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.     

Patterns of fungal diversity and composition along a salinity gradient

Estuarine salinity gradients are known to influence plant, bacterial and archaeal community structure. We sequenced 18S rRNA genes to investigate patterns in sediment fungal diversity (richness and evenness of taxa) and composition (taxonomic and phylogenetic) along an estuarine salinity gradient. We sampled three marshes—a salt, brackish and freshwater marsh—in Rhode Island. To compare the relative effect of the salinity gradient with that of plants, we sampled fungi in plots with Spartina patens and in plots from which plants were removed 2 years prior to sampling. The fungal sediment community was unique compared with previously sampled fungal communities; we detected more Ascomycota (78%), fewer Basidiomycota (6%) and more fungi from basal lineages (16%) (Chytridiomycota, Glomeromycota and four additional groups) than typically found in soil. Across marshes, fungal composition changed substantially, whereas fungal diversity differed only at the finest level of genetic resolution, and was highest in the intermediate, brackish marsh. In contrast, the presence of plants had a highly significant effect on fungal diversity at all levels of genetic resolution, but less of an effect on fungal composition. These results suggest that salinity (or other covarying parameters) selects for a distinctive fungal composition, and plants provide additional niches upon which taxa within these communities can specialize and coexist. Given the number of sequences from basal fungal lineages, the study also suggests that further sampling of estuarine sediments may help in understanding early fungal evolution.     

Molluscan diversity in tidal marshes along the Scheldt estuary (The Netherlands, Belgium)

The mollusc fauna of 64 sites in 31 tidal marshes was surveyed along a salinity gradient from freshwater to marine conditions in the river Scheldt (Belgium–The Netherlands). A total of 10649 specimens involving 31 taxa were identified. Salinity turned out to be a major factor in mollusc assemblages in the Scheldt estuary, but other factors can not be excluded. In the marine part five species were common, compared to the brackish part where only Assiminea grayana was abundant. In the freshwater zone species richness was highest (24). There was a significant correlation between flooding frequency and species richness in the tidal freshwater marsh `Durmemonding'. Finally, the survey confirmed the distribution of the amphibious hygromiid snail Pseudotrichia rubiginosa, a species which in Belgium only occurs in the marshes of the tidal freshwater part of the Scheldt and its tributaries.     

Notes On The Occurrence Of Clathrus Cancellatus,Tournf., In Argyllshire

Background: Estuaries are characterised by salinity gradients and regular flooding events. These environmental factors form stress gradients, along which species composition changes.

Aims: Analyse and compare patterns of plant species diversity along the estuarine salinity and flooding gradients of the Elbe and Connecticut Rivers.

Methods: Vegetation was sampled at three elevations (low, mid, high) in five sites of each marsh type (fresh, brackish, salt) in both estuaries. Patterns of species density (SD) and evenness (E) along the gradients were analysed and compared between the two estuaries with three-factor ANOVAs.

Results: The regional species pool was 33% higher for the Connecticut than for the Elbe. SD of fresh marshes (19 ± 2.2) was more than twice in the Connecticut than in the Elbe. We found an overall increase in SD from low to high elevation and from salt to freshwater marshes in both estuaries. However, SD and E were strongly depressed at intermediate elevations in the Elbe fresh and brackish marshes.

Conclusions: Although diversity patterns in the two estuaries show overall similarities, patterns of SD and E differ, when particular elevational zones and marsh types are compared. We hypothesise this to be due to evolutionary and historical influences on the regional species pools, shaping the impact of local biotic and abiotic processes.     

