Restoration of peat‐forming vegetation by rewetting species‐poor fen grasslands

Questions: Does succession of rewetted species‐poor fen grasslands display similar trends when different water levels, sites and regions are compared? Will restoration targets as peat growth and waterfowl diversity be reached? Location: Valley fen of the river Peene (NE‐Germany) and the Hanság fen (Lake Neusiedler See, NW‐Hungary). Methods: Analysis of permanent plot data and vegetation maps over a period of up to seven years of rewetting. The general relations between newly adjusted water levels and changes in dominance of helophytic key species during early succession are analysed considering four rewetting intensities (water level classes) and eight vegetation types (Phalaris arundinacea type, Carex type, Glyceria maxima type, Phragmites australis type, Typha type, aquatic vegetation type, open water type and miscellaneous type). Results: The initial period of balancing the site conditions and vegetation is characterised by specific vegetation types and related horizontal vegetation structures. Most vegetation types displayed similar trends within the same water level class when different sites and regions were compared. A significant spread of potentially peat forming vegetation with dominance of Carex spp. or Phragmites as desired goal of restoration was predominantly restricted to long‐term shallow inundated sites (water level median in winter: 0–30 cm above surface). Open water patches as bird habitats persisted mainly at permanent inundated sites (water level median in winter > 60 cm above surface). Conclusions: Site hydrology appeared as a main force of secondary succession. Thus the rewetting intensity and restoration targets have to be balanced adequately.     

Quick Target Vegetation Recovery after Restorative Shrub Removal and Mowing in a Calcareous Fen

Removal of shrubs and trees is an important management and restoration practice to promote openness and light‐dependent vegetation in fens, especially as tree cover is increasing in previously open wetlands. The effects of woody vegetation removal on target species have been poorly documented in wetlands up to now. In this study, I investigated the effect of tree and shrub removal (especially of Juniperus communis) on the target vegetation in a partly overgrown and degraded grazed rich fen after 6 years. I also tested whether additional intensified management by mowing could promote initial recovery. Shrub removal resulted in a rapid recovery of species‐rich fen vegetation such that after 6 years brown moss cover more than tripled and target species richness doubled and became similar to values of a reference area in a favorable conservation status. Additional mowing resulted in a much higher abundance of the target rich fen vascular plants. The reasons for the success at this site may be the proximity to well‐developed rich fen vegetation, presence of cattle that dispersed diaspores, and presence of bare, colonizable substrate. Thus, it may be more beneficial to restore and expand already existing sites in a partly favorable status than to restore severely deteriorated sites. Extensive management by woody vegetation removal may be an alternative method to maintain high conservation values of open mires and other wetlands, where grazing or mowing is not necessary or feasible to meet future needs in response to overgrowth caused by global warming.     

Mollusc diversity patterns in Central European fens: hotspots and conservation priorities

Aim To assess mollusc species composition and diversity patterns of treeless fen sites and to find simple environmental parameters that characterize diversity hotspots and priority sites for conservation. Location Western Carpathian Mountains, Europe. Methods Mollusc communities were sampled quantitatively from a homogeneous area of 16 m² in the central part of each of 145 treeless fen sites. Water conductivity and pH, geographical coordinates, altitude and habitat size of the sites studied, mean annual rainfall, mean annual temperature and mean January temperature were compiled for each plot. Nestedness in species composition was tested using the binmatnest program to confirm the ‘nested habitat‐quality hypothesis’. Patterns in species diversity were analysed using the regression trees method to isolate the main predictors of species diversity. Results Nested subset patterns of species composition were found along the gradient of mineral richness. Species distribution was highly nested (T_obs = 11.21, P << 0.001) in mineral‐poor sites (with water conductivity < 300 μS cm^?1, n = 42) and was highly correlated with the site’s mineral richness (r_s = 0.76, P << 0.001). By contrast, species distribution and richness of mineral‐rich sites (c.≥ 300 μS cm^?1, n = 103) were not controlled by mineral richness. Variation in species richness was further explained by January temperature, landscape geomorphology, and total habitat area. The southern mineral‐rich low altitude fens were the most species rich, especially those of larger total area (23 species on average). These 24 sites (17% of all sites) harboured 90% of all recorded species, including all highly endangered ones. Mineral‐rich fens in montane valleys were the second most important group because they hosted the majority of populations of two rare glacial relict species (Vertigo geyeri and Pupilla alpicola). Main conclusions The significant nestedness raises the possibility of conserving the whole fen‐mollusc species pool within the most species‐rich sites. Thus, to select the conservation priority sites, easily available site characteristics for the prediction of species richness are needed. This knowledge can help us maintain fen biodiversity, which has become closely dependent on conservation management practices after the cessation of traditional mowing of fens for haymaking.     

