Nutrient limitations in an extant and drained poor fen: implications for restoration

In a species-rich poor fen (Caricetum nigrae) and a species-poor drained fen, the difference in nutrient limitation of the vegetation was assessed in a full-factorial fertilization experiment with N, P and K. The results were compared to the nutrient ratios of plant material and to chemical analysis of the topsoil. A rewetting experiment with intact sods was carried out in the glasshouse and the results are discussed in view of restoration prospects of drained and degraded peatlands. In the undrained poor fen the above-ground biomass yield was N-limited while the vegetation of the drained fen was K-limited. Experimental rewetting of intact turf samples, taken in the drained site, did not change the biomass yield or the type of nutrient limitation. It was concluded that mire systems which have been subjected to prolonged drainage are inclined to pronounced K-deficiency, probably due to washing out of potassium and harvesting the standing crop. This may hamper restoration projects in degraded peat areas where nature conservation tries to restore species-rich vegetation types with a high nature value.     

Restoration strategies for wet grasslands in Northern Germany

Wet meadows of the Calthion type are strongly endangered ecosystems in Northern Germany due to agricultural intensification and abandonment. Empirical data of biocoenotic structures, hydrologic parameters, nutrient budgets and soil structure are used for an integrated evaluation of differently degenerated wet grasslands on peat soils (mesotrophic and eutrophic Calthion, Lolio-Potentillion, abandoned wet meadows) with respect to their functions in the landscape. Based on the evaluation of the investigated ecosystems two specific management targets are derived: (1) the maintenance and restoration of Calthion systems from degenerated states to re-establish high biocoenotic diversity and to avoid high nutrient leakages; (2) The restoration of the systems functioning as nutrient sinks. The potentials and prerequisites for the development of the investigated systems and suitable management measures concerning these two targets are described, focussing on groundwater dynamics, phytomass production, nutrient losses and the re-establishment of typical plant species. Our results illustrate that referring to target 1 none of the management measures discussed are suitable to restore strongly degenerated Lolio-Potentillion systems to Calthion systems completely. The limited restoration success is caused by partially irreversible changes of the biotic and abiotic properties of the Lolio-Potentillion sites. A re-development of the mesotrophic Calthion from eutrophic Calthion sites or abandoned wet meadows seems to be impossible. From this point of view restoration efforts should mainly focus on the preservation and maintenance of near-natural Calthion ecosystems. Nevertheless, referring to the high portion of Lolio-Potentillion in the landscape of Northern Germany, it is essential to reduce their ecological function as a nutrient source. This could be achieved by moderate rewetting and harvesting. For strongly degenerated grasslands, which can hardly be restored according to target 1, the establishment of eutrophic swamps is a desired development target (target 2). A suitable measure for this target is the drastic rising of the groundwater level. But we must take into account that the current knowledge about the nutrient retention function, resulting from this management measure, is not sufficient to predict this function in detail for former intensively used fen ecosystems. The presented interrelationships and management alternatives are integrated into a knowledge based modelling system to support decision making.     

Fen‐meadow succession in relation to spatial and temporal differences in hydrological and soil conditions

Question: In fen meadows with Junco‐Molinion plant communities, falling groundwater levels may not lead to a boosted above‐ground biomass production if limitation of nutrients persists. Instead, depending on drainage intensity and micro‐topography, acidification may trigger a shift into drier and more nutrient‐poor plant communities. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Long‐term study (1988‐1997) in a fen meadow along a gradient in drainage intensity at different scales. Results: Above‐ground biomass increased only slightly over ten years, despite a lower summer groundwater table. The accountable factors were probably a limited availability of nutrients (K in the higher well‐drained plots, P in the intermediate plots and N in the lower hardly drained plots), plus removal of hay. Junco‐Molinion species increased in dry sites and Parvo‐caricetea species increased in wet sites, presumably primarily because of soil acidification occurring when rainwater becomes more influential than base‐rich groundwater. The extent of the shift in species composition depends primarily on the drainage intensity and secondarily on microtopography. Local hydrological measures have largely failed to restore wetter and more basic‐rich conditions. Conclusions: Acidification and nutrient removal, leaching and immobilization resulted in the succession towards Junco‐Molinion at the cost of Calthion palustris elements. Lower in the gradient this change was reduced by the presence of buffered groundwater in slightly drained sites. To conserve the typical plant communities of the Junco‐Molinion to Calthion gradient in the long term, further acidification must be prevented, for example by inundation with base‐rich surface water.     