High site fidelity and low site connectivity in temperate salt marsh fish populations: a stable isotope approach

Adult and juvenile fish utilise salt marshes for food and shelter at high tide, moving into adjacent sublittoral regions during low tide. Understanding whether there are high levels of site fidelity for different species of coastal fish has important implications for habitat conservation and the design of marine protected areas. We hypothesised that common salt marsh fish species would demonstrate a high site fidelity, resulting in minimal inter-marsh connectivity. Carbon (¹³C) and nitrogen (¹⁵N) stable isotope ratios of larvae and juveniles of five common salt marsh fish (Atherina presbyter, Chelon labrosus, Clupea harengus, Dicentrarchus labrax, Pomatoschistus microps), seven types of primary producer and seven secondary consumer food sources were sampled in five salt marshes within two estuary complexes along the coast of south-east England. Significant differences in ¹³C and ¹⁵N signatures between salt marshes indicated distinct sub-populations utilising the area of estuary around each salt marsh, and limited connectivity, even within the same estuary complex. ¹⁵N ratios were responsible for the majority of inter-marsh differences for each species and showed similar site-specific patterns in ratios in primary producers, secondary consumers and fish. Fish diets (derived from isotope mixing models) varied between species but were mostly consistent between marsh sites, indicating that dietary shifts were not the source of variability of the inter-marsh isotopic signatures within species. These results demonstrate that for some common coastal fish species, high levels of site fidelity result in individual salt marshes operating as discrete habitats for fish assemblages.     

Distribution and speciation of phosphorus along a salinity gradient in intertidal marsh sediments

We examined forms of solid phosphorus fractions in intertidal marsh sediments along a salinity (0–22%.) gradient in a river-dominated estuary and in a marine-dominated salt marsh with insignificant freshwater input. Freshwater marsh sediments had the highest ratio of organic N:P of between 28:1 and 47:1 mol:mol, compared to 211 to 311 molmol in the saltmarshes, which is consistent with a trend toward P-limitation of primary production in freshwater and N-limitation in salt marshes. However, total P concentration, 24.7±11.1mol P g dw^–1 (±1 SD) averaged over the upper meter of sediment, was greatest in the freshwater marsh where bioavailablity of P is apparently limited. In the freshwater marsh the greatest fraction of total P (24–51%.) was associated with humic acids, while the importance of humic-P decreased with increasing salinity to 1–23%. in the salt marshes. Inorganic P contributed considerably less to total sediment P in the freshwater marsh (15–40%.) than in the salt marshes (33–85%.). In reduced sediments at all sites, phosphate bound to aluminum oxides and clays was an important inorganic P pool irrespective of salinity. Inorganic P associated with ferric iron [Fe(III)] phases was most abundant in surface sediments of freshwater and brackish marshes, while Ca-bound P dominated inorganic P pools in the salt marshes. Thus, our results showed that particle-bound P in marsh sediments exhibited changes in chemical association along the salinity gradient of an estuarine system, which is a likely consequence of changes in ionic strength and the availability of iron and calcium.     

Spatial and temporal distribution patterns of the macrozoobenthos assemblage in the salt marshes of Tejo estuary (Portugal)

The present study focuses on the spatial and temporal distribution of the macroinvertebrate community of the salt marsh areas of the Tejo estuary, based on surveys conducted from autumn 1998 to summer 2000. Samples were collected quarterly in five different intertidal areas along an elevation gradient in: mudflats, creek mouths, creeks, pioneer salt marsh areas and middle marsh areas. A total of 36 benthic invertebrate taxa were identified. Insect larvae were the most well represented group, with 10 taxa identified. Oligochaetes and ostracods were the most numerically abundant taxa, whereas bivalves dominated in biomass. Benthic macroinvertebrate assemblages were dominated, both in number and biomass, by deposit feeders. Three distinct macroinvertebrate assemblages were distinguished along the elevation gradient, based on species presence, density and biomass: the unvegetated muddy areas with a macrobenthic assemblage composed mostly by infauna; the salt marsh pioneer areas of Spartina maritima in which several epibenthic taxa were found, as well as endobenthic taxa characteristic of muddy sediment; and the creek margins, with epifauna taxa such as insect larvae and crustaceans and a low abundance of benthic infauna. Total biomass in the unvegetated and Spartina areas was higher during spring and summer mainly due to the increase in biomass of Scrobicularia plana and Hydrobia ulvae. No decreases in the salt marsh macroinvertebrate biomass values were observed during the highest densities of their potential nektonic predators (summer). This fact might indicate that macroinvertebrates are not a limiting resource for the nektonic species, and that the natural biomass increment of these invertebrate species could be masking the predation/disturbance caused by the nektonic species.     

