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.     

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.     

Nutrient limitation and plant species composition in temperate salt marshes

Addition of inorganic nitrogen, phosphorus and potassium in a factorial design in two ungrazed Wadden-Sea salt marshes at low and high elevations showed that nitrogen was the limiting nutrient. No effects of nutrient addition were detected in the 1st year, probably due to a considerable rainfall deficit during the growing season. In the 2nd year, which was more humid, only nitrogen addition caused significant effects in both the low salt marsh dominated by Puccinellia maritima and the high marsh dominated by Festuca rubra. No two-way or three-way interactions with phosphorus or potassium were found. In the low marsh, nitrogen addition had a negative effect on the biomass of Puccinellia, but a positive effect on the biomass of Suaeda maritima and on the total above-ground biomass. Puccinellia was replaced by Suaeda after nitrogen addition, due to shading. In the high salt marsh, no significant effects of fertilizer application on total above-ground biomass were found, due to the weak response of the dominant species Festuca rubra, which accounted for 95% of total biomass. The biomass of Spergularia maritima increased, however, as a response to nitrogen addition.The shoot length of Festuca was positively affected by nitrogen fertilization. It is suggested that stands of Festuca reached maximal biomass at the study site without fertilization and that its growth was probably limited by self-shading. Received: 22 September 1996 / Accepted: 5 April 1997     

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.     

Environmental and biological controls on methyl halide emissions from southern California coastal salt marshes

Methyl bromide (CH₃Br) and methyl chloride(CH₃Cl) emission rates from southernCalifornia coastal salt marshes show largespatial and temporal variabilities that arestrongly linked to biological and environmentalfactors. Here we discuss biogeochemical linesof evidence pointing to vegetation as theprimary source of CH₃Br and CH₃Clemissions from salt marshes. Sediments andmacroalgae do not appear to be major producersof these compounds, based on observations thatthe highest fluxes are not inhibited by soilinundation; their emissions are not correlatedwith those of certain gases produced in soils;and emissions from mudflat- andmacroalgae-dominated sites are relativelysmall. In contrast, the seasonal and spatialvariabilities of methyl halide fluxes in thesesalt marshes are consistent with the productionof these compounds by vascular plants, althoughthe possibility of production by microflora orfungi associated with the salt marsh vegetationis not ruled out. Flux chamber measurements ofemission rates are largely correlated to theoverall plant biomass enclosed in the chamber,but appear also to be highly dependent on thepredominant plant species. Emission ratesfollow a diurnal trend similar to the trends ofambient air temperature and photosyntheticallyactive radiation, but not surface soiltemperature. Diurnal variabilities in thecarbon isotope compositions of CH₃Cl andCH₃Br and their relative ratios ofemissions are consistent with simultaneouslycompeting mechanisms of uptake andproduction.     

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     

Recognition of co-existence pattern of salt marshes and mangroves for littoral forest restoration

Climate-change driven sea level rise causes a increase in salinity in coastal wetlands accelerating the alteration of the species composition. It triggers the gradual extinction of species, particularly the mangrove population which is intolerant of excessive salinity. Thus despite being crucial to a wide range of ecosystem services, mangroves have been identified as a vulnerable coastal biome. Hence restoration strategy of mangroves is undergoing rigorous research and experiments in literature at an interdisciplinary level. From a data-driven perspective, analysis of mangrove occurrence data could be the key to comprehend and predict mangrove behavior along different environmental parameters, and it could be important in formulating management strategy for mangrove rehabilitation and restoration. As salt marshes are the natural salt-accumulating halophytes, mitigating excessive salinity could be achieved by incorporating salt-marshes in mangrove restoration activities. This study intends to find a novel restoration strategy by assessing the frequent co-existence status of salt marshes, with the mangroves, and mangrove associates in different zones of degraded mangrove patches for species-rich plantation. To achieve this, we primarily design a novel methodological framework for the practice of knowledge discovery concerning the coexistence pattern of salt marshes, mangroves, and mangrove associates along with environmental parameters using a data mining paradigm of association rule mining. The proposed approach has the capability to uncover underlying facts and forecast likely facts that could automate the study in the field of ecological research to comprehend the occurrence of inter-species relationships. Our findings are based on published data gathered on the Sundarban Mangrove Forest, one of the world’s most important littoral forests. The existing literature reinforces the findings that include all the sets of frequently co-occurring mangroves, their associates, and salt marshes along the salinity gradient of coastal Sundarbans. A detailed understanding of the occurrence patterns of all these, along with the environmental variables, would be able to promote decision-making strategy. This framework is effective for both academia and stakeholders, especially the foresters/ conservation planners, to regulate the spread of salt marshes and the restoration of mangroves as well.     

