Secondary succession on a high salt marsh at different grazing intensities

Succession occurs on a large part of German salt marshes following abandonment or reduction of grazing. Its speed and effect on the biodiversity of salt marshes has been discussed in the literature. Permanent plot studies show site-dependent differences in successional outcome. If grazing is to be continued, there is uncertainty about the stocking rates that are optimal for the conservation of plant diversity. We present an eight year permanent plot study with different grazing treatments on a high salt marsh at the marsh island of Hallig Langeness, Germany, which is enclosed by a summer dike. The study site is owned by WWF. Successional outcomes on the permanent plots depend on grazing intensity and depth of the groundwater table which is correlated with soil salinity. Intermediate stocking rates support a mixture of halophytes and glycophytes, the relative proportion of which depends on the depth of the groundwater table. Cattle grazing at stocking rates of 0.6 livestock units per ha were found to be optimal for plant diversity conservation. At sites with deeper groundwater tables and no grazing, dominantElymus spp. stands develop and diversity is strongly reduced. If the groundwater table is high, succession following grazing abandonment is retarded and small halophytes prevail.     

Nutrient fluxes within a small north temperate salt marsh

The water exchange between a small (4.1 hectare) salt marsh adjoining the Great Bay Estuary system of New Hampshire, USA was sampled during 16 tidal cycles between July, 1976 and November, 1977. Tidal amplitude, temperature, salinity, nutrient concentrations (ammonia-N, nitrate-N, nitrite-N, orthophosphate-P, total-P, silicates) and suspended particulates were measured. Conspicuous tidal hydrographic patterns were observed. Mean concentrations of nitrate-N and silicates varied with season. The tidal information, combined with volume determinations, was extrapolated to determine the net flux of hydrographic parameters on monthly and yearly bases. Ammonia-N showed a pronounced seasonality of net exchange by regression analyses. Ammonia-N and suspended particulates had a statistically significant import into the marsh. However, the net fluxes of the other materials were not statistically different from O.Published with the approval of the Director of the New Hampshire Agricultural Experiment Station as Scientific Contribution No. 930Scientific Contribution No. 69 of the Jackson Estuarine Laboratory     

Twenty years of salt marsh succession on a Dutch coastal barrier island

After a formerly grazed salt marsh was released from cattle grazing, changes in plant species composition were monitored for 20 yr, using vegetation maps and permanent plots. Three areas, differing in age and nutrient status were compared. The number of plant species and plant communities decreased.Elymus athericus (Elytrigia pungens) became dominant in most plant communities after 5–20 yr on the oldest and most productive salt marsh. In younger areas it took more time forE. athericus to become dominant. At least 7 cm of clay seemed to be a prerequisite for this plant species to increase in dominance. The results from monitoring over decades are discussed in view of the knowledge on succession over centuries as derived from a chronosequence.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.     

滨海盐沼及其植物群落的分布与多样性

滨海盐沼是广泛存在于世界中、高纬度地区的一种湿地生态系统,具有抵御风暴潮灾害、净化污染物和为珍稀濒危生物提供适宜生境等重要的生态和经济价值。滨海盐沼因随高程变化而急剧变化的环境梯度和植物带状分布现象而为生态学者阐释自然界物种的分布机制提供了理想系统。主要概述了滨海盐沼的定义、特点、类型、全球分布以及潮汐作用、土壤盐度等环境因子特征;阐述了不同尺度下滨海盐沼的植物群落分布和多样性特征。在滨海盐沼植物群落的分布特征上,重点阐述了中尺度下的植物带状分布,即植物群落往往在白海向陆渐高的不同高程梯度上表现出显著的分带分布,不同植物各自占据该梯度上的一定区域。通常认为,带状分布是植物竞争和物理性胁迫共同调控的结果,但其在不同地理区域的普适性仍存争议。滨海植物群落多样性往往较低,在中、小尺度上盐沼植物多样性受控于盐度、潮汐等物理性胁迫、植物间相互作用等因子的作用;在大尺度上盐沼植物多样性可能随纬度增大而增加。系统深入地认识滨海盐沼植物群落生态格局和过程,将为气候变化、生物入侵等人类影响下的滨海盐沼生态系统的管理和恢复提供有益经验。     

