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.     

Determinants of expansion for<Emphasis Type="Italic">Phragmites australis</Emphasis>, common reed,in natural and impacted coastal marshes

The rapid spread ofPhragmites australis in the coastal marshes of the Northeastern United States has been dramatic and noteworthy in that this native species appears to have gained competitive advantage across a broad range of habitats, from tidal salt marshes to freshwater wetlands. Concomitant with the spread has been a variety of human activities associated with coastal development as well as the displacement of nativeP. australis with aggressive European genotypes. This paper reviews the impacts caused by pure stands ofP. australis on the structure and functions of tidal marshes. To assess the determinants ofP. australis expansion, the physiological tolerance and competitive abilities of this species were examined using a field experiment.P. australis was planted in open tubes paired withSpartina alterniflora, Spartina patens, Juncus gerardii, Lythrum salicaria, andTypha angustifolia in low, medium, and high elevations at mesohaline (14‰), intermediate (18‰), and salt (23‰) marsh locations. Assessment of the physiological tolerance ofP. australis to conditions in tidal brackish and salt marshes indicated this plant is well suited to colonize creek banks as well as upper marsh edges. The competitive ability ofP. australis indicated it was a robust competitor relative to typical salt marsh plants. These results were not surprising since they agreed with field observations by other researchers and fit within current competition models throught to structure plant distribution within tidal marshes. Aspects ofP. australis expansion indicate superior competitive abilities based on attributes that fall outside the typical salt marsh or plant competition models. The alignment of some attributes with human impacts to coastal marshes provides a partial explanation of how this plant competes so well. To curb the spread of this invasive genotype, careful attention needs to be paid to human activities that affect certain marsh functions. Current infestations in tidal marshes should serve as a sentinel to indicate where human actions are likely promoting the invasion (e.g., through hydrologic impacts) and improved management is needed to sustain native plant assemblages (e.g., prohibit filling along margins).     

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.     

Phenotypic Variation Among Invasive <Emphasis Type="Italic">Phragmites australis</Emphasis> Populations Does Not Influence Salinity Tolerance

Phenotypic variation within species can have community- and ecosystem-level effects. Such variation may be particularly important in ecosystem engineers, including many invasive species, because of the strong influence of these species on their surrounding communities and environment. We combined field surveys and glasshouse experiments to investigate phenotypic variation within the invasive common reed, Phragmites australis, among four estuarine source sites along the east coast of North America. Field surveys revealed variation in P. australis height and stem density among source sites. In a glasshouse environment, percent germination of P. australis seeds also varied across source sites. To test the degree to which phenotypic variation in P. australis reflected genetic or environmental differences, we conducted a glasshouse common garden experiment assessing the performance of P. australis seedlings from the four source sites across a salinity gradient. Populations maintained differences in morphology and growth in a common glasshouse environment, indicating a genetic component to the observed phenotypic variation. Despite this variation, experimentally increased porewater salinity consistently reduced P. australis stem density, height, and biomass. Differences in these morphological metrics are important because they are correlated with the impacts of invasive P. australis on the ecological communities it invades. Our results indicate that both colonization and spread of invasive P. australis will be dependent on the environmental and genetic context. Additional research on intraspecific variation in invasive species, particularly ecosystem engineers, will improve assessments of invasion impacts and guide management decisions in estuarine ecosystems.     

Does<Emphasis Type="Italic">Phragmites</Emphasis> expansion alter the structure and function of marsh landscapes? Patterns and processes revisited

