Impact of local- and regional-scale restoration measures on a vulnerable rich fen in the Naardermeer nature reserve (the Netherlands)

Restoration of rich fens is commonly attempted through local-scale measures, such as removal of sod or blockage of ditches. However, regional-scale restoration measures, that aim to re-establish the original hydrology in which rich fens developed, might have a more long-lasting effect. We investigated the effect of local- and regional-scale restoration measures on a vulnerable rich fen in the Naardermeer nature reserve in the Netherlands. We compared water quality and vegetation composition of the fen before and after the restoration measures, almost 30 years apart. Overall rich fen species increased and although this indicates the desired increased supply of fresh mineral-rich groundwater to the fen, continued succession towards poor fen vegetation has not been prevented in the entire fen. Despite sod layer removal, we observed an increase in a Polytrichum-dominated vegetation in patches that are primarily fed by rainwater. Our findings confirm results from a previous study which showed that brackish palaeo-groundwater is still contributing substantially to the water balance of the fen, especially in periods of precipitation deficit. We conclude that the local- and regional-scale restoration measures have been successful in increasing the abundance of rich fen species in parts of the fen. However, considering the pressures of climate change and high atmospheric N-deposition on the fen, it is uncertain whether rich fen species can be sustained in quite nutrient-poor conditions in the future. Therefore, there is a need for continued management that keeps the nutrient-poor and mineral-rich conditions of the fen intact.

     

Higher acidification rates in fens with phosphorus enrichment

Question: Why is bryophyte succession in eutrophicated fens faster than in natural fens? Location: Mineral‐rich fens in The Netherlands and NW Europe. Methods: Literature review on the ecology of four bryophyte species in various successional types as observed in Dutch fens. Results: Bryophyte succession in eutrophicated fens from the brown moss Calliergonella cuspidata to Sphagnum squarrosum is much faster than in natural fens with species shifts from Scorpidium scorpioides to Sphagnum subnitens. Under P‐poor conditions, the brown moss stage is stabilized as long as mineral‐rich water is supplied. This is because S. scorpioides is tolerant of rainwater, is a strong competitor and can counteract acidification to some extent while S. subnitens is intolerant to groundwater and has low growth rates and low acidification capacity. In contrast, the Sphagnum stage is stable after rapid succession from rich‐fen mosses under P‐rich conditions. Calliergonella cuspidata has suboptimal growth in rainwater, possibly due to ammonium toxicity, while the high growth rates of S. squarrosum in nutrient‐rich and highly acidic groundwater allow early establishment and rapid expansion. Conclusions: If measures to improve fen base status occur in environments of increased nutrient (P) availability, the management may not lead to the desired restoration of brown moss stages, but instead to rapid acidification by S. squarrosum.     

Effects of restoration measures on plant communities of wet heathland ecosystems

Question: Which are the success and failure of restoration measures, particularly sod‐cutting and hydrological measures, in small wetlands on mineral soils in The Netherlands. Location: Twente, in the eastern part of The Netherlands. Methods: Success or failure of restoration measures has been assessed by comparing experimental plots with that in reference plots for (1) species richness, (2) re‐establishment of endangered species and (3) species composition, including life forms and Red List species. In total 119 samples were taken in 42 permanent plots in fen meadows, small‐sedge marshes, wet heathlands, and soft‐water pools. Topsoil samples were analysed for pH, Ca, Mg, Na, K and Cation Exchange Capacity. Gradient analysis was carried out by means of Canonical Correspondence Analysis. Results: Sod‐cutting, in combination with re‐wetting measures (infilling of ditches and drains), led to restoration of communities of soft‐water pools and small sedge marshes within five years. This rapid recovery is probably related to the presence of persistent seed banks of the component species of these communities. Complete restoration of fen meadows took longer (10–15 yr). Apparently, many species of fen meadows have short‐lived seed banks. Sod‐cutting of a degraded wet heathland and a soft‐water pool was only successful temporarily, probably as a consequence of low water tables. Conclusions: To prevent depletion of (persistent) soil seed banks, sod‐cutting in nutrient‐poor wetlands is not recommended in areas where the groundwater regime and the base status of the soil can not be restored to levels required by plant communities of wet heathlands.     

