Dynamics of seasonal bicarbonate supply in a dune slack: effects on organic matter, nitrogen pool and vegetation succession

The seasonal variation in groundwater composition was studied in an old dune slack complex on the Wadden Sea island of Schiermonnikoog that recently had lost practically all rare basiphilous plant species. In order to assess its restoration perspectives the groundwater acid neutralization capacity (ANC _aq ) of various sites was discussed in relation to the decalcification depth, amount of organic matter and amount of nitrogen in the topsoil layer. The decalcification depth reflected the (former) hydrological regime quite well; shallow in the exfiltration zone and deep in the infiltration part of the slack. The pattern of HCO₃ ^–, SO₄ ^2– and pH reflected the functioning of a flow-through lake, even when the slack was not flooded. Relics of low productivity (mesotrophic) vegetation (Samolo-Littorelletum) with rare species were related to low amounts of organic matter and nitrogen in the topsoil layer. At this site the variation in HCO₃ ^– concentration was also relatively low during the year. It was hypothesized that a regular supply of acid buffering components from calcareous soil layers buffers the pH at a high level and slows down the accumulation of organic matter thus creating opportunities for the rare basiphilous plant species. A possible biological control mechanism for these processes was discussed. A reconstruction of the former, present and future eco-hydrological situation was made of the dune slack.     

Alternative stable states in a wet calcareous dune slack in The Netherlands

Abstract. Evidence is presented for the occurrence of alternative stable states in a wet calcareous dune slack on the Frisian island of Texel, The Netherlands. An early pioneer stage (0.5 kgm^?2 total standing crop) and a more productive later successional stage (2.9 kg m^?2) occur side by side, with sharp boundaries between them. The pioneer vegetation has been recorded at the site for more than 62 yr. These features indicate the occurrence of a positive‐feedback mechanism that has led to alternative stable states. Analyses of ground and surface water composition, and decalcification depths, indicated that hydrologically the study site can be characterized as a flow‐through slack, with exfiltration of calcareous groundwater on one side and infiltration of surface water on the other side of the slack. These differences in hydrological conditions have led to distinct differences in environmental conditions within the dune slack. The occurrence of the two successional stages can, however, not be explained by differences in hydrological conditions since both stages occur side by side in the centre of the dune slack. It is, therefore, more likely that biotic interactions are the cause of the vegetation pattern. Three possible mechanisms for feedback processes are discussed: (1) enhanced nitrogen loss; (2) sulfide toxicity and (3) nutrient accumulation in internal cycle.     

Influence of base flow stream bank seepage on riparian zone nitrogen biogeochemistry

We examined the effect of sustained stream bank seepage during base flow conditions on the pore water nitrogen biogeochemistry of two riparian zones in lowland agricultural areas in southern Ontario, Canada. Nitrate, ammonium and dissolved oxygen concentrations in riparian subsurface water over a two-year period showed well-organized spatial patterns along stream bank seepage flow paths that extended seasonally up to 25 m inland. High levels of dissolved oxygen and NO₃⁻ in stream inflow were depleted rapidly at the stream bank interface suggesting the occurrence of aerobic microbial respiration followed by denitrification. A zone of NH₄⁺ accumulation persisted in more anaerobic sediments inland from the bank margin, although the magnitude and intensity of the pattern varied seasonally. A bromide tracer and NO₃⁻ co-injection at the stream bank interface indicated that bank seepage occurred along preferential flow paths in a poorly sorted gravel layer in the two riparian zones. Depletion of NO₃⁻ in relation to co-injected bromide confirmed that the bank margin was a hot spot of biogeochemical activity within the riparian zone. Conceptual models of humid temperate riparian zones have focused on nitrogen biogeochemistry in relation to hillslope to stream hydrologic flow paths. However, our results suggest that sustained stream bank inflow during low flow conditions can exert a dominant control on riparian nitrogen cycling in lowland landscapes where level riparian zones bounded by perennial streams receive limited subsurface inflows from adjacent slopes.     

