Zooplankton species composition in the Oosterschelde (SW Netherlands) before,during and after the construction of a storm-surge barrier

During the period of construction of a storm-surge barrier current velocities decreased strongly and the Eastern compartment of the basin obtained a lagoon-like character. The rotifer Synchaeta spp., already abundant in the neighbouring salt and brackish lakes, profited from this condition. Higher and less fluctuating salinities caused the estuarine character of this compartment to disappear in the post-barrier years. This was reflected in the obscured succession of some Acartia species: the estuarine A. tonsa dominated in pre-barrier- and barrier years during summer, while the marine A. clausi was abundant in the post-barrier Oosterschelde during that time. Also the changed food conditions (Bakker & Vink, 1993) may have played a role in this phenomenon.Further changes in species composition were hardly observed. The main changes were of a quantitative nature (Bakker & Van Rijswijk, 1993; Tackx et al., 1993). A list of the commonly occurring species is given.     

Hypoxic coma as a strategy to survive inundation in a salt-marsh inhabiting spider

Spiders constitute a major arthropod group in regularly inundated habitats. Some species survive a flooding period under water. We compared survival during both submersion and a recovery period after submersion, in three stenotopic lycosids: two salt-marsh species Arctosa fulvolineata and Pardosa purbeckensis, and a forest spider Pardosa lugubris. Both activity and survival rates were determined under controlled laboratory conditions by individually surveying 120 females kept submerged in sea water. We found significant differences between the three species, with the two salt-marsh spiders exhibiting higher survival abilities. To our knowledge, this study reports for the first time the existence of a hypoxic coma caused by submersion, which is most pronounced in A. fulvolineata, the salt-marsh spider known to overcome tidal inundation under water. Its ability to fall into that coma can therefore be considered a physiological adaptation to its regularly inundated habitat.     

The response of salt marsh vegetation to tidal reduction caused by the Oosterschelde storm-surge barrier

In 1986 a sluice gate barrier was completed in the mouth of the Oosterschelde estuary. The barrier has been partially or completely closed during 1986 and the first months of 1987. Consequently the high tides were reduced to such a level that the salt marshes were scarcely flooded. Since April 1987 the barrier has been closed on average during two out of 706 high tides a year. Although the barrier allows tidal exchange the tidal flow has been restricted as a result of the reduced width of the mouth of the estuary. This restriction of the tidal flow caused a 26 cm decrease of mean high water in the eastern part of the estuary. As a result the inundation frequencies of the salt marshes decreased.The response of salt-marsh vegetation to this tidal reduction was analyzed using annual records (1982 till 1990) of species composition in 57 permanent plots in two marshes at the southern shore of the estuary. Analysis of the response of individual species to marsh elevation in the pre and the post-barrier situation revealed that most species moved down the marsh elevation gradient. The first axis of an ordination (DCA-1) was significantly negatively related to inundation frequency. Between 1984 and 1990 all plots were displaced towards a significantly higher ordination score, indicating a trend towards a species composition from higher up the marsh. The position of most plots along DCA-1 remained stable until 1985 and started to increase in 1986 or 1987. The vegetation in plots dominated either by Halimione portulacoides or by Spartina anglica, started to change in 1985. This premature change was attributed to frost damage in January 1985. The initially high rate of change along DCA-1 decreased in 1989 and 1990. This would suggest that the vegetation re-equilibrated with the newly established tidal conditions. Further analysis revealed no significant difference in the relation between inundation frequency and sample score along DCA-1 between 1984 and 1990. This corroborates the view that species composition had re-equilibrated with the tidal conditions. Along DCA-2 the samples were displaced towards a significantly higher score as well. The change was attributed to an increase of the perennial Halimione and the annual Suaeda maritima. The annual Atriplex hastata displayed an increase from 1986 till 1988, but strongly declined in 1990. The transient response of this annual was described by DCA-3.During the pre-barrier phase several attempts had been made to predict the vegetation response to tidal reduction. We developed a multivariate model (CCA) with inundation frequency as a constraining factor to describe the relation between inundation frequency and species composition in the pre-barrier phase in 1984. Next we used this model to predict the score of the samples along CCA-1 in the post-barrier phase from the inundation frequencies in 1990. The actual ordination score in 1990 was calculated from the observed species composition. The predicted ordination scores did not differ significantly from the observed ones. In retrospect it is concluded that the response of the vegetation to the reduction of the tides, as far as the fraction of species composition which is related to inundation frequency is concerned, could have been predicted by our model.