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.     

Nitrogen Retention by Soil Biota; A Key Role in the Rehabilitation of Natural Grasslands?

Environmental stress is the main cause of the decline of species diversity in low‐productive fen meadows in the Netherlands. Attempts to restore species diverse fen meadows e.g. by sod cutting frequently fail. We supposed that unsuccessful efforts are due to ignoring the impact of environmental stress on the performance of soil biota, which play a key role in N‐immobilization and keeping available‐N for primary production low. We investigated both pristine and degraded natural sites and successfully and unsuccessfully restored sites of poor and rich fen meadows. We determined plant species composition, soil chemical properties, N‐pools in soil biota, N‐mineralization rates, and N‐fluxes. In pristine rich and poor fen meadows, mineral‐N was poorly available for primary production due to a strong N‐immobilization by soil biota. Annual N‐immobilization fluxes exceeded by far the annual N‐harvest by primary production. N‐immobilization in pristine fens was higher than in degraded fens. In successfully restored rich fens, net N‐mineralization was lower and N‐immobilization higher than in the unsuccessful category. From our results, we derived the hypothesis that in degraded or in unsuccessfully restored fens the soils internal N‐balance shifted from N‐immobilization to net N‐mineralization, favoring biomass production but disadvantaging plant species diversity. N‐retention driven by an active N‐immobilizing soil biological community, is likely a decisive process for successful recovery of plant species diversity in low productive fen meadows. We recommend that restoration techniques should stimulate a functionally diverse soil fauna, as this may enhance the storage of available nutrients in the soil food web.     

Germination and seedling survival in fens undergoing succession

We studied mechanisms of vegetation change in fens subject to succession from open water to floating mats and finally herbaceous rich-fens. Earlier research showed that these systems are characterized by transient seed banks. Our main question was whether seedlings of later successional fen stages are already present in earlier stages, remaining subordinate in the vegetation until conditions become suitable for them. If, however, conditions during succession change in a way that only a limited set of species can survive as seedlings during each of the successional stages, no seedling bank will exist. The transient character of the seeds would then imply that seeds will not germinate and will subsequently die and that seeds that have germinated in the “wrong” stage will not become established. We hypothesized that: (1) germination and seedling survival of fen species are significantly better in the successional fen stage for which these species are characteristic, (2) as a consequence no seedling banks occur in these fens. In a field experiment, seeds of five characteristic fen species in the standing vegetation of three successional fen stages i.e. raft fen, quaking fen and rich fen were sown in each of these stages in a turf pond in the Tienhoven area, The Netherlands. Germination and seedling survival were measured over two growing seasons together with environmental variables. Germination was higher in the “own” stage for all species groups as was survival for quaking fen species and rich fen species. For both these stages, percentage of germination and survival of four out of five characteristic species were significantly higher in the “own” stage. Germination and survival can be considered stage-dependent and it was concluded that seedling banks do not exist in these fens. Site-specific environmental variables act as a sieve and differentiate on species presence already during early life history stages. We found clues that the environmental sieve acts at the level of nutrient availability, tolerance for high sulphide concentration and light climate. Because of the transient seed bank and absence of a seedling bank in these fen wetlands, successful establishment of species necessitates a continuous dispersal of characteristic species until the environmental conditions permit establishment. This also implies that species of the whole successional sere should be present within dispersal distance.     

