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.     

Wetland dicots and monocots differ in colonization by arbuscular mycorrhizal fungi and dark septate endophytes

As an initial step towards evaluating whether mycorrhizas influence composition and diversity in calcareous fen plant communities, we surveyed root colonization by arbuscular mycorrhizal fungi (AMF) and dark septate endophytic fungi (DSE) in 67 plant species in three different fens in central New York State (USA). We found colonization by AMF and DSE in most plant species at all three sites, with the type and extent of colonization differing between monocots and dicots. On average, AMF colonization was higher in dicots (58±3%, mean±SE) than in monocots (13±4%) but DSE colonization followed the opposite trend (24±3% in monocots and 9±1% in dicots). In sedges and cattails, two monocot families that are often abundant in fens and other wetlands, AMF colonization was usually very low (<10%) in five species and completely absent in seven others. However, DSE colonization in these species was frequently observed. Responses of wetland plants to AMF and DSE are poorly understood, but in the fen communities surveyed, dicots appear to be in a better position to respond to AMF than many of these more abundant monocots (e.g., sedges and cattails). In contrast, these monocots may be more likely to respond to DSE. Future work directed towards understanding the response of these wetland plants to AMF and DSE should provide insight into the roles these fungal symbionts play in influencing diversity in fen plant communities.     

Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.     

Loss of habitat specialists despite conservation management in fen remnants 1995–2006

Many ecosystems of high conservation value have been shaped by human impacts over centuries. Today, traditional management of semi-natural habitats is a common conservation measure in Europe. However, despite traditional management, habitat remnants may still loose specialist species due to surrounding land-use change or atmospheric nitrogen deposition. To detect trends in species density (2-m² plot scale) and habitat quality in calcareous fens in the pre-Alps of Switzerland, we surveyed 36 traditionally managed fens in 1995/97 and again in 2005/06 (five plots per fen). The fens occurred at three altitudinal levels (800–1000, 1000–1200, 1200–1400 m asl) and were either extensively grazed or mown once a year. Despite these traditional management regimes, species density of fen specialists and of all bryophytes decreased during this decade (vascular plant specialists: ?9.4%, bryophyte specialists: ?14.9%, all bryophytes: ?5.7%). Management had no effect on the number of Red-List species and habitat specialists of vascular plants per plot. However, bryophyte species density was more strongly reduced in grazed fens. Species density of vascular plant generalists increased between the two surveys (+8.2%) but not of bryophytes. Among vascular plants, Red-List species decreased from 1.01 to 0.78 species per plot. Furthermore, between the two surveys aboveground plant biomass, mean plant-community indicator values for nutrients and species density of nutrient indicators increased, whereas mean plant indicator values for soil moisture, light and peat, and species density for peat indicators, decreased. We attribute these changes and the loss of specialist species over the past decade mainly to land-use change in the surrounding area and to nutrient inputs. Thus, despite traditional management, calcareous fens in the pre-Alps suffer from ongoing habitat deterioration and endangered plant species remain threatened. For their long-term protection, we suggest to reduce nutrient inputs and, where necessary, to restore hydrology and adjust grazing management.     

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.     

Vegetation along hydrologic, edaphic, and geochemical gradients in a high-elevation poor fen in Canaan Valley, West Virginia

Wetland plant community composition and pattern are regulated by a host of abiotic/environmental gradients and biotic factors. We used multivariate analyses to classify wetland plant communities and determine the relation of hydrologic, edaphic, and geochemical gradients on community composition and spatial distribution among 18 vegetation, hydrology, and soil sampling points in Abe Run, a botanically unique poor fen in northeastern West Virginia. We also examined the interactions of disturbance with the physical environment and species composition. A total of 179 vascular plant species were identified from sample plots. Vegetative composition and structure, dominant soil texture, and hydrology lacked the distinct concentric zonation of northern peatlands; instead, all were patchy and varied considerably over short distances. Graminoid-forb meadows with primarily silt-loam mineral horizons, greater depth to groundwater, and fewer days of inundation characterized the lower reaches of the wetland. These plots were more acidic, with absent or shallow O horizons, and lower concentrations of soil base cations (Ca, Mg, K). In the upper reaches of the wetland, mixed herb-shrub-tree dominated communities were structurally and compositionally more complex; here, organic horizons were much more prominent, peat depth ranged from 80 to 100 cm, and the average depth to water table was 10 cm less than for wells in the lower reaches of the wetland. Plots from upper transects (3–6) tended to have more shrub and tree cover, and higher concentrations of soil base cations. Much of the variability among plots in the upper and lower reaches of the wetland are consistent with beaver inundation of a large portion of the wetland during the 1980s. Multiple-response permutation procedures verified the difference (P < 0.0001) between vegetation of plots in the lower and upper reaches of the watershed. Because fens are connected to groundwater, these habitats are particularly vulnerable to disturbances, particularly those that alter existing land use and land cover. Minimization of disturbances in the surrounding watershed, controls on an excessive deer population, removal of exotic, non-native species, and control on foot traffic are all integral to maintaining the integrity of this high-value wetland.     