Groundwater seepage stimulates the growth of aquatic macrophytes

1. The impact of groundwater seepage on the growth of submerged macrophytes was investigated in experiments on the isoetid Littorella uniflora and the elodeid Myriophyllum alterniflorum both in the laboratory and in the field. Isoetids rely mostly on sediment‐derived CO₂ and nutrients via root uptake, whereas elodeids acquire their inorganic carbon and nutrients from the water column. We thus hypothesised that L. uniflora would respond positively to seeping ground water as it should improve both CO₂ and nutrient supply. 2. Laboratory experiments were conducted by percolating vegetated cores containing natural sediment or technical sand with artificial ground water of high CO₂ concentrations and with either high or low levels of nutrients. Field experiments were conducted in the oligotrophic Lake Hampen, Denmark, with custom‐built seepage‐growth chambers that permitted a near‐natural flow‐through of seeping ground water. Chambers with a solid bottom, and thus no flow‐through of seeping ground water, served as controls in both laboratory and field experiments. In the field, seepage chambers were installed at a site with relatively high seepage fluxes (ground water from forest catchment), at a site with much lower seepage fluxes but with higher nutrient concentrations (ground water from agricultural catchment) and at a reference site with no net discharge or recharge of ground water. 3. Positive growth responses were observed in the field at transects with high groundwater discharge compared to the control chambers with no seepage. No growth response was observed at the reference transect with low or alternating direction of groundwater seepage. The growth rates of L. uniflora in the field were significantly higher in seepage treatments compared to control treatments, and final plant mass was up to 70% higher than that for plants where seepage was excluded. In areas with high groundwater discharge, a strong positive correlation was found between groundwater seepage fluxes, growth rates, and final plant mass for L. uniflora, while there was no such relationship at the reference transect. The growth of M. alterniflorum was also significantly affected by groundwater seepage, but to a lesser degree than L. uniflora. Laboratory experiments generally showed the same trend for both L. uniflora and M. alterniflorum, and the positive influence of seeping ground water was apparently related to increased inorganic carbon supply and, to a lesser degree, improved nutrient availability. 4. Groundwater discharge results in enhanced growth of isoetids and to some extent elodeids inhabiting a groundwater‐fed softwater lake. We propose that the shallow dense vegetation present where most of the discharge takes place acts as a biological filter that retains nutrients that otherwise would end up in the water column and could result in increased algal growth.     

Changes in soil and vegetation during dune slack succession

Many rare plant species occur in Dutch wet dune slacks, particularly in the Junco baltici‐Schoenetum nigricantis. For nature management it is important to understand the processes controlling the presence of these basiphilous early successional communities, which is why we investigated vegetation and soil development during succession in coastal dune slacks. We compared 12 chronosequential stages of 0, 2, 4, 9, 10, 13, 25, 30, 43, 60, 70 and 85 yr in five different dune slack systems. In four of these locations turf had earlier been removed in order to restore the basiphilous pioneer stage. The main variation in the vegetation is related to the acidification/soil enrichment gradient and the salinity/maritime gradient. During succession, organic matter accumulates and acidification takes place. Maritime influence can buffer the soil and postpone the succession of basiphilous pioneer communities for many years. A significant correlation with age was found for 18 variables. Multiple regressions predicted changes in the vegetation (dependent variables: biomass, cover of Salix repens, Calamagrostis epigejos and Schoenus nigricans) as a function of acidification, organic matter accumulation, increase in available P and presence of Na in the soil. We conclude that natural ageing of the vegetation and the associated process of accumulation of biomass drive succession in this hydrosere. The underlying soil processes are acidification and organic matter accumulation. During succession dominance shifts from S. nigricans to S. repens or C. epigejos. Maintenance of the pioneer character of the habitat is only possible by local intervention or by natural or man‐induced dune forming. The effect of sod‐stripping depends on the environmental conditions and, in case of acidification, success is limited. Succession can be postponed by mowing.     

