A comparison of fens in natural and artificial landscapes

Fens depend on inputs of groundwater or surface water. In Western Europe especially soligenous fens, receiving groundwater, are threatened by human hydrological intervention. We demonstrate the impact of artificial versus natural hydrologies on such fens by comparing 3 case areas: the Biebrza valley (reference) and the Gorecht and Vecht river plains (both reclaimed and drained). The patterns found in the fairly undisturbed Biebrza area suggest local water quality is governed by a strong regional groundwater flow emerging in the fen near the valley margins and seeping through it down to the river. Hence water quality gradients are smooth: there is little variation in water type over large distances. The pattern is determined by the natural geomorphology. In the reclaimed Vecht and Gorecht river plains large differences exist at short distance. Regional water flow from the adjacent ridges into the plains is weak here and governed primarily by water management (polders and pumping wells). However, the relations between specific water types and fen species and communities in this artificial pattern are quite similar to those found in the natural landscape. Low-productive rich fens are fed by calcium-rich and base-rich, nutrient-poor groundwater in both cases. While conservation of such rich fens is served best by maintaining the natural groundwater flow, some opportunities for restoration with an artificial hydrology are discussed.     

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.     

Environmental conditions and fen vegetation in three lowland mires

The abiotic conditions and fen vegetation in three lowland mires were analysed. Two of these mires are in the Netherlands. They have deteriorated considerably as a result of human pressure. One mire complex is in Poland. Its hydrology is almost undisturbed. The variation in the water composition in the fens was associated with the variation in the amount of regional groundwater discharge originating from the uplands, and to a lesser extent by the infiltration of polluted water pumped into the fens.The 26 vegetation types ranged from poor fen to eutrophic reedland. Most types were typical for only one region.There were clear differences between the three regions. Both the species composition and the water quality for the vegetation types indicated that the fens in the Vecht river plain are eutrophicated, whereas the fens in Northwestern Overijssel suffer from acidification. The Biebrza valley fens are well developed and are mainly fed by fresh calcareous groundwater.Stepwise logistic regression performed on 28 fen species revealed that the hydrochemical variables explained only a part of the variance: the regional variable area explained a considerable amount of the variance for most species. In 27% of the cases it was possible to fit an optimum curve for the species response to relevant hydrochemical variables. Monotonic curves could be fitted in 64% of the cases; mostly they described the response to variables in a way that is supported by ecological literature. It was concluded that the dataset should be constructed differently to enable the calculation of generally applicable standards.     

Environmental drivers of subalpine and alpine fen vegetation in the Southern Rocky Mountains,Colorado, USA

Fens are widely distributed wetlands worldwide and provide vital habitat for plant and animal species in mountainous regions. Alpine fens are rare in the Rocky Mountains and concentrated in the San Juan Mountains where broad regions at high elevation have relatively level topography and suitable climate to favor peat accumulation. Studies of montane and boreal peatlands have identified water chemistry as a main driver of vegetation composition. This study investigated whether similar drivers of vegetation composition are important for alpine and subalpine fens in the San Juan Mountains of Colorado, USA. Water chemistry variables were most important in structuring subalpine and alpine fen vegetation. However, these variables explained considerably less variation in alpine than subalpine fen vegetation. In addition, lower variance of water chemistry in alpine fens did not lead to lower beta diversity of vegetation in alpine than in subalpine fens. Although alpine and subalpine fen vegetation supports similar beta diversity, key differences occur in the environmental drivers of their vegetation composition.     

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.     

Release of inorganic N,P and K in peat soils; effect of temperature,water chemistry and water level

