The ecology of temperate salt-marsh fucoids. I. Occurrence and distribution of Ascophyllum nodosum ecads

The distribution of several free-living Ascophyllum nodosum ecads, including scorpioides and mackaii, in a temperate salt marsh is described. Morphological characterization of these ecads by the presence or absence of air bladders and reproductive receptacles, and size and shape of fronds, indicated that several free-living forms occur throughout the marsh. Plants resembling the ecad mackaii were more closely associated with exposed areas along the low-tide regions, while scorpioides-type ecads prevailed on the Spartina alterniflora-dominated marsh banks and flats in the middle and upper intertidal regions of the marsh. Maximum biomass of ecads occurred during the spring months in the absence of S. alterniflora, whereas minimum ecad biomass was associated with maximum S. alterniflora densities in the late summer and fall months. Morphological differentiation of dwarf-type ecads was related to environmental components other than exposure to low and/or fluctuating salinities.This research was suported by research grant AG-375 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Seasonal variations in salt-marsh macroalgae photosynthesis. I. Ascophyllum nodosum ecad scorpioides

Photosynthesis in whole plants of the intertidal alga Ascophyllum nodosum ecad scorpiodes was evaluated by measuring ¹⁴C-uptake under a variety of light intensities and approximately monthly intervals during a 15 month study. Photosynthetic rates were determined in terms of dry weight, pigment content and uptake into ethanolsoluble and insoluble fractions. The specimens, naturally acclimated to in situ light intensities and temperatures, exhibited photosynthetic responses to light intensity which differed with time of year. Maximum photosynthetic potential occurred during the spring months and minimum potential occurred during late summer and winter months. Variations in photosynthetic potential were closely related to seasonal changes in field biomass. Both photosynthetic potential and biomass were inversely related to growth patterns of the salt-marsh phanerogam Spartina alternilora.This research was supported by Research Grant AG-375 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Photosynthesis and respiration of exposed salt-marsh fucoids

Photosynthesis and respiration of the salt-marsh fucoids Ascophyllum nodosum ecad scorpioides and Fucus vesiculosus were investigated using an infrared CO₂ gas analyzer under a variety of light intensities, temperatures, and levels of desiccation while the algae were exposed to the atmosphere. Results indicated that net photosynthesis (0.5 to 2.0 mg C/g dry weight/h) saturated rapidly at light intensities (0.1 to 0.2 g cal/cm²/min) which were approximately 10 to 50% of the daily summer maximum intensities for algae found under phanerogam (Spartina alterniflora) canopies. Desiccation exhibited the most pronounced effect on photosynthesis, which increases slightly between 0 and 25% water loss, levels off, and decreases sharply at water losses greater than 50%. Dark respiration (0.1 to 0.3 mg C/g dry weight/h) is also inhibited by desiccation. Both species of algae appear to be broadly adapted to all three parameters investigated.This research was supported by research grants AG-375 and BO 38018 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).Communicated by M.R. Tripp, Newark     

Effects of chronic oil pollution on a salt-marsh grass community

A small, naturally defined saltmarsh in Chesapeake Bay (USA) was repeatedly dosed with a No. 2 fuel oil. The production, decay rates, physical characteristics and hydrocarbon content of the dominant grass species present in the marsh were routinely monitored. Spartina alterniflora was the only grass species displaying effects of the oil dosing; this is attributed to the grass's location in the marsh. A substantial portion of the S. alterniflora in the marsh was killed by the oil dosing; the remaining portion evidenced sublethal effects including delayed development in the spring, increased density, and reduced mean weight per stem. The second annual cohort of shoots, usually produced in late summer and early fall, was suppressed almost entirely. Decay rates of S. alterniflora in the oiled marsh were higher and peaked later than decay rates in a control marsh. Oil which entered the roots and rhizomes of dead S. alterniflora was retained in a relatively undergraded state for at least 7 months after dosing stopped.Virginia Institute of Marine Science Contribution No. 945     

Growth of the salt marsh periwinkleLittoraria irrorata on fungal and cordgrass diets

