Ab P. Grootjans

Hydrological landscape settings of base-rich fen mires and fen meadows: an overview

ABSTRACT Question: Why do similar fen meadow communities occur in different landscapes? How does the hydrological system sustain base-rich fen mires and fen meadows? Location: Interdunal wetlands and heathland pools in The Netherlands, percolation mires in Germany, Poland, and Siberia, and calcareous spring fens in the High Tatra, Slovakia. Methods: This review presents an overview of the hydrological conditions of fen mires and fen meadows that are highly valued in nature conservation due to their high biodiversity and the occurrence of many Red List species. Fen types covered in this review include: (1) small hydrological systems in young calcareous dune areas, and (2) small hydrological systems in decalcified old cover sand areas in The Netherlands; (3) large hydrological systems in river valleys in Central-Europe and western-Siberia, and (4) large hydrological systems of small calcareous spring fens with active precipitation of travertine in mountain areas of Slovakia. Results: Different landscape types can sustain similar nutrient poor and base-rich habitats required by endangered fen meadow species. The hydrological systems of these landscapes are very different in size, but their groundwater flow pattern is remarkably similar. Paleo-ecological research showed that travertine forming fen vegetation types persisted in German lowland percolation mires from 6000 to 3000 BP. Similar vegetation types can still be found in small mountain mires in the Slovak Republic. Small pools in such mires form a cascade of surface water bodies that stimulate travertine formation in various ways. Travertine deposition prevents acidification of the mire and sustains populations of basiphilous species that elsewhere in Europe are highly endangered. Conclusion: Very different hydrological landscape settings can maintain a regular flow of groundwater through the top soil generating similar base-rich site conditions. This is why some fen species occur in very different landscape types, ranging from mineral interdunal wetlands to mountain mires.

Ecological restoration of rich fens in Europe and North America: from trial and error to an evidence-based approach.

Fens represent a large array of ecosystem services, including the highest biodiversity found among wetlands, hydrological services, water purification and carbon sequestration. Land‐use change and drainage has severely damaged or annihilated these services in many parts of North America and Europe; restoration plans are urgently needed at the landscape level. We review the major constraints on the restoration of rich fens and fen water bodies in agricultural areas in Europe and disturbed landscapes in North America: (i) habitat quality problems: drought, eutrophication, acidification, and toxicity, and (ii) recolonization problems: species pools, ecosystem fragmentation and connectivity, genetic variability, and invasive species; and here provide possible solutions. We discuss both positive and negative consequences of restoration measures, and their causes. The restoration of wetland ecosystem functioning and services has, for a long time, been based on a trial‐and‐error approach. By presenting research and practice on the restoration of rich fen ecosystems within agricultural areas, we demonstrate the importance of biogeochemical and ecological knowledge at different spatial scales for the management and restoration of biodiversity, water quality, carbon sequestration and other ecosystem services, especially in a changing climate. We define target processes that enable scientists, nature managers, water managers and policy makers to choose between different measures and to predict restoration prospects for different types of deteriorated fens and their starting conditions.

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.

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.

Zero methane emission bogs: extreme rhizosphere oxygenation by cushion plants in Patagonia

• Vascular wetland plants may substantially increase methane emissions by producing root exudates and easily degradable litter, and by providing a low-resistance diffusion pathway via their aerenchyma. However, model studies have indicated that vascular plants can reduce methane emission when soil oxygen demand is exceeded by oxygen released from roots. Here, we tested whether these conditions occur in bogs dominated by cushion plants. • Root-methane interactions were studied by comparing methane emissions, stock and oxygen availability in depth profiles below lawns of either cushion plants or Sphagnum mosses in Patagonia. • Cushion plants, Astelia pumila and Donatia fascicularis, formed extensive root systems up to 120 cm in depth. The cold soil (< 10°C) and highly decomposed peat resulted in low microbial activity and oxygen consumption. In cushion plant lawns, high soil oxygen coincided with high root densities, but methane emissions were absent. In Sphagnum lawns, methane emissions were substantial. High methane concentrations were only found in soils without cushion plant roots. • This first methane study in Patagonian bog vegetation reveals lower emissions than expected. We conclude that cushion plants are capable of reducing methane emission on an ecosystem scale by thorough soil and methane oxidation.

