Distribution of modern salt-marsh foraminifera in the Albemarle–Pamlico estuarine system of North Carolina, USA: Implications for sea-level research

We described the distributions of foraminifera from ten physiographically distinct salt marshes in the Albemarle–Pamlico estuarine system, North Carolina using 193 surface samples. We defined elevation-dependent ecological zones at individual sites using cluster analysis and detrended correspondence analysis. Additionally, seven principal biozones of salt-marsh foraminifera were identified that have distinctive spatial distributions reflecting a pattern of salinity regimes caused by the current configuration of barrier-island inlets. High salinity sites along the southern Outer Banks are associated with sub-tidal calcareous assemblages, low marshes dominated by Miliammina fusca and high marsh environments defined by Haplophragmoides wilberti, Trochammina inflata and Arenoparrella mexicana. In contrast, lower salinity marshes have Ammobaculites spp. in sub-tidal settings, Miliammina fusca-dominated low marshes and high marsh settings characterized by Jadammina macrescens. Spatial variation of foraminiferal populations and the potential for biozones to migrate in response to changing inlet configuration and salinity, suggests that datasets of modern salt-marsh foraminifera from multiple environments would be appropriate for reconstructing Holocene relative sea level in North Carolina.     

Salt Marsh Restoration in Connecticut: 20 Years of Science and Management

In 1980 the State of Connecticut began a tidal marsh restoration program targeting systems degraded by tidal restrictions and impoundments. Such marshes become dominated by common reed grass (Phragmites australis) and cattail (Typha angustifolia and T. latifolia), with little ecological connection to Long Island Sound. The management and scientific hypothesis was that returning tidal action, reconnecting marshes to Long Island Sound, would set these systems on a recovery trajectory. Specific restoration targets (i.e., pre‐disturbance conditions or particular reference marshes) were considered unrealistic. However, it was expected that with time restored tides would return ecological functions and attributes characteristic of fully functioning tidal salt marshes. Here we report results of this program at nine separate sites within six marsh systems along 110 km of Long Island Sound shoreline, with restoration times of 5 to 21 years. Biotic parameters assessed include vegetation, macroinvertebrates, and use by fish and birds. Abiotic factors studied were soil salinity, elevation and tidal flooding, and soil water table depth. Sites fell into two categories of vegetation recovery: slow, ca. 0.5%, or fast, more than 5% of total area per year. Although total cover and frequency of salt marsh angiosperms was positively related to soil salinity, and reed grass stand parameters negatively so, fast versus slow recovery rates could not be attributed to salinity. Instead, rates appear to reflect differences in tidal flooding. Rapid recovery was characterized by lower elevations, greater hydroperiods, and higher soil water tables. Recovery of other biotic attributes and functions does not necessarily parallel those for vegetation. At the longest studied system (rapid vegetation recovery) the high marsh snail Melampus bidentatus took two decades to reach densities comparable with a nearby reference marsh, whereas the amphipod Orchestia grillus was well established on a slow‐recovery marsh, reed grass dominated after 9 years. Typical fish species assemblages were found in restoration site creeks and ditches within 5 years. Gut contents of fish in ditches and on the high marsh suggest that use of restored marsh as foraging areas may require up to 15 years to reach equivalence with reference sites. Bird species that specialize in salt marshes require appropriate vegetation; on the oldest restoration site, breeding populations comparable with reference marshland had become established after 15 years. Use of restoration sites by birds considered marsh generalists was initially high and was still nearly twice that of reference areas even after 20 years. Herons, egrets, and migratory shorebirds used restoration areas extensively. These results support our prediction that returning tides will set degraded marshes on trajectories that can bring essentially full restoration of ecological functions. This can occur within two decades, although reduced tidal action can delay restoration of some functions. With this success, Connecticut's Department of Environmental Protection established a dedicated Wetland Restoration Unit. As of 1999 tides have been restored at 57 separate sites along the Connecticut coast.     

