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.     

Evaluating salt marsh restoration in Delaware Bay: The response of blue crabs,<Emphasis Type="Italic">Callinectes sapidus</Emphasis>, at former salt hay farms

Marshes are important habitats for various life history stages of many fish and invertebrates. Much effort has been directed at restoring marshes, yet it is not clear how fish and invertebrates have responded to marsh restoration. The blue crab,Callinectes sapidus, uses marsh habitats during much of its benthic life. We investigated the response of blue crabs to marsh restoration by comparing crab abundance (catch per unit effort), mean size and size frequency distribution, sex ratio, and molt stages of crabs in recently restored marshes that were former salt hay farms to that of adjacent reference marshes with similar physical characteristics in the mesohaline portion of Delaware Bay. Field sampling occurred monthly (April–November) in 1997 and 1998 using replicate daytime otter trawls in large marsh creeks and weirs in smaller intertidal marsh creeks. Blue crabs were either equal or more abundant, the incidence of molting was in most months similar, and population sex ratios were indistinguishable in restored and reference marshes, suggesting that the restored areas attract crabs and support their growth. Site location had a greater effect on the sex ratio of crabs such that marshes closer to the mouth of the bay contained a higher percentage of adult female crabs. In each annual growing season (April–July), the monthly increase in crab size and, in some months (June–July), the incidence of molting at the restored sites was greater than the reference sites, suggesting that the restored sites may provide areas for enhanced growth of crabs. These results suggest that blue crabs have responded positively to restoration of former salt hay farms in the mesohaline portion of Delaware Bay.     

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.     

Utilization of <Emphasis Type="Italic">Spartina</Emphasis>- and <Emphasis Type="Italic">Phragmites</Emphasis>-Derived Dissolved Organic Matter by Bacteria and Ribbed Mussels (<Emphasis Type="Italic">Geukensia demissa</Emphasis>) from Delaware Bay Salt Marshes

Phragmites australis has been invading Spartina-alterniflora-dominated salt marshes throughout the mid-Atlantic. Although, Phragmites has high rates of primary production, it is not known whether this species supports lower trophic levels of a marsh food web in the same manner as Spartina. Using several related photochemical and biological assays, we compared patterns of organic matter flow of plant primary production through a key salt marsh metazoan, the ribbed mussel (Geukensia demissa), using a bacterial intermediate. Dissolved organic matter (DOM) was derived from plants collected from a Delaware Bay salt marsh and grown in the laboratory with ¹⁴C-CO₂. Bacterial utilization of plant-derived DOM measured as carbon mineralization revealed that both species provided bioavailable DOM to native salt marsh bacteria. Total carbon mineralization after 19 days was higher for Spartina treatments (36% ¹⁴CO₂ ± 3 SE) compared with Phragmites treatments (29% ±2 SE; Wilcoxon–Kruskal–Wallis rank sums test, P < 0.01). Pre-exposing DOM to natural sunlight only enhanced or decreased bioavailability of the DOM to the bacterioplankton during initial measurements (e.g., 7 days or less) but these differences were not significant over the course of the incubations. Mixtures of ¹⁴C-labeled bacterioplankton (and possibly organic flocs) from ¹⁴C-DOM treatments were cleared by G. demissa at similar rates between Spartina and Phragmites treatments. Moreover, ¹⁴C assimilation efficiencies for material ingested by mussels were high for both plant sources ranging from 74% to 90% and not significantly different between plant sources. Sunlight exposure did not affect the nutritional value of the bacterioplankton DOM assemblage for mussels. There are many possible trophic and habitat differences between Spartina- and Phragmites-dominated marshes that could affect G. demissa but the fate of vascular plant dissolved organic carbon in the DOM to bacterioplankton to mussel trophic pathway appears comparable between these marsh types.     

<Emphasis Type="Italic">Deepwater Horizon</Emphasis> Oil Spill Impacts on Salt Marsh Fiddler Crabs (<Emphasis Type="Italic">Uca</Emphasis> spp.)

