Environmental gradients and herbivore feeding preferences in coastal salt marshes

Current theories of plant-herbivore interactions suggest that plants may differ in palatability to herbivores as a function of abiotic stress; however, studies of these theories have produced mixed results. We compared the palatability of eight common salt marsh plants that occur across elevational and salinity stress gradients to six common leaf-chewing herbivores to determine patterns of plant palatability. The palatability of every plant species varied across gradients of abiotic stress in at least one comparison, and over half of the comparisons indicated significant differences in palatability. The direction of the preferences, however, was dependent on the plant and herbivore species studied, suggesting that different types of stress affect plants in different ways, that different plant species respond differently to stress, and that different herbivore species measure plant quality in different ways. Overall, 51% of the variation in the strength of the feeding preferences could be explained by a knowledge of the strength of the stress gradient and the type of gradient, plant and herbivore studied. This suggests that the prospects are good for a more complex, conditional theory of plant stress and herbivore feeding preferences that is based on a mechanistic understanding of plant physiology and the factors underlying herbivore feeding preferences.     

The impact of grazing on plant communities,plant populations and soil conditions on salt marshes

Grazing an abandoned salt marsh causes retrogressive succession, since mid salt-marsh communities change into lower salt-marsh communities. Grazing and mowing are compared in detail. Both management practices enhance species diversity in an abandoned salt marsh. This can be attributed to the removal of litter. The finding that lower salt-marsh species appear more with grazing than with mowing or abandoning is not related to a higher soil salinity as compared to mowing or abandoning, but probably to locally baring of the soil by grazing animals. Only species of pioneer or unstable environments seem to have a persistent seed bank, for other species seed dispersal seems to be a limiting factor for their establishment.Nomenclature follows Heukels & van Ooststroom (1977) for species; Westhoff & den Held (1969) for syntaxa.Mrs R. Rusthoven analyzed the soil samples, Mr E. Leeuwinga drawed the figures, and Mrs J. O'Brien corrected the English text.     

Methodology for studying exchanges between salt marshes and coastal marine waters

The salt marshes of the Mont St. Michel bay represent a complex system in continuous change, mostly due to the frequent exchanges with the coastal waters through tidal processes. In such ecosystems, water is an important element insofar as it represents the common vector of flows between and among several ecosystem compartments. The purpose of the approach discussed here is to estimate the volume of water coming in and out and to determine the variations of the water quality according to time and nutrients concentrations. The estimation of the water fluxes is dependent on the channel calibration downstream of the watershed. Among the different methods examined, the continuous integrals calibration appears as the best one because the water level changes very quickly.Up to now, estimations of nutrients exchanges in wetlands have been based on rigorously regular field sampling, in consideration of the fact that exchanges occurred mainly during annual spring tides or during spring tides of each cycle of the year. According to our results, it seems that every tide, and portion of a tide, of a monthly and seasonal cycle has some importance and variability, which suggests that all parts of a tide should be considered in estimations of exchanges between wetlands and coastal waters.Corresponding Editor: W. Mitsch     

Nutrient limitation of phytoplankton in solar salt ponds in Shark Bay,Western Australia

The aim of this research was to examine nutrient limitation of phytoplankton in solar salt ponds of varying salinity at Useless Inlet in Western Australia. These ponds use solar energy to evaporate seawater for the purpose of commercial salt production. A combination of techniques involving water column nutrient ratios, comparisons of nutrient concentrations to concentration of magnesium ions and bioassays were used in the investigation. Comparisons of changes in dissolved inorganic nitrogen to phosphorus ratios and concentrations of dissolved inorganic nutrients against changes in concentrations of the conservative cation Mg²⁺ indicated that phytoplankton biomass was potentially nitrogen limited along the entire pond salinity gradient. Nutrient addition bioassays indicated that in low salinity ponds, phytoplankton was nitrogen limited but in high salinity ponds, phosphorus limited. This may be due to isolation of phytoplankton in bioassay bottles from in situ conditions as well as to changes in phytoplankton species composition between ponds, and the variable availability of inorganic and organic nutrient sources. The differences in limiting nutrient between methods indicate that phytoplankton cells may be proximally limited by nutrients that are not theoretically limiting at the pond scale. Dissolved organic nutrients constituted a large proportion of total nutrients, with concentrations increasing through the pond sequence of increasing salinity. From the change in nutrient concentrations in bioassay bottles, sufficient dissolved organic nitrogen may be available for phytoplankton uptake in low salinity ponds, potentially alleviating the dissolved inorganic nitrogen limitation of phytoplankton biomass. Guest Editors: J. John & B. Timms Salt Lake Research: Biodiversity and Conservation—Selected Papers from the 9th Conference of the International Society for Salt Lake Research     