Short-term responses to salinity of an invasive cordgrass

Salinity is one of the main chemical factors in salt marshes. Studies focused on the analysis of salinity tolerance of salt marsh plants are very important, since they may help to relate their physiological tolerances with distribution limits in the field. Spartina densiflora is a South America cordgrass, which has started its invasion of the European coastline from the southwestern Iberian Peninsula. In this work, short-term responses in adult tussocks of S. densiflora from southwestern Spain are studied over a wide range of salinity in a greenhouse experiment. Our results point out that S. densiflora has a high tolerance to salinity, showing high growth and net photosynthesis rates from 0.5 to 20 ppt. S. densiflora showed at the lowest salinity (0.5 ppt) high levels of photoinhibition, compensated by higher levels of energy transmission between photosystems. Adaptative mechanisms, as those described previously, would allow it to live in fresh water environments. At the highest salinity (40 ppt), S. densiflora showed a high stress level, reflected in significant decreases in growth, net photosynthesis rate and photochemical efficiency of Photosystem II. These responses support S. densiflora invasion patterns in European estuaries, with low expansion rates along the coastline and faster colonization of brackish marshes and river banks. This revised version was published online in July 2006 with corrections to the Cover Date.     

Modification of Sediments and Macrofauna by an Invasive Marsh Plant

Invasive grasses have recently altered salt marsh ecosystems throughout the northern hemisphere. On the eastern seaboard of the USA, Phragmites australis has invaded both brackish and salt marsh habitats. Phragmites australis influence on sediments and fauna was investigated along a salinity and invasion-age gradient in marshes of the lower Connecticut River estuary. Typical salinities were about 19–24 ppt in Site I, 9–10 ppt in Site II and 5–7 ppt in Site III. Strongest effects were evident in the least saline settings (II and III) where Phragmites has been present the longest and exists in monoculture. Limited influence was evident in the most saline region (I) where Phragmites and native salt marsh plants co-occur. The vegetation within Phragmites stands in tidal regions of the Connecticut River generally exhibits taller, but less dense shoots, higher above-ground biomass, and lower below-ground biomass than does the un-invaded marsh flora. There were lower sediment organic content, greater litter accumulation and higher sediment chlorophyll a concentrations in Phragmites- invaded than un-invaded marsh habitat. Epifaunal gastropods (Succinea wilsoni and Stagnicola catascopium) were less abundant in habitats where Phragmites had invaded than in un-invaded marsh habitat. Macro-infaunal densities were lower in the Phragmites-invaded than un-invaded habitats at the two least saline sites (II and III). Phragmites stands supported more podurid insects, sabellid polychaetes, and peracarid crustaceans, fewer arachnids, midges, tubificid and enchytraeid oligochaetes, and greater habitat-wide taxon richness as measured by rarefaction, than did the un-invaded stands. The magnitude and significance of the compositional differences varied with season and with site; differences were generally greatest at the oldest, least saline sites (II and III) and during May, when faunal densities were higher than in September. However, experimental design and the 1-year study period precluded clear separation of salinity, age, and seasonal effects. Although structural effects of Phragmites on salt marsh faunas are evident, further investigation is required to determine the consequences of these effects for ecosystem function. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparison of physical characteristics between created and natural estuarine marshes in Galveston Bay,Texas

Five natural and ten created Spartinaalternifloramarshes in the Lower Galveston BaySystem were compared to determine if there weresignificantly different physical characteristicsassociated with each type of marsh. The saltmarsheswere compared on the basis of microhabitats,length-width ratio, area-perimeter ratio, marsh-wateredge ratio, total size of S. alternifloraplantcommunities, fetch distances, angle of exposure,orientation, and elevation. All physicalmeasurements, except for elevation, were obtained fromphotography analyzed with the use of a GeographicInformation System with digital image processingcapabilities. Differences existed between natural andcreated marshes. The natural marsh sites in this studywere characterized by highly undulant marsh-wateredges, island-like S. alternifloraplant stands,concave shorelines, and low elevations. Createdmarshes were characterized by relatively smoothmarsh-water edges, an unbroken shoreline morphology,convex to straight shoreline configurations, andelevations on the edge and inner portions of the marshhigher than those of natural marshes. The lowelevations of the natural marsh appear to be due tocoastal subsidence in the Galveston Bay area alongwith rising sea level. Reticulated marshes andundulant shorelines appear to be caused by consequentdrowning of the natural marshes. High elevations insome of the created marshes are related to erosion ofthe low elevation marsh or deposition of coarsesediments at the marsh-water edge.     