Topographic position and water chemistry of fens in a Dutch river plain

Abstract. On the Vecht river plain (western Netherlands), small fens, remnants of a large mesotrophic wetland bordering a moraine, of 1 to 5 ha are found in a man-made matrix of lakes and pastures. The regional position of the fens, local position of sampling sites, composition of the vegetation and local hydrological variables were measured. Polders in the river plain produce a complex hydrology obscuring the regional zonation between moraine and river. Water supply and species composition are determined more by a site's regional than local position. High-productivity reedlands are abundant close to the river. Carex paniculata reedlands receive large amounts of river water, which gives their fen water a high K⁺ concentration. Low-productivity C. diandra fens and litter fens have their optimum closer to the moraine. C. diandra fens are fed mainly by inflowing nutrient-poor ground- or surface water; litter fens receive primarily rainwater. Nutrients in fen water and in peat are lowest in C. diandra and C. lasiocarpa fens, but do not differ significantly between the communities. In both, iron seems to be more important than calcium in reducing phosphate solubility. Iron richness in the C. diandra fens is caused by present inflows of ground- or surface water, while in C. lasiocarpa fens, which succeed the former, iron richness is the result of historical inflows.     

Drepanocladus trifarius– an example of unsuspected niche widths among mosses

The habitat of the pleurocarpous moss Drepanocladus trifarius is commonly described as mineral‐rich wet fens. We sampled individual D. trifarius shoots at 214 pre‐defined randomly distributed spots in an area of ca 15 km² in a sloping fen in central–western Sweden. We assessed the habitat variation of the sampling spots in this area by means of a multi‐variate analysis (DCA) based on the species identity of the ten shoots adjacent to D. trifarius, and by indicator values for light availability and substrate moisture, acidity, and nutrient availability of the associated species. Both approaches indicated that D. trifarius occurred in well lit and constantly moist, wet or submerged places. Most spots were weakly acid to sub‐neutral, but some spots had distinctly acid conditions. Generally the spots were nutrient‐poor, but a few were nutrient‐ rich. DCA revealed gradients from relatively wet, sub‐neutral to basic towards drier and more acid habitats, and from nutrient‐poor to ‐richer environments. The species obviously exhibits a broader than expected niche width with respect to substrate acidity and nutrients. We suggest that succession, disturbance, or climatic factors may explain the species’ occurrences in different habitats.     

The relationships of vegetation to surface water chemistry and peat chemistry in fens of Alberta,Canada

The relationships between vegetation components, surface water chemistry and peat chemistry from 23 fens in boreal Alberta, Canada, substantiate important differences along the poor to rich fen gradient. Each of the three fen types have their own characteristic species. The extreme-rich fens are characterized by Calliergon trifarium, Drepanocladus revolvens, Scirpus hudsonianus, S. cespitosus, Scorpidium scorpioides, and Tofieldia glutinosa. Moderate-rich fens are characterized by Brachythecium mildeanum, Carex diandra, Drepanocladus vernicosus, D. aduncus, and D. polycarpus. Poor fens are characterized by Carex pauciflora, Drepanocladus exannulatus, Sphagnum angustifolium, S. jensenii, and S. majus. Moderate-rich fens have fewer species in common with poor fens than with extreme-rich fens, while species richness is highest in the moderate-rich fens and lowest in poor fens. Variation in vascular plant occurrence appears to be more associated with nutrient levels, while bryophytes are more affected by changes in acidity and mineral elements. Based on chemical criteria, the three fen types are clearly separated by surface water pH, calcium, magnesium, and conductivity, but are less clearly differentiated by the nitrogen and phosphorus components of the surface waters. Moderate-rich fens are chemically variable both temporally and spatially, whereas poor fens and extreme-rich fens are more stable ecosystems. Whereas components of alkalinity-acidity are the most important factors that distinguish the three fen types in western Canada, nutrient concentrations in the surface waters generally do not differ appreciably in the three fen types.     

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

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 ground water flow pattern is remarkably similar. Paleoecological 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.     

Key variables controlling the vegetation of a cool-temperate mire in northern Japan