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Zygotic dormancy underlies prolonged seed development in Franklinia alatamaha (Theaceae): a most unusual case of reproductive phenology in angiosperms

In Franklinia alatamaha (Theaceae) an extended period of dormancy, associated with winter, separates pollination in the late summer and autumn from observable fruit growth, which occurs during the subsequent summer season. Here, ovule and early seed development were examined to decipher the timing of reproductive events that underlie this unusual phenological pattern. Female gametophytes were found to be mature before pollination. Evidence for double fertilization was observed soon after pollination. Early endosperm development progresses for up to 3 months after fertilization but comes to a standstill at the onset of winter. The zygote becomes dormant shortly after fertilization and does not divide during the autumn or winter. At the start of the following growing season, endosperm development is reinitiated and the first events associated with the formation of an embryo occur. Seed development is completed roughly at the same time as new flowers are opening, a full year after pollination and fertilization. Dehiscence of fruits does not occur until later in autumn. The prolonged zygotic dormancy in Franklinia is exceedingly rare among angiosperms and differs markedly from patterns of delayed fertilization that typically underlie extended periods between pollination and seed maturation in other temperate, perennial species.     

Effects of sea level rise on mangrove Avicennia population growth,colonization and establishment: Evidence from a field survey and greenhouse manipulation experiment

Mangroves will either face longer tidal inundation or retreat landwards in response to on-going accelerated sea level rise. However, little is known about the growth, regeneration or colonization of mangrove seedlings under the different tidal inundation regimes associated with accelerated sea level rise. In the present study, a field survey and a greenhouse mesocosm experiment were conducted to evaluate possible effects of accelerated sea level rise on colonization, establishment and seedling growth of a mangrove pioneer species, Avicennia marina. Avicennia populations at different elevations of the intertidal zone on Xiamen Island in Fujian Province, China showed distinctive patterns in both seedling density and plant age. The seedlings at lower elevations had less annual biomass accumulation and population productivity, but higher shoot to root ratios, suggesting that elevation has positive effects on seedling growth. The greenhouse mesocosm experiment with 1-year-old A. marina seedlings utilized five inundation periods (0, 2, 4, 6 and 12 h in a semidiurnal tidal cycle) and two inundation depths (root immersion and canopy immersion). Both inundation period and depth exerted significant and negative effects on biomass accumulation, photosynthetic rate, leaf electron transportation and water use efficiency. However, the negative effects of canopy immersion were more profound than root immersion. Canopy immersion exacerbated the effects of prolonged inundation, with no seedlings surviving under canopy immersion at the 12-hour treatment. These results suggest that at lower elevations with higher sea level, canopy immersion will have greater negative effects on seedling colonization, establishment and early growth of A. marina. This finding is instrumental in predicting the future dynamics of mangrove forests under increasing sea levels.     

The importance of salt-marsh wetness for seed exploitation by dabbling ducks Anas sp.

The relationship between the inundation of a salt marsh in southeast Denmark not subject to lunar tides and the availability and predation of seeds of the annuals Salicornia spp. and Suaéda maritima by autumn staging dabbling ducks was studied by carrying out exclosure experiments over the course of 2 years. There was a marked difference in the wetness of the salt marsh between the two study years, which resulted in distinct temporal patterns of salt-marsh use by dabbling ducks. In both years, the depletion of seeds of both Salicornia spp. and S. maritima was initiated subsequent to the flooding of the sample transects, which also induced the gradual release of seeds from the plants within the exclosures. Nevertheless, seeds were removed more rapidly in plots visited by dabbling ducks than in the exclosures. The predation of seeds took place as soon as the individual plants had been fully submerged, but before the seeds were released from the plants. The timing of flooding events during early autumn may potentially affect the availability of the salt-marsh seed stock. Therefore, weather conditions may impose critical constraints on the feeding opportunities for dabbling ducks during autumn migration on non-tidal salt marshes.     

The effect of experimental inundation and sediment deposition on the survival and growth of two herbaceous riverbank plant species

Flow and sediment regimes of floodplain rivers around the world have been changed by river regulation and land management, altering the ecological processes structuring riparian plant communities. However, the combined effects of inundation and sedimentation processes on riparian plant survival and growth are poorly understood. The separate and interactive effects of inundation duration and sedimentation depth on survival, growth and vegetative reproduction of two herbaceous clonal river bank plant species, Carex bichenoviana and Calotis scapigera, were investigated in an outdoor laboratory experiment. Plants were grown under three levels of sediment addition and two levels of inundation duration and were harvested after 60 days. Both species survived up to 30 days inundation in the absence of sediment. Growth and vegetative reproduction were increased for C. scapigera, but reduced for C. bichenoviana. Sediment deposition during periods of inundation reduced survival of both species, with C. scapigera unable to survive. Carex bichenoviana survived, but as inundation duration and depth of sediment increased, growth and vegetative reproduction were reduced. Our results indicate sediment deposition during flooding can change the tolerance of riparian plant species to inundation, reducing survival and growth.     