Environmental determinants of Phragmites australis expansion in a New Jersey salt marsh: an experimental approach

Interdependence among disturbance events, ecosystem properties, and biological invasions often make causal relationships difficult to discern. For example, Phragmites australis invasion in mid-Atlantic salt marshes is often associated with disturbances that create well-drained features as well as with low sulfide concentrations, but explanations of these associations have been elusive. We tested experimentally: 1) that disturbances increasing wetland drainage facilitate Phragmites invasion by altering sulfide concentrations and salinity; 2) that translocation allows plants to spread beyond drainage areas; and 3) that plants can then lower edaphic stress through pressure ventilation of the rhizosphere and promote further expansion. At the invasion front, treatments of 1) severing rhizomes to halt translocation and 2) combined severing with clipping dead culms to limit ventilation of the rhizosphere killed most culms, but did not affect pore water chemistry. In already invaded areas, severing and clipping reduced culm height and panicle production, severing alone and in combination with clipping also raised sulfide and ammonium concentrations in the root zone. There were no treatment effects on plant performance or pore water chemistry along mosquito ditches, where sulfide concentrations were negligible. Small-scale hydrological alterations such as ditches appear to provide suitable sites for the establishment of Phragmites because soils are well-drained and are low in free sulfides. Subsequent expansion into more hostile areas occurs through translocation, with well-drained areas acting as sources for essential substances. Once established, the plant increases rhizosphere oxygenation and lowers sulfide concentrations.     

Role of biotic interactions and physical factors in determining the distribution of marsh species along an estuarine salinity gradient

Understanding the mechanisms that shape plant distribution patterns is a major goal in ecology. We investigated the role of biotic interactions (competition and facilitation) and abiotic factors in creating horizontal plant zonation along salinity gradients in the Elbe estuary. We conducted reciprocal transplant experiments with four dominant species from salt and tidal freshwater marshes at two tidal elevations. Ten individuals of each species were transplanted as sods to the opposing marsh type and within their native marsh (two sites each). Transplants were placed at the centre of 9‐m² plots along a line parallel to the river bank. In order to disentangle abiotic and biotic influences, we set up plots with and without neighbouring vegetation, resulting in five replicates per site. Freshwater species (Bolboschoenus maritimus and Phragmites australis) transplanted to salt marshes performed poorly regardless of whether neighbouring vegetation was present or not, although 50–70% of the transplants did survive. Growth of Phragmites transplants was impaired also by competition in freshwater marshes. Salt marsh species (Spartina anglica and Puccinellia maritima) had extremely low biomass when transplanted to freshwater marshes and 80–100% died in the presence of neighbours. Without neighbours, biomass of salt marsh species in freshwater marshes was similar to or higher than that in salt marshes. Our results indicate that salt marsh species are precluded from freshwater marshes by competition, whereas freshwater species are excluded from salt marshes by physical stress. Thus, our study provides the first experimental evidence from a European estuary for the general theory that species boundaries along environmental gradients are determined by physical factors towards the harsh end and by competitive ability towards the benign end of the gradient. We generally found no significant impact of competition in salt marshes, indicating a shift in the importance of competition along the estuarine gradient.     

Seed origin determines the range expansion of the clonal grass Elymus athericus

The recent invasion of clonal grasses to novel habitats poses a threat to biodiversity in various habitats. Elymus athericus, a clonal grass of north-western European salt marshes, is currently increasing in abundance and invading new habitats. In this study, we analyzed controlling factors for seedling establishment of E. athericus in frequently flooded low marsh habitats. Here, biotic and abiotic conditions are very different from the conditions of the parental sites with established populations higher up in the marsh. Hence, we hypothesized that seedling establishment at the expanding low marsh edge would depend on the parental origin (either through maternal effects or heritable local adaptation). We further hypothesized that seedling origin interacts with biotic factors such as herbivory and competition as well as with abiotic factors like inundation frequency. We tested the dependence of seedling survival, growth and vegetative reproduction on these factors in a factorial transplant experiment on Schiermonnikoog. Survival was high, with 77% of the planted seedling surviving until the end of the experiment. Biotic factors had a much stronger effect on seedling growth and mortality than parental origin and were independent of inundation. However, parental origin strongly interacted with herbivory and competition, with seedlings performing better under the conditions that resembled their parental site.We conclude that seedlings of E. athericus, a species that was previously thought to occur only in mid- to high marsh elevation, can establish at a frequently inundated low-marsh sites. Long term survival and further invasion will primarily depend on biotic factors in interaction with seed origin. Our results suggest that next to herbivory, limitation of seeds adapted to colonizing conditions is likely to slow down range expansion.     