Salt stress limitation of seedling recruitment in a salt marsh plant community

Summary Seedling recruitment in salt marsh plant communities is generally precluded in dense vegetation by competition from adults, but is also relatively rare in disturbance-generated bare space. We examined the constraints on seedling recruitment in New England salt marsh bare patches. Under typical bare patch conditions seed germination is severely limited by high substrate salinities. We examined the germination requirements of common high marsh plants and found that except for one notably patch-dependent fugitive species, the germination of high marsh plants is strongly inhibited by the high soil salinities routinely encountered in natural bare patches. Watering high marsh soil in the greenhouse to alleviate salt stress resulted in the emergence of up to 600 seedlings/225 cm². The vast majority of this seed bank consisted of Juncus gerardi, the only common high marsh plant with high seed set. We tested the hypothesis that salt stress limits seedling contributions to marsh patch secondary succession in the field. Watering bare patches with fresh water partially alleviated patch soil salinities and dramatically increased both the emergence and survival of seedlings. Our results show that seedling recruitment by high marsh perennial turfs is limited by high soil salinities and that consequently their population dynamics are determined primarily by clonal growth processes. In contrast, populations of patch-dependent fugitive marsh plants which cannot colonize vegetatively are likely governed by spatially and temporally unpredictable windows of low salinities in bare patches.     

Nutrient limitation and algal blooms in urbanizing tidal creeks

Tidal creeks are commonly found in low energy systems on the East and Gulf Coasts of the United States, and are often subject to intense watershed human development. Many of these creeks are receiving urban and suburban runoff containing nutrients, among other pollutants. During the period 1993-2001, we studied three tidal creeks located in southeastern North Carolina, a rapidly urbanizing area. All three creeks received anthropogenic nutrient loading. Oligohaline to mesohaline stations in upper tidal creek regions had much higher nutrient (especially nitrate-N) concentrations than lower creek areas, and hosted spring and summer phytoplankton blooms that at times exceeded 200 μg chlorophyll a l⁻¹. Phytoplankton biomass during winter was low at all stations in all three creeks. Spring and summer nutrient addition bioassay experiments were conducted to characterize the nutrients limiting phytoplankton growth. Water from high salinity stations in all three creeks always showed significant positive responses to nitrate-N inputs, even at concentrations as low as 50 μg N l⁻¹. Low salinity stations in upper creek areas often showed significant responses to nitrate-N inputs, but on occasion showed sensitivity to phosphorus inputs as well, indicating the influence of anthropogenic nitrate loading. During several experiments, one of the upper stations showed no positive response to nutrient inputs, indicating that these stretches were nutrient replete, and further phytoplankton growth appeared to be light-limited either by phytoplankton self-shading or turbidity. Water from upper creek areas yielded much higher chlorophyll a concentrations in bioassay experiments than did lower creek water. In general, these urbanizing tidal creeks were shown to be very sensitive to nitrogen loading, and provide a physical environment conducive to phytoplankton bloom formation in nutrient-enriched areas. Tidal creeks are important ecological resources in that they are considered to be nursery areas for many species of fish and shellfish. To protect the ecological function of these small, but very abundant estuarine systems, management efforts should recognize their susceptibility to algal blooms and focus on control of nonpoint source nutrient inputs, especially nitrogen.     