Some Aspects of the Reproductive Biology of Limonium vulgare Mill., and Limonium humile Mill.

The germination of two species of salt-marsh plants Limoniumvulgare Mill., and L. humile Mill., is investigated in relationto their ecology. Germination is inhibited by sea water, butgermination even in fresh water is low. Inhibition of germinationby sea water is negatively correlated with water uptake. However,a treatment in sea water stimulates subsequent germination infresh water. The effect of the sea-water treatment appears tobe one of osmotic shock which weakens the seed coat. Variationin response to the treatment is partly genetical and partlyenvironmental. Seeds in different parts of the salt-marsh willbe subject to differing conditions, both during seed developmentand during the interval between seed maturation and seed dispersal.In connection with this the effect of leaching by sea waterand fresh water is considered. The response to a sea-water treatmentvaries with time. Considered in the long term it shows somecyclical variation which might correspond with the tidal cycle.It is shown that seeds of Limonium can survive long periodsin sea water. There is a slow rate of germination during immersion.Most of those seeds which do not germinate in sea water surviveto germinate rapidly in fresh water. The response to oxygenconcentration, light, and temperature is also investigated.Experiments on seedling growth and establishment show the needfor fresh water and a suitable substrate. Mud gives a lowergermination but a higher survival, while sand gives a highergermination but a lower survival. The dispersal of Limoniumseeds in nature is investigated. Seeds of Limonium are mostabundant in the drift line, but even these only represent asmall fraction of the total produced. It is considered thatreproduction by seed is only important when a new area is beinginvaded. Subsequent reproduction is largely by vegetative spread.The ecological implications of the germination experiments arediscussed.     

Manganese as an ecological factor in salt marshes

The plant available manganese concentration (Mn²⁺) of salt-marsh sediments was compared to that of acidic and neutral soils. The mean soil-manganese concentration was higher in the top 1 cm of salt-marsh soil than in the neutral soil and comparable to that of the acidic soil (0–5 cm). A peak in the soil-manganese concentration in the upper marsh was observed one week after the spring tide but this effect was not evident in the lower marsh. Despite these differences, there was no correlation between mean manganese concentration and position on the marsh.The response to manganese of salt-marsh halophytes was studied by measuring growth and root elongation in a range of Mn²⁺ concentrations with and without sodium chloride. Although there was a differential response to manganese between salt-marsh species, manganese resistance was not related to position on the marsh. Most of the species investigated were tolerant of Mn²⁺ at concentrations higher than normally recommended for plant growth. Moreover a salt-marsh ecotype of Festuca rubra was found to have a higher manganese resistance than an inland ecotype of the same species.When sodium chloride was included in the growth medium, salt-marsh plants had a greatly increased resistance to manganese associated with a reduced uptake. This effect is reflected in the tissue-manganese concentration which was lower than in Deschampsia flexuosa although both groups of plants were exposed to a similar range of Mn²⁺ concentrations. It is concluded that sodium chloride markedly reduces the phytotoxicity of manganese in salt marshes.Nomenclature following Clapham, Tutin & Warburg (1968). Flora of the British Isles.The work was carried out while one of us (C. E. Singer) was in receipt of an SERC studentship, which is gratefully acknowledged.     

Reproductive strategies may explain plant tolerance to inundation: A mesocosm experiment using six marsh species

We investigated the impact of inundation duration on sexual and asexual reproduction strategies in six hygrophilic angiosperm species (2 annuals - Ranunculus sardous and Ranunculus ophioglossifolius, 2 rhizomatous - Juncus articulatus and Eleocharis palustris, and 2 stoloniferous species - Glyceria fluitans and Agrostis stolonifera). Plant growth during three inundation durations (natural, +3 weeks, +6 weeks) was assessed in 20 mesocosms. Biomass and reproductive traits were measured at the onset of flowering and at seed set.The six species maintained or increased their total mass in response to increased inundation period, suggesting a good tolerance to inundation stress. No changes were recorded in flowering time for the five species that flowered. The two annual species increased the number of flowers and mass allocated to sexual reproduction with increased duration of inundation, promoting extensive seed production. The response was species-specific in perennial species with an either a positive or a negative Gaussian curve of both traits in response to the increased duration of inundation. The duration of inundation affected ramet production in all perennial species except A. stolonifera, promoting ramet production in two out of the three species. In all species except E. palustris, the length and number of stolons or rhizomes were maintained, suggesting the importance of these connections in the resistance of perennial plants to flooding stress. These species occurring in repeatedly inundated habitats thus appeared to be tolerant to long inundations thanks to both sexual reproduction and vegetative propagation.     