Feeding habits of young predatory fishes in marsh creeks situated along the salinity gradient of the Schelde estuary, Belgium and The Netherlands

Fish and macrobenthos were sampled in four different marshes along the salinity gradient of the Schelde estuary, Belgium/Netherlands, to investigate the importance of marsh creeks as foraging grounds for the dominant, larger fish species. The total density and biomass of all the main macrobenthic taxa (Corophium volutator, Nereis diversicolor, Oligochaeta, Macoma baltica and Heteromastus fliliformis) were measured. The feeding habits of the larger predatory fishes (Platichthys flesus, Dicentrarchus labrax) were investigated. Qualitative and quantitative stomach analyses included the calculation of different indices, showing the niche breadth (as diet diversity) and the niche overlap (as similarity between the predators diet) for this habitat. These analyses showed that the two most important benthic prey species for P. flesus were C. volutator and N. diversicolor. D. labrax preyed upon a wider range of species, including C. volutator, N. diversicolor, Crangon crangon, Carcinus maenas and Orchestia spp. The stomach diversity of D. labrax and P. flesus showed differences between the marshes although there was no consistent pattern in diet composition, reflecting the opportunistic nature of feeding by these large predators. The fullness indices of both flounder and sea bass did not differ significantly along the salinity gradient and the estimated minimum consumption by these predators did not indicate a top-down control of the macrobenthic community. The salt marsh creeks seem to provide excess food for the visiting fish species. The benthic prey was present in very high abundances, which may suggest that the typical nursery species such as C. crangon and C. maenas, and early juveniles of P. flesus, D. labrax and Pomatoschistus microps were not preyed upon significantly. This supports the hypothesis that salt marsh creeks provide good refuge areas for nursery species against predation by larger fish.     

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.     

闽江河口区淡水和半咸水潮汐沼泽湿地土壤产甲烷菌多样性

为认识盐度对河口潮汐沼泽湿地土壤产甲烷菌的影响,应用PCR-RFLP技术及测序分析对闽江河口区淡水-半咸水盐度梯度上分布的4个短叶茳芏潮汐沼泽湿地土壤产甲烷菌群落结构进行研究。闽江河口区短叶茳芏潮汐沼泽湿地土壤产甲烷菌群落结构受盐度影响明显,位于下洋洲和塔礁洲的短叶茳芏潮汐淡水沼泽湿地土壤产甲烷菌的香农-威纳多样性指数值分别为2.81和2.65,位于蝙蝠洲和鳝鱼滩的短叶茳芏潮汐半咸水沼泽湿地土壤产甲烷菌香农-威纳多样性指数值分别仅为2.33和2.27。系统发育分析表明:短叶茳芏沼泽湿地土壤产甲烷菌类群主要有甲烷杆菌目(Methanobacteriales),包括Methanobacterium、Methanobrevibacter和Methanobacteriaceae;甲烷微菌目(Methanomicrobiales),主要有Methanoregula,以及甲烷八叠球菌目(Methanosarcinales),主要有Methanosarcina和Methanococcoides。闽江河口区短叶茳芏潮汐淡水沼泽湿地土壤主要的优势产甲烷菌有Methanoregula、Methanosarcina和Methanobacterium,而短叶茳芏潮汐半咸水沼泽湿地土壤主要的优势产甲烷菌则转化为仅以Methanoregula为主。     

Effect of salinity-altering pulsing events on soil organic carbon loss along an intertidal wetland gradient: a laboratory experiment

