A comparative study of the mollusc communities of two oligohaline intertidal marshes: Spatial and temporal distribution of abundance and biomass

Molluscs were collected monthly for a year from two low salinity (0–9‰) intertidal marshes dominated by the macrophytesJuncus roemerianus orSpartina cynosuroides in St. Louis Bay, Mississippi. TheJuncus marsh had lower soil organic matter, higher pH and was more frequently inundated than theSpartina marsh. Eight species of gastropods were abundant and dominated in the higherSpartina marsh, while three bivalve species were dominant in theJuncus marsh. Of the common species,Succinea ovalis, Vertigo ovata andDeroceras laeve are gastropods of terrestrial origins;Geukensia demissa granosissima (bivalve) andMelampus bidentatus (gastropod) are euryhaline estuarine species and the remaining gastropods (Detracia floridana, Littoridinops palustris, Onobops jacksoni) and bivalves (Polymesoda caroliniana, Cyrenoida floridana) are brackish species. Most species were capable of continuous recruitment (based on size class analysis), but exhibited peak activity in particular seasons. Bivalve abundance correlated to temperature, and gastropod abundance was negatively correlated to soil pH. These correlations reflect the influence of flooding regime at the two sites. Biomass was greater in theJuncus marsh because of the increased presence of the large-bodiedPolymesoda. Polymesoda represented >90% and >50% of the total biomass in theJuncus andSpartina (except summer) marshes respectively but always <-5% of the individuals collected. Gastropod biomass was the same in both marshes. Species diversity (H′) was greater inSpartina except for summer months. TheJuncus marsh always exhibited greater species richness. Evenness (J′) determined seasonal changes in diversity (H′). Similarity values (C_z) were always quite low, with highest values in spring In contrast to faunal studies from Gulf and East Coast salt marshes, we found 1) fewer species, 2) communities comprised of unique species combinations, 3) greatest mean densities in summer, and 4) potentially less productivity by the molluscs of our sites. These mollusc communities exhibit structural characteristics that emphasize the unique ecotonal nature of the oligohaline marshes within which they are found.     

The influence of morphology in determining the decomposition of two salt marsh macrophytes

Annual decomposition rates of Spartina alterniflora height forms and Juncus roemerianus were determined in situ in three North Carolina salt marshes using the litter bag method. The decomposition of Spartina was significantly influenced by size, i.e., height form, with the taller plants which had greater amounts of stem tissue, being more resistant to decay. Instantaneous decay rates for short and medium Spartina were not significantly different at any site, but they were both significantly greater than that of the tall form at two of the three study sites. Juncus decomposed more slowly than Spartina during the first 8 months, but had decomposed as completely as all three height forms of Spartina at two of the study sites by the end of the 13-month study period. Constant submergence appeared to inhibit decomposition since there was twice as much undecomposed plant material remaining in bags placed in tidal creeks as in those on the marsh surface.     

Plant growth and productivity along complex gradients in a Pacific northwest brackish intertidal marsh

Environmental characteristics were measured and recorded in the Skagit Marsh, a brackish intertidal marsh on Puget Sound, Washington. Four transects were placed perpendicular to a known gradient of increasing salinity which began with fresh water at the bank of one of the outlets of the Skagit River and reached a surface water salinity of 22‰ at a point alongshore 5 km north of the outlet. The environmental characteristics which were measured varied along gradients (soil texture, organic carbon in fines, soil column temperature, free soil water salinity) or had a patchy distribution (soil redox potential, soil macro-organic matter). Growth and production vary across the marsh. The maximum aboveground standing crop (1,742 g m^?2 dry weight) was measured at a site with 0–4‰ free soil water salinity, dominated by the sedgeCarex lyngbyei. In more saline areas (8–12‰), the bulrushScirpus americanus was dominant and standing crop values dropped to a third of the maximum. Species performance varied in a complex manner as did the environment.C. lyngbyei had diminished growth and decreased standing crop in areas where salinity was higher.S. americanus was equally productive in low elevation, high salinity sites and in high elevation, low salinity sites. An increase in shoot density for dominant species occurred in saline areas as individual shoot weights and leaf areas decreased. Because species responded differently, environmental variation was magnified in the population and community responses of the marsh vegetation.     