Hydrologic and edaphic constraints on Schoenoplectus acutus,Schoenoplectus californicus,and Typha latifolia in tidal marsh restoration

The demand for an improved knowledge base for planning and management of tidal marsh restoration worldwide has become more fully recognized. In the Sacramento‐San Joaquin Bay Delta, California, U.S.A., concerns have arisen about the degradation of the Delta and key ecosystem services. One restoration method proposed includes intentionally breaching levees that protect agricultural lands to re‐establish a hydrology that encourages tidal marsh development. Our research investigated relevant constraints on vegetation establishment and expansion of key tidal marsh species. We transplanted three macrophyte species (Schoenoplectus acutus, Schoenoplectus californicus, and Typha latifolia) using two transplant types (rhizomes and adults) in locations that varied in hydrologic and edaphic conditions at Liberty Island, a post‐levee breach tidal marsh restoration site. Two years of monitoring revealed that transplanted adults outperformed rhizomes. In addition, S. californicus exhibited greater survival and vegetation expansion. S. californicus vegetation expansion covered a maximum area of approximately 23 m², which is two orders of magnitude (OOM) greater than the maximum area covered by S. acutus (approximately 0.108 m²) and three OOM greater than T. latifolia (approximately 0.035 m²). Results suggest that hydrologic regime and degree of soil compaction are influential in controlling vegetation establishment and expansion. Greater vegetation expansion occurred in transplant sites characterized by a deeper surface layer of non‐compacted soil in conjunction with shorter durations of flooding. Information derived from this study is valuable to restoration planning in the Delta and other tidal marshes worldwide where these species occur, especially in terms of setting restoration goals and trajectories based on site‐specific environmental characteristics.     

Are biotic factors significant in influencing the distribution of halophytes in saline habitats?

The influence of biotic factors on the distribution and establishment of halophytes is being considered in this review. Physicochemical factors, such as salinity and flooding, often are considered to be the determining factors controlling the establishment and zonational patterns of species in salt marsh and salt desert environments. Sharp boundaries commonly are found between halophyte communities even though there is a gradual change in the physicochemical environment, which indicates that biotic interactions may play a significant role in deterining the distribution pattern of species and the composition of zonal communities. Competition is hypothesized to play a key role in determining both the upper and lower limits of species distribution along a salinity gradient. Field and laboratory experiments indicate that the upper limits of distribution of halophytes into less saline or nonsaline habitats is often determined by competition. There appears to be a reciprocal relationship between the level of salt tolerance of species and their ability to compete with glycophytes in less saline habitats. Halophytes are not competitive in nonsaline habitats, but their competitive ability increases sharply in saline habitats. Allelopathic effects have been reported in salt desert habitats, but have not been reported along salinity gradients in salt marshes. Some species of halophytes that are salt accumulators have the ability to change soil chemistry. Chemical inhibition of intolerant species occurs when high concentrations of sodium are concentrated in the surface soils of salt desert plant communities that are dominated by salt-accumulating species. Establishment of less salt-tolerant species is inhibited in the vicinity of these salt-accumulating species. Herbivory is reported to cause both an increase and a decrease in plant diversity in salt marsh habitats. Heavy grazing is reported to eliminate sensitive species and produce a dense cover of graminoids in high marsh coastal habitats. However, in other marshes, grazing produced bare patches that allowed annuals and other low marsh species to invade upper marsh zonal communities. A retrogression in plant succession may occur in salt marshes and salt deserts because of heavy grazing. Intermediate levels of grazing by sheep, cattle, and horses could produce communities with the highest species richness and heterogeneity. Grazing by geese produced bare areas that had soils with higher salinity and lower soil moisture than vegetated areas, allowing only the more salt-tolerant species to persist. Removal of geese from areas by use of inclosures caused an increase in species richness in subarctic salt marshes. Invertebrate herbivores could also inhibit the survival of seeds and the ability of plants to establish in marshes. Parasites could play a significant role in determining the species composition of zonal communities, because uninfected rarer species are able to establish in the gaps produced by the death of parasitized species.     

What confines an annual plant to two separate zones along coastal topographic gradients?