In the Netherlands, fens that are fed by polluted river water are often eutrophic, whereas fens fed by calcium-rich groundwater often are mesotrophic. Differences in trophic status can not always be attributed to differences in the nutrient load of the water. In this paper we try to determine if the inflow of river water in fens, in fact, accelerates the soil nutrient release, thereby creating more eutrophic conditions (‘internal eutrophication’). For this purpose, we compared nutrient release rates (N, P and K) in soil cores fromSphagnum peat andCarex peat saturated with different media, that were artificially created to mimic the three basic water sources: polluted river water, unpolluted calcium-rich groundwater and rainwater. In addition, we studied the effect of temperature and water level on nutrient release rates. The experiments proved thatSphagnum peat released much more P and ammonium thanCarex peat. The strong site effect proved consistent throughout the water chemistry treatments, which indicates that soil quality may be the most important agent determining nutrient release rates. Nevertheless, it was established that water chemistry and water level are of significant influence on nutrient release rates in peat soils. In particular, river water stimulated P release by the peat, most notably in theSphagnum peat. P-release in both soils was only minor when the soils were incubated in clean Ca-rich groundwater. It is suggested that P release is strongly associated with soil chemical processes, and that high P release rates after incubation in river water are due to the high sulphate content of the water. The net release from the soil of ammonium, potassium and phosphate increased with increasing temperature. A freezing treatment significantly increased nutrient availability. The results of the experiments are examined in the context of hydrologic management strategies for the conservation of fens in agricultural landscapes.     

Delineating landscape-scale processes of hydrology and plant dispersal for species-rich fen conservation: the Operational Landscape Unit approach

Restoration and conservation of species-rich nature reserves requires inclusion of landscape-scale connections and transport processes such as hydrologic flows and species dispersal. These are important because they provide suitable habitat conditions and an adequate species pool. This study aimed at identifying the key hydrologic flows and plant dispersal processes affecting a landscape with species-rich fen reserves where restoration measures are carried out to set back succession. It also intended to use this information for delineating the area relevant for conservation planning on an Operational Landscape Unit map. The study was carried out for complexes of fen ponds in former turbaries in the Vechtplassen area, The Netherlands. A number of recent insights on plant dispersal were integrated with knowledge on hydrologic flows in the present approach. The results showed that groundwater discharge to ensure mesotrophic, base-rich conditions, should be enhanced by restoring the groundwater recharge areas NE of the reserves. A nearby lake with suitable water chemistry was also identified as a key source of surface water to feed the fens in dry periods. Water dispersal was identified as important within the fen reserves, whereas dispersal by daily migrating dabbling ducks, typically occurring over 2–3 km, was the most important route connecting the reserves with the surrounding landscape. The delineation of the Operational Landscape Unit for this region provides a basis for conservation and restoration that take fundamental landscape connections and transport processes into account. This unique approach simultaneously considers hydrological transport processes as well as species dispersal in the larger landscape beyond the reserves themselves and therefore leads to greater success of restoration and conservation.     

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.     

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     

Assessing Restoration Perspectives of Disturbed Brook Valleys: The Gorecht Area, The Netherlands

Several landscape restoration alternatives were evaluated in the Gorecht area, a cultivated river plain in the northern part of the Netherlands. A landscape analysis was performed to investigate the hydrological functioning of this area. Groundwater composition in the area was assessed by using distribution patterns of indicative plant species. Results proved to be consistent with an interpolation of actual data of the groundwater composition. Groundwater flow was simulated with hydrological models to explain the observed patterns in water chemistry. It appeared that upwelling Ca-rich groundwater is now absent in the area, contrary to the past situation. Because a constant supply of Ca-rich water is an essential condition for mesotraphent fen vegetation, we concluded that under the present conditions the regeneration prospects for these vegetation types are poor. We suggest that the plan to regenerate groundwater-fed fens be abandoned for a plan to create surface-water-fed marshes.     

Denitrification and dinitrogen fixation in two quaking fens in the Vechtplassen area,The Netherlands

This paper reports laboratory experiments on dinitrogen fixation and denitrification for two small quaking fens (discharge fen and recharge fen) using the acetylene reduction assay and the acetylene inhibition technique, respectively.Nitrogenase activity was detected in peat muck and associated with Alnus glutinosa saplings throughout the study period (May–October 1987), whereas no activity was observed with Sphagnum species. The annual amount of dinitrogen fixed was estimated at 2.1 and 12.7 kg N/ha/y for the recharge fen and the discharge fen, respectively.Denitrification at ambient nitrate levels (0.1 ppm NO₃) was absent in the discharge fen and very low in the recharge fen (0.1 g N/g/d, or 0.3 kg N/ha/y). In nitrate-amended soil samples denitrification rates were 2 to 3 orders of magnitude higher. It is argued that in situ denitrification rates in the fens studied will depend almost entirely on the nitrate supply by precipitation. Denitrification rates associated with precipitation are estimated at 1.1 kg N/ha/y for both fens.     