Seasonal abundance and diversity of spiders in two intertidal marsh plant communities

The spiders of two Mississippi marsh communities were studied from January 1982 through March 1983. Monthly collections were made in two adjacent marsh plant zones dominated bySpartina cynosuroides (L.) Roth andJuncus roemerianus Scheele respectively. A total of 38 species of spiders (36 inSpartina, 33 inJuncus) representing 13 families were collected. The dominant species in theSpartina zone includedPirata mayaca Gertsch,Lycosa watsoni Gertsch (Lycosidae),Clubiona saltitans Emerton,Scotinella formica (Banks) (Clubionidae),Floricormus sp. (Linyphiidae),Dictyna sylvania Chamberlin & Ivie (Dictynidae),Paramaevia hobbsae (Barnes) (Salticidae), andAgelenopsis barrowsi Gertsch (Agelenidae). The dominant species in theJuncus zone includedLycosa watsoni, Pirata mayaca, Clubiona saltitans andSarinda hentzi (Banks) (Salticidae). Density, biomass, species richness and equitability peaked in May in theJuncus zone and in June in theSpartina zone. Peak levels of density and biomass corresponded to the reproductive activity of the common species, while diversity patterns were attributable to the reproductive activity of the less common species. Mean values of density and biomass over the study period were 84.8 spiders per m² and 155.6 mg per m² in theSpartina zone and 39.4 spiders per m² and 133.0 mg per m² inJuncus zone. The Juncus zone was flooded more frequently, contained less litter, and supported lower overall density and diversity of spiders.     

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.     

Identification of important primary producers in a Chesapeake Bay tidal creek system using stable isotopes of carbon and sulfur

The use of multiple stable isotopes in the study of trophic relationships in temperate estuaries has usually been limited to euhaline systems, in which phytoplankton, benthic microalgae, andSpartina alterniflora are major sources of organic matter for consumers. Within large estuaries such as Chesapeake Bay, however, many species of consumers are found in the upper mesohaline to oligohaline portions. These lower salinity wetlands have a greater abundance of macrophytes that use C3 photosynthesis to fix carbon, in addition toS. alterniflora, which fixes carbon via the C4 photosynthetic pathway. In a broad survey of the biota and sediments of a brackish tidal creek tributary to Chesapeake Bay, combined δ¹³C and δ³⁴S measurements disclosed a balanced contribution to secondary production from phytoplankton, C3 macrophytes,Spartina sp., and benthic microalgae. Surface sediment δ¹³C suggested that the organic matter from C3 plants was derived both from allochthonous sources (terrestrial runoff) and from autochthonous production (marsh macrophytes). Unlike most estuarine systems studied to date, which are dominated by algae (phytoplankton and benthic microalgae) and C4 macrophytes, C3 plants are of greater importance in the diets of consumers in this low-salinity creek system.     

Exchange of microbial biomass between aSpartina alterniflora marsh and the adjacent tidal creek

Salt marsh systems of the southeastern United States are characterized by extensive stands ofSpartina alterniflora. These marshes may influence the concentrations of material suspended in flooding and receding tidal waters. The ability of aSpartina alterniflora-dominated marsh to influence the concentration of suspended microbial biomass was investigated through the use of a 142-m long flume. The flume extended through stands of tall-, medium-, and short-heightSpartina. Water passing through the tallSpartina lost a considerable portion of microbial biomass. Initial samples from medium-heightSpartina were collected from water that had already passed through the tall grass. These samples contained 20 to 70% less microbial biomass than did water entering the tallSpartina. Calculations of mass transport suggest that the tallSpartina zone of marsh acts as a sink for microbial biomass while the short-heightSpartina tends to export biomass (to the tallSpartina zone). The marsh as a whole acts as a sink for microbial biomass. Transport estimates from 32 individual tide cycles were modeled to obtain an annual estimate of transport. As a consequence of high variability among individual transport estimates, no annual transport estimate could be distingushed from a net-zero transport.     

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.     