We investigated the roles of flooding, salinity, and plant competition in creating a bimodal zonation pattern of the marsh dominant annual plant, Suaeda salsa, along coastal topographic gradients on the Pacific coast of northern China. In two consecutive years, we manipulated salinity and flooding, salinity, and competition for S. salsa seedlings that had been transplanted into the mudflat, the high marsh, and the upland, respectively. S. salsa plants that had been transplanted into the mudflat were completely eliminated in the non-elevated treatments whereas they performed much better in the 10 cm elevated treatments, regardless of salinity treatments. Although the performance of S. salsa transplanted into the high marsh did not differ between the fresh (watered) and the salt (control) treatments, S. salsa seedling emergence in the high marsh was nearly completely inhibited in the salt treatments. In contrast, a large number of S. salsa seedlings did emerge in the fresh treatments. S. salsa transplanted into the upland performed well when neighbors were removed, whereas it appeared to be strongly suppressed when neighbors were present. These data indicated that flooding, salinity, and competition all played a role in determining the zonation pattern of S. salsa. Furthermore, the importance of salinity was found to vary with life-history stage. Based on the results from these field manipulative experiments, we suggest that the marsh plant zonation paradigm may hold true for plant distributions along landscape-scale topographic gradients from mudflats to uplands in general. The relative importance of flooding, salinity, and competition, however, may vary at different elevations within a site and between sites. Handling editor: Pierluigi Viaroli     

Blocking Phragmites australis reinvasion of restored marshes using plants selected from wild populations and tissue culture

This study tested a vegetation strategy for controlling Phragmites australis invasion into brackish marshes as an alternative to the current technique of repeated herbicide sprays followed by burning. This strategy involves blocking P. australis by planting desired plants selected from wild populations and/or tissue culture regenerants at key points on the major routes of P. australis invasion. The planting of native species was conducted at three sites in a herbicide-treated P. australis marsh near Salem, NJ. Wild population selections of three upland marsh shrubs, Myrica cerifera, Baccharis halimifolia, and Iva frutescens, as well as two grass species, Spartina alterniflora and Spartina patens, and two rushes, Juncus gerardi and Juncus roemerianus, were planted according to their normal zonation positions. Tissue culture regenerated plants of the two grasses and two rushes, and the sedge species Scirpus robustus, were also planted. Plant growth at each site was monitored each year after planting for up to 3 years. Most plants of B. halimifolia, I. frutescens, J. roemerianus, and S. patens demonstrated a consistent vigorous growth at all three sites, whether or not the plants were collected from wild populations or were tissue culture regenerants. These multi-layered walls of plants demonstrated effectiveness in controlling the P. australis by restricting or inhibiting its spread. Upon screening 48 regenerated plants of S. patens at one of the three sites, we found that some regenerants showed enhanced characteristics for blocking P. australis, such as greater expansion and a high stem density. The availability of the tissue culture-regenerated plants of the native marsh species makes it possible to select lines from local genotypes that have desirable characteristics for wetland restoration projects, such as blocking P. australis reinvasion.     

Utilization of a citizen monitoring protocol to assess the structure and function of natural and stabilized fringing salt marshes in North Carolina

Narrow fringing salt marshes dominated by Spartina alterniflora occur naturally along estuarine shorelines and provide many of the same ecological functions as more extensive marshes. These fringing salt marshes are sometimes incorporated into shoreline stabilization efforts. We obtained data on elevation, salinity, sediment characteristics, vegetation and fish utilization at three study sites containing both natural fringing marshes and nearby restored marshes located landward of a stone sill constructed for shoreline stabilization. During the study, sediment accretion rates in the restored marshes were approximately 1.5- to 2-fold greater than those recorded in the natural marshes. Natural fringing marsh sediments were predominantly sandy with a mean organic matter content ranging between 1.5 and 6.0%. Average S. alterniflora stem density in natural marshes ranged between 130 and 222 stems m⁻², while mean maximum stem height exceeded 64 cm. After 3 years, one of the three restored marshes (NCMM) achieved S. alterniflora stem densities equivalent to that of the natural fringing marshes, while percentage cover and maximum stem heights were significantly greater in the natural than in the restored marshes at all sites. There was no significant difference in the mean number of fish, crabs or shrimp captured with fyke nets between the natural and restored marshes, and only the abundance of Palaemonetes vulgaris (grass shrimp) was significantly greater in the natural marshes than in the restored ones. Mean numbers of fish caught per 5 m of marsh front were similar to those reported in the literature from marshes adjacent to tidal creeks and channels, and ranged between 509 and 634 fish net⁻¹. Most of the field data and some of the sample analyses were obtained by volunteers as they contributed 223 h of the total 300 h spent collecting data from three sites in one season. The use of fyke nets required twice as many man-hours as any other single task. Vegetation and sediment parameters were sensitive indicators of marsh restoration success, and volunteers were capable of contributing a significant portion of the labor needed to collect these parameters. The U.S. Government's right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Constraints on Salt Marsh Development Following Managed Coastal Realignment: Dispersal Limitation or Environmental Tolerance?