Exotic plant invasion in the highly degraded upland landscape of Hong Kong, China

Plant communities in the continental tropics have suffered less from exotic plant invasions than their oceanic island counterparts. Most studies have focused on near-pristine communities. By contrast, we examine the resistance of semi-natural continental plant communities in Hong Kong, which have been suffering from chronic and massive human impacts. We compiled a list of all naturalized non-native species recorded in Hong Kong and then sampled the plant communities for exotic species along roadsides, a stream through semi-natural vegetation, and in semi-natural vegetation away from both roads and streams on Tai Mo Shan, Hong Kong’s highest peak (957 m). Similar surveys were repeated in other areas of Hong Kong. More than 162 naturalized exotic plant species have been recorded in Hong Kong. On Tai Mo Shan, 29 exotic species were recorded in roadside vegetation, with the diversity but not percentage cover declining significantly with altitude. Fifteen exotic species were found along the stream, including two not found along the roadside. Only six exotic species were found away from roads and streams, all in unshaded areas disturbed by feral cattle. In all surveys, no exotics were found in closed woody vegetation or in open areas without feral cattle, except for one species. The shade-tolerant tropical Asian tree Syzygium jambos was found invading along some streamsides without anthropogenic disturbance. Despite centuries of massive human impacts, exotic plant invasions in Hong Kong are still largely confined to habitats that suffer from chronic human disturbance. Feral cattle promote invasion where people are absent, but this problem still seems potentially reversible. Only Syzygium jambos is of possible current conservation concern.     

An ecosystems approach to base-rich freshwater wetlands,with special reference to fenlands

A survey of base-rich wetlands in The Netherlands is presented. The main area of their occurrence is the low-lying Holocene part of the country, until some thousand years ago a large and coherent wetland landscape: the Holland wetland. The development of various parts of the Holland wetland into marshes, fens and bogs can be understood from hydrological relations in mire basins, as recognized in the distinction of primary, secondary and tertiary mire basin stages. Presently, the remnants of the Holland wetland are separate base-rich wetlands. The succession of their vegetation reflects various abiotic conditions and human influences. Three main developmental periods are distinguished as regards these factors. The first, geological period of mire development is seen as a post-glacial relaxation, with the inertia due to the considerable mass of wetland as a stabilizing factor. Biological “grazing” influences, as an aspect of utilization by humans, converted base-rich wetlands to whole new types in the second, historical period. Presently, mass and harvesting have decreased in importance, and actual successions in terrestrializing turbaries seem to reflect rapidly changing environmental conditions. Human control could well become the most important factor in the future development of wetland nature. The present value of open fen vegetation strongly depends on the continuation of the historical harvesting. The development of wooded fen may help to increase the mass of wetland in the future. Best results in terms of biodiversity are expected when their base state is maintained through water management. The vegetation and hydrology of floating fens in terrestrializing turbaries is treated in some more detail. Various lines and phases in the succession are distinguished. Open fen vegetation at base-rich, yet nutrient-poor sites is very rich in species threatened elsewhere. The fast acidification of certain such fens is attributed to hydrological and management factors. This acidification is illustrated in the profile of a floating raft sample. At the scale of these small fens, the elemental structure comprising base-rich fen, transitional fen and bog vegetation, is not as stable as it was in the large Holland wetland. A critical role seems to be played by the supply of bases with the water influx. The changing base state is supposed to change the nutrient cycling to such an extent that it would be correct to call this trophic excitation of the ecosystem, rather than just eutrophication. Eutrophication indicates a quantitative reaction to an increased nutrient supply, the internal system being unaltered. The drainage of fens, resulting in an increased productivity of the vegetation, provides another example of excitation, to the effect that the functional system is dramatically changed internally.     