Vegetation study as a generator for population biological and physiological research on salt marshes

The paper deals with some views on the phytocoenose in relation to the functioning of vegetation and its plant-species populations in space and time. From these viewpoints the study of vegetation is seen as a field of tension between the organismic and reductionistic approaches. Both have their value, provided any dogmatism is avoided and either can be applied to the other.In the field of vegetation structure characteristic features of life-form spectra and species distribution, inversion phenomena in zonation, and community architecture in relation to production and decomposition are discussed. In this connection some remarks are made on habitat and niche differentiation with respect to the phytocoenose concept.Vegetation dynamics are discussed in relation to the introduction of Spartina anglica, the frequency of flooding by the tides, different environmental disturbances caused by heavy winter frost, rainfall and hot and dry periods, as well as to human interferences for agricultural and civil-technical purposes.It is suggested that salt-marsh plants may have found refuge areas in inland habitats as well as on more southerly coastal sites during glaciations.Nomenclature follows Tutin et al. (1964–1980).The author wishes to thank his collaborators Messrs B. P. Koutstaal and W. de Munck for much fieldwork, and Mrs M. J. van Leerdam-de Dreu and Messrs A. A. Bolsius and J. A. van den Ende for the preparation of text and figures.Communication No. 302 of the Delta Institute for Hydrobiological Research.     

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.     

海岸带盐沼生态系统卤代甲烷释放研究进展

卤代甲烷是破坏臭氧层的主要物质,也是重要的痕量温室气体和有机污染物.目前,其源汇格局和全球收支存在很大的不确定性.受海陆两相交互作用影响的海岸带盐沼是卤代甲烷重要的自然来源.本文综述了卤代甲烷自然源汇的研究现状、海岸带盐沼卤代甲烷的释放规律以及主要影响因素.鉴于当前研究中存在的问题,其后的研究需在以下几个方面进一步加强: 1)较长时间尺度、更大区域范围的盐沼卤代甲烷释放规律及源汇评估研究;2)利用目前已比较成熟的稳定同位素技术,更精确地定量不同盐生植物种类以及各种生物类型对盐沼卤代甲烷的贡献比率;3)关注潮水对其产生的直接或间接影响,进一步研究潮汐涨落过程、淹水时长等对盐沼卤代甲烷释放的影响;4)人类活动和气候变化对盐沼卤代甲烷释放的影响.     

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.     

Rafting on wrack as a mode of dispersal for plants in coastal marshes

Here, I examine the potential for dead plant material, or wrack, to act as a vector of dispersal for plants among coastal marshes along the Palmer River in New England, USA. Three hundred mats of wrack floating along a 2.5 km section of river spanning tidal fresh, brackish and salt marshes were sampled for propagules of plants from May 1997 to October 1998. Thirty species of plants and 2 species of algae were associated with the wrack, and 24 of the plant species were viable under greenhouse conditions. Common and abundant species of tidal fresh, brackish and salt marshes were found in the wrack, as well as introduced and invasive species. Wrack contained plants suited to different modes of dispersal, including water, wind and adherence to animals. Measurements of the movement of individual mats of wrack along a 200 m section of the river indicated that wrack travelled at speeds ranging from 1.1 to 2.6 km h⁻¹, which translates to potential dispersal distances ranging from 6.5 to 15.9 km for one ebb or flood tide of 6 h duration. Tagged mats of wrack tracked for one ebb or flood tide travelled at least 2.5 km along the river. These results suggest that wrack may be an important agent of plant dispersal among coastal marshes in estuaries.     