Nutrient limitation in Crater Lake,Oregon

Experiments were carried out to determine what nutrient (or nutrients) was primarily responsible for limiting phytoplankton productivity in ultraoligotrophic Crater Lake. The experiments included in situ and laboratory nutrient addition bioassays utilizing the natural phytoplankton community, Selenastrum capricornutum bottle assays, photosynthetic responses, photosynthetic carbon metabolism, and response of dark uptake of ¹⁴CO₂ with the addition of NH ₄ ⁺ . The results suggested that a trace metal(s) or its availability was the primary factor limiting the epilimnetic phytoplankton productivity. Nitrogen was extremely low, and quickly became limiting with the addition of trace metals and a chelator. Iron is the most likely candidate as the limiting nutrient. Trace metals and nitrogen are also both important in limiting phytoplankton at 100 m, a depth where biologically mediated turnover of nutrients seems to be more important.     

Ecophysiological adaptations of coastal halophytes from foredunes and salt marshes

Ecophysiological strategies of coastal halophytes from foredunes and salt marshes are discussed. A comparison is made of the factors that limit growth in salt marshes and sand dunes. In salt marshes, zonation and succession are primarily governed by variation in soil salinity, which strongly depends on inundation with seawater. Results are described of experiments which aim at separating salinity and inundation effects on growth, osmotic and mineral relations in a comparison of salt-marsh halophytes. The growth response of plants cannot simply be correlated (and causally explained) with the concentration of Na, Cl, and K in the tissues. Also, the compatible osmotic solutes proline and methylated quaternary ammonium compounds may accumulate both in species with a positive response to increased salinity and in species with a growth reduction under seawater inundation. More likely inadequate adaptation of the plants water potential with these components is partly the cause of retarded growth. Disfunctioning of the plant in this respect may be at three levels: (a) total water potential of the plant, (b) (loss) of turgor pressure potential; (c) regulation at the cellular level. The ecological importance of some factors in seawater other than sodium chloride is considered. In coastal sand dunes airborne rather than soil salinity limits plant growth, together with the effects of abrasion, sand accretion, drought and the poor nutrient status of the dune sand. Adaptations of sand-dune species to these factors may consist of: large seeds with storage tissue germinating in the dark and seedling growth enough to emerge through the accreted sand. Aerial parts must be resistant to mechanical damage (high wind speed and abrasion), possibly by a sclerophyllous and tough structure. Efficient nutrient uptake, translocation and retranslocation seem to help survive sand-dune species in a nutrient-poor rooting medium.     

Population dynamics of annual Salicornia species in the tidal salt marshes of the Oosterschelde,The Netherlands

1. The population dynamics of two Salicornia species from the Bergen op Zoom salt marsh (south-west Netherlands) was examined. Based on the results of several field studies three preliminary life tables were constructed, two for S. procumbens agg. populations growing respectively on the mud flats and in the salt marsh, and one for S. europaea agg. living in the upper marsh.
2. The life cycles are described and quantified in terms of eight phases and the transition probabilities between them, starting from a notional individual representative of each population.
3. The models depicting the life cycle of S. procumbens show a mean offspring number of 4.26 individuals per parent for the mud-flat population and 0.18 for the salt-marsh population. The S. europaea model gives an output of 0.44 individuals per parent. These results reflect the fluctuations in population size observed in sample plots over the years 1976–78.
4. Comparison of the transition probabilities reveals that on the mud flats most S. procumbens individuals die during pollination and seed germination, while the population in the salt marsh proper is thinned especially during the seed phase in winter time and during the growth from established seedlings to maturation. S. europaea behaves in a similar but less pronounced way to S. procumbens in the salt marsh.
5. Probabilities for one flower or one seed to produce a mature flowering plant were calculated, and were compared with those found in the literature. They are roughly of the same order of magnitude as the probabilities for other annual species, but much higher than those reported for biennial species.

     

Comparison of niche breadths and overlaps of halophytes on salt marshes of differing diversity

Comparisons were made of niche breadths and niche overlaps of halophytes growing in two emergent saltmarsh communities differing in the number of their constituent species and in diversity. Niche parameters were calculated along the major niche dimension of tidal inundation which was assessed using three related measurements, namely height on the marsh, number of submergences per year, and longest period of continuous exposure. This reduces the danger of measuring niches on one resource dimension along which species exhibit non-linear responses. Results obtained using these three measurements were very similar. Mean niche overlaps were significantly smaller, and most species had lower niche breadths, on the marsh with more diverse vegetation. These results agree with Pianka's (1974) niche overlap hypothesis; they indicate that competitive interactions between species are acting with different intensity on the two marshes, and that such interactions are important in determining the distribution limits of both low-and high-marsh species in the emergent salt-marsh community.Nomenclature follows Clapham, Tutin & Warburg (1962). Acknowledgements. The authors gratefully acknowledge a travel grant from the British Council enabling them to attend the conference. The work was conducted whilst P.J.R. was in receipt of a Natural Environment Research Council studentship.     