Abstract. In the cool-temperate Bibi Mire, Hokkaido, Japan, valley fens and flood-plain fens have quite different vegetation. The main variables controlling the vegetation were all hydrological: mean water level, water level fluctuation and surface water flow. Chemical factors such as electrical conductivity, dissolved oxygen and related peat decomposition were less important. The pH was about neutral and has little effect. The flood-plain fen developed under fluctuating water table conditions. The dominant species are Calamagrostis langsdotffii and Carex pseudocuraica. When temporal inundation occurs in the rainy or typhoon seasons, the submergence stimulates bud germination of the stoloniferous C. pseudocuraica, which can rapidly elongate its stolons upward. Some large floating peat mats occurred in the flood-plain fen zone. On these mats some Alnus japonica saplings establish and patches of alder forest can arise. Here the water level was higher than in the peripheral alder forest zone. The valley fen is dominated by Carex lasiocarpa var. occultans and/or C. limosa. It is formed under stable water table conditions in the inundated parts of the mire -where the non-inundated wet areas are dominated by alder trees. In the area where the surface water is flowing, these two fen sedges grow in deeper water since the high oxygen content is considered to compensate the flooding stress.     

Testing a relict distributional pattern of fen plant and terrestrial snail species at the Holocene scale: a null model approach

Aim The term relict refers to a formerly widespread species currently occurring in refugia that provide a persistent combination of specific ecological conditions. In peatlands, direct palaeoecological evidence of relict status exists for some plant species and, in the case of calcareous sediments, for some snail species. We tested whether some species are significantly linked to old calcareous fens at the millennial scale independent of the effect of recent fen area. We focused on three organism groups – vascular plants, bryophytes and land snails – that differ in the degree of preservation of their remains in calcareous fen sediments and in their dispersal ability. Location Western Carpathians (Slovakia and the Czech Republic). Methods The sample sites comprised 47 well‐preserved calcareous fens, from which we compiled complete recent species lists, measured the area and analysed radiocarbon‐dated samples from the deepest sediment and from the beginning of complete deforestation, as indicated by plant and snail fossils. Using the species co‐occurrences in large data sets, we identified calcareous fen specialists and compared their recent distribution patterns against a null model that controlled for the effect of fen area. Results Two land snail species, eleven vascular plant species and no bryophyte species have statistically significant affinities with old fens, independent of the effect of recent fen area. For one bryophyte and one snail, the effects of age and area are not distinguishable. Main conclusions The results for land snails, being abundantly preserved and easily determinable in calcareous fen deposits, are in full accordance with the direct macrofossil evidence. This suggests that our approach indirectly revealed a relict distribution of the species. Identification of species that are significantly linked to ancient localities at the millennial scale has great potential in palaeoecology for the detection of stands with old sediments, and in nature conservation as a tool for the identification of long‐term‐persisting rare species that infrequently colonize young sites and thus deserve priority in the protection of their habitats. From a theoretical perspective, limited dispersal from old to new localities of the same habitat can contribute to spatial effects in biotic assemblages, even at relatively fine scales.     

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 restoration of species-rich heathland communities in the Netherlands

The wet heathland communities of the Ericetum tetralicis and the Cirsio-Molinietum have declined in the Netherlands due to acidification, eutrophication and lowering of the water table. To investigate the prospects of restoration of both communities, the effects of sod cutting and hydrological measures on vegetation and soil chemistry were studied in two nature reserves where these plant communities occurred decades ago. The combination of sod cutting and hydrological measures has restored several rare, groundwater dependent heathland communities. Sod cutting has restored the Ericetum tetralicis, but not the Cirsio-Molinietum. This might be due to the absence of viable seeds of characteristic species of the Cirsio-Molinietum and/or the absence of optimal site conditions, especially high phosphorus concentrations in the top soil. The high phosphorus concentrations might be a consequence of high mineralization rates and/or prolonged inundation with iron-poor water and the decreased flux of iron-rich groundwater into the topsoil. Restoration of the Cirsio-Molinietum only seems possible when sod cutting is carried out together with hydrological measures that counter prolonged inundation and reinforce the discharge of base and iron-rich groundwater.     