Vegetation development in dune slacks: the role of persistent seed banks

Abstract. The soil seed bank composition was determined at four sites in the dune slack ‘Koegelwieck’ on the Dutch Wadden Sea island of Terschelling. At three different sites in the slack, where sod-cutting experiments down to the mineral sand had been carried out, the established vegetation and seed bank were assessed after 5, 9 and 39 yr of undisturbed development, respectively. In addition, a fourth site in the slack was investigated, where vegetation development had proceeded for 80 yr since plant colonization of bare soil and where nowadays a vegetation dominated by Calamagrostis epigejos and Salix repens occurs. Together these four sites can be regarded as a chronosequence of dune slack formation. Clear time sequences were detected in the seed bank data. Many late successional species showed a significant increase in the number of seeds during the succession. Some of the early successional basiphilous pioneer species such as Anagallis minima, Centaurium littorale, Littorella uniflora, Radiola linoides and Samolus valerandi, showed either a decrease during the time of succession or an optimum in the vegetation while remaining present in the seed bank in low but detectable numbers. They could, therefore, play a role in re-establishment of the vegetation after sod-cutting. One of the target species, Schoenus nigricans, established within a few years after removal of the sod. However, no seeds of this species have been detected in the soil below either of the successional stages. Based on the species disappearance from the established vegetation and based on the independent data of Thompson et al. (1997) an estimation of seed longevity could be made for several Red List species of wet dune slacks.     

Nutrient limitation and vegetation changes in a coastal dune slack

Abstract. Basiphilous pioneer species are among the most endangered plant species in The Netherlands. They find most of their refuges in young coastal dune slacks, especially on the Wadden Sea islands. For the purpose of nature management it is important to know which processes control the presence of basiphilous pioneer communities, and to learn about the nature of slacks harbouring the concerning successional sequences. In a large dune slack on the Island of Terschelling, we assessed soil nutrient status and tested for nutrient limitation in four chronosequential stages: 2, 6, 37 and ca. 80 yr of age. Stage 2 harboured a basiphilous pioneer vegetation; in the stages 3 and 4 a dense vegetation of dwarf shrubs and grasses occurred. Soil organic matter and nutrient concentrations in each stage were measured in 1991. In 1992 and 1993 fertilizers were applied to all stages to detect nutrient limitation. Rates of accumulation of organic matter, nutrients and above-ground biomass were estimated. When interpreted as successional stages, the different stages represent a sequence as expected on the basis of general successional theory. There was a peak in yearly nutrient accumulation between the 6- and 37-yr old stage and a steady state after ca. 80 yr. Between the first two and the latter two stages a shift occurred from allogenic to autogenic succession which correlated with a shift in emphasis from available nutrients to light availability as limiting resources. Basiphilous pioneer species suffered only deficiency of nitrogen, probably because of their low phosphorus requirements. It is concluded that in dune slack habitats, in addition to a low nutrient availability in general, a very low phosphorus availability favours basiphilous pioneer species to species showing co-limitation of nitrogen and phosphorus as found in some grasses and dwarf shrubs. A comparison between the effects of lime addition and the effects of nitrogen and phosphorus additions suggests that, in the early stages, soil buffering increases the availability of nitrogen and inhibits the availability of phosphorus. Sod cutting is an effective technique for restoring basiphilous pioneer vegetation, when slacks are acidified only superficially and buffering-mechanisms can be reactivated. Yearly mowing and removing of standing crop may prolong the life-span of basiphilous pioneer vegetation, when soil acidification has not yet dropped below pH 6.     

Groundwater input affecting plant distribution by controlling ammonium and iron availability

Question: How does groundwater input affect plant distribution in Alnus glutinosa (black alder) carrs? Location: Alder carrs along the river Meuse, SE Netherlands. Methods: Three types of site, characterized by groundwater flow, were sampled in 17 A. glutinosa carrs. Vegetation and abiotic data (soil and water chemistry) were collected and analysed using a Canonical Correspondence Analysis. Based on the results, a laboratory experiment tested the effect of groundwater input (Ca²⁺) on pore water chemistry (NH₄⁺ availability). Results: Environmental factors indicating groundwater input (Ca²⁺ and Fe²⁺), correlating with the NH⁺₄ concentration in the pore water, best explained the variation in plant distribution. NH₄⁺ availability was determined by Ca²⁺ input via the groundwater and subsequent competition for exchange sites in the sediment. As a result, nutrient‐poor seepage locations fully fed by groundwater were dominated by small iron resistant plants such as Caltha palustris and Equisetum fluviatile. More nutrient‐rich locations, fed by a combination of groundwater and surface water, allowed the growth of taller iron resistant plant species such as Carex paniculata. Nutrient‐rich locations with stagnating surface water were hardly fed by groundwater, allowing the occurrence of fast growing and less iron tolerant wetland grasses such as Glyceria fluitans and G. maxima. Conclusion: Groundwater input affects plant composition in A. glutinosa carrs along the river Meuse by determining nutrient availability (ammonium) and concentrations of toxic iron.     