We assess the probability and importance of different spatial distributions ofPhragmites australis (Trin Ex Steud) within brackish tidal marshes of the mid-Atlantic United States coast. The comparative impact ofPhragmites expansion on the larger coupled marsh-estuary system may partially be a function of the landscape area dominated byPhragmites, the landscape position occupied byPhragmites, the landscape pattern created byPhragmites expansions, and the resulting impact on tidal drainage networks. We find evidence thatPhragmites establishment can occur at many landscape positions, and thatPhragmites spread within a marsh can occur via colonization (new patches), linear clonal growth (along a preferred axis), or circular clonal growth (non-directional, random spread). Early intervals ofPhragmites spread were dominated by colonization for all sites except for Piermont Marsh (which appeared to be dominated by linear clonal growth) and Lang Tract (which appeared to be dominated by circular clonal growth). Although 46–100% of new patches ofPhragmites occurred within 5 m of drainages, at only one site (Piermont Marsh, New York) didPhragmites populations remain concentrated along creek banks. Except for Iona Island, New York, which appears to be in an early stage ofPhragmites invasion, patch dynamics at all sites showed an increase followed by a decrease in patch number, as independent patches became established, expanded, and coalesced. We also found some evidence for a loss of first order streams at later stages ofPhragmites invasions in several sites (Hog Island, Lang Tract, Silver Run).     

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.     

A framework for evaluating potential ecological effects of implementing biological control of<Emphasis Type="Italic">Phragmites australis</Emphasis>

The increasing abundance ofPhragmites australis in many North American coastal and inland wetlands is of great concern to many scientists and land managers. While some considered the species native, others considered it an introduced invasive species. Regardless of its origin, control measures using mechanical, physical, and chemical means are implemented widely. With the exception of increased salinity, reductions in the abundance ofP. australis are usually temporary and control measures need to be repeated frequently increasing the cost and the potential for nontarget effects. The failure of traditional control attempts resulted in increased interest to develop a biological control program. Recent research suggests that genetically distinct populations (introduced and native) occur in North America. The native genotypes appear noninvasive and declining while the introduced genotypes are invasive replacing native genotypes where they co-occur. Accidentally introduced European herbivores specialized onP. australis are spreading across North America and can be extremely abundant in the Northeast. Introduction of additional specialized herbivores as biological control agents from the European range ofP. australis, if approved and successful, will result in changes in plant communities, wetland food webs, and management practices. The potential changes associated with introduction of biological control agents to wetland plant communities and their food webs may be more beneficial than current management strategies allowing the introducedP. australis genotypes and its associated herbivores to spread.     

Exploring the Effects of Shoreline Development on Fringing Salt Marshes Using Nekton,Benthic Invertebrate,and Vegetation Metrics

Fringing marshes are important but often overlooked components of estuarine systems. Due to their relatively small size and large edge to area ratio, they are particularly vulnerable to impacts from adjacent upland development. Because current shoreland zoning policies aim to limit activities in upland buffer zones directly next to coastal habitats, we tested for relationships between the extent of development in a 100-m buffer adjacent to fringing salt marshes and the structure of marsh plants, benthic invertebrates, and nekton communities. We also wanted to determine useful metrics for monitoring fringing marshes that are exposed to shoreline development. We sampled 18 fringing salt marshes in two estuaries along the coast of southern Maine. The percent of shoreline developed in 100-m buffers around each site ranged from 0 to 91 %. Several variables correlated with the percent of shoreline developed, including one plant diversity metric (Evenness), two nekton metrics (Fundulus heteroclitus %biomass and Carcinus maenas %biomass), and several benthic invertebrate metrics (nematode and insect/dipteran larvae densities in the high marsh zone) (p?<?0.05). Carcinus maenas, a recent invader to the area, comprised 30–97 % of the nekton biomass collected at the 18 sites and was inversely correlated with Fundulus %biomass. None of these biotic metrics correlated with the other abiotic marsh attributes we measured, including porewater salinity, marsh site width, and distance of the site to the mouth of the river. In all, between 25 and 48 % of the variance in the individual metrics we identified was accounted for by the extent of development in the 100-m buffer zone. Results from this study add to our understanding of fringing salt marshes and the impacts of shoreline development to these habitats and point to metrics that may be useful in monitoring these impacts.     

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.     