Restoration of a Landfill Site in Berlin, Germany by Spontaneous and Directed Succession

We report on spontaneous and directed succession on a dry sandy landfill site of low fertility at Berlin‐Malchow, Germany. Changes in species composition and cover were followed on unmown and mown permanent plots of 2 × 2 m size through 5 years of vegetation development. Species richness on unmown plots was relatively constant during the time of observation, with 20 to 25 species per 4 m². Total cover of unmown plots continuously increased from approximately 10% in the first year to 80% in the fifth year. There are no clearly discernible sequential successional stages until present. The species composition includes species of all life forms, which colonized the site immediately after the initiation of the succession process representing the initial floristic composition type of vegetation development. However, perennial grasses and herbs gradually increased in cover up to approximately 40%. Woody plants were also present from the first year of succession and increased up to more than 20% cover in the fifth year, forming a shrub layer (>0.5 m) after the second year. Mowing significantly increased species richness, which was evident from the third year onward. This effect was mainly due to the reduction of the tall perennial grass Calamagrostis epigejos. Solidago canadensis and woody species were also significantly affected (lower cover and height), whereas short perennial herbs like Plantago lanceolata and Trifolium repens benefited from mowing.     

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.     

Differential and reversible responses of common fen meadow species to abandonment

Abstract. We studied the effects of abandonment on two common fen plant species. In mown and a chronosequence of abandoned fen meadows spanning 35 yr, we measured fitness traits of the sedge Carex davalliana and the forb Succisa pratensis. Cessation of mowing had little effect on fitness traits and seed production of C. davalliana, but seedling density decreased more than threefold. Population density of S. pratensis decreased with increasing community biomass, but was not affected by the cessation of mowing. However, flowering frequency increased threefold and seed production was 20% higher in fallow meadows. Consequently, seedling density of S. pratensis increased nearly threefold after abandonment. However, these changes were not dependent on the age of the fallow. In a common garden and germination experiment, we found no differences in either species between plants from fallows and mown fen meadows, except for the height of the flowering stalk of S. pratensis. The combined results from the common garden experiment and the field studies indicate that changes in fitness traits observed in fallows were mostly phenotypic and likely to be reversible. If other species react in similar ways, there is a high potential for re‐establishing traditional fen meadow communities from fallows by mowing.     

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.     

Measures to develop a rich-fen wetland landscape with a full range of successional stages

Species-rich plant communities characteristic for succession from mesotrophic open water to fen are very rare in The Netherlands. These vegetation types used to occur in turf ponds in the low lying peatland area, created by peat dredging and filled with water due to seepage of mesotrophic, well-buffered groundwater. One of the goals of the National Nature Policy Plan is to create new opportunities for the initial terrestrialization communities through ecological engineering, e.g., restoration and creation of open water habitats. Restoration of the abiotic conditions in acidified floating fen communities is only possible by a combined measure of removal of the Sphagnum-layer and superficial drainage of surplus rain water. New turfponds have been excavated. This study showed that the abiotic conditions (i.e., water depth and water chemistry) are favorable for the development of aquatic communities characteristic of mesotrophic conditions. The aquatic plant species found in the new ponds also point in this direction, e.g., Chara major and Ch. delicatula are very abundant as are seven Potamogeton species. It is concluded that a constant discharge of groundwater and a good connectivity between the ponds and the existing remnants of plant communities desired in the area are essential for the conservation and development of these species-rich plant communities.     

Mechanisms for Dominance in An Early Successional Old Field by the Invasive Non-Native Lespedeza Cuneata (Dum. Cours.) G. Don

Researchers studying invasive plants often concentrate their efforts on predictive models thought to allow invasive plants to dominate native landscapes. However, if an invasive is already well established then experimental research is necessary to provide the information necessary to effectively manage the species. Prescribing appropriate management strategies without prior experimental research may not only be ineffective but also may squander limited resources or have the unintended consequence of furthering spread. Lespedeza cuneata(Dum. Cours.) G. Don. is a well-established invasive plant of old fields and tall-grass prairie in the US. Managers suspect this species shades-out native plants and this is proposed as its primary mechanism for dominance. Using field experiments we tested probable factors allowing the species to establish itself and, once established, interfere in old field plant communities. We also examined the effects of two common anthropogenic disturbances (mowing and nutrients) on L. cuneata growth and establishment. When L. cuneata was treated (clipping, herbicide and stem pull-back) there was a significant increase in species richness and native species cover. Stem density and canopy cover of L. cuneata increased significantly with mowing frequency but decreased with nutrient input. We suggest that mowing benefits L. cuneata while also hindering woody competition. Results also indicate L. cuneata is less prevalent on nutrient enriched soils than on unamended soil. Lespedeza cuneata appears to suppress native plants by shading them out and it can subsequently take over grassland communities. Since it has a varying response to human induced disturbances and may actually benefit from mowing, land managers should be cautious when utilizing this as a management tool.     