Tidal and diurnal influence on food consumption of a salt marsh killifish Fundulus heteroclitus

Feeding patterns during four 24-h periods, sampled at 3-h intervals, were investigated for the mummichog Fundulus heteroclitus, in a Delaware, USA tidal marsh. Two factors potentially influencing feeding patterns, time of day and tide height, were examined. On 2 of the sampling periods a low tide occurred in the morning, while on the other 2 sampling periods a high tide occurred in the morning. Results are reported as g-dry wt. of food per g-dry wt. of fish. F. heteroclitus is primarily a daytime feeder that most actively feeds at high tide, regardless of whether or not the high tide inundates marsh surface areas. When tide height was sufficient to inundate the marsh surface, fish invaded these areas and consumed prey characteristic of the marsh surface. F. heteroclitus is an important link in energy transfers between the marsh surface and subtidal systems, enhancing its own energy supplies by consuming marsh surface prey whenever available.     

Primary production and heterotrophy of a pennate and a centric salt marsh diatom

Radionuclide labeled sodium bicarbonate and glucose were used to measure the rates of primary production and glucose uptake by Nitzschia obtusa var. undulata and Cymbella pusilla. The presence of glucose in the medium enhanced primary production in both diatom species. Uptake of glucose in C. pusilla was photoinhibited. A maximum of 9.3 nmol C mg dry wt^-1 h^-1 of glucose was taken up in the light without the presence of NaHCO₃ in the medium and with NaHCO₃ present the value was 18 nmol C mg dry wt^-1 h^-1. The ratio of inorganic carbon fixed to the total carbon budget (autotrophy and heterotrophy) was 0.51 for N. obtusa var. undulata and 0.62 for C. pusilla.     

Use of the “Acetylene blockage” technique for assaying denitrification in a salt marsh

The acetylene blockage technique was evaluated for measurement of denitrification in salt-marsh sediments (near Halifax, Nova Scotia, Canada). N₂O in the gas phase of closed Spartina alterniflora marsh-sediment systems was analyzed with use of a thermal conductivity gas chromatograph sensitive to approximately 0.1 nmoles ml^-1 gas. No N₂O was detected for unfertilized sediment samples taken through the growing season and incubated in sealed buckets with 10% C₂H₂. For sediment samples amended with nitrate and for enrichments, initial rates of N₂O evolution were higher in the presence of 10% C₂H₂ than in the absence of C₂H₂, but after longterm incubation N₂O was consumed in some samples containing C₂H₂ as well as in samples without C₂H₂. In addition, total gaseous nitrogen (N₂ and N₂O) production in the absence of C₂H₂ was higher than in the presence of C₂H₂. Acetylene appears to be an inconsistent inhibitor of N₂O reduction in salt-marsh sediments. The usefulness of the acetylene-denitrification technique in this habitat is, therefore, questionable.     

Diel flux of dissolved carbohydrate in a salt marsh and a simulated estuarine ecosystem