The growth of the salt marsh periwinkleLittoraria irrorata (collected from Sapelo Island, Georgia in 1991, initial shell length 6.2 to 11.5 mm) on various diets was measured. Growth was highest on a diet of standing-dead leaves ofSpartina alterniflora. Periwinkles provided with marsh sediment, yellow-green, sterile, or bacteria-colonized leaves lost organic mass. Fungal-colonized leaves and pure mycelia of fungi common on standing-dead leaves allowed intermediate growth. Growth onS. alterniflora-based diets was negatively correlated with the phenolics content of the food, and positively correlated with its lipid content. No correlation was found between growth and protein content. The digestibility ofS. alterniflora leaves, estimated with the acid-insoluble ash technique, was highest when yellow-green leaves were used. Colonization by fungi or bacteria caused it to decline. ForS. alterniflora-based diets, growth rates were positively correlated with the amount of time spent on the food.     

Root respiration and oxygen flux in salt marsh grasses from different elevational zones

Plants growing in waterlogged environments are subjected to low oxygen levels around submerged tissues. While internal oxygen transport has been postulated as an important factor governing flooding tolerance, respiration rates and abilities to take up oxygen under hypoxic conditions have been largely ignored in plant studies. In this study, physiological characteristics related to internal oxygen transport, respiration, and oxygen affinity were studied in low intertidal marsh species (Spartina alterniflora and S. anglica) and middle to high intertidal species (S. densiflora, S. patens, S. foliosa, a S. alterniflora × S. foliosa hybrid, S. spartinae, and Distichlis spicata). These marsh plants were compared to the inland species S. pectinata and the crop species rice (Oryza sativa), corn (Zea mays), and oat (Avena sativa). Plants were grown in a greenhouse under simulated estuarine conditions. The low marsh species S. anglica was found to transport oxygen internally at rates up to 2.2 μmol O₂ g fresh root weight⁻¹ h⁻¹. In contrast, marsh species from higher zones and crop species were found to transport significantly less oxygen internally, although rice plants were able to transport 1.4 μmol g⁻¹ h⁻¹. Under hypoxic conditions, low marsh species were better able to remove dissolved oxygen from the medium compared to higher marsh species and crops. The oxygen concentration at which respiration rates declined due to limited oxygen (P _crit) was significantly lower in low marsh species compared to inland and crop species; P _crit ranged from <4 μM O₂ in the low marsh species S. anglica up to 20 μM in the inland species corn. Flooding-sensitive crop species had significantly higher aerobic respiration rates compared to flooding-tolerant species in this study. Crop species took up 3.6–6.7 μmol O₂ g⁻¹ h⁻¹ while all but one marsh species took up <3.5 μmol O₂ g⁻¹ h⁻¹. We conclude that oxygen transport, aerobic demand, and oxygen affinity all play important and interrelated roles in flood tolerance and salt marsh zonation.     

Macroalgae as habitat patch islands for Scutellidium lamellipes (Copepoda: Harpacticoida) and Ampithoe tea (Amphipoda: Gammaridae)

Scutellidium lamellipes and Ampithoe tea live in patches formed by the brown alga Pelvetia fastigiata. The distributions and abundance of both crustaceans were examined on P. fastigiata obtained during 1975 and 1976 from 7 collecting and 6 experimental sites in the mid-intertidal zone at La Jolla, California (USA). At each collecting site, the abundance of each crustacean generally increases with increasing plant size. Following defaunation and transplantation of algae, abundances of both crustaceans on individual plants appear to be governed by equilibria determined by in situ reproduction, immigration and emigration. Immigration rates of both animal species decrease significantly with increasing distance from areas occupied by P. fastigiata. During short-term experiments, neither crustacean was able to establish colonies on defaunated plants placed across a 30 m P. fastigiata-free zone and both species would have to reproduce to persitst on plants transplanted to that site. More frequent immigration and reproduction (and, for S. lamellipes at least, decreasing emigration rates with increasing patch size) would increase abundances and decrease probabilities of local extinction on larger and/or less distant plants. However, even within sites with isolated plants, the smallest plants often are not inhabited by either crustacean. The densitics of both species differ with plant size and distribution. Densities of S. lamellipes are greatest within a large aggregation of P. fastigiata, whereas those of A. tea are greatest on moderate-sized isolated plants at mid-tide levels.     