Understanding the Groundwater Hydrology of a Geographically-Isolated Prairie Fen: Implications for Conservation

The sources of water and corresponding delivery mechanisms to groundwater-fed fens are not well understood due to the multi-scale geo-morphologic variability of the glacial landscape in which they occur. This lack of understanding limits the ability to effectively conserve these systems and the ecosystem services they provide, including biodiversity and water provisioning. While fens tend to occur in clusters around regional groundwater mounds, Ives Road Fen in southern Michigan is an example of a geographically-isolated fen. In this paper, we apply a multi-scale groundwater modeling approach to understand the groundwater sources for Ives Road fen. We apply Transition Probability geo-statistics on more than 3000 well logs from a state-wide water well database to characterize the complex geology using conditional simulations. We subsequently implement a 3-dimensional reverse particle tracking to delineate groundwater contribution areas to the fen. The fen receives water from multiple sources: local recharge, regional recharge from an extensive till plain, a regional groundwater mound, and a nearby pond. The regional sources deliver water through a tortuous, 3-dimensional “pipeline” consisting of a confined aquifer lying beneath an extensive clay layer. Water in this pipeline reaches the fen by upwelling through openings in the clay layer. The pipeline connects the geographically-isolated fen to the same regional mound that provides water to other fen clusters in southern Michigan. The major implication of these findings is that fen conservation efforts must be expanded from focusing on individual fens and their immediate surroundings, to studying the much larger and inter-connected hydrologic network that sustains multiple fens.     

Measuring the efficiency of fen restoration on carabid beetles and vascular plants: a case study from north‐eastern Germany

The aim of the study was to assess the effects of fen rewetting on carabid beetle and vascular plant assemblages within riverine fens along the river Peene in north‐eastern Germany. Drained (silage grassland), rewetted (restored formerly drained silage grassland), and near‐natural (fairly pristine) stands were compared. Eighty‐four beetle species (7,267 individuals) and 135 plant species were recorded. The richness of vascular plant species and the number of endangered species were highest on near‐natural fens. Fourteen years of rewetting did not increase plant species numbers compared with drained fens. For carabid beetles, however, species richness and the number of stenotopic species were highest on rewetted fens. Rewetting caused the replacement of generalist carabids by wetland specialists, but did not provide suitable habitat for specialist fen carabids or for plant species of oligo‐ or mesotrophic fen communities. Therefore, raising the water table on fens with nutrient‐rich, degraded peat was not sufficient for restoring species assemblages of intact fens, although water level was the most important environmental factor separating species assemblages. Our study illustrated that insects and plants may respond differentially to restoration, stressing the need to consider different taxa when assessing the efficiency of fen restoration. Furthermore, species assemblages of intact fens could not be restored within 14 years, highlighting the importance of conserving pristine habitat.     

Persistence of high elevation fens in the Southern Rocky Mountains,on Grand Mesa,Colorado, U.S.A.