Salinity changes resulting from storm surge, tides, precipitation, and stormwater run-off are common in coastal wetlands. Soil microbial communities respond quickly to salinity changes, altering the rate of soil organic carbon (SOC) loss and associated biogeochemical processes. This study quantified the impact of salinity-altering pulses on SOC loss, defined as microbial respiration (CO₂ flux) at high and low tide, CH₄ flux, and dissolved OC (DOC) release, in 3 intertidal wetlands (Jacksonville, FL, USA). Intact soil cores from a freshwater tidal, brackish, and salt marsh were exposed to simulated tides and 3 salinity pulsing events during a 53-day laboratory experiment. Soil and water physio-chemical properties, nutrient release, and microbial indicators were measured. Microbial respiration was the dominate pathway of SOC loss (>97 %). Soil hydraulic conductivity was greater in brackish and salt marshes and was critical to overall soil respiration. High tide CO₂ flux was greatest in the freshwater marsh (58 % of SOC loss) and positively correlated with DOC concentration; low tide CO₂ flux was greatest in brackish and salt marshes (62 and 70 % of SOC loss, respectively) and correlated with NH₄ ⁺ and microbial biomass. The freshwater marsh was sensitive to brackish pulses, causing a 112 % increase in respiration, presumably from accelerated sulfate reduction and N-cycling. SOC loss increased in the salt marsh pulsed with freshwater, suggesting freshwater run-off may reduce a salt marsh’s ability to keep-pace with sea level rise. Increased inundation from storm surges could accelerate SOC loss in freshwater marshes, while decreasing SOC loss in brackish and salt marshes.     

Nitrogen further promotes a dominant salt marsh plant in an increasingly saline environment

Aims Human alterations of the environment are combining in unprecedented ways, making predictions of alterations to natural communities a difficult and pressing challenge. Estuarine systems have been subject to a high degree of modification, including increased nitrogen (N) inputs and altered salinity, factors important in shaping estuarine plant communities. As human populations increase and the climate changes, both N and salinity levels are likely to increase in these coastal marshes. Our objective was to evaluate the interactive effects of N and salinity on US West Coast salt marsh species; in particular, the performance of the dominant species Sarcocornia pacifica (pickleweed) alone and in mixed species assemblages. We expected increased salinity to favor S. pacifica but that N enrichment could help maintain greater species richness through use of N in salinity tolerance mechanisms.Methods We crossed treatments of N (added or not) and salinity (salt added or not) in a field experiment at a salt marsh in the San Francisco Estuary, California, USA, in each of three habitats: (i) monotypic pickleweed on the marsh plain, (ii) monotypic pickleweed along channels and (iii) mixed assemblages along channels. In a greenhouse experiment, we crossed treatments of N (added or not) and salinity (at three levels to simulate brackish to saline conditions) in (i) pots of pickleweed only and (ii) the same species mix as in the field.Important findings N addition doubled S. pacifica biomass and branching in both channel and marsh plain habitats regardless of salinity and greatly increased its dominance over Distichlis spicata and Jaumea carnosa in mixed assemblages along channels. In the greenhouse, S. pacifica biomass increased 6- to 10-fold with N addition over the range of salinities, while D. spicata and J. carnosa biomass increased with N addition only at lower salinity levels. Thus, while localized management could influence outcomes, expected overall increases in both N and salinity with human population growth and climate change are likely to enhance the production of S. pacifica in US West Coast marshes while reducing the diversity of mixed species assemblages. This decline in diversity may have implications for the resilience of marshes already subject to multiple stressors as the climate changes.     

Fifteen Years of Vegetation and Soil Development after Brackish-Water Marsh Creation