The effects of drainage on groundwater quality and plant species distribution in stream valley meadows

Conditions in fen meadows in Dutch stream valleys are influenced by both deep (Ca²⁺-rich) and shallow (Ca²⁺-poor) groundwater flows. The distribution patterns of phreatophytic (groundwater-influenced) plant species showed distinct relationships with the distribution of different groundwater types. Large fluctuations in the ionic composition of groundwater were observed in the upper peat layers of drained areas. Ca²⁺-rich groundwater was replaced by precipitation water to a considerable depth (1–1.5 m). These fluctuations in groundwater composition were less pronounced in undrained areas.It was observed that characteristic species of fen meadow communities and rare phreatophytic species were restricted to areas with high water tables, which were nourished by either Ca²⁺-poor or by Ca²⁺-rich groundwater. Few species showed a preference for drained areas, where replacement of groundwater types gave rise to the occurrence of an intermediate groundwater type, which was thought to be the result of an increased acidification of the top soil (increased influence of infiltration water).It was argued that the endangered species were best preserved in areas with an undisturbed discharge of natural groundwater flows.Abbreviations EC = Electrical conductivity     

Hydrology of Dutch Cirsio‐Molinietum meadows: Prospects for restoration

Abstract. Fen meadows (Cirsio dissecti‐Molinietum) are seriously threatened by desiccation, acidification and eutro‐phication. In The Netherlands several projects were launched to restore damaged fen meadows. This review describes how successes and failures of these restoration projects depend on hydrological systems. Six hydrological systems have been distinguished, which all provide the site conditions required by this community. Nowadays, the best developed fen meadows are found in the higher Pleistocene landscape of The Netherlands, where they depend on base‐rich groundwater discharging from local or large groundwater systems. Fen meadows of the lower Holocene landscape usually occur in man‐made surface water systems. Almost all stands have been severely deteriorated. Restoration of fen meadows in the Pleistocene landscape is promising when the hydrology is only slightly disturbed or when hydrological measures are taken in combination with sod cutting. Restoration prospects of fen meadows in the Holocene landscape are low. Until now a complete regeneration of Cirsio‐Molinietum meadows has not been realized. Restoration measures failed to restore high pH values in the top soil. It is hypothesized that viable seeds of many target species lack in the soil seed bank. In addition, the dispersal capacities of these species seem to be limited.     

Shrub removal facilitates recovery of wetland species in a rewetted fen

Drainage and shrub expansion are the main threats to the biodiversity of fens and fen meadows, whereas rewetting and the removal of shrubby species are frequently applied restoration measures. We examine whether removal of shrubs enhances recovery of target species in a degraded fen subjected to moderate rewetting. The study was located in the drained fen Ca?owanie (central Poland), where remnants of open fen communities and willow-invaded fens exist in former turf-pits, surrounded by degraded meadows on dried peat. All these three habitat types were included in a monitoring grid, which covered an area of 2.2 ha. Within 55 quadrats of 20 m × 20 m we monitored occurrence of 52 species, i.e. two groups of target species (fen indicators and wet meadow indicators) and indicators of two failure scenarios (degraded fen indicators and eutrophic wetland indicators), during six years following shrub removal, rewetting and re-application of conservational mowing, using a 3-step ordinal abundance scale. NMDS ordination revealed a gradual convergence of shrub removal plots and reference plots. We noticed significant effects of year and habitat type on all indicator groups, but only fen indicators have shown a clear (increasing) trend within shrub removal plots. Degraded fen indicators (ruderal and opportunistic species) initially expanded on shrub removal plots, but this effect disappeared in the following years. We conclude that shrub removal enhances establishment of target species in a moderately drained and then rewetted fen and attribute this effect to lowered competition for light. However, given high costs of this method and long-lasting problems with shrub resprouts, we recommend applying shrub removal only to recently overgrown sites, which still retained high botanical diversity. Heavily degraded fen meadows did not react on the increase of moisture, which indicates that more advanced restoration measures, such as top soil removal are needed there.     