Methane release from a brackish intertidal salt-marsh embayment of Chesapeake Bay,Maryland

Gaseous methane loss from a brackish, intertidal salt marsh sediment was measured in April, June, August, and October 1977. Twenty-four sediment cores were taken on each date. Annual loss of methane carbon from the mud flats was 10.7 g CH₄?C per m² per year, a value closer to freshwater values than marine systems. Loss of methane fromSpartina peat was low.     

Spatial and temporal variations in terrestrially-derived organic matter from sediments of the Delaware estuary

Terrestrially-derived organic matter in sediments of the Delaware Estuary originates from riverine transport of soils and fresh litter, sewage and industrial wastes, and marsh export of organic matter. The quantity, composition, and spatial distribution of terrigenous organic matter in sediments was determined by elemental (C and N), lignin, and stable carbon isotope analyses. Sediments in the upper Delaware Estuary had low organic carbon content and high lignin content. In contrast, sediments in the lower Delaware Estuary had high organic carbon content and low lignin content. There was a slight decrease in the proportion of syringyl and cinnamyl phenols relative to vanillyl phenols between the upper estuary and lower estuary. Differences in lignin and stable carbon isotope compositions between sediments of the Delaware Estuary and sediments of the Broadkill River estuary (an adjoining salt-marsh estuary) supported previous observations that marshes do not export substantial quantities of organic matter to estuaries. Additional results suggested that lignin-rich sediments were concentrated in the upper estuary, most likely in the zone of high turbidity. Furthermore, algal material diluted lignin-rich sediments, particularly in the lower estuary. The weaker algal signal in bottom sediments compared to that in suspended particulate matter suggested algal material was decomposed either in the water column or at the sediment-water interface. Physical sorting of sediments prior to deposition was also indicated by observations of compositional differences between the upper and lower estuary bottom sediments. Finally, seasonal variations in primary productivity strongly influenced the relative abundance of terrestrial organic matter. In fall, however, the proportion of lignin was greatest because of a combination of greater inputs of terrestrially-derived organic matter, lower river discharge, and a decrease in algal biomass.     

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.     

Analysis of tidal marsh vegetation patterns in two Georgia estuaries using aerial photography and GIS

Aerial photographs and GIS analysis were used to map the distribution of tidal marsh vegetation along the salinity gradients of the estuaries of the Altamaha and Satilla Rivers in coastal Georgia. Vegetation maps were constructed from 1993 U.S. Geological Survey Digital Orthophoto Quarter Quads, 1∶77,000-scale color infrared photographs taken in 1974 and 1∶24,000-scale black and white photographs taken in 1953, Changes between years were identified using a GIS overlay analysis. Four vegetation classifications were identified and groundtruthed with field surveys: salt marsh (areas containing primarilySpartina alterniflora), brackish marsh (Spartina cynosuroides andS. alterniflora), Juncus (Juncus roemerianus), and fresh marsh (Zizania aquatica, Zizaniopsis miliacae, and others). There was no evidence for an upstream shift in marsh vegetation along the longitudinal axis of either estuary over the time frame of this analysis, which implies there has not been a long-term increase in salinity. Although the inland extent of each marsh zone was further upstream in the Satilla than the Altamaha, they corresponded to similar average high tide salinities in each estuary: areas classified as salt marsh occurred from the mouth up to where average high tide salinity in the water was approximately 15 psu;Juncus ranged from 21 to 1 psu; brackish marsh ranged from 15 to 1 psu; and fresh marsh was upstream of 1 psu. Approximately 63% of the 6,786 ha of tidal marsh vegetation mapped in the Altamaha and 75% of the 10,220 ha mapped in the Satilla remained the same in all 3 yr.Juncus was the dominant classification in the intermediate regions of both estuaries, and shifts between areas classified asJuncus and either brackish or salt marsh constituted the primary vegetation change between 1953 and 1993 (87% of the changes observed in the Altamaha and 95% of those in the Satilla). This analysis suggests that the broad distribution of tidal marsh vegetation along these two estuaries is driven by salinity, but that at the local scale these are dynamic systems with a larger number of factors affecting the frequently changing borders of vegetation patches.     