Is succession in wet calcareous dune slacks affected by free sulfide?

Abstract. Consequences of sulfide toxicity on succession in wet calcareous dune slacks were investigated. Sulfide may exert an inhibitory effect on dune slack plants, but several pioneer species exhibit ROL (Radial Oxygen Loss) and thereby protect themselves against free sulfide. Under oxic conditions free sulfide will be oxiginated to harmless sulfate. However, successive species when not capable of ROL may be sensitive to free sulfide and cannot invade the area. Therefore, the occurrence of free sulfide may have a stabilizing effect on the pioneer vegetation. Data on the vertical distribution of oxygen, redox and sulfide were collected in mesocosms with Littorella uniflora or Carex nigra, with and without microbial mats and compared to control mesocosms. Also, in situ data were collected in a dune slack on the Frisian Island of Texel. In the mesocosms, free sulfide was detected only at nighttime in C. nigra populated mesocosms and in unvegetated units, but not in L. uniflora vegetated mesocosms. In the field, sulfide and redox profiles showed distinct differences between the groundwater exfiltration and infiltration site of the dune slack. At the exfiltration site, sulfide was only occasionally found; in contrast, measurable amounts of free sulfide were regularly found at the infiltration site of the slack. Since Phragmites australis dominates in the infiltration site of the slack, the results suggest that free sulfide accelerate the succession, rather than slowing it down by the exclusion of some plant species.     

Oxygen supply and the adaptations of animals in groundwater

1. The first part of this review focuses on the oxygen status of natural groundwater systems (mainly porous aquifers) and hyporheic zones of streams. The second part examines the sensitivity of groundwater organisms, especially crustaceans, to low oxygen concentrations (< 3.0 mg L^?1 O₂). 2. Dissolved oxygen (DO) in groundwater is spatially heterogeneous at macro- (km), meso- (m) and micro- (cm) scales. This heterogeneity, an essential feature of the groundwater environment, reflects changes in sediment composition and structure, groundwater flow velocity, organic matter content, and the abundance and activity of micro-organisms. Dissolved oxygen also exhibits strong temporal changes in the hyporheic zone of streams as well as in the recharge area of aquifers, but these fluctuations should be strongly attenuated with increasing distance from the stream and the recharge zone. 3. Dissolved oxygen gradients along flow paths in groundwater systems and hyporheic zones vary over several orders of magnitude (e.g. declines of 9 × 10^?5 to 1.5 ×10^?2 mg L^?1 O₂ m^?1 in confined aquifers and 2 × 10^?2 to 1 mg L^?1 O₂ m^?1 in parafluvial water). Several factors explain this strong variation. Where the water table is close to the surface, oxygen is likely to be consumed rapidly in the first few metres below the water table because of incomplete degradation of soil-generated labile dissolved organic carbon (DOC) in the vadose zone. Where the water table is far from the surface, strong oxygen depletion in the vicinity of the water table does not occur, DO being then gradually consumed as groundwater flows down the hydraulic gradient. In unconfined groundwater systems, oxygen consumption along flow paths may be compensated by down-gradient replenishment of DO, resulting either from the ingress of atmospheric oxygen or water recharge through the vadose zone. In confined groundwater systems, where replenishment of oxygen is impossible, the removal time of DO varies from a few years to more than 10 000 years, depending mainly on the organic carbon content of the sediment. Comparison of the hyporheic zones between systems also revealed strong differences in the removal time and length of underground pathways for DO. This strong variability among systems seems related to differences in contact time of water with sediment. 4. Although groundwater macro-crustaceans are much more resistant to hypoxia than epigean species, they cannot survive severe hypoxia (DO < 0.01 mg L^?1 O₂) for very long (lethal time for 50% of the population ranged from 46.7 to 61.7 h). In severe hypoxia, none of the hypogean crustaceans examined utilized a high-ATP yielding metabolic pathway. High survival times are mainly a result of the combination of three mechanisms: a high storage of fermentable fuels (glycogen and phosphagen), a low metabolic rate in normoxia, and a further reduction in metabolic rate by reducing locomotion and ventilation. It is suggested here that the low metabolic rate of many hypogean species may be an adaptation to low oxygen and not necessarily result from an impoverished food supply. 5. An interesting physiological feature of hypogean crustaceans is their ability to recover from anaerobic stress and, more specifically, rapidly to resynthesize glycogen stores during post-hypoxic recovery. A high storage and rapid restoration of fermentable fuels (without feeding) allows groundwater crustaceans to exploit a moving mosaic of suboxic (< 0.3 mg L?1 O₂), dysoxic (0.3–3.0 mg L^?1 O₂) and oxic (> 3 mg L^?1 O₂) patches. 6. It is concluded that although hypogean animals are probably unsuited for life in extensively or permanently suboxic groundwater, they can be found in small or temporarily suboxic patches. Indeed, their adaptations to hypoxia are clearly suited for life in groundwater characterized by spatially heterogeneous or highly dynamic DO concentrations. Their capacity to survive severe hypoxia for a few days and to recover rapidly would explain partly why ecological field studies often reveal the occurrence of interstitial taxa in groundwater with a wide range of DO.     