Small headwater fens at high elevations exist in the dry climatic regime of western Colorado, despite increasing demands for water development since the 1800’s. Fens on Grand Mesa have accumulated plant material as peat for thousands of years due to cold temperatures and consistently saturated soils. The peatlands maintain unique plant communities, wildlife habitat, biodiversity, and carbon storage. We located and differentiated 88 fens from 15 wet meadows and 2 marshes on Grand Mesa. Field work included determining vegetation, soils, moisture regimes, and impacts from human activities. All fens were groundwater-supported systems that occurred in depressions and slopes within sedimentary landslide and volcanic glacial till landscapes. Fens occupied 400 ha or less than 1 % of the 46,845 ha research area and ranged in size from 1 to 46 ha. Peat water pH in undisturbed sites ranged from 4.3 to 7.1. Most fens had plant communities dominated by sedges (Carex) with an understory of brown mosses. Variation in vegetation was controlled by stand wetness, water table level, organic C, conductivity (EC), and temperature °C. Fen soils ranged from 13.6 to 44.1 % organic C with a mean of 30.3 %. Species diversity in fens was restricted by cold short growing seasons, stressful anaerobic conditions, and disturbance. Multivariate analysis was used to analyze relationships between vegetation, environmental, and impact variables. Stand wetness, water table level, OC, electrical conductivity (EC), and temperature were used to analyze vegetation variance in undisturbed fens, wet meadows, and marshes. Vegetation composition in impacted fens was influenced by flooding, sedimentation, stand wetness, water table level, OC, EC, and temperature. Hydrologically modified fens supported 58 plant species compared to 101 species in undisturbed fens. Analysis of historical 1936–2007 aerial photographs and condition scalars helped quantify impacts of human activities in fens as well as vegetation changes. Fourteen fens had evidence of peat subsidence, from organic soil collapse, blocks of peat in the margins, soil instability, and differences in surface peat height between the fen soil surface and the annually flooded soil surface. Of 374 ha of fens in the Grand Mesa study area, 294 ha (79 %) have been impacted by human activities such as ditching, drainage, flooding, or vehicular rutting. Many fens had little restoration potential due to severe hydrological and peat mass impacts, water rights, or the cost of restoration.     

Flooding and groundwater dynamics in fens in eastern Poland

Abstract. Dynamics in hydrology and water chemistry in the Biebrza mires (Poland) were examined by means of a sampling survey that was repeated four times between 1987 and 1992. The dynamics in the vertical stratigraphy of water types in the peat profile are considerable from close to the mire surface to a depth of 50 cm. Water composition in the root zone correlated best with vegetation types during extremely dry or wet conditions. In the root zone of groundwater-fed rich fens with Caricetum limoso-diandrae and Calamagrostietum strictae vegetation, specific groundwater types evolve from the interaction of discharging groundwater from below the root zone and the temporal influence of precipitation and evapotranspiration. The Caricetum limoso-diandrae is fed by the continuous discharge of nutrient-poor, relatively mineral-rich water. The site conditions in the Calamagrostietum strictae are determined by occasional flooding and the presence of discharging mineral-poor groundwater in the lower part of the root zone. In the Caricetum limoso-diandrae and the Calamagrostietum strictae the maximum variations in water level were 56 and 86 cm, respectively. The composition of shallow groundwater of the Betuletum humilis/Caricetum rostratodiandrae fen is diluted most compared to other vegetation types by rainwater in wet periods. In periods of prolonged drought it has a water type that is affected by evapotranspiration and peat mineralisation. The water level varies by only 33 cm. In the Magnocaricion and Glycerietum maximae in the floodplain the water composition is determined by spring flooding of the river and the natural draw-down that occurs in the following summer. Here, maximum variations in water level were 108 and 117 cm, respectively.     

Nutrient dynamics in small mesotrophic fens surrounded by cultivated land

Summary Release of inorganic N and P in the organic soils of three small quaking fens in The Netherlands was studied by means of in situ incubation of the peat soil in plastic bottles. One of the fens had higher biomass production and lower species richness than the other two. The former fen is located in an area with downward groundwater percolation, whereas the latter fens are situated in an area of upward seepage of groundwater rich in calcium and bicarbonate.Mineralization of N proved to be slow in the low-productive fens, and 6 times faster in the highly productive fen. In the latter fen the amount of N mineralized during a certain period exceeded by far the amount accumulated in that period in the above-ground parts of the vegetation.The release of inorganic P was extremely slow in the two low-productive fens and rapid in the highly productive fen. The release rates were not related to the total P content of the soils. The slow P release in the low-productive fens may be due to the continuous inflow of groundwater rich in Ca, Al and Fe compounds.It can be concluded that the soil is a potentially large source of inorganic N and P in quaking fens. The importance of the soil versus other N and P sources is greater in nutrient-rich fens.     