Food resources of postlarval brown shrimp (Penaeus aztecus) in a Texas salt marsh

Field and laboratory experiments were conducted to identify the sources of food in the natural diet of postlarval brown shrimp (Penaeus aztecus Ives). A series of enclosures placed in East Lagoon (29°20N; 94°45W) on Galveston Island, Texas, USA, in May 1985, were used to evaluate the individual and combined contribution of Spartina alterniflora detritus, epiphytes of S. alterniflora, plankton, and demersal fauna in terms of differences in shrimp growth and carbon assimilation (stable carbon-isotope analysis). Demersal fauna (harpacticoid copepods, amphipods, tanaids and polychaete annelids), and plankton (>0.095 mm) accounted for approximately 53 and 47% of the growth of the postlarvae (11 to 22 mm rostrum-telson length), respectively, while the autochthonous plant substrates, S. alterniflora detritus and epiphytes, contributed little. Laboratory experiments confirm that a mixed diet consisting of both animal protein and phytoplankton promotes maximum growth. Our results indicate that plankton may be an important allochthonous source of carbon contributing to the growth and development of shrimp in the salt marsh.     

Effects of Invasive Cordgrass on Presence of Marsh Grassbird in an Area where It Is Not Native

The threatened Marsh Grassbird (Locustella pryeri) first appeared in the salt marsh in east China after the salt marsh was invaded by cordgrass (Spartina alterniflora), a non‐native invasive species. To understand the dependence of non‐native Marsh Grassbird on the non‐native cordgrass, we quantified habitat use, food source, and reproductive success of the Marsh Grassbird at the Chongming Dongtan (CMDT) salt marsh. In the breeding season, we used point counts and radio‐tracking to determine habitat use by Marsh Grassbirds. We analyzed basal food sources of the Marsh Grassbirds by comparing the δ¹³C isotope signatures of feather and fecal samples of birds with those of local plants. We monitored the nests through the breeding season and determined the breeding success of the Marsh Grassbirds at CMDT. Density of Marsh Grassbirds was higher where cordgrass occurred than in areas of native reed (Phragmites australis) monoculture. The breeding territory of the Marsh Grassbird was composed mainly of cordgrass stands, and nests were built exclusively against cordgrass stems. Cordgrass was the major primary producer at the base of the Marsh Grassbird food chain. Breeding success of the Marsh Grassbird at CMDT was similar to breeding success within its native range. Our results suggest non‐native cordgrass provides essential habitat and food for breeding Marsh Grassbirds at CMDT and that the increase in Marsh Grassbird abundance may reflect the rapid spread of cordgrass in the coastal regions of east China. Our study provides an example of how a primary invader (i.e., cordgrass) can alter an ecosystem and thus facilitate colonization by a second non‐native species. Efectos de Spartina alterniflora Invasora Sobre Locustella pryeri en un Área Donde No Es Nativa     

Flax Pond ecosystem study: Exchanges of phosphorus between a salt marsh and the coastal waters of Long Island Sound

The exchanges of phosphorus between the Flax Pond, a tidal Spartina alterniflora marsh on the north shore of Long Island (USA) and Long Island Sound were measured over 18 months. Phosphate was exported from the marsh from May through December and imported during the remainder of the year. Organic phosphorus appears to be accumulated in all seasons, but the yearly phosphorus budget of the marsh is approximately balanced despite the accumulation of about 6 mm of sediment annually.     