Nitrate Accumulation and Leaching in Surface and Ground Water Based on Simulated Rainfall Experiments

To evaluate the process of nitrate accumulation and leaching in surface and ground water, we conducted simulated rainfall experiments. The experiments were performed in areas of 5.3 m² with bare slopes of 3° that were treated with two nitrogen fertilizer inputs, high (22.5 g/m² NH₄NO₃) and control (no fertilizer), and subjected to 2 hours of rainfall, with. From the 1st to the 7th experiments, the same content of fertilizer mixed with soil was uniformly applied to the soil surface at 10 minutes before rainfall, and no fertilizer was applied for the 8th through 12th experiments. Initially, the time-series nitrate concentration in the surface flow quickly increased, and then it rapidly decreased and gradually stabilized at a low level during the fertilizer experiments. The nitrogen loss in the surface flow primarily occurred during the first 18.6 minutes of rainfall. For the continuous fertilizer experiments, the mean nitrate concentrations in the groundwater flow remained at less than 10 mg/L before the 5th experiment, and after the 7th experiment, these nitrate concentrations were greater than 10 mg/L throughout the process. The time-series process of the changing concentration in the groundwater flow exhibited the same parabolic trend for each fertilizer experiment. However, the time at which the nitrate concentration began to change lagged behind the start time of groundwater flow by approximately 0.94 hours on average. The experiments were also performed with no fertilizer. In these experiments, the mean nitrate concentration of groundwater initially increased continuously, and then, the process exhibited the same parabolic trend as the results of the fertilization experiments. The nitrate concentration decreased in the subsequent experiments. Eight days after the 12 rainfall experiments, 50.53% of the total nitrate applied remained in the experimental soil. Nitrate residues mainly existed at the surface and in the bottom soil layers, which represents a potentially more dangerous pollution scenario for surface and ground water. The surface and subsurface flow would enter into and contaminate water bodies, thus threatening the water environment.     

Hydro-ecological analysis of the Biebrza mire (Poland)

Vegetation composition and structure of 58 sites along gradients in the valley mire of Biebrza, Poland, are related to physical and chemical variables of groundwater and peat. The three most prominent hydrochemical processes in the valley are (a) dissolution of calcite; (b) dissolution of iron, manganese and aluminium; and (c) enrichment with nitrogen and potassium. Major factors determining these processes are vertical flow of the groundwater and river flooding.Within the rheophilous zone of the mire, calcium-richness of the shallow groundwater and base-saturation of the peat are caused by upward seepage of groundwater originating from adjacent higher grounds. This groundwater movement keeps the larger part of the mire saturated with calcium.Good correlations exist between hydrochemistry and vegetation patterns. Groundwater-fed sites support a characteristic rich fen vegetation (Caricetum limoso-diandrae) with a low biomass production. The flood-plain vegetation consists of highly-productive communities of Glycerietum maximae and Caricetum elatae. In a belt in the Upper Basin where neither flooding nor upward seepage occurs, succession, probably caused by intensified drainage, leads to a dwarf-shrub vegetation (Betuletum humilis; poor fen).     

