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.     

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.     

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.     

Restoration of coastal dune slacks in the Netherlands

In order to stop the continuous decline of typical dune slack communities along the Dutch coast, restoration projects have been carried out since 1952. Restoration measures consisted of re-introducing traditional management techniques in dune slacks, such as mowing, grazing and sod removal, or constructing artificial dune slacks to compensate for lost biodiversity elsewhere. An analysis of successful and unsuccessful projects showed that constructing new dune slacks was not very successful for maintaining new populations of endangered dune slack species, since such projects were often carried out in areas where seed banks were depleted, while hydrological conditions and seed dispersal mechanisms were sub-optimal. The construction of sand dikes to prevent sea intrusion in large beach plains was, unintentionally, a temporary success for the establishment of many Red List species, although such measures often disrupted natural dune slack formation. Successful sites were all characterised by a regular discharge of calcareous groundwater provided by local or regional hydrological systems, where not very long ago populations of typical dune slack plants were present. Under such conditions, sod removal was a successful measure to create pioneer stages which were relatively stable, due to a very slow accumulation of organic matter in the topsoil. It is argued that new and more flexible coastal defence strategies can provide new opportunities for natural and relatively stable pioneer stages of dune slack formation, suitable for the long term preservation of endangered dune slack species.     

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.     

The relationship between vegetation dynamics and water table in tropical dune slacks

Abstract. Inundation of tropical dune slacks is an irregular phenomenon produced by consecutive years with high precipitation and also by intense tropical storms. Our hypothesis was that the distribution, structure and composition of vegetation in tropical dune slacks have changed over time as a result of various factors, principally their topographical position with respect to the water table, frequency and duration of inundation, and historical fluctuations in climate. We tested this hypothesis in two different slacks that represent the slack community at the study site. Inundation caused changes in species composition and distribution, as well as in vegetation structure. In one slack, inundation was considered as disturbance because its intensity increased when areas remained inundated for longer periods of time, causing death of vegetation cover and favouring invasion of more aggressive, secondary species (Pluchea odorata, Ambrosia artemisiifolia, Panicum maximum). When inundation lasted less than three months, rhizomatous herbs remained (Cyperus articulatus, Lippia nodiflora, Hydrocotyle bonariensis) inhibiting the succession process. The results suggest that the different degrees of inundation and the irregular frequency and intensity of inundation in the slacks under study, have favoured different successional tendencies.     

Dune slack restoration in Dutch mainland coastal dunes

The 45,000 ha Dutch dunes are relatively well protected but they suffer from severe desiccation. During the last decade a lot of knowledge has been obtained about the steering processes of eco-hydrology in dunes resulting in well-founded schemes to restore wet dune slack communities. In two case studies a large-scale restoration of the landscape was necessary. Since 1987 in the North-Holland Dune Reserve the groundwater catchment has been reduced (from 8 million m³ to 2 million m³ per year) and the resulting development monitored. In another case, large-scale intervention on 35 ha of the Amsterdam Water Supply Dunes in 1995 resulted in the restoration of active sand drift with opportunities for the establishment of pioneer vegetation. In small parts of the area, the surface has stabilised as a result of vegetation growth, but elsewhere drifting sand has spread and covered previously stable surfaces. Many wet to moist dune slacks developed and results relating to geomorphology, hydrology and vegetation are presented. Finally, in 1996 yet another project of 50 ha was carried out in the Meijendel dunes, in which infiltration ponds were removed to allow vegetation succession to develop from scratch. Results of these three case studies will be presented. If only hydrological restoration is carried out, sod-cutting was found to be necessary to restore the whole series of vegetation types of dune slacks. In two cases an area was denuded. The seed bank and surrounding slacks seem to play an important role in the success of restoration, this is especially true for the pioneer vegetation types. Future monitoring will prove whether critical (Red List) plant species will return.     