Deterministic processes have limited impacts on foliar fungal endophyte communities along a savanna-forest successional gradient

Patterns and drivers of succession provide insight into the mechanisms that govern community assembly, but remain poorly understood for microbial communities. We assess whether successional trends of trees are mirrored by foliar endophyte communities of three tree species across a deterministic woody successional gradient. Additionally, we test the relative contribution of abiotic predictors, biotic factors, and spatial distance between sites in predicting composition and richness of endophyte communities. Unlike the tree community, endophyte communities showed no consistent evidence of deterministic succession. Host identity was the most important factor structuring endophyte community composition; within hosts, spatial distance from the indigenous forest and between samples was important, while environmental predictors had small and inconsistent effects. Much variation in endophyte composition remained unexplained. In contrast, endophyte richness was well-explained by predictor variables. Host identity was most important in predicting endophyte richness, while the effect of other predictors on richness differed between host species. We conclude that deterministic succession in trees did not result in deterministic succession in endophyte communities; instead community assembly was most strongly influenced by host identity; while within hosts, neutral processes may be more important for endophyte assembly than deterministic factors.     

Opposing patterns of zooplankton diversity and functioning along a natural stress gradient: when the going gets tough,the tough get going

Salinity represents a major structuring factor in aquatic habitats which strongly affects species richness. We studied the relationships among species richness, density and phylogenetic diversity of zooplankton communities along a natural salinity gradient in astatic soda pans in the Carpathian Basin (Hungary, Austria and Serbia). Diversity and density showed opposing trends along the salinity gradient. The most saline habitats had communities of one or two species only, with maximum densities well above 1000 ind l^?1. Similarity of communities increased with salinity, with most of the highly saline communities being dominated by one highly tolerant calanoid copepod, Arctodiaptomus spinosus, which was at the same time the only soda‐water specialist. Salinity obviously constrained species composition and resulted in communities of low complexity, where few tolerant species ensure high biomass production in the absence of antagonistic interactions. The pattern suggests that environmental stress may result in highly constrained systems which exhibit high rates of functioning due to these key species, in spite of the very limited species pool.     

Relationship between light availability and species richness during fen grassland succession

The complex relation between standing crop and species richness in herbaceous plant communities is still one of the most intensively discussed topics in vegetation ecology. In this study, we focus on the extent to which light availability is able to explain this relationship in fen grasslands. We analyse changes of light availability (measured as relative irradiance, RI), standing crop and species richness during fen grassland succession. Our study include communities representing differences in drainage intensity and nutrient status (‘land use intensification sere’) and successional stages developing after abandonment (‘abandonment seres’). Both in the land use intensification sere and in the abandonment seres, we recorded an increase in standing crop reaching highest values of approx. 800 g m^‐2. RI, species richness and number of small‐growing species declined in the successional seres. RI was highest in stands of low‐productive communities, where light compensation points (5% RI) were already attained at soil surface and light saturation points (30% RI) at soil surface or at 15 cm height. Horizontal heterogeneity of RI at soil surface and at 30 cm height decreased in the abandonment seres, but not at 30 cm height in the land use intensification sere. Furthermore, we detected a signiñcant negative relation between standing crop and RI. Species richness declined with increasing standing crop and thus consequently also with decreasing RI. This result points out that light competition might be of importance for this pattern. The number of small‐growing species decreased exponentially with an increase in standing crop (and decrease in RI). It can be assumed that low light availability negatively affects small‐growing species at standing crop values higher than 400–500 g m². As the maintenance of small‐growing species and the improvement of habitat conditions for their establishment are important aims of nature conservation in fen grasslands, management strategies should be orientated at maintaining standing crops that are not higher than these values.     

Patterns in woody species diversity, richness and partitioning of diversity in forest communities of tropical deciduous forest biome

The regressive succession model hypothesizes tropical savanna-woodlands to be a degraded stage of primary deciduous forests. Species diversity, richness and evenness of woody species in savanna-woodlands, secondary deciduous forests and mature deciduous forests of central India were compared to test if the regressive succession explained pattern in species richness, diversity, functional diversity and basal area. At the plot scale (0.1 ha) secondary deciduous forests and savanna-woodlands had similar species diversity, a pattern not consistent with the regressive model of deciduous forest succession, and mature deciduous forests had greater species diversity and richness (p<0.05). When examined at a larger scale or community scale by pooling all plots within a community type, the trend in diversity persisted even with greater effort allocated to sampling of secondary deciduous forests. Species richness at the community scale was greatest in secondary deciduous forest as expected from species area relationship. The communities shared 28 woody species but the species composition was significantly different between the communities. I suggest that conservation of tropical deciduous forests based on regressive succession model is problematic.     

