In a greenhouse experiment, we exposed Spartina foliosa, the native cordgrass in California salt marshes, to two levels of diuron. After plant exposure to diuron for 28 days, all spectral reflectance indices and virtually all fluorescence parameters indicated reduced pigment and photosynthetic function, verified as reduced CO₂ assimilation. Diuron exposure was not evident, however, in plant morphometry, indicating that reflectance and fluorescence were effective indicators of sub-lethal diuron exposure. Several indices (spectral reflectance index ARI and fluorescence parameters EQY, Fo, and maximum rETR) were sensitive to diuron concentration. In field trials, most of the indices as well as biomass, % cover, and canopy height varied predictably and significantly across a pesticide gradient. In the field, ARI and Fo regressed most significantly and strongly with pesticide levels. The responses of ARI and Fo in both the laboratory and the field make these indices promising as sensitive, rapid, non-destructive indicators of responses of S. foliosa to herbicides in the field. These techniques are employed in remote sensing and could potentially provide a link between landscapes of stressed vegetation and the causative stressor(s), which is crucial for effective regulation of pollution.     

Growth of the marsh elder<Emphasis Type="Italic">Iva frutescens</Emphasis> in relation to duration of tidal flooding

Iva frutescens is a common shrub at the upland fringe of salt marshes throughout the East and Gulf Coasts of North America. Within a marsh, its location and relative size are governed largely by the degree of flooding by seawater.Iva’s wide distribution and restricted location within salt marshes may make it a useful indicator of overall conditions of the marshes. This work was designed to provide basic information on the age and growth ofI. frutescens, especially as they relate to the degree of flooding that is needed in order to investigateIva’s potential as an indicator. Cross-sections of older stems (living and standing dead) from salt marshes in Rhode Island, United States, were examined in order to age stems and estimate their growth rate from cumulative increase in woody tissue. Most stems were six yr old or less, suggesting that aboveground structures live for only a few years. Stem diameter correlated with growth rate and aboveground biomass. Elevation at the root zone was used to estimate the duration that plants were flooded, which was negatively correlated with stem diameter. The most robust plants came from sites that were flooded only up to 6–7% of the total time during the growing season. No plants were found in areas flooded more than 30% of the time.