Hydrological variability, organic matter supply and denitrification in the Garonne River ecosystem

1. Groundwater nitrate contamination has become a worldwide problem as increasing amounts of nitrogen fertilisers are used in agriculture. Alluvial groundwater is uniquely juxtaposed between soils and streams. Hydrological connections among these subsystems regulate nutrient cycling. 2. We measured denitrification using an in situ acetylene‐block assay in a nitrate‐contaminated portion of the Garonne River catchment along a gradient of surface water–ground water mixing during high (snowmelt) and low flow. 3. During high flow (mid‐April to early June) the water table rose an average of 35 cm and river water penetrated the subsurface to a great extent in monitoring wells. Denitrification rates averaged 5.40 μgN₂O L^?1 min^?1 during the high flow period, nearly double the average rate (2.91 μgN₂O L^?1 min^?1) measured during base flow. This was driven by a strong increase in denitrification in groundwater under native riparian vegetation. Nitrate concentrations were significantly lower during high flow compared with base flow. Riparian patches had higher dissolved organic carbon concentrations that were more aromatic compared with the gravel bar patch closest to the river. 4. Multiple linear regression showed that the rate of denitrification was best predicted by the concentration of low molecular weight organic acids. These molecules are probably derived from decomposition of soil organic matter and are an important energy source for anaerobic respiratory processes like denitrification. The second best predictor was per cent surface water, reflecting higher denitrification rates during spring when hydrological connection between surface water and ground water was greatest. 5. Our results indicate that, while denitrification rates in Garonne River alluvium were spatially and temporally variable, denitrification was a significant NO₃ sink during transport from the NO₃‐contaminated floodplain to the river. DOC availability and river–floodplain connectivity were important factors influencing observed spatial and temporal patterns.     

Indirect N2O emissions from shallow groundwater in an agricultural catchment (Seine Basin, France)

Production and accumulation of nitrous oxide (N₂O), a major greenhouse gas, in shallow groundwater might contribute to indirect N₂O emissions to the atmosphere (e.g., when groundwater flows into a stream or a river). The Intergovernmental Panel on Climate Change (IPCC) has attributed an emission factor (EF_5g) for N₂O, associated with nitrate leaching in groundwater and drainage ditches—0.0025 (corresponding to 0.25% of N leached which is emitted as N₂O)—although this is the subject of considerable uncertainty. We investigated and quantified the transport and fate of nitrate (NO₃ ^?) and dissolved nitrous oxide from crop fields to groundwater and surface water over a 2-year period (monitoring from April 2008 to April 2010) in a transect from a plateau to the river with three piezometers. In groundwater, nitrate concentrations ranged from 1.0 to 22.7 mg NO₃ ^?–N l^?1 (from 2.8 to 37.5 mg NO₃ ^?–N l^?1 in the river) and dissolved N₂O from 0.2 to 101.0 μg N₂O–N l^?1 (and from 0.2 to 2.9 μg N₂O–N l^?1 in the river). From these measurements, we estimated an emission factor of EF_5g = 0.0026 (similar to the value currently used by the IPCC) and an annual indirect N₂O flux from groundwater of 0.035 kg N₂O–N ha^?1 year^?1, i.e., 1.8% of the previously measured direct N₂O flux from agricultural soils.     

Saline seepage and vertical distribution of oxygen in a brackish ditch

Summary Seepage caused salinity gradients in a 120 cm deep ditch adjacent to Lake Veere in the Dutch Delta area. The water was mixed by strong horizontal currents, forced by a pumping engine. The restoration of vertical chloride and oxygen gradients was studied.During summer when the level of Lake Veere was raised by 70 cm, a rapid seepage into the ditch was observed, ranging from 6 to 10 mm.h^–1. This caused an anoxic saline layer. At the anaerobic-aerobic interface 1.4 to 2.4 mg O₂.l^–1.h^–1 of the oxygen present in the upper water column was taken up. Sulfide oxidation required one quarter of this amount of oxygen.During winter the level of Lake Veere was lowered again. Normally seepage was absent then, and chloride entered the water column by diffusion; the mud chlorinity decreased from 15 to 3 Cl^– on its way to the ditch. Only after maximum discharge of polder water a moderate seepage of less than 4.4 mm.h^–1 restored a chloride gradient; about 1.6 mg O₂.l^–1.h^–1, produced by benthic diatoms, had to be supplied to the reducing materials in the seepage water in less than 12 hours.A saline seepage was simulated in the laboratory. These simulation tests showed that seepage below 1 mm.h^–1, as well as benthic respiration only, had a small impact on the vertical oxygen profile in the water column. Bottom oxygen demand increased with seepage velocity. At a strong seepage the water column could become entirely anaerobic.Communication no. 226.     