Seed Banks in Arid Wetlands with Contrasting Flooding,Salinity and Turbidity Regimes

Aquatic plant communities in arid zone wetlands underpin diverse fauna populations and ecosystem functions yet are relatively poorly known. Erratic flooding, drying, salinity and turbidity regimes contribute to habitat complexity, creating high spatial and temporal variability that supports high biodiversity. We compared seed bank density, species richness and community composition of aquatic plants (submergent, floating-leaved and emergent) among nine Australian arid zone wetlands. Germinable seed banks from wetlands within the Paroo and Bulloo River catchments were examined at nested scales (site, wetland, wetland type) using natural flooding and salinity regimes as factors with nondormant seed density and species richness as response variables. Salinity explained most of the variance in seed density (95%) and species richness (68%), with flooding accounting for 5% of variance in seed density and 32% in species richness. Salinity-flooding interactions were significant but explained only a trivial portion of the variance (<1%). Mean seed densities in wetlands ranged from 40 to 18,760 m⁻² and were highest in wetlands with intermediate levels of salinity and flooding. Variability of densities was high (CVs 0.61–2.66), particularly in saline temporary and fresh permanent wetlands. Below salinities of c. 30 g l⁻¹ TDS, seed density was negatively correlated to turbidity and connectivity. Total species richness of wetlands (6–27) was negatively correlated to salinity, pH and riverine connectivity. A total of 40 species germinated, comprising submergent (15 species), floating-leaved or amphibious (17 species), emergent (6 species) and terrestrial (6 species) groups. Charophytes were particularly important with 10 species (five Chara spp., four Nitella spp. and Lamprothamnium macropogon), accounting for 68% of total abundance. Saline temporary wetlands were dominated by Ruppia tuberosa, Lamprothamnium macropogon and Lepilaena preissii. Variable flooding and drying regimes profoundly altered water quality including salinity and turbidity, producing distinctive aquatic plant communities as reflected by their seed banks. This reinforces the importance of hydrology in shaping aquatic biological communities in arid systems.     

Restoration of a species‐rich fen‐meadow after abandonment: response of 64 plant species to management

Abstract. Eleven years of abandonment of a species‐rich fen‐meadow under undisturbed environmental conditions resulted in transformation into areas with tall herb‐, sedge‐ and rush‐dominated communities and areas with Alnus thicket. Species cover was measured in permanent plots in both community types and succession was monitored during 14 yr of restoration following reintroduction of management. The annual increase in accumulated species number followed a log‐log‐time linear regression during 10 yr of grazing management. The expected number of years taken before this annual rate was equal to annual extinction, i.e. a stable situation according to species density, was up to six. The response of 64 species to management was evaluated through paired statistical tests of changes in cover and frequency over time. In total, 55 species could each be allocated to one unique response model (monotone or non‐monotone, concave models) independently of the importance value used (cover or frequency) and type of management (grazing following felling or mowing and mowing without grazing). Species which increased in response to grazing had the most persistent seed banks and CR‐strategies, while species decreasing in response to grazing had less persistent seed banks and CS‐strategies. Some of the species which increased due to grazing followed a model with a local maximum in cover and frequency. The results are discussed in relation to management of species with high cover value during restoration succession.     

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.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Big game or big loss? High deer densities are threatening woody plant diversity and vegetation dynamics

Land-use change and current game management have favored an increasing population of wild ungulates (especially deer) in many regions of the Northern Hemisphere. Here, we assess the impact of high deer densities (>30 ind km^?2) on the highly diverse woody vegetation of Mediterranean ecosystems, where big game have been favored for the last decades. We examine whether prolonged deer browsing (over 30 years) affected plant composition, diversity and dynamics of the original (non-browsed) vegetation. Deer browsing led to an average decrease of 30.4 % in woody plant diversity (species richness), due to a lack of regeneration for the most preferred plant species. Species belonging to early stages of succession (mostly Labiatae and Cistaceae) were non-preferred by deer. Conversely, the most preferred species belonged mainly to late stages of plant succession. Deer impact on Mediterranean shrublands is causing biotic homogenization of plant communities and is threatening vegetation dynamics by forcing it to return to earlier succession stages. Strict deer population control favouring larger trophies but lower offspring numbers together with an adequate habitat management (increasing grass and acorn availability) would be the most efficient measures to reverse this diversity loss. Restoration work seems only appropriate for the most vulnerable species. We highlight the need of sampling deer-free areas with low or null historical browsing to assess the real impact of deer on woody plant diversity and vegetation dynamics.