Vegetation fluctuation in mediterranean dune ponds in relation to rainfall variation and water extraction

Abstract. The structure of dune ponds hygrophytic vegetation has a spatial organisation in belts around the pond basin, closely related to water level and flooding regime. Doñana National Park has an important representation of temporal dune ponds, which are subjected to rainfall fluctuations and may be suffering the impact of water extraction from the neighbouring tourist resort. Permanent transects in a vegetation complex of five dune ponds have been monitored during a eight year period (1990-1997). This period was characterised by a number of dry years (annual rainfall around 300 mm), located between two wet cycles (800-900 mm). Transects were analysed in 1990 (wet period), 1994 (dry) and 1997 (wet) by hierarchical agglomera-tive clustering. During the dry period hygrophytic species showed regression, with a high mortality of some species like Ulex minor, while the xerophytic species advanced to lower areas. Seedlings of some xerophytic species colonised the dry surface of the pond basin. The lowering of the water table varied in the different ponds, ranging from 312 to 190 cm depending on topography and the distance to the pumping area. The new period of flooding during 1995-96 and 1996-97 cycles provided the opportunity for hygrophytic spe cies to re-establish themselves in their original places. This study suggest that changes in vegetation are caused by the interaction between weather conditions and human disturbance (water extractions). In our example man-made disturbance is more marked during the dry periods while wet periods tend to obscure the effects of water extractions. From a management perspective, long-term monitoring of water table and vegetation structure is revealed as a key procedure to the management of land-water ecotones on pressured areas and threatened habitats.     

Restoration of Wet Dune Slacks on the Dutch Wadden Sea Islands: Recolonization After Large-Scale Sod Cutting

The effects of sod cutting were studied in a dune area on the Dutch Wadden Sea Island of Texel. Sod cutting was carried out in a range of different dune slacks in order to restore dune slack vegetation with many endangered Red List species. Sod cutting removed approximately 96% of the soil seed bank. Species abundant in the seed bank, notably Juncacea, also had a high frequency in the vegetation that established during the first year after the restoration measures. Many other species not registered in the seed bank or in the former vegetation also appeared. Species richness in the monitored plots exceeded that of uncut reference plots after a few years. Colonization rates were higher than extinction rates in most plots, indicating that a stable state has not been reached after 5 years. Differences in species richness between slacks appeared to be related to the occurrence of source areas nearby and availability of dispersal agents, such as flooding and animals.     

Accumulation rates of soil organic matter in wet dune slacks on the Dutch Wadden Sea islands

Background and aims

A long-term monitoring program (ranging from 16 to 77 years) on the Dutch Wadden Sea Islands provided well-documented examples of vegetation succession in wet dune slacks. We used this opportunity to study soil organic matter (SOM) accumulation in relation to vegetation succession. The aim of this paper is to identify the factors which regulate accumulation rates of SOM in wet dune slacks.

Methods

We used several soil chronosequences using data from the monitoring program together with data from a long-term research activity and more recent measurements. We used several soil chronosequences using data from the monitoring program together with data from a long-term research (up to 150 years) and more recent measurements. Field measurements included pH, soil organic matter, above ground standing crop and water levels. Water level regimes (inundation duration and mean minimum water level), were simulated using a hydrological model. Capable of simulating inundation duration and water-level fluctuations, this model used field measurements collected over more than 5 years, as well as precipitation and evapotranspiration data collected over a period of 25 years.

Results

Sampling two synchronic chronosequences showed that SOM accumulations increased linearly during the first 50–60 years and then levelled off. Sampling various diachronic chronosequences over time showed a wide variation in accumulation rates. Slacks with low productive species, such as Littorella uniflora, showed low accumulation rates (0.02–0.08 kg/m²/year), and persisted even over a period of more than 90 years. In contrast, slacks dominated by high productive species, such as Phragmites australis, showed ten times higher accumulation rates (0.17–0.26 kg/m²/year) over a similar time period and comparable annual inundation periods (176–240 days). A multiple linear regression showed that variation in SOM accumulation rates was best explained by above-ground biomass of the vegetation.

Conclusions

We conclude that the rate of SOM accumulation in wet dune slacks is primarily controlled by plant above-ground biomass. Both above-ground biomass and SOM accumulation can remain very low over a long period of time when dune slacks are flooded during most of the year and plants with adaptive traits are able to maintain vegetation succession at a pioneer stage.     

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.     