Vegetation Dynamics in Rhode Island Salt Marshes During a Period of Accelerating Sea Level Rise and Extreme Sea Level Events

Sea level rise is a major stressor on many salt marshes, and its impacts include creek widening, ponding, vegetation dieback, and drowning. Marsh vegetation changes have been associated with sea level rise across southern New England, but most of these studies pre-date the current period of rapidly accelerating sea level rise coupled with episodic events of extreme increases in water levels. Here, we combine data from two salt marsh monitoring and assessment programs in Rhode Island that were designed to assess marsh responses to sea level rise and use these data to document temporal and spatial patterns in marsh vegetation during the current period of extreme water level increases. Vegetation monitoring at two Narragansett Bay salt marshes confirms the ongoing decline of the salt meadow species Spartina patens during this period as it becomes replaced by Spartina alterniflora. Bare ground resulting from vegetation dieback was significantly related to mean high water levels and led to the rapid conversion of mixed Spartina assemblages to S. alterniflora monocultures. A broader spatial assessment of RI marshes shows that S. alterniflora dominance increases at lower elevation marshes toward the mouth of Narraganset Bay. Our data provide additional evidence that S. patens continues to decline in southern New England marshes and show that losses can accelerate during periods of extreme high water levels. Unless adaptive management actions are taken, we predict that marshes throughout RI will continue to lose salt meadow habitat and eventually resemble lower elevation marshes that are already dominated by S. alterniflora monocultures.     

Common reed grass,<Emphasis Type="Italic">Phragmites australis</Emphasis>, expansion into constructed wetlands: Are we mortgaging our wetland future?

The use of constructed wetlands to replace natural wetlands has become a widespread management tool. Because of the inherent disturbed nature of these sites, constructed wetlands are susceptible to colonization by undesirable plant species. Vegetated communities in 15 constructed wetland sites ranging in age from 1 to 12 yr and in size from 0.4 to 5.3 ha were surveyed using differential global positioning system (GPS) technology in 1994. These sites were re-surveyed in 2000. Colonization of the sites byPhragmites australis expanded from 73% of the sites in 1994 to 80% of the sites in 2000. The total area colonized byP. australis within the sites increased from 3.47 to 4.96 ha. In some sites, the area ofP. australis decreased, which appears to be correlated with an increase in scrub-shrub vegetation (0.986, p=0.014). Similar to results from the previous study, sites that are surrounded by subtidal perimeter ditches have significantly lessP. australis than those sites without perimeter ditches (p=0.019).P. australis expansion rates within the sites varied from 0.1 to 5.6 yr^?1. Colonization of constructed wetland sites byP. australis should be a continued concern of resource managers. Activities such as planting scrub-shrub species on the upland-wetland berm and construction of subtidal perimeter ditches should be considered as methods to reduce the probability of invasion.     

Wetland Loss Patterns and Inundation-Productivity Relationships Prognosticate Widespread Salt Marsh Loss for Southern New England

Tidal salt marsh is a key defense against, yet is especially vulnerable to, the effects of accelerated sea level rise. To determine whether salt marshes in southern New England will be stable given increasing inundation over the coming decades, we examined current loss patterns, inundation-productivity feedbacks, and sustaining processes. A multi-decadal analysis of salt marsh aerial extent using historic imagery and maps revealed that salt marsh vegetation loss is both widespread and accelerating, with vegetation loss rates over the past four decades summing to 17.3 %. Landward retreat of the marsh edge, widening and headward expansion of tidal channel networks, loss of marsh islands, and the development and enlargement of interior depressions found on the marsh platform contributed to vegetation loss. Inundation due to sea level rise is strongly suggested as a primary driver: vegetation loss rates were significantly negatively correlated with marsh elevation (r ²?=?0.96; p?=?0.0038), with marshes situated below mean high water (MHW) experiencing greater declines than marshes sitting well above MHW. Growth experiments with Spartina alterniflora, the Atlantic salt marsh ecosystem dominant, across a range of elevations and inundation regimes further established that greater inundation decreases belowground biomass production of S. alterniflora and, thus, negatively impacts organic matter accumulation. These results suggest that southern New England salt marshes are already experiencing deterioration and fragmentation in response to sea level rise and may not be stable as tidal flooding increases in the future.     