Does an ant-dispersed plant,Viola reichenbachiana,suffer from reduced seed dispersal under inundation disturbances?

Many plant species use ants as seed dispersers. This dispersal mode is considered to be susceptible to disturbances, but the effect of natural, small-scale disturbances is still unknown. We investigated how small-scale disturbances due to inundation affect seed dispersal in Viola reichenbachiana, a dominant myrmecochorous herb in riparian forests. Inundation disturbances were high in depressions and low on hillocks of the forest floor. We found that V. reichenbachiana was similarly abundant at highly and less disturbed sites, contrary to other, non ant-dispersed species. We also found that the motivation of ants to disperse seeds was higher at highly disturbed sites. Nevertheless, the number of seeds dispersed was similar at highly disturbed and weakly disturbed sites and seedlings of V. reichenbachiana were equally abundant. We conclude that inundation disturbances do not interfere with mutualistic seed dispersal by ants in V. reichenbachiana, and suggest that this may possibly contribute to the persistence of V. reichenbachiana under inundation.     

Impact of nest relocation on the reproductive success of Loggerhead Turtles,Caretta caretta,in the Göksu Delta,Turkey (Reptilia: Cheloniidae)

When the nests of marine turtles are at a risk of inundation, relocation of the nests are often used in the conservation measures. Here, I determined the effect of nest relocation on Loggerhead Turtle (Caretta caretta) egg hatching success during the 2013 and 2014 nesting seasons in the Göksu Delta, Mersin, Turkey. I compared natural and relocated clutches, including those relocated before and after inundation, and evaluated 102 (94.6%) and 63 (81.1%) of survived nests in 2013 and 2014 respectively. Relocated nests experienced a 30% decrease in hatching success and a more prolonged incubation period compared to nests left in situ. Egg failure in nests relocated before and after inundation was similar in early-stage embryos, whereas it was three-fold higher in mid-stage embryos and two-fold lower in late-stage embryos. Thus, there was no significant difference in overall hatching success between the two relocation types. Moreover, there was no effect of delayed relocation of nests after inundation on hatching success. Possible impacts specific to the nesting site should be considered and explored before using nest relocation as a conservation tool. The relocation approach is recommended for nests at a high risk of inundation when the loss of nests is inevitable.     

Conditions for seedling establishment and probable function of seed in the clonal sedge Carex rugulosa Kük. in riverside marshes

Carex rugulosa Kük. forms large meadows in moist zones along estuarine riversides. These meadows are usually maintained by rhizomatous ramet production (clonal growth), and the appearance of seedlings is uncommon. We studied the conditions required for seedling establishment. In areas in which clonal ramets were dense (906–1050/m²), and the relative light intensity at ground level was low (0.8–3.8%), seed germination was entirely suppressed. In contrast, many seedlings (288–513/m²) were observed in areas in which clonal ramets had been considerably reduced (13–269/m²) by continuous inundation in the previous year. In these areas, the relative light intensity had increased (20.5–79.3%). It was inferred that seeds resisted the prolonged inundation that killed the ramets, and that germination was induced under these open conditions. These results suggest that the main functions of sexual reproduction in C. rugulosa are recovery after severe damage to vegetation using seeds from the seed bank and the dispersal of seeds to areas without dense vegetation. Most seedlings died when the marshy meadow was temporarily inundated by heavy rain. However, transplanted seedlings survived better at artificially elevated sites that were not submerged. This shows that inundation during the seedling stage impedes seedling survival. Therefore, seeds can contribute to recovery after ramet decline or colonization to open area when (i) the rainfall is not heavy to cause seedling submergence or (ii) seeds are dispersed to higher area which is not inundated.     