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.     

Effects of Small-Scale Armoring and Residential Development on the Salt Marsh-Upland Ecotone

Small-scale armoring placed near the marsh-upland interface to protect single-family homes is widespread but understudied. Using a nested, spatially blocked sampling design on the coast of Georgia, USA, we compared the biota and environmental characteristics of 60 marshes adjacent to either a bulkhead, a residential backyard with no armoring, or an intact forest. We found that marshes adjacent to bulkheads were at lower tidal elevations and had features typical of lower elevation marsh habitats: high coverage of the marsh grass Spartina alterniflora, high density of crab burrows, and muddy sediments. Marshes adjacent to unarmored residential sites had higher soil water content and lower porewater salinities than the armored or forested sites, suggesting that there may be increased freshwater input to the marsh at these sites. Deposition of Spartina wrack on the marsh-upland ecotone was negatively related to elevation at armored sites and positively related at unarmored residential and forested sites. Armored and unarmored residential sites had reduced densities of the high marsh crab Armases cinereum, a species that moves readily across the ecotone at forested sites, using both upland and high marsh habitats. Distance from the upland to the nearest creek was longest at forested sites. The effects observed here were subtle, perhaps because of the small-scale, scattered nature of development. Continued installation of bulkheads in the southeast could lead to greater impacts such as those reported in more densely armored areas like the northeastern USA. Moreover, bulkheads provide a barrier to inland marsh migration in the face of sea level rise. Retaining some forest vegetation at the marsh-upland interface and discouraging armoring except in cases of demonstrated need could minimize these impacts.     

Composition and sources of extractable organic matter from a sediment core in Lake Kivu,East African rift valley

Lake Kivu is a gas-charged East African rift lake with currently anoxic bottom water. The extractable compounds and residual organic matter of a short sediment core have δ¹³C values typical of lacustrine microbial detritus. The total extracts consist primarily of polar compounds such as n-alkanoic acids, hydroxyalkanoic acids, triterpenoids, steroids and monosaccharides, with minor amounts of n-alkanes and n-alkanols. These tracer compounds and δ¹³C values indicate that the organic matter in the surficial and deeper sedimentary record was dominated by bacterial sources. The sapropelic sediment between these horizons contains organic matter from primarily algal with lesser bacterial input. Terrestrial organic markers are minor in all samples. The major fractions of the compounds in the total extracts were oxidized in the upper water column prior to transit through the anoxic bottom water to sedimentary deposition. The sapropelic horizon may reflect lake water turnover with ventilation or hydrothermal activity and consequently increased algal blooms.     

Rates, patterns, and impacts ofPhragmites australis expansion and effects of experimentalPhragmites control on vegetation, macroinvertebrates, and fish within tidelands of the lower Connecticut River

Phragmites expansion rates (linear at 1–3% yr⁻¹) and impacts of this expansion on high marsh macroinvertebrates, aboveground production, and litter decomposition fromPhragmites and other marsh graminoids were studied along a polyhaline to oligohaline gradient. These parameters, and fish use of creeks and high marsh, were also studied inPhragmites control sites (herbicide, mowing, and combined herbicide/mow treatments).Phragmites clones established without obvious site preferences on oligohaline marshes, expanding radially. At higher salinities,Phragmites preferentially colonized creekbank levees and disturbed upland borders, then expanded into the central marsh. Hydroperiods, but not salinities or water table, distinguishedPhragmites-dominated transects. Pooled samples ofPhragmites leaves, stems, and flowers decompose more slowly than other marsh angiosperms;Phragmites leaves alone decompose as or more rapidly than those of cattail. AbovegroundPhragmites production was 1,300 to 2,400 g m⁻² (about 23% of this as leaves), versus 600–800 g m⁻² for polyhaline to mesohaline meadow and 1,300 g m⁻² for oligohaline cattail-sedge marsh. Macroinvertebrates appear largely unaffected byPhragmites expansion or control efforts; distribution and densities are unrelated to elevation or hydroperiod, but densities are positively related to litter cover. Dominant fish captured leaving flooded marsh wereFundulus heteroclitus andAnguilla rostrata; both preyed heavily on marsh macroinvertebrates.A. rostrata andMorone americana tended to be more common inPhragmites, but otherwise there were no major differences in use patterns betweenPhragmites and brackish meadow vegetation. SAV and macroalgal cover were markedly lower within aPhragmites-dominated creek versus one withSpartina-dominated banks. The same fish species assemblage was trapped in both plus a third within the herbicide/mow treatment. Fish biomass was greatest from theSpartina creek and lowest from thePhragmites creek, reflecting abundances ofF. heteroclitus. Mowing depressedPhragmites aboveground production and increased stem density, but was ineffective for control.Phragmites, Spartina patens, andJuncus gerardii frequencies after herbicide-only treatment were 0.53-0.21; total live cover was <8% with a heavy litter and dense standing dead stems. After two growing seasonsAgrostis stolonifera/S. patens/J. gerardii brackish meadow characterized most of the herbicide/mow treatment area;Phragmites frequency here was 0.53, contributing 3% cover. Both values more than doubled after four years; a single treatment is ineffective for long-termPhragmites control.     