Analyzing and quantitatively evaluating the organic matter source at different ecologic zones of tidal salt marsh, North Jiangsu Province, China

In order to explore the effect of different ecological zones and their above plants in the organic matter cycling of the whole tidal salt marsh, indicators such as total organic carbon (TOC), total nitrogen (TN), C/N ratio, δ¹³C and δ¹⁵N of surface, core sediments, and plants of tidal salt marshes in North Jiangsu Province are analyzed. Subsequently, distribution regularities of these measurement indicators are discussed, and the biogeochemistry processes between sediments and plants are also analyzed. Lastly, the organic matter sources of different ecologic zones in tidal salt marsh are evaluated, and the organic matter accumulations in different ecologic zones induced by their plants are also compared. These results indicate that TOC, TN, C/N ratio and δ¹³C showed obvious zonal distribution. The organic matter sources are dominated by marine input in the silt flat, artemisia schrenkiana flat, and the transition zone between silt and spartina alterniflora flat, and are controlled by terrigenous input in spartina alterniflora flat. Spartina alterniflora plays an important role in the accumulation of organic matter in the whole tidal salt marshes ecosystem. In the study area, the annually increased TOC, organic matter and TN in the spartina alterniflora, artemisia schrenkiana and reed flats reach 6,451, 12,043 and 536 t, respectively. The amount of TOC, organic matter and TN accumulated in the spartina alterniflora flat is more than that in other ecological zones, which shows that the spartina alterniflora flat exert a non-replaceable effect on the material cycle and exchange in the whole tidal salt marshes ecosystem.     

Artificial wounding decreases plant biomass and shoot strength of the brown seaweed Ascophyllum nodosum (Fucales, Phaeophyceae)

Brown seaweeds can serve as habitat and food for a number of mesoherbivore species, but the effect of mesoherbivore grazing on the fitness of individual seaweed plants is not well documented. Here we investigate how mechanical wounding, which mimics the grazing damage made by the herbivorous gastropod Littorina obtusata, affects the biomass change, apical growth, and individual shoot strength of the fucoid seaweed species Ascophyllum nodosum both in natural field populations and in the laboratory. We did not detect a significant effect of wounding on apical shoot growth in the laboratory, but wounding decreased the plant biomass in half of the studied A. nodosum populations in the field. Furthermore, we found that the strength of individual A. nodosum shoots was significantly lower when compared with unwounded control shoots initially after wounding. However, after 11 days incubation in the laboratory, the strength of the wounded shoots had increased almost to the level of the control shoots, although there was still a significant difference in the force required to break the shoots. These results indicate that wounding can increase the tissue loss (and thereby decrease the fitness) of A. nodosum plants in natural populations, probably through weakening of individual shoots. However, A. nodosum appears to have a mechanism to strengthen the shoots after mechanical wounding, and therefore, the timing of damage (e.g. prior to a storm) probably determines the amount of plant tissue lost through wave-induced forces.     

Effect of diurnal variations in natural irradiance on the apical length growth and light saturation of growth in five species of benthic macroalgae

The diurnal variation in apical length growth rate of five benthic macroalgae from the Swedish West Coast (Fucus spiralis L., F. vesiculosus L., F. serratus L., Ascophyllum nodosum (L.) Le Jolis, and Chondrus crispus Stackhouse) was measured during 1.5 to 2.0-h intervals for 3 to 5 d in September 1984. All species showed a maximum in growth rate in the morning, followed by a continuous decline throughout the rest of the day (most pronounced in F. serratus, A. nodosum and C. crispus). By comparing the growth rates with the average irradiance during the same interval, saturation irradiances for length growth in natural daylight regime were estimated; 60 to 70 Wm^-2 for F. spiralis and F. vesiculosus, 90 to 100 Wm^-2 for A. nodosum, 30 to 40 Wm^-2 for F. serratus and 10 to 20 Wm^-2 for C. crispus. Average growth rates during the dark hours ranged from 33% of average day growth rates in A. nodosum to 63% in C. crispus, respectively.     

Seasonal variations in salt-march macroalgae photosynthesis. II. Fucus vesiculosus and Ulva lactuca

Photosynthesis in whole plants of the salt-marsh algae Fucus vesiculosus and Ulva lactuca was evaluated by ¹⁴C-uptake under a variety of light intensities at approximately mately monthly intervals during a 15-month study. Photosynthetic capacity in both species was closely related to seasonal irradiation patterns and changes in field biomass. Maximum photosynthesis occurred in the spring and summer months. Photosynthesis on a dry weight basis was higher in U. lactuca, while photosynthesis on a chlorophyll a basis was equal in both species. Photosynthetic capacity was inversely related to pigment content. Maximum chlorophyll a concentrations occurred during the winter. Frond profile studies in F. vesiculosus indicated that apices always exhibited greatest photosynthetic capacity. Uptake of ¹⁴C into ethanol-soluble and insoluble fractions was different in each species. F. vesiculosus showed greater activity in the ethanol-soluble fraction while U. lactuca exhibited greater activity in ethanol-insoluble fractions.This research was supported by Research Grant AG-375 from the National Science Foundation and, in part, by the State University of New York Research Foundation and the Energy Research and Development Administration (ERDA).     