The influence of vegetation,salinity, and inundation on seed banks of oligohaline coastal marshes

Sea level rise may alter salinity and inundation regimes and create patches of open water in oligohaline coastal marshes, potentially affecting the composition and germination of seed bank species. We conducted seedling emergence experiments to: (1) examine the effects of standing vegetation on the seed banks of three oligohaline marsh communities in coastal Louisiana (dominated by Paspalum vaginatum Sw., Sagittaria lancifolia L., or Spartina patens (Ait.) Muhl., respectively); and (2) investigate the effects of salinity and inundation regime on germination of seed bank species. We also studied the effect of a temporary increase in salinity (to simulate a salt water intrusion event) on the viability of buried seeds. We found that the presence or absence of vegetation within a community affected the abundance of some species in the seed bank but had little effect on species composition. Also, the seed banks of the three communities exhibited considerable overlap in species composition and had similar species richness (10–11) and diversity (antilog Shannon-Weaver diversity index = 6.5–7.1), despite differences in vegetation type. Higher salinities and flooding reduced seedling emergence for most species; few species emerged at salinities above four parts per thousand (ppt), and only Sagittaria lancifolia and Eleocharis parvula germinated well under flooded conditions. A temporary increase in salinity did not affect species richness or seedling emergence of most species. Our results suggest that differences in vegetation may have little effect on the composition of seed banks of oligohaline marshes. However, higher salinities and greater depth and duration of inundation (anticipated as global sea level continues to rise) may decrease recruitment of seed bank species, reducing their abundance in oligohaline marsh communities.     

For everything a season: Smoke-induced seed germination and seedling recruitment in a Western Australian Banksia woodland

Abstract The influence of factors associated with fire on seed germination of Australian native species is generally well documented, but examples involving the use of smoke as a fire analogue for ecological research remain limited. The role of season of treatment in the efficacy of smoke as a promotive germination agent was investigated over two growing seasons using natural soil stored seedbanks in Banksia woodland near Perth, Western Australia. Smoke was applied to unburnt sites in the autumn, winter and spring of 1994. Germinant emergence and seedling survival of 37 species representing 18 families was monitored in both unburnt sites and in adjacent, recently burnt sites until the second spring after treatment (October 1995). Recruitment from seed was found to be profoundly affected by the season in which dormancy breaking treatment had been applied. The promotive effect extended beyond the initial year of application. For the majority of the species investigated, application of smoke to unburnt sites in autumn promoted a significantly greater germination response than treatment in winter or spring. In only three cases (introduced annuals, the Fabaceae and Hibbenia amplexicaulis) did autumn smoke treatment not yield better germination than in summer-burnt counterparts. However, in almost half of the cases examined, proportions of seedlings surviving past their first summer after emergence in burnt areas were consistently greater than those in smoked or untreated sites. Most notably, no seedlings emerging during the spring of the first year of study survived into the following summer. Implications of the results with respect to future seed bank research and management of native vegetation are discussed.     

Vertical migrations in two hygrophilous species of micro‐snails in relation to time of the year and habitat type

The aim of this study was to investigate vertical migration behaviors in two species of hygrophilous micro‐snails, Vertigo moulinsiana and Vertigo angustior, in relation to the time of the year (spring and summer) at two sites that differ in ground water level (periodically inundated site and non‐inundated site). The study shows different patterns of vertical migrations in the studied species. Vertigo angustior demonstrated a strong affinity to the litter layer (a weak tendency for vertical movements), independent of the time of year and site studied. By contrast, V. moulinsiana showed a clear tendency for vertical migrations, which differed depending on the time of year and site. These differences may be related to the spatial segregation of microhabitats used by these two species at the sites studied and to differences in the ability to resist inundation. Vertigo angustior is associated with microhabitats which are not subjected to prolonged inundation and tolerates a brief submersion. The periodic character of vertical migrations may suggest the effect of endogenous factors related to reproduction in V. moulinsiana. Some plasticity of this behavior in relation to habitat conditions demonstrated by this species may be an adaptation to live in unpredictably flooded environments.     