Groundwater chemistry and vegetation of gradients from rich fen to poor fen in the Naardermeer (the Netherlands)

In the eastern part of the Naardermeer peatlands (the Netherlands) a regional calcium-rich groundwater flow discharges (here often called the seepage area), whereas in the western part infiltration takes place. The ecological consequence of this hydrological pattern is reflected by the pattern in reedland communities. In the seepage area, which is characterized by Thelypteris-reedlands including many rare and endangered species (Caricion davallianae, Calthion palustris), there is a complex gradient of water types. The lime potential in the peat soil is clearly influenced by the hydrological gradient. In the ombrotrophic (poor fen) part of the gradient (containing species of Caricion curto-nigrae) the lime potentials are low and the groundwater contains low amounts of dissolved ions. The rare and endangered species (Caricion davallianae) are restricted to a small area with high lime potentials which is nourished by regional calcium-rich groundwater. High lime potentials were also measured in eutrophic reedlands influenced by brackish groundwater. Several species which generally occur in wet meadows even show a preference for this brackish environment.In a part of the seepage area succession from rich fen to poor fen and Alnus wood has taken place over a period of 40 years. This development has been caused by the diminishing amount of fresh seepage due to a lowering of the water levels in the surrounding area.The characteristics of poikilotrophic zones (contact zones between water flows) are discussed in relation to their significance for the preservation of endangered marsh species.Abbreviations EC₂₅= Electrical conductivity measured at 25 °C     

Effects of temporary desiccation on the mobility of phosphorus and metals in sulphur-rich fens: differential responses of sediments and consequences for water table management

In the Netherlands, permanent damming of sulphate (SO₄ ^2–)-rich surface water, in order to rewet desiccated wetlands, has resulted in stagnation and eutrophication of surface water. Permanent damming of surface water prevents periodic drought during summer and leads to suppressed iron (Fe)-rich groundwater input and to stimulated SO₄ ^2– reduction, all likely stimulating depletion of reducible Fe in the sediment. A laboratory experiment was conducted to assess the importance of temporary desiccation, its differential effects on various sediment types and the consequences for water table management. Permanent high SO₄ ^2–-rich surface water tables above sediments that are indirectly affected by shallow groundwater flows, resulted in severe eutrophication. The effect of temporary desiccation on phosphorus (P) mobilization and immobilization strongly depended on the sediment Fe and P pools in combination with the buffering capacity of the sediment. Desiccation of sediment that is indirectly affected by shallow groundwater flows, led to a long-lasting reduction in phosphate (o-PO₄ ^3–) release from the sediment because a reduced Fe pool is present, resulting in the release of Fe upon oxidation. Formerly dry sediments that have not been influenced by groundwater for a long time do not possess such a reduced Fe pool and desiccation did not reduce P-release from these sediments resulting in considerable eutrophication of the water layer immediately after rewetting. In sediment of seepage zones that are directly and permanently influenced by deeper groundwater, reduced Fe and calcium levels are so high that o-PO₄ ^3– was effectively immobilized under oxidized as well as reduced conditions. The results indicate that restoration of desiccated wetlands can not be achieved by simply retaining water by means of constructed dams. If water retention is artificially increased, temporary drops in water level during the summer are necessary to recharge the reducible P-binding Fe pool in large zones of the wetlands in order to prevent eutrophication.     