Arbuscular mycorrhizal colonization of halophytes in Central European salt marshes

 Halophytes from both coastal and inland Central European salt marshes were examined for colonization by arbuscular mycorrhizal (AM) fungi. Plants from different families were strongly colonized but the degree of colonization varied with the individual plant and apparently during the vegetation period, too. Members of the typical non-mycorrhizal families like Armeria maritima of the Plumbaginaceae and Salicornia europaea of the Chenopodiaceae were found to be colonized, particularly in the drier salt marshes. High numbers of Glomus spores were found in the saline soils, especially those of the inland locations examined. Approximately 80% of these spores were from Glomus geosporum as shown by a typical restriction fragment length polymorphism (RFLP) pattern of the amplified internal transcribed spacer regions. The present study demonstrates that RFLP analysis is useful when screening habitats for the occurrence of mycorrhizal fungi which can be identified only with difficulty by morphological criteria. Accepted: 25 August 2000     

Sediment nutrient accumulation and nutrient availability in two tidal freshwater marshes along the Mattaponi River, Virginia, USA

Sediment deposition is the main mechanism of nutrient delivery to tidal freshwater marshes (TFMs). We quantified sediment nutrient accumulation in TFMs upstream and downstream of a proposed water withdrawal project on the Mattaponi River, Virginia. Our goal was to assess nutrient availability by comparing relative rates of carbon (C), nitrogen (N), and phosphorus (P) accumulated in sediments with the C, N, and P stoichiometries of surface soils and above ground plant tissues. Surface soil nutrient contents (0.60–0.92% N and 0.09–0.13% P) were low but within reported ranges for TFMs in the eastern US. In both marshes, soil nutrient pools and C, N, and P stoichiometries were closely associated with sedimentation patterns. Differences between marshes were more striking than spatial variations within marshes: both C, N, and P accumulation during summer, and annual P accumulation rates (0.16 and 0.04 g P m^–2 year^–1, respectively) in sediments were significantly higher at the downstream than at the upstream marsh. Nitrogen:P ratios <14 in above ground biomass, surface soils, and sediments suggest that N limits primary production in these marshes, but experimental additions of N and/or P did not significantly increase above ground productivity in either marsh. Lower soil N:P ratios are consistent with higher rates of sediment P accumulation at the downstream site, perhaps due to its greater proximity to the estuarine turbidity maximum.     

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.     

The influence of rainfall variability on the species composition of a northern California salt marsh plant assemblage

Natural variability in the species composition of salt marsh plant assemblage was studied for 4 years (1986–90) at Bolinas Lagoon, California (USA). The study was conducted during a period in which little or no physical disturbance occurred in the marsh. During the study, Bolinas Lagoon experienced highly variable rainfall, both within and between years. In years with average or below average winter and spring rainfall, the cover of Salicornia virginica increased relative to that of the other species. During the one year with much higher than average spring rainfall, the cover of Salicornia declined and rare species increased in relative abundance. These patterns suggest that under typical marsh conditions, the plant assemblage experiences stress due to low availability of freshwater and high soil salinity. Under such conditions, the stress-tolerant Salicornia dominates the assemblage. If, however, there is abundant rainfall early in the growing season, the other species are released from stress and are able to increase in cover. The marsh of Bolinas Lagoon appears to be resistant to change and able to tolerate stress. Few studies have examined natural variability in species in undisturbed marshes and more of such studies must be made in order to understand whether or not marshes are in general resistant to change caused by rainfall variability.Abbreviations MLLW = mean lower low water     