A multidisciplinary evaluation of restoration measures in a degraded Cirsio-Molinietum fen meadow

. An integrated multidisciplinary approach including descriptive and experimental research of environmental and biotic factors was used to evaluate an attempt to restore a species-rich fen meadow, phytosociologically belonging to the Cirsio-Molinietum. Expert judgement and preliminary research indicated that soil acidification, due to drainage in the surrounding area, caused the degeneration of the fen meadow. Flooding the meadow with surface water was applied in order to stop acidification. A small irrigated wetland (helophyte filter) was constructed to reduce nutrient availability in the surface water. Sod cutting was applied in a small area as well. The two measures were evaluated through: (1) measurements of hydrological and soil factors, (2) seed bank analysis, (3) recording species composition in permanent plots, (4) assessing the type and extent of nutrient limitation using a full-factorial fertilization experiment, (5) assessing the effect of liming on vegetation biomass, and (6) recording the growth of three introduced fen meadow species in the liming experiment. The research was carried out in a degraded site with intact vegetation, a degraded site where the top soil was removed by sod cutting and a reference site with intact Cirsio-Molinietum vegetation. Groundwater levels in the degraded and the stripped site differed slightly from those of the reference site but they were within the range of Cirsio-Molinietum requirements. Groundwater quality slightly improved in the stripped site. Top-soil removal considerably reduced the macro-nutrient contents and caused a stronger P-deficiency. No increase in soil pH was observed immediately after top-soil removal. A small increase in soil pH was found after five years in the stripped site, indicating a slight recovery from acidification. Liming after top-soil removal had little effect on the soil pH after six months but stimulated the growth of established species such as Agrostis canina. Few viable seeds of characteristic fen meadow species were present in the seed banks of both degraded and well developed fen meadow sites. The species composition after top soil removal reflected the seed bank of the degraded fen meadow and establishment of characteristic fen meadow species was poor, even after five years. The introduced species (Carex hostiana, C. panicea, Cirsium dissectum) survived during the first year in both the degraded and the stripped site. Liming initially stimulated the growth of two species, but in the second year this effect disappeared. After two years, none of the introduced species survived at the stripped site. The restoration measures failed so far, probably because acidification could not be counteracted satisfactorily. Finally, it was concluded that our multidisciplinary approach enables us to draw proper conclusions and that it is suitable for other restoration attempts as well.

Restoration of the Cirsio dissecti-Molinietum in The Netherlands : Can we rely on soil seed banks?

Abstract Vegetation and soil seed banks of a threatened Atlantic fen meadow community were studied using recent phytosociological records and seedling emergence from soil samples. Similarly managed but differently degraded stands that suffered different levels of species impoverishment were compared. The actual vegetation was related to a set of phytosociological references representing the subassociations of the community. DCA positions of reference relevés from the different subassociations were overlapping, suggesting that in all references many common species occur. Recent records were positioned in-between the seed bank samples and the references. The soil seed banks of all stands were dominated by ordinary species. Most character species had at most sparse seed banks and no seedlings of locally extinct character species, mentioned in historic floristic records, were detected. In contrast species of pioneer and small-sedge communities as well as those of heathlands were abundant in the seed banks. Based on the vertical distribution of seeds in the soil layers most fen meadow species were classified into transient or short-term persistent seed bank types. We concluded that complete restoration of the Cirsio dissecti-Molinietum without reintroduction is only likely in stands that were degraded only a few years ago. On the other hand, the presence of viable seeds of Nanocyperion and Parvocaricetea species is promising for the restoration of these communities even after decades. Recreation of pioneer habitats by sod cutting will preserve these species. Nomenclature: van der Meijden (1996) for taxa; Schaminée et al. (1996) for syntaxa. Abbreviations: DCA = Detrended Correspondence Analysis; EXV = Eexterveld; ULE = Ule Krite; WYL = Wyldlannen; WTS = Wijnjeterperschar.

Possible positive-feedback mechanisms: plants change abiotic soil parameters in wet calcareous dune slacks

In this paper the results are presented from a mesocosm study of the effects of typical dune slack plants on the soil solution nutrient contents. In dune slack succession, early successional species often show radial oxygen loss (ROL) whereas their successor species do not show ROL. ROL has impact on abiotic soil parameters and therefore, affect the competitiveness of both species. Mesocosms with Littorella uniflora and Carex nigra, used as respectively a ROL and a non-ROL species, showed remarkable differences in soil solution parameters. Special attention was given to nitrogen, as it is the limiting resource in dune slack succession. Mesocosms with L. uniflora showed a higher nitrate content in the soil than mesocosms with C. nigra and the control. Moreover, estimating the nitrogen balance, a significantly higher fraction of nitrogen was missing in L. uniflora (57%) than in C. nigra (5%). The enhanced nitrogen loss in mesocosms with L. uniflora could act as a positive-feedback mechanism for early successional stages that slows down the vegetation development in early stages of dune slack succession towards the more-productive later stages. The mechanism could even lead to alternative stable states in dune slack succession.

Hydrology of Dutch Cirsio‐Molinietum meadows: Prospects for restoration

. 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.

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 kg m –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. Nomenclature:van der Meijden (1996) for phanerogams; Schaminée et al. (1995) for syntaxa.