Impact of Fertilization on a Salt Marsh Food Web in Georgia

We examined the response of a salt marsh food web to increases in nutrients at 19 coastal sites in Georgia. Fertilization increased the nitrogen content of the two dominant plants, Spartina alterniflora and Juncus roemerianus, indicating that added nutrients were available to and taken up by both species. Fertilization increased Spartina cover, height, and biomass and Juncus height, but led to decreases in Juncus cover and biomass. Fertilization increased abundances of herbivores (grasshoppers) and herbivore damage, but had little effect on decomposers (fungi), and no effect on detritivores (snails). In the laboratory, herbivores and detritivores did not show a feeding preference for fertilized versus control plants of either species, nor did detritivores grow more rapidly on fertilized versus control plants, suggesting that changes in herbivore abundance in the field were driven more by plant size or appearance than by plant nutritional quality. Community patterns in control plots varied predictably among sites (i.e., 17 of 20 regression models examining variation in biological variables across sites were significant), but variation in the effects of fertilization across sites could not be easily predicted (i.e., only 6 of 20 models were significant). Natural variation among sites was typically similar or greater than impacts of fertilization when both were assessed using the coefficient of variation. Overall, these results suggest that eutrophication of salt marshes is likely to have stronger impacts on plants and herbivores than on decomposers and detritivores, and that impacts at any particular site might be hard to distinguish from natural variation among sites.     

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

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

A compound-specific n-alkane δC and δD approach for assessing source and delivery processes of terrestrial organic matter within a forested watershed in northern Japan

We measured molecular distributions and compound-specific hydrogen (δD) and stable carbon isotopic ratios (δ¹³C) of mid- and long-chain n-alkanes in forest soils, wetland peats and lake sediments within the Dorokawa watershed, Hokkaido, Japan, to better understand sources and processes associate with delivery of terrestrial organic matter into the lake sediments. δ¹³C values of odd carbon numbered C₂₃-C₃₃n-alkanes ranged from −37.2‰ to −31.5‰, while δD values of these alkanes showed a large degree of variability that ranged from −244‰ to −180‰. Molecular distributions in combination with stable carbon isotopic compositions indicate a large contribution of C3 trees as the main source of n-alkanes in forested soils whereas n-alkanes in wetland soil are exclusively derived from marsh grass and/or moss. We found that the n-alkane δD values are much higher in forest soils than wetland peat. The higher δD values in forest samples could be explained by the enrichment of deuterium in leaf and soil waters due to increased evapotranspiration in the forest or differences in physiology of source plants between wetland and forest. A δ¹³C vs. δD diagram of n-alkanes among forest, wetland and lake samples showed that C₂₅-C₃₁n-alkanes deposited in lake sediments are mainly derived from tree leaves due to the preferential transport of the forest soil organic matter over the wetland or an increased contribution of atmospheric input of tree leaf wax in the offshore sites. This study demonstrates that compound-specific δD analysis provides a useful approach for better understanding source and transport of terrestrial biomarkers in a C3 plant-dominated catchment.