Acetylene reduction (nitrogen fixation) in a Delaware,USA salt marsh

Acetylene reduction (nitrogen fixation) was measured in several vegetational areas in a Delaware, USA salt marsh. Samples were collected for 1 yr and the results showed a seasonally variable pattern of acetylene reduction at all stations. Peak rates were generally recorded during the later summer and early fall (September–October). The seasonality was influenced mainly, although not exclusively, by the soil temperature. In addition, samples collected in short Spartina alterniflora stands exhibited rates which were up to 20-fold higher than those found in samples from tall S. alterniflora stands. Over 50% of the total yearly ethylene production occurred from mid-August until the beginning of December at the tall and short S. alterniflora stations. Maximum activity occurred at 5 cm depth for all stations. Surface activity accounted for only 3–4% of the total measured in the top 20 cm. Addition of glucose or mannitol resulted in considerable increases in activity, thus suggesting that heterotrophic acetylene reduction is carbon and/or energy limited. The results obtained in this study indicate that the measured rates are only potential rates and that considerable caution must be used in extrapolating from acetylene reduction rates to nitrogen fixation rates in situ.     

Differing distributions of potentially competing amphipods,copepods and gastropods among specimens of the intertidal alga Pelvetia fastigiata

Possible competitive interactions are inferred from the distributions and abundances of species within three invertebrate groups (Hyale spp., Ampithoe spp. and thallusdwelling species) among unmanipulated and experimental specimens of the brown alga Pelvetia fastigiata collected between 1974 and 1976, inclusive, at La Jolla, California, USA. During the fall, when the vagile amphipods Hyale grandicornis and H. frequens were relatively abundant, they were found mostly on specimens of P. fastigiata at upper mid-intertidal and mid-tide levels, respectively. H. grandicornis often dispersed to mid-tide levels, but the two species cooccurred on few plants. Although H. frequens infrequently occupied P. fastigiata in the spring, H. grandicornis still generally sheltered in upper mid-intertidal plants, in part, because the net reproduction rates of H. grandicornis were low when space was available. The tubiculous amphipod A. tea usually was the only Ampithoe species on P. fastigiata; however, A. lindbergii and A. pollex sometimes dispersed to and released broods on this alga, particularly during the early summer when abundances of A. tea were relatively low. Similarly, net immigration rates of other thallus-dwelling copepods were relatively high when abundances of Scutellidium lamellipes were low. Moreover, several copepods were most abundant on different specimens of small isolated plants, and thallus-dwelling gastropods on larger plants at sites with relatively low densities of S. lamellipes on P. fastigiata; such copepods and gastropods generally were not abundant on aggregated plants sheltering large numbers of S. lamellipes. Such differences suggest that competition for food or space is among the factors causing negative correlations in the densities of Ampithoe spp. and thallusdwelling species on P. fastigiata.     

Response and resilience of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> to sudden dieback