The Deepwater Horizon oil spill was the largest marine oil spill in US waters to date and one of the largest worldwide. Impacts of this spill on salt marsh vegetation have been well documented, although impacts on marsh macroinvertebrates have received less attention. To examine impacts of the oil spill on an important marsh invertebrate and ecosystem engineer, we conducted a meta-analysis on fiddler crabs (Uca spp.) using published sources and newly available Natural Resources Damage Assessment (NRDA) and Gulf of Mexico Research Initiative (GoMRI) data. Fiddler crabs influence marsh ecosystem structure and function through their burrowing and feeding activities and are key prey for a number of marsh and estuarine predators. We tested the hypothesis that the spill affected fiddler crab burrow density (crab abundance), burrow diameter (crab size), and crab species composition. Averaged across multiple studies, sites, and years, our synthesis revealed a negative effect of oiling on all three metrics. Burrow densities were reduced by 39 % in oiled sites, with impacts and incomplete recovery observed over 2010–2014. Burrow diameters were reduced from 2010 to 2011, but appeared to have recovered by 2012. Fiddler crab species composition was altered through at least 2013 and only returned to reference conditions where marsh vegetation recovered, via restoration planting in one case. Given the spatial and temporal extent of data analyzed, this synthesis provides compelling evidence that the Deepwater Horizon spill suppressed populations of fiddler crabs in oiled marshes, likely affecting other ecosystem attributes, including marsh productivity, marsh soil characteristics, and associated predators.     

Valuation of continuous and intermittent<Emphasis Type="Italic">Phragmites</Emphasis> control

Quantifying the relative value ofPhragmites australis andSpartina alterniflora habitat is important to evaluate the benefits and risks of different attempts to addressPhragmites expansion on the U.S. eastern seaboard. Two contrasting approaches commonly used to restore tidal marsh habitats invaded byPhragmites communities involve sprayingPhragmites with herbicide only when its coverage of a particular marsh area is near or close to 100%. Alternatively, after the first application, herbicide is annually applied on any surviving patches ofPhragmites present in a mosaic of other marsh vegetation. A model is introduced to evaluate the relative habitat value of these control regimes, here termed the Intermittent and Continuous. Compared to the Intermittent approach, the area of herbicide application in the Continuous approach is higher in the first 6 yr, but lower the reafter. The cumulative gain in habitat quality after 20 yr in either approach is sensitive to the presumed relative values ofPhragmites versusSpartina habitat, and may even be negative if they are nearly equal. Annual applications of herbicide to patches ofPhragmites appears to generate more habitat value and with less herbicide than occasional applications whenPhragmites cover is at is maximum.     

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.     

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.     

Reproductive success of wading birds using<Emphasis Type="Italic">Phragmites</Emphasis> marsh and upland nesting habitats

Colonial nesting of long-legged wading birds (Ciconiiformes) in the coastal northeastern U.S. is limited primarily to islands, which provide isolated habitats that are relatively free of ground predators. Estuarine wetlands in this heavily developed region, including foraging wetlands and fringe marshes surrounding nesting islands, are often dominated byPhragmites australis. On Pea Patch Island in Delaware Bay, site of one of the largest and most enduring mixed-species heron colonies on the East Coast, wading birds nest inPhragmites marsh habitat as well as in adjacent upland shrubs and trees. BecausePhragmites is aggressively managed in Delaware Bay, we investigated the relative habitat value of marsh and upland nesting sites for the purpose of developing recommendations for marsh and wildlife management. Utilization of marsh habitat by nesting birds ranged from 27–82% during 1993–1998. Two species (great blue heronArdea herodias and great egretA. alba) never nested inPhragmites, four species (little blue heronEgretta caerulea, snowy egretE. thula, cattle egretBubulcus ibis, and black-crowned night-heronNycticorax nycticorax) nested in approximately equal proportions in both habitats, and one species (glossy ibisPlegadis falcinellus) was largely confined to marsh nesting. Productivity (egg and nestling production) varied between habitats for some species. Cattle egrets produced larger clutches and had higher hatching rates inPhragmites compared to upland habitat. Little blue herons were more successful in the uplands. Managers should retainPhragmites marsh at colony sites, such as Pea Patch Island, where it provides critical habitat for nesting wading birds both as substrate for nesting and buffer habitat to control human disturbance.     

Comparison of fish and macroinvertebrate use of<Emphasis Type="Italic">Typha angustifolia,Phragmites australis</Emphasis>, and treated<Emphasis Type="Italic">Phragmites</Emphasis> marshes along the lower Connecticut River