The role of cattle in maintaining plant species diversity in wet dune valleys

The succession of species-rich wetland vegetation in dune valleys into species-poor dwarf shrub vegetation was followed by means of permanent vegetation plots, in which the cover of vascular plant, moss and lichen species were recorded over a period of up to 33 years. Low density cattle grazing is an effective substitute for rabbits in stalling this succession, thus preserving the local plant species diversity. The influence of direct rabbit grazing pressure on the vegetation was studied in exclosures. The differences were significant, but not long-lasting after the exclosures had been opened again to the rabbits. Cladina lichen-rich Empetrum-heathland in the dunes is a stage in a probably cyclic succession that might be triggered by grazing. The total vascular plant, moss and lichen diversity of the dune ecosystem can be maintained by a combination of extensive cattle grazing and a regular but limited re-creation of pioneer situations.     

Changes in normalized difference vegetation index of deserts and dunes with precipitation in the middle Heihe River Basin

AimsPrecipitation is the major water source for desert ecosystems, with its temporal dynamics significantly driving the changes of ecosystem structure and function in desert regions. The objectives of this study are to evaluate the changes in normalized difference vegetation index (NDVI) with seasonal precipitation and different climate years in two cover types (desert and dune).MethodsBased on the daily rainfall dataset of 2000-2012 in the middle Heihe River Basin in Northwest China and the NDVI extracted from the moderate resolution imaging spectroradiometer (MODIS) images, we performed linear regression analysis to examine the correlation of NDVI_INT (accumulated NDVI) with precipitation in two cover types (desert and dune). Two measures of the precipitation are P_c (cool-season precipitation from last September to late February) and P_w (warm-season precipitation in between March and August), while NDVI was characterized with NDVI_INT-pys (previous-year during the different climate years (dry: 2001-2003, wet: 2004-2007)) and the different periods of a growing season (i.e. whole growing season from May through September, early growing season in between May and June, and late growing season in between July and September).Important findings We found that: (1) the determinants of growing season NDVI_INT and their order were NDVI_INT-pys > P_c > P_w for the deserts, while the order was P_w > NDVI_INT-pys for the dunes. The determinants and their order of NDVI_INT in early growing season were NDVI_INT-pys > P_c for the desert, while they were NDVI_INT-pys > P_c = P_w for the late growing season. However, for the dunes, NDVI_INT of the early and late growing season appeared determined by NDVI_INT-pys and P_w, respectively. (2) During the dry and wet periods, precipitation was not a significant factor influencing NDVI_INT for the desert and dune. However, significant increases in NDVI_INTwere observed at dune under wet condition. With the wet years continued, the length of the wet years become an important determinant of NDVI_INT at both cove types, particularly at dune. In addition, it appeared that different changes in NDVI with precipitation existed between the two cover types, but with very similar effects of time-lag. These findings provide useful references for further understanding the mechanisms of NDVI changes with precipitation.     

Enhanced nitrogen loss may explain alternative stable states in dune slack succession

Ecological theory emphasizes competitive interactions between plant species when explaining primary succession in plants. Ecosystem processes, such as nutrient accumulation, are often regarded as independent, steering successional changes without being affected by the interacting plant species. We present experimental evidence that plant species in wet dune slack systems are able to affect ecosystem processes in their favor by reducing the rate of nitrogen accumulation by coupled nitrification–denitrification, promoting their competitive position. We compared denitrification rates of two early successional species having radial oxygen loss (ROL) with two non-ROL late successional species in a mesocosm experiment. The denitrification rates were significantly higher in mesocosms planted with early successional species Littorella uniflora (P<0.001) and Schoenus nigricans (P<0.05), relative to the rates found in the presence of non-ROL species, Carex nigra and Calamagrostis epigejos . We analyzed the consequences of enhanced nitrogen loss on the competition between an early and a late successional species by means of a simple theoretical model. Our analysis revealed that early successional species capable of ROL might retard successional changes and lock the ecosystem in an unproductive state for an extended period of time. We emphasize that understanding of ecosystem processes is crucial in successful conservation of high biodiversity vegetation in wet dune slacks.     