The halophilous vegetation of the Orumieh lake salt marshes,NW. Iran

The halophilous vegetation of the Orumieh lake salt marshes has been studied, using the Braun-Blanquet method. Vegetation types have been defined by physiognomic-floristic system. The following six main groups of communities are recognized: (1) Semi-woody shrub and perennial halophytic communities (Class Halocnemetea strobilacei) including 6 associations and 5 subassociations, (2) Annual halophytic communities (Class Thero-Salicornietea) including 5 associations and 1 subassociation, (3) Salt marsh brushwood communities (Class Tamaricetea) including 4 associations, (4) Rush and herbaceous perennial halophytic communities (Class Juncetea maritimi) including 7 associations and 1 subassociation, (5) Rush and herbaceous perennial halotolerant communities (Class Agrostietea stoloniferae) including 5 associations, (6) Hydrophilous halotolerant communities (Class Phragmitetea) including 3 associations and 2 subassociations. The soil of these communities has been analysed and their habitats are described and discussed.     

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.     

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.     

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.     

滨海盐沼湿地蓝色碳汇功能、形成机制及其增汇潜力

盐沼湿地具有很高的碳捕获与存储能力, 是缓解全球变暖的有效蓝色碳汇(蓝碳)。未来气候变暖和海平面上升可能增加盐沼湿地的固碳能力, 其蓝碳功能越来越受到国际社会的重视。该文重点围绕盐沼湿地蓝碳形成的关键过程、光合碳分配过程及影响机制、碳沉积埋藏特征及其来源解析、盐沼湿地土壤碳库稳定性及其微生物机制、盐沼湿地蓝碳过程动态模拟及其增汇潜力等5个方面进行综述。在此基础上, 针对当前研究的不足, 提出今后的研究中需要进一步探究盐沼湿地植被海陆梯度分布格局对碳吸收能力和碳分配的影响, 土壤有机碳沉积和埋藏速率及其对全球变化的响应, 盐沼湿地土壤碳库的稳定性及其横向碳流动, 气候变化和海平面上升背景下盐沼湿地蓝碳模拟与增汇潜力评估, 以及盐沼湿地蓝碳的增汇技术和途径。以期为深入理解盐沼湿地蓝碳形成过程与机制, 预测全球变化背景下盐沼湿地蓝碳功能的潜在变化趋势和制定蓝碳增汇途径提供理论支持, 助力碳达峰、碳中和目标实现。     

Below-ground biomass in healthy and impaired salt marshes

Twelve salt marshes in south Louisiana (USA) were classified as either impaired or healthy before a summer sample collection of above- and below-ground biomass and determination of sediment accretion rates. The above-ground biomass of plant tissues was the same at both impaired and healthy salt marshes and was not a good predictor of marsh health. However, below-ground root biomass in the upper 30cm was much lower in the impaired marshes compared to the healthy marshes. Compromises to root production apparently occur before there is an obvious consequence to the above-ground biomass, which may quickly collapse before remedial action can be taken. The subsequent change in vertical position of the marsh surface may be equivalent to many years of accretion, and be irreversible within decades without considerable effort. These results are consistent with the hypothesis that it is the plants below-ground accumulation of organic matter, not inorganic matter that governs the maintenance of salt marsh ecosystem in the vertical plane. Reversing the precursor conditions leading to marsh stress before the collapse of the above-ground biomass occurs is therefore a prudent management objective and could be easier than restoration.