In search of a hydrological explanation for vegetation changes along a fen gradient in the Biebrza Upper Basin (Poland)

The understanding of succession from rich fen to poorer fen types requires knowledge of changes in hydrology, water composition, peat chemistry and peat accumulation in the successional process. Water flow patterns, water levels and water chemistry, mineralisation rates and nutrient concentrations in above-ground vegetation were studied along a extreme-rich fen-moderate-rich fen gradient at Biebrza (Poland). The extreme-rich fen was a temporary groundwater discharge area, while in the moderate-rich fen groundwater flows laterally towards the river. The moderate-rich fen has a rainwater lens in spring and significant lower concentrations of calcium and higher concentrations of phosphate in the surface water. Mineralisation rates for N, P and K were higher in the moderate-rich fen. Phosphorus concentrations in plant material of the moderate-rich fen were higher than in the extreme-rich fen, but concentrations of N and K in plant material did not differ between both fen types. Water level dynamics and macro-remains of superficial peat deposits were similar in both fen types.We concluded that the differences observed in the moderate-rich and the extreme-rich fens were caused by subtile differences in the proportion of water sources at the peat surface (rainwater and calcareous groundwater, respectively). Development of an extreme-rich fen into a moderate-rich fen was ascribed to recent changes in river hydrology possibly associated with a change in management practices. The observed differences in P-availability between the fen types did not result in significantly different biomass. Moreover, biomass production in both fen types was primarily N-limited although P-availability was restricted too in the extreme-rich fen. Aulacomnium palustre, the dominant moss in the moderate-rich fen, might be favoured in competition because of its broad nutrient tolerance and its quick establishment after disturbance. It might outcompete low productive rich fen species which were shown to be N-limited in both fens. We present a conceptual model of successional pathways of rich fen vegetation in the Biebrza region.     

Experimental restoration of a fen plant community after peat mining

Question: Which restoration measures (introduction of donor diaspore material, application of straw mulch, alteration of residual peat depths) contribute to the establishment of a fen plant community on minerotrophic surfaces after peat mining? Location: Rivière‐du‐Loup peatland, southern Québec, Canada at 100 m a.s.1. Methods: The effectiveness of introducing fen plants with the application of donor diaspore material was tested. The donor diaspore material, containing seeds, rhizomes, moss fragments, and other plant propagules, was collected from two different types of natural fens. We tested whether the application of straw mulch would increase fen species cover and biodiversity compared to control plots without straw mulch. Terrace levels of different peat depths (15 cm, 40 cm, and 56 cm) were created to test the effects of different environmental site conditions on the success of re‐vegetation. Results: Applying donor seed bank from natural fens was found to significantly increase fen plant cover and richness after the two growing seasons. Straw mulch proved to significantly increase fen plant richness. The intermediate terrace level (40 cm) had the highest fen plant establishment. Compared to reference sites, the low terrace level (15 cm) was richer in base cations, whereas the high terrace level (56 cm) was much drier. Conclusions: The application of donor diaspore material was demonstrated as an effective technique for establishing vascular fen plants. Further re wetting measures are considered necessary at the restoration site to create a fen ecosystem rather than simply restoring some fen species.     

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.     

Recent changes in the plant composition of wetlands in the Jura Mountains

Aim

To assess vegetation changes in montane fens and wet meadows and their causes over 38 years.

Location

Wetlands, Jura Mountains (Switzerland and France).

Methods

Plots were inventoried in 1974 and re‐located in 2012 (quasi‐permanent plots) on the basis of sketches to assess changes in plant communities. The 110 plots belonged to five phytosociological alliances, two in oligotrophic fens (Caricion davallianae, Caricion fuscae) and three in wet meadows (Calthion, Molinion, Filipendulion). Changes between surveys were assessed with NMDS, and changes in species richness, Simpson diversity, species cover and frequency and the causes of these changes were evaluated by comparing ecological indicator values.

Results

Changes in species composition varied between alliances, with a general trend towards more nutrient‐rich flora with less light at ground level. Species diversity declined, with a marked decreasing trend for the typical species of each alliance. These species were partly replaced by species belonging to nitrophilous and mesophilous grasslands. However, no trend towards drier conditions was detected in these wetlands. The largest changes, with an important colonization by nitrophilous species, occurred in the Swiss sites, where grazing was banned 25 years ago. As a result of floral shifts, many plots previously belonging to fens or wet mesotrophic meadows shifted to an alliance of the wet meadows, generally Filipendulion. Moreover, communities showed a slight trend towards more thermophilous flora.