Dynamics of nitrous oxide in groundwater at the aquatic–terrestrial interface

Few data are available to validate the Intergovernmental Panel on Climate Change's (IPCC) emission factors for indirect emissions of nitrous oxide (N₂O). In particular the N₂O emissions resulting from nitrogen leaching and the associated groundwater and surface drainage (EF5-g) are particularly poorly characterized. In situ push–pull methods have been used to identify the fate of NO₃⁻ in the groundwater. In this study, we adapted a previously published in situ denitrification push–pull method to examine the fate of ¹⁵N₂O introduced into the subsoil–groundwater matrix. Enriched ¹⁵N₂O was manufactured, added to groundwater via a closed system in the laboratory, and then introduced into the groundwater–subsoil matrix in an upland-marsh transition zone of a salt marsh and a forested alluvial riparian zone. Conservative tracers (SF₆ and Br⁻) and ¹⁵N₂O were injected into the groundwater and left for 1–4 h after which the groundwater was sampled. Added ¹⁵N₂O behaved in a conservative manner at one site while the other site showed variability with some injections showing significant consumption (3–8 μg N₂O-¹⁵N kg⁻¹ soil day⁻¹) of ¹⁵N₂O. Our results show that the fate and dynamics of N₂O in groundwater are complex and variable and that these dynamics should be considered in the development of improved IPCC inventory calculations.     

Vertical distribution, biomass, production and turnover of fine roots along a topographical gradient in a sandy shrubland

In an artificial Salix gordejevii Chang et Skv. plantation of the Horqin sandy land, we investigated vertical distribution (in 0–100 cm depth), biomass (FRD), fine root production (FRP), fine root length density (FRLD) and turnover of fine roots (<2 mm diameter) at three sites (dune top, midslope and bottom of dune) along leeward slopes. Meanwhile, the correlation between FRP and soil available resources was analyzed. Our results indicate that more than 65% of total fine root biomass is distributed in 0–40 cm depth, and the patterns are different at three sites. The mean monthly FRD ranges from 227 to 324 g·m^?2, and they follows the order: dune top > midslope > bottom of dune. Ingrowth cores were harvested after 2, 3, 4, 5, 6 and 8 months of installation. At the first five sampling times, FRP and FRLD (0–40 cm) follows the same order with FRD along the topographical gradient, while FRP harvested after 8 months does not follow the same tendency, they are 348, 402 and 356 g·cm^?2 in dune top, midslope and bottom of dune, respectively. Fine root turnover ranges from 1.04–1.92 year^?1, and fine root turnover (20–40 cm) increases from dune top to bottom of dune along the topographical gradient. Correlation analysis between FRP and soil available resources indicates that only mean soil volumetric water content significantly correlates with annual FRP, which suggests that soil water content might be more crucial for shrub growth than fertility along the topographical gradient.     

Fluctuating deposition of ocean water drives plant function on coastal sand dunes

Sea‐level rise will alter the hydrology of terrestrial coastal ecosystems. As such, it becomes increasingly important to decipher the present role of ocean water in coastal ecosystems in order to assess the coming effects of sea‐level rise scenarios. Sand dunes occur at the interface of land and sea. Traditionally, they are conceived as freshwater environments with rain and ground water as the only water sources available to vegetation. This study investigates the possibility of ocean water influx to dune soils and its effect on the physiology of sand dune vegetation. Stable isotopes are used to trace the path of ocean water from the soil to the vegetation. Soil salinity, water content and δ¹⁸O values are measured concurrently with stem water and leaf tissue of eight species during the wet and dry season and from areas proximal and distal to the ocean. Our results indicate the dune ecosystem is a mixed freshwater and marine water system characterized by oceanic influence on dune hydrology that is spatially heterogeneous and fluctuates temporally. Ocean water influx to soil occurs via salt spray in areas 5–12 m from the ocean during dry season. Accordingly, vegetation nearest to the sea demonstrate a plastic response to ocean water deposition including elevated integrated water use efficiency (δ¹³C_leaf) and uptake of ocean water that comprised up to 52% of xylem water. We suggest physiological plasticity in response to periodic ocean water influx may be a functional characteristic common to species on the leading edge of diverse coastal habitats and an important feature that should be included in modeling coastal ecosystems. Rising sea level would likely cause a repercussive landward shift of dune species in response to encroaching maritime influences. However, human development would restrict this process, potentially causing the demise of dune systems and the protection from land erosion they provide.