Vegetation change in an ombrotrophic mire in northern England after excluding sheep

Abstract. The role of sheep grazing on vegetation change in upland mires removed from livestock farming and surrounded by conifer plantation was investigated with a grazing trial at Butterburn Flow in northern England. Paired grazed and ungrazed plots from central and peripheral locations were compared over 14 yr. Vegetation data from 34 mires in Kielder Forest provided an ordination framework within which vegetation trends were investigated. A gradient from dry moorland/hummock to wet mire/hollow vegetation dominated this framework and may reflect hydrological variability and structural vegetation differences between the mires. Some species were significantly affected by change in grazing intensity and there were differences between the edge and the centre of the mire. Overall vegetation change depended upon the grazing management and the position of the plots such that the removal of sheep grazing decreased the cover of species typical of wet ombrotrophic conditions, but only at the periphery of the mire. The vegetation in one plot became very similar to that of mires elsewhere in Kielder Forest where sheep were removed several decades ago. Cessation of grazing on upland mires is likely to lead to slow structural and species change in vegetation at the mire edge with a long‐term loss of ombrotrophic species. The nature conservation significance of these changes will depend upon whether or not management objectives target natural conditions or wish to maximize ombrotrophic vegetation. The context of external factors such as climate and pollution may, however, be more important in determining site condition on the wettest mires.     

黑河中游荒漠生态系统归一化植被指数对降水的响应

降水是荒漠生态系统主要的水分来源, 是植被结构和功能变化的根本驱动力。该研究以黑河中游砾质荒漠(砾漠)和沙质荒漠(沙漠)为研究对象, 基于2000-2012年中分辨率成像光谱仪(MODIS)获取的归一化植被指数(NDVI)数据以及日降水数据, 运用多元线性回归法, 分析了砾漠和沙漠植被生长季(5-9月)、生长季早期(5-6月)和晚期(7-9月)累积NDVI (NDVI_INT, INT表示某时间段的累积值)对冷季降水(P_c, 前一年9月至当年2月累积降水)、暖季降水(P_w, 当年3月至8月累积降水)、前一年生长季NDVI_INT (NDVI_INT-pys)以及干湿气候期(干旱期: 2001-2003; 湿润期: 2004-2007年)的响应。研究结果表明: (1)砾漠植被生长季NDVI_INT年际变化的影响因素排序为NDVI_INT-pys > P_c > P_w, 沙漠植被NDVI_INT则为P_w > NDVI_INT-pys; 砾漠生长季NDVI_INT早期NDVI_INT年际变化的影响因素排序为NDVI_INT-pys > P_c, 晚期则为NDVI_INT-pys > P_c = P_w; 而沙漠生长季早期NDVI_INT年际变化的影响因素为NDVI_INT-pys, 晚期是P_w。(2)在干湿气候期内, 降水量并非是影响荒漠NDVI_INT变化的关键因子。干湿气候期交替时, 砾漠NDVI_INT较沙漠增加明显; 湿润期内, 湿润期持续的长短是影响两种生境植被NDVI_INT的关键因子, 以沙漠较为明显。黑河中游砾漠和沙漠植被生产力对冷暖季降水及干湿气候期响应具有明显的差异, 但总体显示出荒漠植被生产力对降水响应具有滞后性特征。以上结论可为揭示荒漠植被生产力对降水的响应机理提供参考。     

Effects of prescribed fire on the composition of woody plant communities in southeastern Texas

Abstract. The response to prescribed burning of plant communities ranging from dry to wet habitats was monitored using permanent plots sampled from 1989 to 1993. Temporal controls for fire effects were provided by matched sets of plots protected from fire by newly constructed fire breaks. Changes in species composition were studied by ordination of strata of trees (> 5 cm DBH), small trees (2–5 cm DBH), large saplings (1–2 cm DBH), and small saplings and seedlings (50–140 cm tall). Results show that changes occurred largely in the small tree stratum, in which xeric species increased in importance. Although there were changes in sapling and seedling strata, no clear direction of change was recognized. Fire had little effect on the tree stratum. Of the seven community types under study, three types, sandhill, upland pine, and upperslope pine-oak, were most strongly affected, as indicated by post-fire change in positions of samples representing these communities in ordination space and reduction in understory species abundance. Samples representing the other four mesic and wet communities showed little or no change in their positions. These short-term results indicate that changes in vegetation resulting from fire were small and were mostly restricted to the dry types in which possible compositional change is expected to occur. This differential effect of fire suggests that the influence of fire is secondary to that of topographical and soil gradients in determining vegetation pattern under current fire regimes. Fire seems to reinforce an overall vegetation gradient controlled by soil in southeastern Texas.     

Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993

Abstract. Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage. Species richness (yearly mean and cumulative species number) and change in species richness differed between vascular plants and cryptogams, and between forest types. The number of vascular plant species decreased in pine forest in dry years; bryophyte species number increased in spruce forest. Statistically significant vegetation change, as tested by constrained ordination (CCA) with time as the constraining variable, is demonstrated for most one-year periods and for the five-year period in most forest types. Vegetation change along identified gradients, measured as plot displacement along DCA ordination axes, also occurred. The magnitude of year-to-year vegetation change was related neither to forest type nor to one-year period; different responses to climatic and environmental change were observed in each forest type. The largest average displacement observed, from medium-rich spruce forest towards poor spruce forest, was interpreted as a long-term trend. Humus-layer pH decreased by ca. 0.25 units from 1988 to 1993, most strongly in medium-rich spruce forest where exchangeable Ca decreased and Al and Mn increased strongly. Our study supports the hypothesis that vascular plants show a long-term and broad-scale response to soil acidification. Change in bryophyte composition is linked to some very long growing-seasons. Detailed analysis of short-term vegetation dynamics enhances the interpretation of long-term changes and stresses the complementarity of univariate and multivariate methods in the analysis of vegetation change.     

Wet dune slacks: decline and new opportunities

For a number of infiltrated coastal dune areas it is discussed to what extent artificial infiltration for the public water supply affects the quality of soil, groundwater and vegetation around pools and ponds, and what its effect is on the vegetation. Further, the results of investigations into the quality of vegetation, soil and water of a number of non-infiltrated, less affected dune areas are presented. The emphasis is on changes in groundwater flow pattern and on changes in the chemical composition of groundwater on the vegetation of wet dune slacks. Finally, recommendations for the management of wet dune slacks are presented. It can be concluded that the introduction of nutrients through infiltration causes an abundance of nitrophilous herbaceous vegetation along the banks of all infiltration ponds and most dune pools. Of the three investigated macro-nutrients, nitrate, potassium and phosphate, the latter shows the most significant correlation with the composition, cover and biomass of the vegetation. The moist biotopes of non-infiltrated dunes have largely disappeared because of desiccation, mainly as a consequence of water withdrawal, afforestation and coastal erosion. Relatively unaffected dune slacks can be found in the dunes on the Dutch Wadden Sea islands and a small number of dune areas on the mainland. In most areas, however, a serious decline in many rare species has been observed during the past twenty years because of eutrophic and acid precipitation, often in combination with disturbances of the groundwater regime.     

Short dry spells in the wet season increase mortality of tropical pioneer seedlings

Variation in plant species performance in response to water availability offers a potential axis for temporal and spatial habitat partitioning and may therefore affect community composition in tropical forests. We hypothesized that short dry spells during the wet season are a significant source of mortality for the newly emerging seedlings of pioneer species that recruit in treefall gaps in tropical forests. An analysis of a 49-year rainfall record for three forests across a rainfall gradient in central Panama confirmed that dry spells of ≥10 days during the wet season occur on average once a year in a deciduous forest, and once every other year in a semi-deciduous moist and an evergreen wet forest. The effect of wet season dry spells on the recruitment of pioneers was investigated by comparing seedling survival in rain-protected dry plots and irrigated control plots in four large artificially created treefall gaps in a semi-deciduous tropical forest. In rain-protected plots surface soil layers dried rapidly, leading to a strong gradient in water potential within the upper 10 cm of soil. Seedling survival for six pioneer species was significantly lower in rain-protected than in irrigated control plots after only 4 days. The strength of the irrigation effect differed among species, and first became apparent 3–10 days after treatments started. Root allocation patterns were significantly, or marginally significantly, different between species and between two groups of larger and smaller seeded species. However, they were not correlated with seedling drought sensitivity, suggesting allocation is not a key trait for drought sensitivity in pioneer seedlings. Our data provide strong evidence that short dry spells in the wet season differentially affect seedling survivorship of pioneer species, and may therefore have important implications to seedling demography and community dynamics.