The contribution of rewetting to vegetation restoration of degraded peat meadows

Question: What is the contribution of a rise in groundwater level to vegetation restoration of degraded peat meadows compared to abandonment only? Location: Abandoned peat meadows in the central part of The Netherlands. Methods: Comparison of species composition and species abundance of vegetation and seed banks of reference and rewetted peat meadows, using plant trait and seed bank analysis. Results: Vegetation of rewetted meadows shared on average only 27% of their species with the reference meadow, while this was 50% on average for species in the seed bank. Rewetted meadows had a lower total number of species and a lower number of wet grassland and fen species present in the vegetation, but had higher species richness per m², although evenness was not affected. Rewetting increased the dominance of species of fertile and near neutral habitats, but did not result in an increase of species of wet or waterlogged habitats. Re-wetted meadows were dominated by species relying mainly on vegetative reproduction and species with a low average seed longevity compared to the reference meadow. Conclusion: Rewetting was not effective as a restoration measure to increase plant species diversity or the number of wet grassland and fen species in the vegetation. If no additional restoration management is applied, the seed bank will be depleted of seeds of species of wet grassland or fen habitats, further reducing the chances of successful vegetation restoration.     

The Influence of Rewetting on Vegetation Development and Decomposition in a Degraded Fen

The rewetting technique border irrigation was installed in a degraded fen peatland in northeastern Germany. Because of the prevailing site conditions, the technique resulted in two different rewetting variants (surface irrigation and temporary inundation) at the study site. This paper reports on the practicability of this technique and the influence of rewetting on vegetation development, decomposition processes and soil nutrient availability, and the possibilities for renewed peat accumulation. The technique proved to be suited for rewetting fen sites with a continuous slope, deep peat layer with low hydraulic conductivity, and upstream water recharge facilities. A subsidence of the ground‐water levels during the summer months, however, could not be avoided in dry years. The vegetation changed slowly from species‐poor grassland into typical fen plant communities, despite rewetting and soil tillage. Species richness, however, was higher in the surface irrigation than in the temporary inundation variant. A sufficient water supply proved to be absolutely necessary to retard decomposition processes because higher decomposition of root materials (i.e., higher k values) occurred under temporary inundated conditions. Generally, the higher water content in the soil after rewetting led to a lower nitrate‐N–to–ammonium‐N ratio in the topsoil in both rewetting variants. In the surface irrigation variant the mineral nitrogen content (Nmin) of the topsoil decreased from 7.8 to 4.4 g N/m², which is also correlated with the increase in water content of the soil. The low Nmin levels of fens which were never deeply drained (0.9–2.8 g N/m²), however, were not reached within the observation period of 3 years.     

Impact of rewetting on the vegetation of a cut‐away peatland

Abstract. We tested whether rewetting improved environmental conditions during peatland restoration and promoted colonization and development of mire vegetation. Vegetation change was monitored in a cut‐away peatland one year before, and four years after, rewetting. Colonizers before rewetting were perennials, mostly typical of hummocks or bare peat surfaces. The main variation in vegetation was related to variation in the amounts of major nutrients and water table level. The wettest site with the highest nutrient level had the highest total vegetation cover and diversity, as well as some species typical of wet minerotrophic mires. Raising the water table level above, or close to, the soil surface promoted development of wet minerotrophic vegetation. Diversity initially decreased because of the disappearance of hummock vegetation but started to recover in the third year. Eriophorum vaginatum and Carex rostrata were both favoured, and bryophytes typical of wet habitats colonized the site. Moderate rewetting promoted the development of Eriophorum vaginatum seedlings and an increase in the cover of tussocks. Bryophytes typical of disturbed peat surfaces spread. In the control site development continued slowly towards closed hummock vegetation. The study showed that raising the water level to, or above, soil surface promotes conditions wet enough for a rapid succession towards closed mire vegetation.     