Aboveground biomass, macro‐organic matter (MOM), and wetland soil characteristics were measured periodically between 1983 and 1998 in a created brackish‐water marsh and a nearby natural marsh along the Pamlico River estuary, North Carolina to evaluate the development of wetland vegetation and soil dependent functions after marsh creation. Development of aboveground biomass and MOM was dependent on elevation and frequency of tidal inundation. Aboveground biomass of Spartina alterniflora, which occupied low elevations along tidal creeks and was inundated frequently, developed to levels similar to the natural marsh (750 to 1,300 g/m²) within three years after creation. Spartina cynosuroides, which dominated interior areas of the marsh and was flooded less frequently, required 9 years to consistently achieve aboveground biomass equivalent to the natural marsh (600 to 1,560 g/m²). Aboveground biomass of Spartina patens, which was planted at the highest elevations along the terrestrial margin and seldom flooded, never consistently developed aboveground biomass comparable with the natural marsh during the 15 years after marsh creation. MOM (0 to 10 cm) generally developed at the same rate as aboveground biomass. Between 1988 and 1998, soil bulk density decreased and porosity and organic C and N pools increased in the created marsh. Like vegetation, wetland soil development proceeded faster in response to increased inundation, especially in the streamside zone dominated by S. alterniflora. We estimated that in the streamside and interior zones, an additional 30 years (nitrogen) to 90 years (organic C, porosity) are needed for the upper 30 cm of created marsh soil to become equivalent to the natural marsh. Wetland soil characteristics of the S. patens community along upland fringe will take longer to develop, more than 200 years. Development of the benthic invertebrate‐based food web, which depends on organic matter enrichment of the upper 5 to 10 cm of soil, is expected to take less time. Wetland soil characteristics and functions of created irregularly flooded brackish marshes require longer to develop compared with regularly flooded salt marshes because reduced tidal inundation slows wetland vegetation and soil development. The hydrologic regime (regularly vs. irregularly flooded) of the “target” wetland should be considered when setting realistic expectations for success criteria of created and restored wetlands.     

Estuarine gradients and the growth and development of Agapanthia villosoviridescens, (Coleoptera), a stem-borer of the salt marsh halophyte Aster tripolium

Summary Phytophagous insects of estuarine salt marshes which live inside their host plants are not directly exposed to estuarine gradients. Host plant quality, however, may change along the estuary as a result of the direct effects of these gradients; as a consequence growth and development of endophagous insects may be influenced. The results of a study of the life cycle of Agapanthia villosoviridescens (Coleoptera, Cerambycidae), a stem-borer of the halophyte Aster tripolium, on three salt marshes along the Westerschelde estuary (the Netherlands) are in line with this hypothesis. It was shown that in upstream direction (1) mean larval weights were consistently higher during the entire period of larval development; (2) the percentage of late instars on a given sampling date generally was higher; (3) the percentage of larvae which underwent successful metamorphosis increased. Furthermore, (4) dry weight of the imagos was highest on the least saline marsh. The effects of estuarine gradients on the Aster host plants was indicated by differences in growth and chloride content between the populations of the three marshes. The non-overlapping geographic distribution of Agapanthia villosoviridescens and its host plant Aster tripolium on the Westerschelde salt marshes may be related to the effects of estuarine gradients on the suitability of the host plant.     

Effects of regular salt marsh haying on marsh plants, algae, invertebrates and birds at Plum Island Sound, Massachusetts

The haying of salt marshes, a traditional activity since colonial times in New England, still occurs in about 400 ha of marsh in the Plum Island Sound estuary in northeastern Massachusetts. We took advantage of this haying activity to investigate how the periodic large-scale removal of aboveground biomass affects a number of marsh processes. Hayed marshes were no different from adjacent reference marshes in plant species density (species per area) and end-of-year aboveground biomass, but did differ in vegetation composition. Spartina patens was more abundant in hayed marshes than S. alterniflora, and the reverse was true in reference marshes. The differences in relative covers of these plant species were not associated with any differences between hayed and reference marshes in the elevations of the marsh platform. Instead it suggested that S. patens was more tolerant of haying than S. alterniflora. Spartina patens had higher stem densities in hayed marshes than it did in reference marshes, suggesting that periodic cutting stimulated tillering of this species. Although we predicted that haying would stimulate benthic chlorophyll production by opening up the canopy, we found differences to be inconsistent, possibly due to the relatively rapid regrowth of S. patens and to grazing by invertebrates on the algae. The pulmonate snail, Melampus bidendatus was depleted in its δ¹³C content in the hayed marsh compared to the reference, suggesting a diet shift to benthic algae in hayed marshes. The stable isotope ratios of a number of other consumer species were not affected by haying activity. Migratory shorebirds cue in to recently hayed marshes and may contribute to short term declines in some invertebrate species, however, the number of taxa per unit area of marsh surface invertebrates and their overall abundances were unaffected by haying over the long term. Haying had no impact on nutrient concentrations in creeks just downstream from hayed plots, but the sediments of hayed marshes were lower in total N and P compared to references. In sum, haying appeared to affect plant species composition but had only short-term affects on consumer organisms. This contrasts with many grassland ecosystems, where an intermediate level of disturbance, such as by grazing, increases species diversity and may stimulate productivity. From a management perspective, periodic mowing could be a way to maintain S. patens habitats and the suite of species with which they are associated.     