<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.     

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.     

<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.     

Growth and Movements of Mummichogs (<Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>) Along Armored and Vegetated Estuarine Shorelines

Alteration of estuarine shorelines associated with increased urbanization can significantly impact biota and food webs. This study determined the impact of shoreline alteration on growth and movement of the estuarine fish Fundulus heteroclitus in a tributary of the Delaware Coastal Bays. Fundulus heteroclitus is abundant along the east coast of the USA, and is an important trophic link between marsh and subtidal estuary. The restricted home range of F. heteroclitus allowed discrete sampling, and fish growth comparisons, along 35–65-m long stretches of fringing Spartina alterniflora and Phragmites australis marsh, riprap, and bulkhead. Fundulus heteroclitus were tagged with decimal Coded Wire Tags. Of 725 tagged F. heteroclitus, 89 were recaptured 30–63 days later. Mean growth rate (0.06–0.15 mm day^?1 across all shoreline types) was greatest at riprap, lowest at Spartina and Phragmites, and intermediate at bulkhead, where growth was not significantly different from any other shoreline. This suggests that discernible environments exist along different shoreline types, even at the scale of tens of meters. No difference in movement distance was detected at different shoreline types; most individuals displayed a high degree of site fidelity. Forty-seven percent were recaptured within 5 m of their tagging location, although alongshore movements up to 475 m were recorded. Estimates of relative F. heteroclitus productivity, using relative density data from a concurrent study, were highest along Spartina and Phragmites, intermediate at riprap, and lowest at bulkhead. Therefore, despite greater growth rates along riprap than at vegetated shores, armoring reduces abundance sufficiently to negatively impact localized productivity of F. heteroclitus.     

Functional Performance of Three Invasive <Emphasis Type="Italic">Marenzelleria</Emphasis> Species Under Contrasting Ecological Conditions Within the Baltic Sea

A 4-week laboratory experiment investigated the behaviour (survival and bioirrigation) and impact of the invasive polychaetes Marenzelleria viridis, M. neglecta and M. arctia on sediment-water solutes exchange, porewater chemistry, and Fe and P interactions in high-salinity sandy sediment (HSS) and low-salinity muddy sediment (LSM) from the Baltic Sea. M. viridis showed deep burrowing with efficient bioirrigation (11 L m^?2 day^?1) and high survival (71%) in HSS, while M. arctia exhibited shallow burrowing with high bioirrigation (12 L m^?2 day^?1) and survival (88%) in LSM. M. neglecta behaved poorly in both ecological settings (bioirrigation, 5–6 L m^?2 day^?1; survival, 21–44%). The deep M. viridis bioirrigation enhanced total microbial CO₂ (TCO₂) production in HSS by 175% with a net efflux of NH₄⁺ and PO₄^3?, at rates 3- to 27-fold higher than for the other species. Although the shallow and intense bioirrigation of M. arctia in LSM stimulated microbial TCO₂ production to some extent (61% enhancement), the nutrient fluxes close to zero indicate that it effectively prevented the P release. Porewater Fe:PO₄^3? ratios revealed that the oxidizing effect of M. arctia bioirrigation increased the PO₄^3? adsorption capacity of LSM twofold relative to defaunated controls while no buffering of PO₄^3? was detected in M. viridis HSS treatment. Therefore, the different behaviour of the three species in various environments and the sharp contrast between M. viridis and M. arctia effects on C, N and P cycling must be considered carefully when the ecological role of Marenzelleria species in the Baltic Sea is evaluated.     