Restoration of brook valley meadows in the Netherlands

Until recently, restoration measures in Dutch brook valley meadows consisted of re-introducing traditional management techniques, such as mowing without fertilisation and low-intensity grazing. In the Netherlands, additional measures, such as rewetting and sod cutting, are now carried out on a large scale to combat negative influences of drainage and acidifying influences by atmospheric deposition. An analysis of successful and unsuccessful projects shows that restoration of brook valley meadows is most successful if traditional management techniques are applied in recently abandoned fields that had not been drained or fertilised. Large-scale topsoil removal in former agricultural fields that had been used intensively for several decades is often unsuccessful since seed banks are depleted, while hydrological conditions and seed dispersal mechanisms are sub-optimal. In areas with an organic topsoil, long-term drainage had often led to irreversible changes in chemical and physical properties of the soil. Successful sites were all characterised by a regular discharge of calcareous groundwater provided by local or regional hydrological systems, and, where not very long ago, populations of target species existed. On mineral soils, in particular, sod removal in established nature reserves was a successful measure to increase the number of endangered fen meadow species. It is argued that attempts to restore species-rich meadows should be avoided on former agricultural fields, where pedological processes have led to almost irreversible changes in the soil profile and where soil seed banks have been completely depleted. From a soil conservation point of view, such areas should be exploited as eutrophic wetlands that are regularly flooded.     

Nutrient limitation and nutrient‐driven shifts in plant species composition in a species‐rich fen meadow

Question: We studied the development and persistence of the effects of nutrient pulses on biomass production and species composition in a fen meadow. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Single pulse fertilization with N and P in a factorial design on an undrained central and a drained margin site in a species‐rich fen meadow (Cirsio dissecti‐Molinietum). Biomass production and species composition were monitored during four years. Results: At the central site, N addition boosted biomass production, but only during one year. The species composition was not changed. P fertilization increased the biomass production and changed the species composition from a vegetation dominated by Carex panicea to a grassland community with abundant Holcus lanatus, but not before the second year. At the margin site, P fertilization changed the species composition in a similar way, but biomass production was not increased. N fertilization had no effect. At both sites the P induced shift in species composition persisted for four years although the P effect declined during the experiment. Conclusions: The biomass responses show that N was limiting in the central site. Another nutrient, besides N and P (probably K) must have been limiting in the marginal site. The fast decline of the N effect on biomass is ascribed to increased denitrification and biomass removal. The delay in the P effect on biomass and species composition and the persistence of the P effect on species composition are ascribed to fast immobilisation and subsequent slow release of fertilizer P in the peat soil. Recurrence of the P pulses is expected to cause permanent changes in species composition.     

Species trait shifts in vegetation and soil seed bank during fen degradation

Fens in Central Europe are characterised by waterlogged organic substrate and low productivity. Human-induced changes due to drainage and mowing lead to changes in plant species composition from natural fen communities to fen meadows and later to over-drained, degraded meadows. Moderate drainage leads to increased vegetation productivity, and severe drainage results in frequent soil disturbances and less plant growth. In the present article, we analyse changes in plant trait combinations in the vegetation and the soil seed bank as well as changes in the seed bank types along gradient of drainage intensity. We hypothesize that an increase in productivity enhances traits related to persistence and that frequent disturbance selects for regeneration traits. We use multivariate statistics to analyse data from three disturbance levels: undisturbed fen, slightly drained fen meadow and severely drained degraded meadow. We found that the abundance of plants regenerating from seeds and accumulating persistent seed banks was increasing with degradation level, while plants reproducing vegetatively were gradually eliminated along the same trajectory. Plants with strong resprouting abilities increased during degradation. We also found that shifts in trait combinations were similar in the aboveground vegetation and in soil seed banks. We found that the density of short-term persistent seeds in the soil is highest in fen meadows and the density of long-term persistent seeds is highest in degraded meadows. The increase in abundance of species with strong regeneration traits at the cost of species with persistence-related traits has negative consequences for the restoration prospects of severely degraded sites.     