Similarity between seed banks and above‐ground vegetation along a salinity gradient

Abstract. Zonation of above‐ground vegetation often occurs in salt marshes along salinity and moisture gradients. The above‐ground vegetation and seed bank in four physiognomically different vegetation zones in a salt marsh were compared to determine their level of similarity using percent similarity as a distance measure. 10‐m transects were established along a salinity gradient through four different vegetation zones; a Salicornia zone, a Salicornia‐Atriplex zone, an Atriplex zone and an Atriplex‐Hordeum zone. A UPGMA cluster analysis demonstrated that the above‐ground vegetation was not usually highly correlated with the seed bank composition of zonal communities. Since seeds of these annual salt marsh species occurred in all zones, the levels of salt stress may be the main factor determining which species were found in the above‐ground vegetation.     

Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient

An invasive variety of Phragmites australis (Poaceae, common reed), the M haplotype, has been implicated in the spread of this species into North American salt marshes that are normally dominated by the salt marsh grass Spartina alterniflora (Poaceae, smooth cordgrass). In some European marshes, on the other hand, Spartina spp. derived from S. alterniflora have spread into brackish P. australis marshes. In both cases, the non-native grass is thought to degrade the habitat value of the marsh for wildlife, and it is important to understand the physiological processes that lead to these species replacements. We compared the growth, salt tolerance, and osmotic adjustment of M haplotype P. australis and S. alterniflora along a salinity gradient in greenhouse experiments. Spartina alterniflora produced new biomass up to 0.6 M NaCl, whereas P. australis did not grow well above 0.2 M NaCl. The greater salt tolerance of S. alterniflora compared with P. australis was due to its ability to use Na(+) for osmotic adjustment in the shoots. On the other hand, at low salinities P. australis produced more shoots per gram of rhizome tissue than did S. alterniflora. This study illustrates how ecophysiological differences can shift the competitive advantage from one species to another along a stress gradient. Phragmites australis is spreading into North American coastal marshes that are experiencing reduced salinities, while Spartina spp. are spreading into northern European brackish marshes that are experiencing increased salinities as land use patterns change on the two continents.     

Habitat selection of wintering passerines in salt marshes of the German Wadden Sea

