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.     

Shading by Phragmites australis: a threat for species-rich fen meadows?

Abstract. Common reed (Phragmites australis Trin.) has spread in fen meadows on the Swiss Plateau during the last decades. An increased dominance of this tall grass may reduce the plant species richness and displace rare or endangered species because of the additional shade. To investigate whether this has actually happened and whether shading by Phragmites was likely to be responsible for the reduction, the plant species composition was surveyed in 241 plots (4 m²) with differing above-ground biomass of Phragmites (ABP). Species richness and the occurrence of characteristic fen species were negatively related to site productivity (total above-ground biomass), but correlations with ABP were generally weaker. The main change associated with increasing ABP within species-rich fens (alliances Molinion and Caricion davallianae) was an increase of species characteristic of the more species-poor alliance Filipendulion. Thus, Phragmites did apparently not play a disproportionate role in the reduction of species richness in the plant communities studied. Selective clipping experiments and light measurements also indicated that shading by Phragmites does not have a strong effect upon the performance of other species in the community, at least in the short term. The reason appeared to lie in the later seasonal growth of Phragmites compared with the other species. These results suggest that the direct impact of shading by Phragmites on the species richness of fen meadows is probably lower than has been assumed. However, possible long-term or indirect effects still need to be investigated.     

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.     

The relation between vegetation and soil chemistry gradients in a ground water discharge fen

Abstract. The soil chemistry of a headwater valley fen is influenced by local ground water discharge that supplies base cations and alkalinity to the fen. An irrigation canal just upward of the fen is the source of this alkalinity. The ecological consequences of this artificial system are studied both on the soil and vegetation level. Rich-fen species of the alliance Caricion davallianae are connected to soil water alkalinity and soil base status. They depend directly on the alkaline ground water discharge. In addition, the local input of this water causes a gradient-rich pattern from poor to rich fen, and it is therefore concluded that it is responsible for the presence of intermediate fen vegetation too. High nutrient levels in the irrigation water have not influenced the fen until now. This case study illustrates the possibility for rich fen restoration after acidification. Irrigation with alkaline water is efficient if excess nutrients can be removed.     

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.

     

Hydro-ecological analysis of the fen system Lieper Posse,eastern Germany

The Lieper Posse is a calcareous, ground water-fed fen system, situated in the end-moraine area of northeastern Germany. Nowadays the fen is partly drained and the vegetation adversely affected; hence regeneration measures are considered. Here we report a pilot survey of the actual vegetation cover in relation to the hydrological conditions of the system. The vegetation in the central part of the fen system can be assigned to the Caricetum lasiocarpae, with small areas of Eleocharitetum quinqueflorae. The southern part of the system includes a Ledo-Pinetum sylvestris bog. Along the edges eutrophic forest types are present. Peat analysis revealed that the system started as an open lake and subsequently changed into a rheophilous mire, ‘percolating mire’. The southern bog has only recently come into being. Hydrological investigations showed that the original mineral-rich ground water had to a large extent been replaced by acid rainwater; this is likely due to the construction of a ditch before 1850. The effects of this drainage on hydrological conditions and some aspects of soil fertility are discussed. Possible consequences for the restoration of calciphilous vegetation types are being considered. It is concluded that the vegetation development is predictable only to a limited extent, which is mainly due to a lack of knowledge on hysteresis effects in both vegetation and soil.     

Are the characeae able to indicate the origin of groundwater in former river channels?

The macroscopic algae Characeae are usually assumed to occur in waterbodies supplied by groundwater with low phosphate content, but the indicative value of the species is seldom defined in bibliography. Former braided channels of the Rhône river are supplied with groundwater originating from the main channel (seepage) or from hillslope aquifer. The aim of the present paper was to determine if it possible to use the Characeae as indicators of physicochemical characteristies of water in order to assess the origin of groundwater supplying former river channels. Four former braided channels of the Rhône River colonized by Characeae were investigated, and the physico-chemical characteristics of i) the channels, ii) the groundwater and iii) the river were measured over a period of several months. Species are arranged along a gradient of conductivity, alkalinity, ammonium and phosphate content of the water. Charophyte species can indicate the origin of groundwater, either seepage or hillslope nutrient-poor aquifer, and integrate both the average value of the chemical parameter, and their variations. C. hispida occurs in a nutrient-poor channel mainly supplied by highly calcareous groundwater coming from hillslope aquifer. Chara major has requirements close to those of C. hispida, but is more tolerant to periodic inputs of nutrients. C. vulgaris and N. syncarpa both tolerate mesotrophic waters originating from both hillslope aquifer and seepage, and C. globularis is associated to a channel mainly supplied by mesotrophic to eutrophic river seepage.     