Naturalization of exotic plant species in north‐eastern North America: trends and detection capacity

Aim To reconstruct historical naturalization trends of exotic vascular plants in Quebec and Massachusetts, two regions that experienced fluctuations in their herbarium specimen collection efforts. We hypothesized that the observed trends are linked to the collection effort and that they differ according to the motive for plant introduction. Methods A checklist of naturalized exotic plant taxa for Quebec, including oldest proof of naturalization and motive for introduction, was built from herbaria, journals, dissertations, theses, bulletins, websites and unpublished records. For Massachusetts, we used the checklist of Sorrie (2005; Rhodora, 107 , 284–329). The collection effort was estimated from a pool of species with specimens from the five largest Quebec herbaria and from the Harvard University Herbarium. Results Naturalization trends of exotic plants in Quebec are similar to those observed in Massachusetts. A large part of the plants naturalized in the 17th, 18th or 19th centuries, with very little naturalization during the last 30–90 years. In general, there is a significant statistical link between herbarium specimen collection effort and the discovery of new naturalized plant taxa. Very few utilitarian plants naturalized in either region during the 20th century. In Quebec, a high number of ornamental plants naturalized during the last 100 years. Main conclusions The link between herbarium specimen collection effort and the discovery of new naturalized plants is real, although not as straightforward as it seems. Our analysis suggests that at least part of the decline in the number of new naturalized exotic plants observed in Quebec and Massachusetts during the last 30–90 years is a direct consequence of the low interest for traditional floristic studies. However, the possibility of a real decline cannot be ruled out. We nevertheless provide here one of the first pieces of evidence of the potential consequences of the decline of local plant collection for environmental management and especially for early detection systems of new invaders.     

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.     

Oribatid mites (Acari,Oribatei) of three fens in the northern part of European Russia

Oribatids from three oxbow fens in Kostroma, Kirov, and Arkhangelsk provinces were studied. A total of 2800 adult mites belonging to 74 species were collected. The similarity of the faunas and communities in these fens is rather low, the Jaccard and Naumov coefficients constituting 22–37% and 11–22%, respectively. From the central part of each fen to its periphery, the diversity of oribatids increased from 4–6 to 25–28 species, and abundance, from 59–123 to 451-737 ind./l. Some oribatid species demonstrated similar preferences within all the fens studied. The oribatid faunas of fens and raised bogs are compared, and their characteristic species are distinguished.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Experimental test of the Invasional Meltdown Hypothesis: an exotic herbivore facilitates an exotic plant,but the plant does not reciprocally facilitate the herbivore

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Effects of topsoil removal,seed transfer with plant material and moderate grazing on restoration of riparian fen grasslands

Question: How do moderate grazing, topsoil removal and hay transfer affect species diversity and abundance on a eutrophic fen grassland site? Location: Northern Germany. Method: A three-factorial field experiment with the factors grazing, topsoil removal and hay transfer of diaspore-rich material was established in 2001. Soil nutrients and seed bank were analysed at the beginning of the experiment, species composition and vegetation development was monitored for four years (2002–2005). Results: Topsoil removal had a significant effect on the abundance of different plant species groups: resident vegetation of agricultural grasslands was suppressed, while clonal reed species were facilitated in recolonising the area. The establishment of regionally rare and endangered species of nutrient-poor fens and wet meadows introduced with hay was achieved mainly on plots with topsoil removal, with the exception of Rhinan-thus angustifolius, which also established on plots with intact topsoil. Effects of grazing after four years of experiments were of minor influence on species composition. Conclusion: The establishment of target plant species of nutrient-poor fens is most successful when both an adequate number of viable diaspores and suitable sites for germination and establishment are available. In our experiment this was achieved by the combination of topsoil removal and hay transfer. We recommend this combination, together with continuous management (grazing/cutting), for further restoration in fen grasslands.     

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.     

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.     

Novel chemistry of invasive exotic plants

Of the many exotic plants that have become naturalized in North America, only a small proportion are pests capable of invading and dominating intact natural communities. In the present study, we tested the hypothesis that the most invasive plants are phytochemically unique in their new habitats. A comparison of exotic plant species that are highly invasive in North America with exotics that are widespread, but non-invasive revealed that the invasive plants were more likely to have potent secondary compounds that have not been reported from North American native plants. On average, the compounds found in the invasive plants were reported from fewer species, fewer genera and fewer families than those from non-invasive plants. Many of the unique phytochemicals from invasive plants have been reported to have multiple activities, including antiherbivore, antifungal, antimicrobial and allelopathic (phytotoxic) effects, which may provide the plants with several advantages in their new environments.     