Since 1965 large areas of lower Connecticut River tidelands have been converted from high diversity brackish meadow andTypha angustifolia marsh to near monocultures ofPhragmites australis. This study addresses the impact ofPhragmites invasion on fish and crustacean use of oligohaline high marsh. During spring tides from early June through early September 2000, fishes and crustaceans leaving flooded marsh along 3 km of the Lieutenant River, a lower Connecticut River tributary, were captured with Breder traps at 90 sites, equally distributed amongPhragmites, Typha, and treated (herbicide and mowing)Phragmites areas. Pit traps, 18 per vegetation type in 2000 and 30 each inPhragmites andTypha in 2001, caught larvae and juveniles at distances of up to 30 m into the marsh interior. There were no significant differences in fish species compositions or abundances among the vegetation types. Size distributions, size specific biomasses, and diets ofFundulus heteroclitus, the numerically dominant fish, were also similar. The shrimpPalaemonetes pugio was more abundant inPhragmites than in other types of vegetation, whereas the fiddler crabUca minax was least numerous inPhragmites. Mean numbers ofF. heteroclitus andP. pugio caught per site event were positively correlated with increasing site hydroperiod. Significantly moreF. heteroclitus were captured along the upper reach of the river where marsh elevations were lower than farther downstream. MoreF. heteroclitus and fewerP. pugio andU. minax were captured during the day than at night. A relatively small number of larval and juvenileFundulus sp. were captured in pit traps, but consistently fewer inPhragmites than inTypha, suggesting thatTypha and brackish meadow marshes may provide better nursery habitat. Vegetation was sampled along a 30 m transect at each trap site in 2000. Plant species diversity was greatest in treatedPhragmites areas and lowest inPhragmites sites.     

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.     

The influence of introduced European green crabs (<Emphasis Type="Italic">Carcinus maenas</Emphasis>) on habitat selection by juvenile native blue crabs (<Emphasis Type="Italic">Portunus pelagicus</Emphasis>)

The European green crab (Carcinus maenas) is a highly successful marine invader, having established populations in a number of areas outside its natural range in the last 100 years. In South Australia,C. maenas can be abundant on intertidal mud flats, which are used by juveniles of the native blue swimmer crab (Portunus pelagicus) and could have the potential to cause substantial negative effects on this species. The influence of adult blue and green crabs on habitat selection by juvenile blue crabs was tested to determine if they responded to both predators in a similar fashion. The presence of predators did not influence habitat selection by juvenile blue crabs in either laboratory or field experiments, but juvenile behavior in the selected habitat did differ between the two adult species. Many more crabs buried themselves beneath the substrate when adult conspecifics were present than when adult green crabs were present. Burying in the presence of adult green crabs was no more frequent than when predators were absent. It remains to be determined if this makes juvenile blue crabs more vulnerable to predation by green crabs than by adults of their own species, or if the difference in response is because green crabs pose a different or lesser threat.     

The impact of yellow shore crabs,<Emphasis Type="Italic">Hemigrapsus oregonensis</Emphasis>, on Early Benthic Phase dungeness crabs,<Emphasis Type="Italic">Cancer magister</Emphasis>, in intertidal oyster shell mitigation habitat

Dungeness crab (Cancer magister) megalopae recruit to northeastern Pacific coastal estuaries, and settle into intertidal and subtidal habitats where they molt into Early Benthic Phase (EBP) crabs, and are dependent on epibenthic structure for shelter from predation. Given the importance of shell refuge to their post-settlement ontogeny, the U.S. Army Corps of Engineers began constructing intertidal plots of oyster shell in the Grays Harbor estuary, Washington, to enhance recruitment and mitigate losses of subtidalC. magister entrained and killed during extensive dredging efforts. When shell habitat was newly constructed, settlement and survival ofC. magister were high, and expectations for the mitigation project were met. During the first several years (1992-1997), plots greater than 1 yr postconstruction were colonized by yellow shore crabs,Hemigrapsus oregonensis (often≥75 crabs m⁻², and abundance of EBPC. magister was reduced to nearly zero. While some predation of settling megalopae byH. oregonensis does occur, the relationship between these species is characterized by density-dependent competitive interactions. Laboratory observations of competition for shell habitat indicate thatH. oregonensis are dominant over EBPC. magister and can evictC. magister from refuge spaces. Field experiments show that high densities of the former cause, the latter to emigrate from shell, and suggest detection and avoidance of areas with high densities ofH. oregonensis by settlingC. magister megalopae. More recently (1998-2001), abundance ofH. oregonensis has declined dramatically within plots of oyster shell, apparently due to recruitment failure, and patterns ofC. magister abundance and production have returned to levels consistent with original expectations of the mitigation project. Both intraspecific and interspecific competition for space are significant factors effecting population, regulation ofC. magister when they are strongly dependent on refuge from predation. Efficacy of constructed oyster shell plots as a mitigation tool may hinge on the population dynamics of a species previously considered of little consequence to the target species.     