The Effects of Integrated Marsh Management (IMM) on Salt Marsh Vegetation,Nekton, and Birds

An integrated marsh management (IMM) project in an urbanized watershed on Long Island, New York, USA, aimed to mitigate salt marsh degradation and to reduce mosquito production by an innovative combination of restoration and open marsh water management methods. The grid ditch network at two treatment marshes was replaced with naturalized tidal channels and ponds. Effects of the hydrologic alterations were monitored utilizing a before–after–control–impact approach. The treatment marshes experienced a number of beneficial outcomes including a fourfold reduction in the invasive Phragmites australis and increased native vegetation cover in the most degraded portions of the marsh, increased abundance and diversity of marsh killifish and estuarine nekton species, higher shorebird and waterfowl densities, and increased avian species diversity. The successful implementation of IMM concept led to improved marsh health and diminished mosquito production. Therefore, this study may serve as a template for similar large-scale integrated salt marsh restoration projects.     

Isotopic and Elemental Composition of Marine Macrophytes as Biotracers of Nutrient Recycling Within a Coastal Lagoon in Baja California,Mexico

Nutrient sources of San Quintin Bay, a coastal lagoon affected by coastal upwelling off Baja California (Mexico), were traced using generalized additive (mixed) models (GAMM) to the stable nitrogen isotopic composition, C:N and N content of two co-occurring macrophytes (the macroalgae Ulva spp. and the seagrass Zostera marina). The geochemical tracers followed a spatial trend that partly responded to the long-term nutrient gradient from the ocean towards the interior of the bay. N content in Z. marina and Ulva spp. decreased linearly (while C:N increased) towards the middle section of the bay to concentration levels that indicate potential N limitation for growth. Concurrently midway into the bay (6–9 km), the δ¹⁵N of both macrophytes showed a gradual enrichment in ¹⁵N reflecting progressive denitrification. The spatial pattern of δ¹⁵N and the decrease in C:N of the macrophytes towards the innermost section of the bay indicated an additional nonoceanic source of dissolved nitrogen in this zone. The similarity of the δ¹⁵N pattern of Z. marina and Ulva spp. implies that their δ¹⁵N composition is mainly controlled by the availability of N, in spite of the physiological differences between taxa. A better fit of GAMM to N content and C:N was obtained for Z. marina than for Ulva spp. indicating that the former delineate more steadily and smoothly the influence of upwelling along the spatial gradient. Nonetheless, Ulva spp. may be analyzed in combination with Z. marina to characterize the environmental conditions at the time of sampling.     

Spatial and Temporal Variation in Brackish Wetland Seedbanks: Implications for Wetland Restoration Following <Emphasis Type="Italic">Phragmites</Emphasis> Control

Chesapeake Bay tidal wetlands are experiencing a broad-scale, aggressive invasion by the non-native, clonal grass Phragmites australis. The grass is often managed with herbicides in efforts to restore native plant communities and wildlife habitat. Management efforts, however, can act as a disturbance, resulting in increased light availability, potentially fostering reinvasion from soil seedbanks. If native vegetation establishes quickly from seedbanks, the site should have greater resiliency against invasion, while disturbed sites where native plants do not rapidly establish may be rapidly colonized by P. australis. We surveyed the soil seedbank of three vegetation cover types in five Chesapeake Bay subestuaries: areas where P. australis had been removed, where P. australis was left intact, and with native, reference vegetation. We determined the total germination, the proportion of the seedbank that was attributable to invasive species, the richness, the functional diversity, and the overall composition of the seedbanks in each of the cover types (i.e., plots). After 2 years of herbicide treatment in the P. australis removal plots, vegetation cover type impacted the total germination or the proportion of invasive species in the seedbank. In contrast, we also found that seedbank functional composition in tidal brackish wetlands was not influenced by vegetation cover type in most cases. Instead, plots within a subestuary had similar seedbank functional composition across the years and were composed of diverse functional groups. Based on these findings, we conclude that plant community recovery following P. australis removal is not seed-limited, and any lack of native vegetation recruitment is likely the result of yet-to-be-determined abiotic factors. These diverse seedbanks could lead to resilient wetland communities that could resist invasions. However, due to the prevalence of undesirable species in the seedbank, passive revegetation following invasive plant removal may speed up their re-establishment. The need for active revegetation will need to be assessed on a case-by-case basis to ensure restoration goals are achieved.