Species richness and composition patterns of clitellate (Annelida) assemblages in the treeless spring fens: the effect of water chemistry and substrate

Spring fens are isolated treeless wetlands of a high conservation value. Their environmental conditions are strongly related to their groundwater chemistry, which controls species distribution within various groups of organisms. Clitellates, a dominant group of non-insect aquatic fauna, however, have never been studied in these habitats. It is unclear from previous studies to what extent the distribution of aquatic non-insect taxa reflects water chemistry rather than the substrate structure. We studied 34 spring fens sampled in 17 isolated sites in the Western Carpathian Mountains to determine mainly the effects of water chemistry and substrate structure on variation in species richness and composition of clitellate assemblages as examples of the non-insect fauna. A total of 34 taxa were found, with 3–15 taxa collected per sample. Species richness was negatively correlated with water mineral concentration measured as water electric conductivity (r = −0.57, P < 0.001) and positively with TOC (r = 0.60, P < 0.001). Surprisingly, the lowest number of taxa was found in calcareous fens and richness increased towards Sphagnum-fens. There was a species turnover related to changes in mineral richness and substrate characters. The main change of species composition was promoted by changes in substrate structure. The second gradient of species composition was linked with the amount of nutrients, moisture, and dominance of sphagna, and was associated with an increase of eurytopic species in fens with high nutrient availability. It was difficult to separate the effects of water chemistry and substrate on clitellate species distributions owing to the fact that variation in tufa precipitation and vegetation was driven by water chemistry changes. This study presented the first quantitative data on fen clitellate assemblages, which appear to have an unusual pattern of species richness. In contrast to plants and molluscs, calcareous fens appeared to be a harsh environment for clitellate species. Only few specialized species, mainly Trichodrilus strandi, were able to establish viable populations. The significant effect of water chemistry on clitellate distribution patterns raises questions about the direct influence of water chemistry on non-insect aquatic taxa, which have previously been considered to be mostly determined by substrate characteristics.     

Peat and water chemistry at Big Run Bog,a peatland in the Appalachian mountains of West Virginia,USA

At Big Run Bog, aSphagnum-dominated peatland in the unglaciated Appalachian Plateau of West Virginia, significant spatial variation in the physical and chemical properties of the peat and in surface and subsurface (30 cm deep) water chemistry was characterized. The top 40 cm of organic peat at Big Run Bog had average values for bulk density of 0.09 g · cm^–3, organic matter concentration of 77%, and volumetric water content of 88%. Changes in physical and chemical properties within the peat column as a function of depth contributed to different patterns of seasonal variation in the chemistry of surface and subsurface waters. Seasonal variation in water chemistry was related to temporal changes in plant uptake, organic matter decomposition and element mineralization, and to varying redox conditions associated with fluctuating water table levels. On the average, total Ca, Mg, and N concentrations in Big Run Bog peat were 33, 15, and 1050 mol · g^–1, respectively; exchangeable Ca and Mg concentrations were 45 and 14 eq · g^–1 , respectively. Surface water pH averaged 4.0 and Ca⁺⁺ concentrations were less than 50 eq · L^–1 . These chemical variables have all been used to distinguish bogs from fens. Physiographically, Big Run Bog is a minerotrophic fen because it receives inputs of water from the surrounding forested upland areas of its watershed. However, chemically, Big Run Bog is more similar to true ombrotrophic bogs than to minerotrophic fens.     

Construction of fens with and without hydric soils

The objective of this study was to show that temperate zone fens could be constructed on a surface of glacial gravel or a surface without preformed hydric soil. A reference study examined the construction of a fen on remnants of a buried fen. Each construct was replicated in four sections planted with seeds, greenhouse-grown stock or natural fen plant communities. An artesian well supplied high carbonate water through the subsurface. After 8 years, each fen contained 71 species in common, the gravel-based fen had 28 additional species and the soil-based fen had 21 additional species. On the third year, the gravel-based fen began producing calcareous peat composed of brown mosses and by the eighth year a 10 cm layer of peat covered most of the surface. The gravel-based fen produced short and less densely distributed plants than the soil-based fen but the gravel-based fen had a greater number of rare plants. While most plantings had a high degree of survival the naturally derived plugs were the most successful and the seed produced the fewest survivors. We show that, given continuous groundwater supply, it is possible to produce a high diversity fen-like environment on either substrate.