We measured an array of biophysical and spectral variables to evaluate the response and recovery of Spartina alterniflora to a sudden dieback event in spring and summer 2004 within a low marsh in coastal Virginia, USA. S. alterniflora is a foundation species, whose loss decreases ecosystem services and potentiates ecosystem state change. Long-term records of the potential environmental drivers of dieback such as precipitation and tidal inundation did not evidence any particular anomalies, although Hurricane Isabel in fall 2003 may have been related to dieback. Transects were established across the interface between the dieback area and apparently healthy areas of marsh. Plant condition was classified based on ground cover within transects as dieback, intermediate and healthy. Numerous characteristics of S. alterniflora culms within each condition class were assessed including biomass, morphology and spectral attributes associated with photosynthetic pigments. Plants demonstrated evidence of stress in 2004 and 2005 beyond areas of obvious dieback and resilience at a multi-year scale. Resilience of the plants was evident in recovery of ground cover and biomass largely within 3 y, although a small remnant of dieback persisted for 8 y. Culms surviving within the dieback and areas of intermediate impact had modified morphological traits and spectral response that reflected stress. These morphometric and spectral differences among plant cover condition classes serve as guidelines for monitoring of dieback initiation, effects and subsequent recovery. Although a number of environmental and biotic parameters were assessed relative to causation, the reason for this particular dieback remains largely unknown, however.     

Transgenic Spartina alterniflora for phytoremediation

Perennial monoculture forming grasses are very important natural remediators of pollutants. Their genetic improvement is an important task because introduction of key transgenes can dramatically improve their remediation potential. Transfer of key genes for mercury phytoremediation into the salt marsh cordgrass (Spartina alterniflora) is reported here. S. alterniflora plays an important role in the salt marsh by cycling of␣elements, both nutrients and pollutants, protects the coastline from erosion, is a keystone species in the␣salt marsh supporting a large food web, which in turn supports a significant segment of economy, including tourism, has an impact on cloud formation and consequently on global weather, and is thus an ecologically important species relevant for our life-support systems. Embryogenic callus of S. alterniflora was co-inoculated with a pair of Agrobacterium strains LBA4404 carrying the organomercurial lyase (merB) and mercuric reductase (merA) genes, respectively, in order to co-introduce both the merA and the merB genes. Seven stable geneticin resistant lines were recovered. The presence of merA and merB genes was verified by PCR and Southern blotting. All but one transgenic lines contained both the merA and the merB sequences proving that co-introduction into Spartina of two genes from separate Agrobacterium strains is feasible and frequent, although the overall frequency of transformation is low. Northern blotting showed differences in relative expression of the two transgenes among individual transformants. The steady-state RNA levels appeared to correlate with the phenotype. Line #7 showed the highest resistance to HgCl₂ (up to 500 μM), whereas line #3 was the most resistant to phenylmercuric acetate (PMA). Wild-type (WT) callus is sensitive to PMA at 50 μM and to HgCl₂ at 225 μM.     

Effects of tidally mediated litter moisture content on decomposition of Spartina alterniflora and S. patens

Relationships between flooding frequency, flooding duration, litter moisture levels, and litter decay rates were investigated across the natural hydrologic gradient common to intertidal salt marshes. The effects on litter decay of natural and experimental alterations of litter moisture content were assessed in both field litterbag experiments (conducted in a southern New Jersey salt marsh from 1989 to 1990) and laboratory incubations (1990). Overall, tidally mediated litter moisture content was the dominant factor controlling litter decay throughout the vegetated marsh. Rates of carbon loss were most closely related to litter moisture levels (r=0.84), which were directly related to flooding frequency (r=0.66) and duration (r=0.63). Litter moisture levels were related to elevation within the tidal range due to increasing surface levation from creekbank to high marsh (ca. 54 cm) and height of litter above the sediment surface. Carbon losses from litter of short and tall form Spartina alterniflora Loisel. and S. patens (Aiton) Muhl. along the marsh elevation gradient indicate that while some of the variations in decay rates may be due to litter type, litter moisture accounted for most of the observed variation between marsh zones and within each litter type. Mousture levels are also affected by the water retention capacity of each litter type, which may also secondarily influence decay rates. Short-term incubations of litter indicated that CO₂ evolution was positively related to moisture content with negligible respiration at moisture levels below 15% (fresh mass), increasing to a maximum between 65 and 75% depending upon litter type. Since most Spartina spp. litter remains above the marsh surface where it maintains a lower moisture content than surface litter, the use of surface litterbags may overestimate rates of carbon loss in some systems. In addition, since changes in elevation of only a few centimeters had significant effects on both litter moisture levels and decay rates, slight changes in tidal regime may have important consequences for organic matter cycling in salt marshes by affecting litter decomposition processes.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.