Status of arbuscular mycorrhizal fungi (AMF) in the Sundarbans of India in relation to tidal inundation and chemical properties of soil

The arbuscular mycorrhizal status of fifteen mangroves and one mangrove associate was investigated from 27 sites of three inundation types namely, diurnal, usual springtide and summer springtide. Roots and rhizospheric soil samples were analysed for spore density, frequency of mycorrhizal colonization and some chemical characteristics of soil. Relative abundance, frequency and spore richness of AMF were assessed at each inundation type. All the plant species except Avicennia alba exhibited mycorrhizal colonization. The study demonstrated that mycorrhizal colonization and spore density were more influenced by host plant species than tidal inundation. Forty four AMF species belonging to six genera, namely Acaulospora, Entrophospora, Gigaspora, Glomus, Sclerocystis and Scutellospora, were recorded. Glomus mosseae exhibited highest frequency at all the inundation types; Glomus fistulosum, Sclerocystis coremioides and Glomus mosseae showed highest relative abundance at sites inundated by usual springtides, summer springtides and diurnal tides, respectively. Spore richness of AMF was of the order usual springtide > diurnal > summer springtide inundated sites. The mean spore richness was 3.27. Diurnally inundated sites had the lowest concentrations of salinity, available phosphorus, exchangeable potassium, sodium and magnesium. Statistical analyses indicated that mycorrhizal frequency and AMF spore richness were significantly negatively correlated to soil salinity. Spore richness was also significantly negatively correlated to available phosphorus. The soil parameters of the usual springtide inundated sites appeared to be favourable for the existence of maximum number of AMF. Glomus mosseae was the predominant species in terms of frequency in the soils of the Sundarbans.     

Role of Brycinus lateralis (Teleostei: Alestidae) in dispersal and germination of Nymphaea nouchali (Angiospermae: Nymphaeaceae) seeds on a seasonal floodplain of the Okavango Delta,Botswana

Seed passage through the gut of vertebrates can be important for seed dispersal, but might influence seed viability. The ability of seeds to germinate after ingestion by seed-eating fish is important for the population dynamics of some plant species, and significant in the evolution of plant–fish interactions. Certain fish in the Okavango Delta, Botswana, are fruit- and seed-eaters and could act as seed dispersers. We sampled 14 fish species in 2013, finding Nymphaea nouchali var. caerulea seeds in the digestive tracts of eight, most commonly in the striped robber Brycinus lateralis. Seeds extracted from the gut of this species had an overall mean germination success of 11.7%. This fish species might well be a legitimate seed disperser, having a positive effect on seed dispersal from parent plants in the Okavango Delta. The current study represents one of the first investigations of the likelihood of seed dispersal by fish on the African continent.     

Seasonal Dependent Effects of Flooding on Plant Species Survival and Zonation: a Comparative Study of 10 Terrestrial Grassland Species

Past research has provided compelling evidence that variation in flooding duration is the predominant factor underlying plant species distribution along elevation gradients in river floodplains. The role of seasonal variation in flooding, however, is far from clear. We addressed this seasonal effect for 10 grassland species by testing the hypothesis that all species can survive longer when flooded in winter than when flooded in summer. We carried out an inundation experiment under simulated conditions of summer and winter flooding in the greenhouse. The results showed that all species survived longer under winter floods than under summer floods. However, responses upon flooding were species-specific. All summer flood-tolerant species had high tolerance for winter floods as well, but summer flood sensitive species survived either a little longer, or dramatically longer when flooded under simulated winter conditions. Next, we examined whether winter or summer survival best predicted the lower distribution limits of the species as measured in a natural flooding gradient after an extremely long winter flood. We found a strong significant relationship between the lower distribution limits of species in the field and their tolerance to summer floods, although we measured the lower limits 14 years after the latest major summer flood. In contrast, no such significant relationship existed with species tolerance to winter floods. Some relatively intolerant species occurred at much higher floodplain elevations as was expected from their tolerance to winter inundation in the experiments. This suggests that zonation patterns as created by occasional summer floods may be maintained for a long time, probably due to the limited ability of species to re-colonise lower positions in the floodplain.     

Policy planning in the Oosterschelde estuary

During the execution of the Delta project the awareness of the environmental values of estuaries was growing strongly. This awareness has led to a reconsideration of the original proposal to close off the Oosterschelde, and to an alternative view of the management of the estuarine areas and of the civil engineering structures. Now, the completed storm-surge barrier and two compartmentation dams, together with the reinforced dikes, guarantee a sufficient safety against flooding.After the decision to build a storm-surge barrier it was soon recognized that a coherent plan would be needed for the management and development of the Oosterschelde. A policy plan was accordingly produced in which the broad outlines of the policy were set out regarding the various potential uses of the estuary and the possible means of effecting them. Because of the periodic improvement and adjustment of the policy plan an environmental research programme was set up. Also, environmental research played a significant role in the decision-making processes for the completion of the barrier and the compartmentation dams as well as for the management of the storm-surge barrier.The coping-stone of the Delta Project marks the beginning of a new period in the water management of the Netherlands: the period of integrated water management.