Salt marshes restored through managed coastal realignment (MR) often develop slowly and show persistent differences in vegetation from natural marshes. Development might be constrained by the availability of propagules or poor suitability of the abiotic environment for their establishment. To distinguish between these factors, we compared vegetation colonization and environmental conditions at a salt marsh created by MR at Brancaster, Norfolk, UK, with five reference marshes, varying in age from 30 to circa 6,000 years. After 5 years, plant communities of the MR site remained different from those in mature reference marshes. In contrast, the communities of the youngest reference marsh were not significantly different from mature reference marshes. At the MR site, abundance of perennial and later‐successional species was low and large areas remained unvegetated. These differences are unlikely to be due to dispersal limitation, because 76% of the species from the local species pool colonized the site within 5 years. Although the annuals Salicornia europaea and Suaeda maritima were abundant by year 2, they were not ubiquitous until the end of the study. Tidal elevations of the MR site were suitable for vegetation development, but soil redox potentials were lower than that at the reference sites. Reducing conditions in the MR site appear to be the major cause of vegetation differences from the reference marshes, as they are associated with an abundance of bare ground and a small range of vegetation clusters. Measures to avoid low sediment redox potentials may have a great benefit in some salt marsh restoration projects.     

A comparison of Phragmites australisin freshwater and brackish marsh environments in North America

This paper compares the available North Americanliterature and data concerning several ecologicalfactors affecting Phragmites australisin inlandfreshwater, tidal fresh, and tidal brackish marshsystems. We compare aboveground productivity, plantspecies diversity, and sediment biogeochemistry; andwe summarize Phragmiteseffects on faunalpopulations in these habitats. These data suggest thatPhragmitesaboveground biomass is higher thanthat of other plant species occurring in the samemarsh system. Available data do not indicate anysignificant difference in the aboveground Phragmitesbiomass between marsh types, nor doesthere appear to be an effect of salinity on height.However, Phragmitesstem density wassignificantly lower in inland non-tidal freshwatermarshes than in tidal marshes, whether fresh orbrackish. Studies of the effects of Phragmiteson plant species richness suggest that Phragmitesdominated sites have lower diversity.Furthermore, Phragmiteseradication infreshwater sites increased plant diversity in allcases. Phragmitesdominated communities appearto have different patterns of nitrogen cyclingcompared to adjacent plant communities. Abovegroundstanding stocks of nitrogen (N) were found to behigher in Phragmitessites compared to thosewithout Phragmites. Porewater ammonium(NH₄ ⁺) did not differ among plant covertypes in the freshwater tidal wetlands, but inbrackish marshes NH₄ ⁺was much higher inSpartinaspp. than in neighboring Phragmitesstands. Faunal uses of Phragmitesdominated sites in North America were found to vary bytaxa and in some cases equaled or exceeded use ofother robust emergent plant communities. In light ofthese findings, we make recommendations for futureresearch.     

Microscale Spatial Distribution of Phragmites australis (Common Reed) Invasion into Spartina patens (Salt Hay)-dominated Communities in Brackish Tidal Marsh

Over the last century, Phragmites australis (common reed) has been expanding rapidly from the marsh–upland boundary into Spartina patens (salt hay)-dominated high marsh communities of the eastern US coast. Whereas direct and indirect human disturbances and changes in hydrology or salinity are likely to influence rates of spread at the landscape scale, the susceptibility of specific plant communities to invasion also influence rates of Phragmites expansion at the local scale. I measured microscale (0.25 m²) spatial patterns of culms (emerging buds and mature stems) in October 1993 at both expanding and stable boundaries of Phragmites populations within a S. patens-dominant matrix. In both expanding and stable plots, Phragmites culms were observed more frequently than expected on hummocks that were created by S. patens tussock-forming root structure. Culm density within a plot was correlated with the percent hummock cover within a plot. Further, Phragmites culms, particularly mature stems, were concentrated along the perimeter of the hummocks. Because the culms were not evenly distributed between hummocks and hollows, I suggest that invasion rates of Phragmites are limited in S. patens communities by microscale differences in hummock availability. The pattern of emergence suggests that expanding rhizomes of Phragmites encounter both competition with S. patens roots on the hummocks and physiological stressors (salinity, anoxia, sulfide concentrations) in the hollows.     