Conclusions

The investigated wetlands in the Jura Mountains have suffered mainly from eutrophication due to land‐use abandonment and N deposition, with a loss of typical species. Areas with constant land use (grazing or mowing) showed less marked changes in species composition. The most important action to conserve these wetlands is to maintain or reintroduce the traditional practices of extensive mowing and livestock grazing in the wetlands, especially in areas where they were abandoned 25 years ago. This previous land‐use change was intended to improve fen conservation, but it was obviously the wrong measure for conservation purposes.     

Stream characteristics and their implications for the protection of riparian fens and meadows

1. Running waters, including associated riparian areas, are embraced by international legal frameworks outlining targets for the preservation, protection and improvement of the quality of the environment. Interactions between stream and river processes and riparian habitats have not received much attention in the management of stream ecosystems, and integrated measures that consider both the ecological status of streams and rivers (sensu EU Water Framework Directive, WFD) and the conservation status of riparian habitats and species (sensu EU Habitats Directive, HD) are rare. 2. Here, we analysed the influence of stream size, morphology and chemical water characteristics for the distribution of water‐dependent terrestrial habitat types, i.e. alkaline fens, periodically inundated meadows and meadows in riparian areas in Denmark using an extensive data set covering a total of 254 stream reaches. A species‐based classification model was used to translate species lists into a standardised interpretation of habitat types protected by the HD in Denmark. 3. No size dependency was found regarding the distribution of fen and meadow vegetation. Instead, the distribution of fen and meadow vegetation was strongly affected by the morphology of the streams. Alkaline fens, periodically inundated meadows and meadows occurred six, five and four times, respectively, less frequently along channelised compared with natural stream reaches. Our results indicate that stream channelisation strongly interfered with the natural hydrology of riparian areas, affecting conditions needed to sustain protected fen and meadow communities. 4. We also found that water chemistry strongly influenced the occurrence of fen and meadow vegetation in riparian areas. The probability of finding fen and meadow vegetation was reduced when total phosphorus (TP) concentration exceeded 40–50 μg P L^?1, whereas meadow vegetation responded less strongly to TP. 5. Our findings highlight the importance of restoring hydrology of riparian areas to improve conditions for fen and meadow vegetation, but also that the water chemistry should be considered when measures that increase hydrological connectivity result in an increase in the probability of flooding.     

Accelerated vegetation succession but no hydrological change in a boreal fen during 20 years of recent climate change

Northern mires (fens and bogs) have significant climate feedbacks and contribute to biodiversity, providing habitats to specialized biota. Many studies have found drying and degradation of bogs in response to climate change, while northern fens have received less attention. Rich fens are particularly important to biodiversity, but subject to global climate change, fen ecosystems may change via direct response of vegetation or indirectly by hydrological changes. With repeated sampling over the past 20 years, we aim to reveal trends in hydrology and vegetation in a pristine boreal fen with gradient from rich to poor fen and bog vegetation. We resampled 203 semi‐permanent plots and compared water‐table depth (WTD), pH, concentrations of mineral elements, and dissolved organic carbon (DOC), plant species occurrences, community structure, and vegetation types between 1998 and 2018. In the study area, the annual mean temperature rose by 1.0°C and precipitation by 46 mm, in 20‐year periods prior to sampling occasions. We found that wet fen vegetation decreased, while bog and poor fen vegetation increased significantly. This reflected a trend of increasing abundance of common, generalist hummock species at the expense of fen specialist species. Changes were the most pronounced in high pH plots, where Sphagnum mosses had significantly increased in plot frequency, cover, and species richness. Changes of water chemistry were mainly insignificant in concentration levels and spatial patterns. Although indications toward drier conditions were found in vegetation, WTD had not consistently increased, instead, our results revealed complex dynamics of WTD as depending on vegetation changes. Overall, we found significant trend in vegetation, conforming to common succession pattern from rich to poor fen and bog vegetation. Our results suggest that responses intrinsic to vegetation, such as increased productivity or altered species interactions, may be more significant than indirect effects via local hydrology to the ecosystem response to climate warming.