The concentrations of total dissolved carbohydrate (TCHO), monosaccharide (MCHO) and polysaccharide (PCHO) were followed over a total of ten diel cycles in a salt marsh and a 13 m³ seawater tank simulating an estuarine ecosystem. Their patterns are compared to those for total dissolved organic carbon (DOC), CO₂, pH, O₂, chlorophyll a, phaeopigments and solar radiation. During 5 of the 6 marsh studies, PCHO underwent periods of sustained accumulation starting in the late morning or early afternoon and continuing into the early evening. These periods possibly represent release of recently synthesized PCHO from phototrophs. similar patterns were not found in the tank although direct associations between TCHO and phaeopigment dynamics suggest that zooplankton excretion was an important source of dissolved carbohydrate. The numbers of planktonic bacteria determined in one tank study increased rapidly during a late morning PCHO pulse and varied inversely with PCHO throughout the afternoon and evening, indicating that they were able to respond rapidly and control natural substrate concentrations on a time scale of a few hours. MCHO fluctuated to a much lesser extent than PCHO at both locations with levels possibly maintained near the bacterial uptake threshold or in a closely regulated steady state. TCHO concentration changes over 2-to 3-h sampling intervals suggest very rapid net system release and uptake with summer rates frequently exceeding 30 g C l^-1h^-1 in the marsh and 20 g C l^-1 h^-1 in the tank.     

Latitudinal variation in top-down and bottom-up control of a salt marsh food web

The shrub Iva frutescens, which occupies the terrestrial border of U.S. Atlantic Coast salt marshes, supports a food web that varies strongly across latitude. We tested whether latitudinal variation in plant quality (higher at high latitudes), consumption by omnivores (a crab, present only at low latitudes), consumption by mesopredators (ladybugs, present at all latitudes), or the life history stage of an herbivorous beetle could explain continental-scale field patterns of herbivore density. In a mesocosm experiment, crabs exerted strong top-down control on herbivorous beetles, ladybugs exerted strong top-down control on aphids, and both predators benefited plants through trophic cascades. Latitude of plant origin had no effect on consumers. Herbivorous beetle density was greater if mesocosms were stocked with beetle adults rather than larvae, and aphid densities were reduced in the "adult beetle" treatment. Treatment combinations representing high and low latitudes produced patterns of herbivore density similar to those in the field. We conclude that latitudinal variation in plant quality is less important than latitudinal variation in top consumers and competition in mediating food web structure. Climate may also play a strong role in structuring high-latitude salt marshes by limiting the number of herbivore generations per growing season and causing high overwintering mortality.     

Macroinfauna of a Southern New England salt marsh: seasonal dynamics and production

The animal-habitat relationships and seasonal dynamics of the benthic macroinfauna were investigated from November 1986 to October 1988 in the Great Sippe-wissett salt marsh (Massachusetts, USA). Total macrofaunal abundance varied seasonally, displaying a peak in late spring and early summer, then declining sharply during late summer and recovering briefly in fall before collapsing in winter. Three macroinfaunal assemblages were found in the marsh, distributed along gradients of environmental factors. These included a sandy non-organic sediment assemblage, a sandy organic sediment assemblage and a muddy sediment assemblage. The species groups characteristic of unstable sandy non-organic sediments included the polychaetes Leitoscoloplos fragilis, Aricidea jefreyssi, Magelona rosea and Streptosyllis verrilli, the oligochaete Paranais litoralis, and the crustacean Acanthohaustorius millsi. Sandy organic sediments were characterized by the polychaetes Marenzelleria viridis, Capitella capitata, Neanthes succinea, N. arenaceodonta, Polydora ligni and Heteromastus filiformis, the oligochaete Lumbricillus sp., and the mollusc Gemma gemma. In muddy sites, the polychaete Streblospio benedicti and the oligochaetes Paranais litoralis and Monopylephorus evertus were the dominant species. Secondary production of benthic macroinfauna in each of these habitats was estimated. The highest values of biomass and production were recorded in the sandy organic sediments. Secondary production was estimated to be 1850 kJ m^-2 yr^-1 in sandy organic areas, but only 281 kJ m^-2 yr^-1 in sandy non-organic areas and 113 kJ m^-2 yr^-1 in muddy areas. This results in an area-weighted average production of 505 kJ m^-2 yr^-1 for the unvegetated areas of the marsh. The Great Sippewissett salt marsh has an area of 483800 m², the total secondary production of the macroinfauna for the whole unvegetated area of the marsh was estimated as 4651 kg dry wt yr^-1, expressed as somatic growth. This production value seems consistent with production data obtained for other intertidal North Atlantic environments.     