Red list plants: colonization by arbuscular mycorrhizal fungi and dark septate endophytes

Since information concerning the mycorrhization of endangered plants is of major importance for their potential re-establishment, we determined the mycorrhizal status of Serratula tinctoria (Asteraceae), Betonica officinalis (Lamiaceae), Drosera intermedia (Droseraceae) and Lycopodiella inundata (Lycopodiaceae), occurring at one of two wetland sites (fen meadow and peat bog), which differed in soil pH and available P levels. Root colonization by arbuscular mycorrhizal fungi (AMF) and dark septate endophytes (DSE) was quantified. Colonization by AMF appeared to be more frequent in the fen meadow than in the peat bog, and depended on the host plant. Roots of S. tinctoria and B. officinalis were well colonized by AMF in the fen meadow (35–55% root length) and both arbuscules and vesicles were observed to occur in spring as well as in autumn. In the peat bog, L. inundata showed a low level of root colonization in spring, when vesicles were found frequently but no arbuscules. In roots of D. intermedia from the peat bog, arbuscules and vesicles were observed, but AMF colonization was lower than in L. inundata. In contrast, the amount of AMF spores extracted from soil at the peat bog site was higher than from the fen meadow soil. Spore numbers did not differ between spring and autumn in the fen meadow, but they were higher in spring than in autumn in the peat bog. Acaulospora laevis or A. colossica and Glomus etunicatum were identified amongst the AMF spores extracted from soil at the two sites. S. tinctoria and B. officinalis roots were also regularly colonized by DSE (18–40% root length), while L. inundata was only rarely colonized and D. intermedia did not seem to be colonized by DSE at all.     

Effectiveness of Turf Stripping as a Measure for Restoring Species-Rich Fen Meadows in Suboptimal Hydrological Conditions

Most species‐rich fen meadows in nature reserves in The Netherlands are acidified due to weaker upwelling of base‐rich groundwater. The present study investigated whether and why turf stripping combined with superficial drainage might promote the long‐term recovery of such meadows and restore the nutrient‐poor, buffered conditions they require. In a field experiment, we analyzed changes in vegetation composition, soil parameters, and soil water chemistry in stripped plots of degraded Cirsio‐Molinietum vegetation over 12 years. After the first five years, many species from the target communities occurred in stripped plots. Both vegetation and soil data showed positive effects of turf stripping on the acid‐buffering capacity. Because sulfate concentration in the soil water decreased over time, whereas the bicarbonate concentration increased, we inferred that there was internal alkalinization driven by sulfate reduction in low‐lying stripped plots. However, the succession toward more acidophilus plant communities, in both control and stripped plots, indicated gradual acidification. This may be caused by a continuing weakening of the upward seepage of base‐rich groundwater as shown by declining calcium concentrations in the soil water. Though turf stripping exposed a nutrient‐poor soil layer with a greater acid‐buffering capacity, these positive effects might not be sufficient to combat the ongoing acidification in the long term.     

Restoration experiments in middle European wet terrestrial ecosystems: an overview

Most wetlands in the central European lowlands have been severely altered by cultivation. As a consequence they no longer fulfil their function as habitats for specialised species and communities, nor as retention areas for water and solid materials. Therefore, a number of renaturation experiments are in progress, which intend to develop and test strategies and measures to improve this defect in landscape diversity. For this purpose experiments on re-wetting, nutrient depletion of eutrophicated areas and re-establishment of typical wetland plant species and phytocoenoses have been performed. An ecological development concept defining the aims and describing their feasibility precedes such experiments. Preliminary results indicate that the reconstruction of the former state (regeneration) is impossible within reasonable time spans. In drained raised bogs overgrown with heather, as well as in those which have been industrially exploited, the primary aim must always be to restore efficient peat formation as far as possible; as a rule one succeeds only with well-growing and nutrient-demanding fen and transitional bog species. In cultivated fens the aim is to reduce peat loss. As a first step this is accomplished by converting arable fields and sown meadows into permanent grassland, if possible with reduced fertilization and low mowing or grazing frequencies, and accompanied by rewetting during winter. Many experiments have sought to impoverish eutrophicated fen soils and introduce typical fen species by sowing or planting, so well tested techniques are available. However, the total prevention of peat loss is only possible by permanent rewetting throughout the year, so that peat accumulation can start again. Only in this way could fens regain their former function as sinks in landscape processes.     