黄海和东海营养盐分布及其对浮游植物的限制

根据1997～1999年黄海和东海4个季节的现场调查资料,分析探讨了黄海、东海的营养盐分布特征及其对浮游植物生长的限制状况．结果表明,在长江口以东及其东北部海域终年存在一个范围很大的营养盐高值区．分析表明,这些营养盐主要来自长江冲淡水的扩展及苏北沿岸流的输送．此外,还获得了1998年长江流域特大洪水期间,迄今被观测到的长江冲淡水中营养盐的最大扩展范围,计算并研究了黄海、东海上层水中Si／N／P比值,结果表明,黄东海上层水中Si／N比值较高,Si不会成为黄东海浮游植物生长的限制因子;但在南黄海南部尤其是西南部、东海近岸及长江口以东海域,N／P比值很高(>30),说明与一般海洋环境的情况不同,黄海、东海有很大一部分海域浮游植物的生长受磷限制,而不是受氮限制．     

Nutrient limitation of two saccharolytic clostridia; secretion, sporulation, and solventogenesis

Abstract Growth of the cellulolytic acidogen Clostridium strain C7 undergoing progressive nutrient limitation has been compared with that of the solventogen Clostridium beijerinckii . On the basis of cellulase secretion, differentiation of dissimilatory metabolism, and sporulation, different survival strategies by the two clostridia in progressive nutrient limitation can be discerned. In addition, the metabolic differentiation to butanol production in Clostridium beijerinckii can be specifically associated with the sporulation stage in which the forespore is enclosed by double membranes but not by a spore coat.     

Integrated management of artificially created salt marshes in the Wadden Sea of Schleswig-Holstein, Germany

In the about 10,000 km² large Wadden Sea 353 km² of salt marshes exist. The great majority of these are artificial, i.e. developed through accretion enhancement techniques. Salt marshes in the Wadden Sea have a high coastal defence value as well as ecological significance. High salt marshes in front of sea walls reduce the wave-impact on the dikes and, therewith, the danger of breaching. At the same time, the salt marshes have a high ecological value, e.g., as resting places for migratory birds. In consequence, for coastal defence, salt marshes should be created and/or maintained. For nature protection, natural dynamic processes (incl. erosion) should prevail. To anticipate this possible conflict among coastal defence and environmental administration, in Schleswig-Holstein an interdepartmental working group established a salt marsh management concept. For information of the interested parties and for monitoring of the concept, an integrated salt marsh board was established, consisting of representatives from public administration and local interest groups (NGO's, dike and water boards, municipalities). Five years after implementation of the concept, this board established a first progress report. Although the concept was, in the beginning, met with great scepticism, five years of practical experience demonstrate that it functions well. The salt marshes are stable or growing, i.e. the management measures are successful. As a planning instrument, the concept is unambiguous, i.e. provides clear guidelines on how to manage the salt marshes. Finally, as an instrument of participation, the salt marsh board is valued positively by all members.     

Environmental factors affecting bacterioplankton and phytoplankton dynamics in confined Mediterranean salt marshes (NE Spain)

Environmental factors accountable for bacterioplankton or phytoplankton biomass dominance were analysed in a confined Mediterranean salt marsh (Empordà Wetlands, NE Spain). Two basins located in the same salt marsh, and with differences in size and catchment's area were compared, during four characteristic situations of the hydroperiod. Since bacterio- or phytoplankton relationships may be affected by other factors such as diel variations or vertical differences in nutrient composition and distribution, high frequency fluctuations due to these factors were also taken into account. Differences in catchment area appeared to be the more plausible explanation of differences in nutrient and organic carbon accumulation among basins, since during confinement basins essentially accumulate the allochthonous nutrient and organic matter supplies that previously entered by runoff. DOC (Dissolved Organic Carbon) favoured the bacterioplankton biomass increase, but also was the main variable significantly affecting phytoplankton biomass. Basins showed marked differences in bacterio- and phytoplankton dominances. Relationships between phytoplankton and bacterioplankton were positive, negative or not significant, depending on the basin and on the period of the year. The phytoplankton mixotrophic capabilities, both phagotrophy and osmotrophy, and their production of UV-screening compounds, as sunscreen, may explain the significant correlation between DOC and phytoplankton biomass, and the significant effect of phytoplankton on bacterioplankton found in these ecosystems.     

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.