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.     

Exploring community assembly through an individual-based model for trophic interactions

Traditionally, the dynamics of community assembly has been analyzed by means of deterministic models of differential equations. Despite the theoretical advances provided by such models, they are restricted to questions about community-wide features. The individual-based modeling offers an opportunity to link bionomic features to patterns at the community scale, allowing us to understand how trait-based assembly rules can arise by dynamical processes. The present paper introduces an individual-based model of community assembly, and discusses some of the major advantages and drawbacks of this approach. The model was framed to deal with predation among size-structured populations, incorporating allometric constraints to energetic requirements, movement, life-history features and interaction relationships among individuals. A protocol of assembly procedure is proposed, in which a period of intense species introductions is followed by a period without introductions. The resultant communities did not present any pattern of trait over-dispersion, meaning that the multivariate distances of bionomic features among co-occurring species were neither larger nor more regular than expected in a random collection of species. It suggests a weak influence of interspecific interactions in the model environment and individualistic rules of coexistence, driven mainly by the spatial structure. This highlights that trait over-dispersion and resource partitioning should not be considered a necessary condition for coexistence, even in communities entirely structured by internal processes like predation and competition.     

Nitrogen exchange between a southeastern USA salt marsh ecosystem and the coastal ocean

The salt marsh ecosystem at North Inlet, South Carolina, USA consistently exported dissolved inorganic nitrogen via tidal exchange with the coastal Atlantic Ocean. Concentrations centrations of NH ₄ ⁺ and NO ₃ ^- +NO ₂ ^- displayed distinct tidal patterns with rising values during ebb flow. These patterns suggest the importance of biogeochemical processes in the flux of material from the salt marsh. NH ₄ ⁺ export peaked during the summer (15 to 20 mg m^-2 tide^-1) during a net balance of tidal water exchange. Remineralization of NH ₄ ⁺ within the salt marsh system appears to be contributing to the estimated annual net export of bout 4.7 g NH ₄ ⁺ -N m^-2 yr^-1. NO ₃ ^- +NO ₂ ^- exports were higher in the fall and winter of 1979 (2 to 4 mg N m^-2 tide^-1). The winter export coincided with a considerable net export of water with no distinctive concentration patterns, suggesting a simple advective export. However, the fall peak of NO ₃ ^- +NO ₂ ^- export occurred during a period of net water balance in tidal exchange and an insignificant freshwater input from the western, forested boundary. During the summer and fall, tidal concentration patterns were particularly apparent, suggesting that nitrification within the salt marsh system was contributing to the estimated annual net export of ca 0.6 g NO ₃ ^- +NO ₂ ^- -N m^-2 yr^-1.Contribution No. 637 from the Belle W. Baruch Institute of Marine Biology and Coastal Research     

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.     

Effects of nitrogen addition on the growth of the salt marsh grassElymus athericus

Effects of nitrogen addition on the growth of the salt marsh grassElymus athericus were studied under greenhouse conditions. The addition of inorganic nitrogen (in the form of nitrate or ammonium and ranging from 0–24 g N/m²) stimulated the growth ofElymus athericus at the highest addition. Addition of nitrogen led to an increase of the soil nitrate concentrations both in the nitrate and ammonium treated soil in the first period of the experiment, whereas no differences were present at the end of the experiment. Ammonium in the ammonium treatments was transformed to nitrate within 15 days. In another experiment the values of the stable isotope nitrogen-15—expressed as δ¹⁵N-in nitrogen compounds used as fertilizer, in salt marsh soil and ofElymus athericus were measured. The δ¹⁵N of the N-compounds added (between ?3.2 and +2.6‰) were lower than the soil (ca.+10‰) and plants (ca.+8‰). During growth in water culture the δ¹⁵N of the leaves, stems and roots ofElymus athericus decreased from +9‰ to ?1‰. The latter value was close to the °¹⁵N of the N-compounds used in the water solution. Addition of N-compounds in soil culture, however, did not lead to such a decrease of the °¹⁵N ofElymus athericus. The difference in δ¹⁵N between soil nitrogen and the N-compounds added may be too small to be used successfully in ecological studies of nitrogen fluxes in the salt marsh environment.     