Salvaged-Wetland Soil as a Technique to Improve Aquatic Vegetation at Created Wetlands in Wyoming,USA

Aquatic plants usually establish following wetland creation from a variety of mechanisms including animal transport, inflows from nearby wetlands, wind dispersal, and seed banks if they are available. However, at created wetlands that are isolated from natural wetlands, aquatic plant communities may not establish even after 10 or more years. One method of improving the establishment of aquatic plants is through the use of salvaged-marsh soils. Using this method, wetland soil from a donor site is collected and spread across the basin of the created wetland. When the proper hydrologic regime is reached at the created site, the seed bank from the donor soil is then present to take advantage of the uncolonized site. Over 1500 wetlands have been created in northeast Wyoming, USA from bentonite mining and most of them have not developed submersed and emergent plant communities due to isolation from plant sources. Our goal was to evaluate the effectiveness of using salvaged-wetland soil as a tool for improving plant growth at created wetlands. Our study took place at 12 newly created wetlands that were isolated from other wetlands by >5 km. Six wetlands were treated as reference wetlands, with no introductions of seeds or propagules. At the other six wetlands we spread ≈10–15 cm of salvaged soil from a donor wetland during the winter of 1999–2000. To identify the potential plants in donor soil, we collected 10 random samples from the donor wetlands and placed them within wetland microcosms in a greenhouse where they were treated to either moist-soil conditions (water at or just below the soil line) or submersed conditions (water levels maintained at 15–30 cm). Treatment wetlands were evaluated for plant growth during the fall of 2000 and 2001, whereas the greenhouse samples were grown for two growing seasons then harvested. Our results show that using salvaged wetland soil increases: (1) the number of plant species present at a wetland over time, (2) the total vegetation coverage in a treated wetland over time, and (3) the total plant biomass in a treated wetland. The species pool available in the salvaged wetland soil was limited to 10 obligate wetland species, but several of them are considered valuable to waterfowl and other wildlife. Furthermore, salvaged-wetland soil could be useful for ameliorating poor substrate conditions (i.e., bentonite) and improving conditions for the establishment of additional species. One concern with this technique is the introduction of invasive or exotic species that could form monocultures of undesirable plants (e.g., cattail [Typha spp.]); introducing more desirable species during the application of salvaged soil could reduce this probability. We believe incorporating salvaged-wetland soil during basin construction could be used to increase the value and productivity of created wetlands in this region.     

Diversity of wetland vegetation in the Bulgarian high mountains, main gradients and context-dependence of the pH role

We fill a gap in understanding wetland vegetation diversity and relationship with environmental determinants in Bulgarian high mountains. A total of 615 phytosociological samples were taken from springs, mires, wet meadows and tall-forb habitats throughout Bulgaria, of which 234 relevés are from mire and spring vegetation above timberline. The vegetation was classified by TWINSPAN and the resulting vegetation types were reproduced by the formal definitions using the combination of Cocktail species groups based on phi-coefficient of joint co-occurrence of the species. Nine vegetation types of springs and fens have been clearly delimited above the timberline. All vegetation types include Balkan endemic species, the representation of which varies. Fens generally harbour more Balkan endemics than do springs, with the exception of species-poor high-altitude Drepanocladetum exannulati. The gradient structure of the vegetation was revealed by DCA and by CCA with forward selection of environmental factors. The major determinants of vegetation variation strongly differ above and below the timberline and likewise between springs and fens. The base-richness gradient controls the floristic variation of Bulgarian submontane fens, whereas the complete data set including both submontane and subalpine fens is governed by the altitude gradient from lowland and basin fens to subalpine fens rich in Balkan endemics. When focusing on sites above the timberline only, the first DCA axis separates fens from springs without organic matter. The major species turnover in springs follows the variation in water pH and mineral content in water, whereas fen vegetation variation is primarily controlled by succession gradient of peat accumulation. Altitude remains an important factor in all cases. Weak correlation between water pH and conductivity was found. This correlation was even statistically insignificant in fens above the timberline. Water pH is not influenced by mineral richness in Bulgarian high mountains, since it is buffered by decomposition of organic matter in fens. In springs, pH reaches maximum values due to strong aeration caused by water flow. The plant species richness decreases significantly with increasing altitude. The increase of species richness towards circumneutral pH, often found in mires, was not confirmed in Bulgarian high mountains. The correlation between species richness and pH was significant only when arctic-alpine species and allied European high-mountain species were considered separately. The richness of boreal species was independent on pH. Some of them had their optima shifted to more acidic fens as compared to regions below the timberline. Our results suggest that subalpine spring and fen vegetation should be analysed separately with respect to vegetation-environment correlations. Separate analysis of fens below and above timberline is quite appropriate.     

Shrub removal facilitates recovery of wetland species in a rewetted fen

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