Responses of Salt Marsh Ecosystems to Mosquito Control Management Practices along the Atlantic Coast (U.S.A.)

Open marsh water management (OMWM) of salt marshes modifies grid‐ditched marshes by creating permanent ponds and radial ditches in the high marsh that reduce mosquito production and enhance fish predation on mosquitoes. It is preferable to using pesticides to control salt marsh mosquito production and is commonly presented as a restoration or habitat enhancement tool for grid‐ditched salt marshes. Monitoring of nekton, vegetation, groundwater level, soil salinity, and bird communities before and after OMWM at 11 (six treatment and five reference sites) Atlantic Coast (U.S.A.) salt marshes revealed high variability within and among differing OMWM techniques (ditch‐plugging, reengineering of sill ditches, and the creation of ponds and radial ditches). At three marshes, the dominant nekton shifted from fish (primarily Fundulidae species) to shrimp (Palaemonidae species) after manipulations and shrimp density increased at other treatment sites. Vegetation changed at only two sites, one with construction equipment impacts (not desired) and one with a decrease in woody vegetation along existing ditches (desired). One marsh had lower groundwater level and soil salinity, and bird use, although variable, was often unrelated to OMWM manipulations. The potential effects of OMWM manipulations on non‐target salt marsh resources need to be carefully considered by resource planners when managing marshes for mosquito control.     

A Marine Plant (Spartina Anglica) Invades Widely Varying Habitats: Potential Mechanisms of Invasion and Control

We report on the habitat dependent invasion and control pattern of the English cordgrass, Spartina anglica C. E. Hubbard, in Puget Sound, Washington. In 36 years, the plant has successfully invaded 73 sites, affecting 3311 ha of marine intertidal habitat, which if allowed to solidly fill, would equal 400ha. Invasion and control both depend on habitat type. Mudflats and low salinity marshes have significantly more solid area of S. anglica than do high salinity marshes and cobble beaches. Control efforts since 1997 have resulted in a 13% decline of the grass. We find that high salinity marshes have the greatest decline ( 70%), low salinity marshes have the lowest decline ( 10%), and mudflat ( 29%) and cobble beaches ( 21%) have intermediate losses. We hypothesize that invasion success and control are dependent on a relatively complex interplay between habitat physical conditions and species interactions.     

Predicting invasion dynamics of four alien Pinus species in a highly fragmented semi-arid shrubland in South Africa

This study explored the determinants of spread of four alien Pinus species and the ability of models to predict invasion dynamics in a complex fragmented landscape. The role of environmental factors, natural and anthropogenic disturbance in relation to invasion history was assessed for different stages in the invasion process using a Geographic Information System. Pines escaped from plantations over the past 30 years and spread into the natural semi-arid shrubland (renosterveld). The pattern of spread was compared with a simulated random distribution using two different techniques, a standard logistic regression, and a new recursive modelling approach (Formal Inference-based Recursive Modelling; FIRM). FIRM analysis improved the accuracy of predictions and revealed interactive effects of variables hidden by the logistic regression analysis. More than 80% of isolated pine individuals were found in 20% of the habitat classified as suitable by the models. Soil pH was the most important predictor for the distribution of isolated trees, whereas the establishment of dense pine stands was largely determined by fire history. Differences in invasive behaviour could be explained by species attributes such as limited dispersal for P. canariensis, and better drought-tolerance for P. halepensis. Sixty-five percent of the current pine distribution was accurately predicted by the spatial distribution of the first trees to have invaded. Such models could be used to predict potential spread of invasive plants and gain a better understanding of the main factors driving the invasion process. However, the spread of invasive species in fragmented landscapes, strongly modified by human activities, is very complicated, and the spread remains difficult to predict in the long term. The dynamics of invasion are discussed in relation to changes in land use and disturbance regime.