Mineralization and Decomposition Rates in Restored Pine Fens

Growing public interest in conserving peatlands has created a need for restoration and rapid indicators of progress in peat formation. Vegetation and hydrological indicators are commonly assessed, but changes in mineralization and decomposition rates might better indicate when peat formation is underway in restored peatlands. In Finland, we investigated differences in mineralization and decomposition in the upper peat layer of five undrained and eight drained Pinus‐dominated fens from 2006 to 2009. Forestry‐drained fens were restored in 2007 by harvesting either whole trees or only stems, and by damming and filling ditches. Before restoration, net N mineralization rate was slightly higher in the drained than in undrained fens, whereas soil pH and Betula leaf litter decomposition rate were lower. After restoration, net N mineralization rate was similar for the undrained and restored fens, except near ditches after stem harvest. Also, soil pH and decomposition rate of Betula leaf litter became similar for undrained and restored fens. We conclude that whole tree harvest is a more suitable method for peatland restoration than stem harvest and that mineralization and decomposition rates are suitable indicators for peat formation after restoration.     

Photosynthetic traits of Sphagnum and feather moss species in undrained,drained and rewetted boreal spruce swamp forests

In restored peatlands, recovery of carbon assimilation by peat‐forming plants is a prerequisite for the recovery of ecosystem functioning. Restoration by rewetting may affect moss photosynthesis and respiration directly and/or through species successional turnover. To quantify the importance of the direct effects and the effects mediated by species change in boreal spruce swamp forests, we used a dual approach: (i) we measured successional changes in moss communities at 36 sites (nine undrained, nine drained, 18 rewetted) and (ii) photosynthetic properties of the dominant Sphagnum and feather mosses at nine of these sites (three undrained, three drained, three rewetted). Drainage and rewetting affected moss carbon assimilation mainly through species successional turnover. The species differed along a light‐adaptation gradient, which separated shade‐adapted feather mosses from Sphagnum mosses and Sphagnum girgensohnii from other Sphagna, and a productivity and moisture gradient, which separated Sphagnum riparium and Sphagnum girgensohnii from the less productive S. angustifolium, S. magellanicum and S. russowii. Undrained and drained sites harbored conservative, low‐production species: hummock‐Sphagna and feather mosses, respectively. Ditch creation and rewetting produced niches for species with opportunistic strategies and high carbon assimilation. The direct effects also caused higher photosynthetic productivity in ditches and in rewetted sites than in undrained and drained main sites.     

Long‐term effects of drainage and initial effects of hydrological restoration on rich fen vegetation

Questions: What vegetational changes does a boreal rich fen (alkaline fen) undergo during a time period of 24 years after drainage? How is plant species richness affected, and what are the changes in composition of ecological groups of species? Is it possible to recover parts of the original flora by rewetting the rich fen? Which are the initial vegetation changes in the flora after rewetting? What are the major challenges for restoration of rich fen flora after rewetting? Location: Eastern central Sweden, southern boreal vegetational zone. Previously rich fen site, drained for forestry purposes during 1978–1979. The site was hydrologically restored (rewetted) in 2002. Method: Annual vegetation survey in permanent plots during a period of 28 years. Results: There were three successional stages in the vegetational changes. In the first stage there was a rapid (< 5 years) loss of rich fen bryophytes. The second step was an increase of sedges and early successional bryophytes, which was followed by an increase of a few emerging dominants, such as Molinia caerulea, Betula pubescens and Sphagnum spp. After rewetting, there are indications of vegetation recovery, albeit at slow rates. Depending on, for instance, initial species composition different routes of vegetation change were observed in the flora after drainage, although after 24 years, species composition became more homogenous and dominated by a few species with high cover. Conclusion: Major changes have occurred after changes in the hydrology (drainage and rewetting) with a severe impact on the biodiversity among vascular plants and bryophytes. Several rich fen bryophytes respond quickly to the changes in water level (in contrast to vascular plants). The recovery after rewetting towards the original rich fen vegetation is slow, as delayed by substrate degradation, dispersal limitation and presence of dominant species.