Behaviour of horses and cattle at two stocking densities in a coastal salt marsh

Livestock grazing has been practiced in salt marshes in the Wadden Sea area since 600 B.C. Currently livestock grazing is also applied for conservation management. However, effects of such grazing management on salt marshes are likely to vary depending on the species of livestock and stocking density due to differences in the behaviour of the animals. Yet, little is known about the behaviour of different livestock species and stocking densities grazing in salt marshes. We studied the grazing behaviour of horses and cattle by focal observation in an experiment with four different grazing treatments on a coastal salt marsh. In all treatments we recorded diet choice, movement and grazing activity, and spatial distribution. Livestock species shared an overlap in diet choice. Yet, horses more often foraged on the short grass Puccinellia maritima, while the cattle diet contained a higher amount of Aster tripolium. Horses travelled longer distances per day and spent more time grazing than cattle. Spatial distribution of cattle was significantly clustered, while horses showed a random distribution utilizing the whole area. Animal behaviour differs between livestock species and stocking densities with respect to diet choice, activity and spatial distribution.     

Pigment composition and size distribution of phytoplankton in a confined Mediterranean salt marsh ecosystem

Pigment composition and size distribution of phytoplankton were analysed in a group of Mediterranean salt marshes, where hydrology is dominated by sudden inputs during sea storms, followed by long periods of confinement. These marshes are characterized by a low inorganic–organic nutrient ratio, and inorganic nitrogen is especially scarce due to denitrification. Nutrients were the main factor affecting phytoplankton biomass, while zooplankton grazing did not control either phytoplankton community composition, or their size distribution. The relative abundance of the different phytoplankton groups was analysed by correspondence analysis using the pigment composition measured by high-performance liquid chromatography (HPLC) and analysed with the CHEMTAX programme. In this analysis, phytoplankton pigment composition was correlated with two nutrient gradients. The first gradient was the ratio of nitrate–total nitrogen (TN), since the different phytoplankton groups were distributed according to their eco–physiological differences in nitrogen uptake. The second gradient was correlated with total nutrient loading. Biomass size distributions frequently showed a lack of intermediate sized nanophytoplankton (2.5–4 μm in diameter), and the importance of this lack of intermediate sizes correlated with dinoflagellate biomass. These results suggested that in confined environments, where nutrients are mainly in an organic form, dinoflagellates take advantage of their mixotrophy, by competing and grazing on smaller phytoplankters simultaneously.     

Phytoplankton in a temperate-zone salt marsh: Net production and exchanges with coastal waters

Phytoplankton production and associated variables were measured in Flax Pond, a 52 ha salt marsh on the north shore of Long Island, New York, from July 1972 to October 1973. Measurements made up to five times per day, once per week, yielded a mean annual net primary production, determined by the ¹⁴C technique, of 20.5 mg C/m³/h; daily means were as high as 60.0 mg C/m³/h. However, when productivity was calculated for the entire marsh ecosystem, the shallow water in the salt marsh produced only 11.7 g C/m² of marsh/year. There was a net flux of phytoplankton from the coastal waters into the marsh; during the summer up to 0.2 g chlorophy 11/m² of marsh was carried in with the tides daily and remained in the marsh. Analysis of the productivity data, as well as variables associated with productivity (pH, standing crop, nutrients, extinction coefficient), indicated that the aquatic portion of the marsh behaved more as a net consumer rather than a net producer of phytoplankton.Research carried out at Brookhaven National Laboratory, supported by the U.S. Atomic Energy Commission and the National Science Foundation under Grant No. AG-375.