N- and P-addition inhibits growth of rich fen bryophytes

A greenhouse experiment was set up to investigate if infrequently and frequently occurring species respond differently to simulated habitat changes. Two frequently occurring (Bryum pseudotriquetrum and Calliergonella cuspidata) and two infrequently occurring (Hamatocaulis vernicosus and Paludella squarrosa) rich fen bryophytes were grown in mixed culture and subjected to rainwater or groundwater and three levels of N, ammonium nitrate (0, 1 and 3?mg?N?L^–1) and P, potassium phosphate (0, 0.05 and 0.1?mg?P/L). All species responded negatively to higher N-levels and three of the four species responded negatively to rainwater and higher P-levels. C. cuspidata had highest relative growth rate (RGR) in all treatments, and the infrequently occurring species had lower RGR and were more negatively affected by high levels of N than the frequently occurring species. A negative effect of rainwater seemed to be caused by higher background levels of N in rainwater compared to groundwater. We found a negative effect of high initial bryophyte density in three of the four species indicating density-dependent inhibition between species. We suggest that maintenance of oligotrophic conditions by recharge of nutrient-poor groundwater is vital for conservation of infrequently as well as frequently occurring rich fen bryophytes.     

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.     

Hydro-ecological analysis of the Biebrza mire (Poland)

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

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     

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.     

Hydro-geochemical analysis of a degraded dune slack

A hydrochemical analysis of the groundwater composition was carried out in a degraded dune slack complex in order to assess the prospects for regeneration of low productivity, basiphilous vegetation types, typical of natural dune slacks. Two hundred and fifty water samples, most of them obtained from minifilters installed in deep borings to a depth of 24 meter below the soil surface, were analyzed for macro-ions, EC₂₅ and pH. The hydrochemical results were compared with simulated flow paths and with the results of isotope analysis (³H and ¹⁸O). The simulation of the groundwater flow paths showed that the inflow of seepage water was only possible when the slack was flooded. Under such conditions groundwater exfiltrates in the upgradient part of the slack, proceeds as surface water to finally pass through the anaerobic slack bottom in the downgradient infiltration part of the slack. The CO₂ producing plant roots and the microbial population in the bottom sediments create a groundwater type, which is highly reduced and aggressive toward calcareous substrates in the subsoil. The distribution of the ³H and ¹⁸O isotopes was in accordance with the simulated groundwater flow pattern, illustrating a modest seepage intensity in the flow-through lake. A lowering of the water table in the surroundings of the slack during the last decades had weakened the above mentioned hydrological mechanism to the extent that at present the bicarbonate groundwater can only enter the slack during short periods in the wet season. These results suggest that the decline of endangered basiphilous slack species in the slack is caused by a decrease in acid buffering capacity of the slack.     

Multiple effects of land‐use changes impede the colonization of open water in fen ponds

Question: Dutch fen areas have become embedded in intensively used landscapes, resulting in biodiversity loss. Hence, plant species that colonize open water inducing the formation of species‐rich floating peat mats have disappeared. Despite many restoration efforts, they have not returned. Is natural succession towards floating mats impeded by site conditions, dispersal limitations or changed biotic interactions? Location: Six Dutch fen reserves: De Deelen, De Weerribben, De Wieden, Westbroek, Molenpolder and Terra Nova. Methods: In 62 fen ponds we determined plant species richness and expansion into open water. We related these to habitat quality (chemical composition of soil and surface water, pond morphology), dispersal potential (distance to remnant populations, likelihood of dispersal) and biotic interactions (presence of muskrats [Ondatra zibethicus L.] and the keystone species Stratiotes aloides). Results: Factor analysis showed that plants expanded further into open water and bank vegetation had higher species richness in areas with older ponds and lower muskrat densities. Locally, high turbidity hampered colonization. Whenever the water was clear, colonization was higher in shallow ponds, and in deep ponds only if Stratiotes was present. Species richness was negatively correlated to nutrient availability in soil and positively correlated to hydrological isolation (decreased sulphate concentrations). We also found that species richness was higher in sheltered banks. Conclusions: Multiple habitat characteristics (turbidity, water depth, nutrient and sulphate concentrations) and the influence of muskrats and Stratiotes all play a role in the lack of restoration success in Dutch fen ponds. Dispersal limitations seem to be overruled by habitat limitations, as colonization often fails even when sufficient propagule sources are present, or when connectivity is high.     

Groundwater and surface water hydrology of a small groundwater-fed fen

The hydrology of a small quaking fen was investigated by measuring all components of the water budget. Water-level measurements indicated that the fen is a focus for groundwater discharge, and that there is a lateral (subsurface) flow from the fen toward surrounding areas during most of the year. The hydrology, however, is strongly influenced by man, as water tables are maintained during dry periods by pumping water into the area. This action results in a reversal of flow patterns as surface water from a ditch infiltrates the fen and groundwater recharge occurs.Two approaches for calculation of groundwater and surface water terms were compared. The two procedures differ in the way that vertical and horizontal hydraulic conductivity of the soil is calculated. Hydraulic conductivity estimated from in situ tube experiments was one to two orders of magnitude lower than hydraulic conductivity estimates from numerical water budget analyses.Four one-month periods were selected for calculations of complete water budgets. Results of calculations of water flow were compared in respect to the overall error in the water budget (calculated as percent difference between inputs and outputs). Water budgets calculated from data on hydraulic conductivity gave high residual errors (56.% ± 16.3%). When hydraulic conductivity was derived from numerical analyses, the errors were very small (2.8% ± 3.1%) and the difference between inputs and outputs over a period of 127 days was only 2.6 mm (0.45%).     