Quantifying Tidal Movements of the Shore Crab <Emphasis Type="Italic">Carcinus maenas</Emphasis> on to Complex Epibenthic Bivalve Habitats

Many subtidal predators undertake regular tidal migrations into intertidal areas in order to access abundant prey. One of the most productive habitats in soft bottom intertidal systems is formed by beds of epibenthic bivalves such as blue mussels (Mytilus edulis) and Pacific oysters (Crassostrea gigas). In the Dutch Wadden Sea, these bivalves might face substantial predation pressure by the shore crab (Carcinus maenas), which increased considerably in numbers during the last 20 years. However, the quantification of this species on bivalve beds is challenging, since most methods common for quantifying animal abundance in marine habitats cannot be used. This study investigated the potential of two methods to quantify the abundance of C. maenas on 14 epibenthic bivalve beds across the Dutch Wadden Sea. The use of the number of crabs migrating from subtidal towards intertidal areas as a proxy of abundance on bivalve beds yielded unreliable results. In contrast, crabs caught with traps on the beds were correlated with the abundance assessed on the surrounding bare flats by beam trawl and therefore provided usable results. The estimates, however, were only reliable for crabs exceeding 35 mm in carapace width (CW). The application of these estimates indicated that crab abundances on bivalve beds were influenced by the biogenic structure. Beds dominated by oysters attracted many large crabs (> 50-mm CW), whereas abundances of medium-sized crabs (35–50-mm CW) showed no relationship to the oyster occurrence. The combination of traps and trawls is capable of quantifying crab abundance on bivalve beds, which offers the possibility to study biotic processes such as predator-prey interactions in these complex structures in more detail.     

<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.     

Blue Crab (<Emphasis Type="Italic">Callinectes sapidus</Emphasis> Rathbun, 1896) Settlement at Three Georgia (USA) Estuarine Sites

The blue crab, Callinectes sapidus Rathbun, 1896, represents the second most important fishery for coastal Georgia; yet, little is known about environmental forces that affect planktonic postlarval settlement in the region. Here, we describe a study to examine the physical mechanisms responsible for blue crab settlement in the extensive salt marsh system of coastal Georgia. Bottom and surface samplers were placed at three sites along a salinity gradient from a low-salinity site in the Altamaha River to a high-salinity area of the Duplin River, Sapelo Island, GA, USA during 2005. Megalopae and juvenile monitoring occurred from July through December. The majority of both megalopae (86.8%) and juvenile (89.3%) blue crabs were recovered in bottom samplers at the low-salinity Altamaha River site during August and early September. Few megalopae were collected at the surface of the Altamaha River or at the two higher-salinity sites in the Duplin and North Rivers. Downwelling winds were unable to explain all settlement events; however, winds with an onshore component regularly preceded settlement events. The use of a multiple-regression model revealed a lagged relationship (r = 0.5461, $ lag = 0–2 days $ lag = 0–2 days ) between wind events, temperature, salinity, maximum tidal height, and settlement.     

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.     

Effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> on infaunal zonation patterns,tidal behavior,and risk of mortality

Rhythmic movements in response to tidal cycles are characteristic of infaunal inhabitant of intertidal soft-bottoms, allowing them to remain in the area with best living conditions. The effect of bioturbators as modifier of local environmental conditions and thus of gradients in intertidal habitats, has not been investigated yet. The Atlantic estuarine intertidal areas are dominated by the burrowing crabChasmagnathus granulatus that generates strong environmental heterogeneity by affecting the physical-chemical characteristics of the sediment. The comparison between intertidal areas with and without crab shows that sediments in the crab beds remain more humid, softer, and homogeneous across the intertidal and along the tidal cycle than areas without crabs. The densities of infauna were higher at high intertidal zones in crab beds than in similar areas without crabs. Infaunal organisms performed vertical movements into the sediment following the tidal cycle that were always of higher magnitude in habitats without crabs. Infaunal species tend to spend most of the time buried into the sediment in the crab bed. Migratory shorebirds use the Atlantic estuarine environments as stopover or wintering sites. They feed (mainly on polychaetes) in the low intertidal zones of both habitats (with and without crabs), but they also feed in the upper intertidal of the crab bed; polychaete per capita mortality rate is higher in the upper part of the crab bed. Environmental heterogeneity produced by crab disturbance has an effect on the infaunal behavior, risk of mortality, and the zonation pattern. This is another example of the ecosystem engineering ability of a burrowing intertidal species.     

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.