The salt marshes of the Wadden Sea are important wintering areas for some species of granivorous passerines, which have declined considerably since the 1960s. We investigated the habitat choice of all wintering passerines in eight study areas in German salt marshes with special consideration of human impact on these habitats. Granivorous species that almost exclusively winter in salt marshes, Shorelark (Eremophila alpestris), Snow Bunting (Plectrophenax nivalis) and Twite (Carduelis flavirostris) were concentrated in the lower salt marsh vegetation and in the driftlines, while all other species preferred the high upper salt marsh communities, although Rock Pipits (Anthus petrosus littoralis) fed in muddy areas along ditches. Shorelarks switched habitat in conditions where seeds were scarce to feed instead on arthropods in upper salt marshes. Intensively sheep-grazed upper salt marshes resemble lower salt marshes in their vegetation and were therefore mainly visited by Shorelarks, Snow Buntings and Twites. In winter, the driftline is preferred by the two former species, while in autumn and spring more birds foraged in the salt marshes. Twites prefer to feed mainly on seeds of Salicornia. Areas with S. europaea are visited mainly in late autumn and early winter, while areas with S. stricta are used throughout the winter because of a steady supply of seeds. Several years after embankment, polders are hardly used any more by the lower salt marsh species as the habitat changes into freshwater marshes. Large embankment projects since the early 1960s have included salt marshes and intertidal flats, and the resultant loss of habitat is responsible for the decline of lower salt marsh species. For other passerine species the effects of reclamation are unknown. The effects of intensified grazing on the wintering populations of Shorelark, Snow Bunting and Twite are still unresolved. Although grazing supports lower salt marsh vegetation, the seed production per plant is much lower there and some important seed producers hardly occur. Since grazing was reduced and embankment projects have been stopped, the salt marsh areas (especially lower salt marshes) have increased and so have the wintering populations of Shorelark, Snow Bunting and Twite. For the other species, the consequences of habitat changes are unknown, although it is suggested that reduced grazing will support them. Reducing the human impact on salt marshes will, in the long run, probably lead to a natural salt marsh with much variety in elevation and in its corresponding vegetation and bird communities. Meanwhile, management by grazing might be required in parts of the salt marshes.     

Zonation of spiders (Araneae) and carabid beetles (Coleoptera: Carabidae) in island salt marshes at the North Sea coast

The specific communities of spiders and carabid beetles of island salt marsh habitats of the East Frisian Island chain at the German North Sea coast were investigated. During the vegetation periods of 1997 and 1998 three pitfall trapping transects were installed on the islands of Borkum and Wangerooge. Within the salt marshes, transects extended from 0 m to 175 m. Elevation gradients varied between 10 cm and 232 cm above MHT (mean high tide). On Borkum, 35 traps were exposed in two transects, on Wangerooge 25 traps were placed in one transect. Three to five elevations above MHT were investigated per transect, each one with five traps. Highest species numbers were recorded in the higher elevated salt marshes. In contrast, highest activity values were noticed in the medium elevated salt marshes. Within both groups, spiders and carabids, four communities were distinguished by indirect gradient analysis. Indicator species were assigned to the different communities that were mainly assorted to different elevations of the salt marshes. Thus, the communities of both taxa corresponded well to the vegetational formations. The importance of sea level rise for structuring the communities of salt marsh arthropods is discussed. Overall, still great uncertainties exist on how arthropod communities and salt marshes themselves will develop.     

Salt marshes along the coast of The Netherlands

The area of salt marshes does no longer increase. The recent erosion coincides with a rise in MHT-level in the last 25 years. Despite the decrease in area, sedimentation continues, especially in the lower salt marsh, which acts as a sink of nitrogen. Assimilation and mineralization of nitrogen are in balance in most plant communities along the gradient from lower to higher salt marshes. Mineralization of nitrogen increases towards the higher salt marsh, whereas the above-ground production and the mean nitrogen content of plants decrease. There is a positive correlation between quality of food plants in salt marshes and breeding success of Brent geese in the arctic tundra. Sedimentation on mainland salt marshes can compensate for the expected sea level rise. This is not the case for island salt marshes, if the relative sea level rise is more than 0.5–1.0 cm yr⁻¹. The natural succession on salt marshes results in an accumulation of organic material, which is related to the dominance of single plant species. It is not clear to which extent this process is enhanced by eutrophication from acid deposition and seawater. Human exploitation of unprotected salt marshes is old and heavy in the system of mound settlements. Reclamation rates by dikes in the last centuries were higher than the rate of area increase. Grazing by cattle as a management practice results in both a higher plant species-richness and community diversity than abandoning; hay-making is intermediate, but shows less structural diversity than grazing with low stocking density. The invertebrate fauna is favoured by a short period of abandoning, but eventually characteristic salt marsh invertebrates are replaced by inland species. Many bird species prefer grazed salt marshes. The final section gives some perspectives. Provided that no further embankments take place the optimal nature management option for plants and animals is a vegetation pattern, which includes areas with a low canopy (grazed) and areas with a tall canopy.