Differential recovery of above‐ and below‐ground rich fen vegetation following fertilization

Abstract. For seven years we studied the recovery of vegetation in a Belgian P limited rich fen (Caricion davallianae), which had been fertilized with nitrogen (200 g.m^?2) and phosphorus (50 g.m^?2) in 1992. The vegetation in this fen has low above‐ground biomass production (< 100 g m^?2) due to the strong P limitation. Above‐ground biomass was harvested from 1992 to 1998 and P and N concentrations measured. In 1998, below‐ground biomass was also harvested. The response to fertilization differed markedly between below‐ and above‐ground compartments. Above‐ground, P was the single most important factor that continued to stimulate growth 7 yr after fertilization. Below‐ground, N tended to decrease live root biomass and increase dead root biomass and seemed to have a toxic effect on the roots. In the combined NP treatment the stimulating effect of P (an increase of live root biomass) was countered by N. The 1998 soil analysis showed no difference in soil P in the plots. Thus, Fe hydroxides are not capable of retaining P in competition with fen vegetation uptake. However, higher capture of P in root Fe coatings from N plots may partially explain this negative N effect. The results suggest that N root toxicity will be amplified in strongly P limited habitats but that its persistence will be less influenced by P availability. This mechanism may be a competitive advantage for N₂ fixing species that grow in strongly P limited wetlands.     

Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow

Theeffects of light competition and litter on seedling recruitment and theperformance of established individuals were examined in Violapalustris. This polycarpic perennial plant was a common component ofspecies-rich fen meadows in northwestern Germany until the middle of thiscentury, but today is considered to be regionally endangered. From summer 1996until summer 1998 a bi-factorial field experiment combining three standing croptreatments (mowing, thinning, control) with two litter layer treatments (litterremoval, no litter removal) was carried out in an abandoned fen meadow toinvestigate the effects of these factors both on V.palustris and on aboveground species composition and diversity.MANOVAs (multivariate analysis of variance) revealed significant treatmenteffects for the performance of V. palustris related toexperimental manipulation. Mowing increased vital attributes including thenumber of rosette leaves, the percentage of individuals with chasmogamous andcleistogamous flowers and the number of rhizomes. In contrast, the length of theleafstalk, mean leaf area and the length of rhizomes were negatively associatedwith mowing. Litter removal significantly increased the number of V.palustris seedlings. The removal of the litter layer already resultedin an increase in aboveground species richness in the first year of theexperiment. After three years of experimental manipulation, both abovegroundspecies richness and diversity were positively associated with mowing.Redundancy Analysis showed that typical wet meadow species (Cardaminepratensis, Lychnis flos-cuculi, Lotuspedunculatus) and species of mesotrophic fens (Violapalustris, Agrostis canina, Potentillapalustris) were positively influenced by mowing. It can be concludedthat V. palustris recruitment and growth is promoted by aregular disturbance regime on fen meadows. The conservation of many typicalspecies of wet meadows and fens in northwestern Germany depends on moderategrazing or cutting. These management types prevent successional changes whichlead to an increase in standing crop and the development of a litter layer, bothof which can cause local extinction.     

Insect conservation in Michigan prairie fen: addressing the challenge of global change

Prairie fen is a globally rare, groundwater dependent peatland community restricted to discrete portions of the glaciated north central USA. Prairie fen harbours a diverse flora composed of sedge wetland and tallgrass prairie species, which in turn support a diversity of rare insects. In Michigan, USA over 20% of the state’s insects of conservation concern are associated with prairie fen, including the globally imperilled Mitchell’s satyr butterfly, Neonympha mitchellii mitchellii (Lepidoptera: Nymphalidae). Here we investigate how global change drivers, including land use change, climate change, and invasive species, may interact to threaten this important community. Specifically, we examine how characteristics of prairie fen habitats—e.g., formation and distribution—interact with the biology of rare fen insects to suggest appropriate short to long term conservation strategies. Our results suggest that prairie fen associated insects are rare for a variety of reasons, including host plant specialization, habitat specialization, and shifting landscape context that limits opportunities for dispersal. We recommend that current conservation efforts focus on stabilization and restoration of existing prairie fens, coupled with directed surveys to monitor population change in insects of concern, and restoration of the landscape matrix to facilitate metapopulation dynamics. In the future, due to the severely fragmented nature of Michigan landscapes, captive rearing and assisted migration may be necessary to conserve some prairie fen insect species. Overall, the effective conservation of fen associated insects will require a shared vision by multiple actors and a willingness to purse that vision over a long time frame.