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Abstract: ‘Ecosystem engineers’ have a critical role in the structure and function of natural communities and need to be considered as focal species to enable successful conservation or restoration of ecosystems. Through introduction and/or managing a single species, an entire community can be influenced. In our study we show that Pedicularis palustris, an endangered hemiparasite in large parts of Europe, can act as an ‘ecosystem engineer,’ speeding up the restoration of undrained fen-meadow ecosystems colonised by species-poor and resistant Carex acuta vegetation. The parasitic behavior of Pedicularis palustris on tall sedges significantly alters local plant diversity, biomass production and soil characteristics. Our experiments show that, under the right hydrological conditions and mowing management practice, several target species for nature conservation can benefit from the gaps it creates in above-ground Carex acuta vegetation and its dense below-ground root system. The more prominent presence of mosses and lower density of the topsoil stimulates the recovery of mesotrophic transition mire with active peat formation. Since this habitat type is specifically protected under the European Habitat Directive, (re)introduction of Pedicularis palustris in similar sites within its geographical distribution range in Europe may be a helpful tool to achieve the imposed obligations for appropriate management and restoration. [Copyright &y& Elsevier]

Abstract: Clonality resulting from the growth of specialized organs is common among plants in wetland habitats. We hypothesize that different wetland habitats select for different attributes of clonal traits. This hypothesis is based on studies of individual species but has not been previously tested at the level of habitat. We compared the functional diversity of clonal growth traits of plants in bogs, fens, wet heathlands, floodplains, river beds, open fresh water habitats, salt marshes, and open marine habitats. Clonal traits (including number of offspring, lateral spread, persistence of connections between ramets, and shoot life span) were analysed with multivariate techniques using species frequency data and with permutation tests using presence/absence data. Based on species frequencies, clonal plants in aquatic habitats (open fresh water habitats, open marine habitats, and river beds) were characterized by the abundant production of freely dispersible propagules, annual shoots, and splitting clones. Species of daily flooded salt marshes were characterized by bi-annual connections between ramets and medium dispersability. In contrast, plants in permanently wet bogs were characterized by polycyclic shoots and low offspring production. The specificity of river beds and open freshwater habitats was also confirmed by permutation tests, which gave equal weight to rare and abundant species. However, species in all other wetland habitats were characterized by the entire range of clonal traits, suggesting weak environmental filtering of analyzed traits by habitat at the present scale. [Copyright &y& Elsevier]

Abstract: The majority of fens in Europe have been transformed for agricultural purposes and have disappeared or become degraded. Fen meadows that developed under low-intensity management of fens also have become degraded. In this paper, we consider the available restoration methods, biotic constraints for restoration and new prospects and approaches for the restoration of severely degraded fens. Due to irreversible changes in landscape settings, hydrology, soil and trophic conditions, a full restoration to natural mires is unlikely. Yet, an improvement of the ecosystem functions and revival of biodiversity in degraded fens is possible. A restoration of semi-natural meadows is one of the alternative targets. Important for restoration efforts to succeed are a sufficient reduction of nutrient levels and preventing acidification. In general, a combination of topsoil removal and seed transfer is an effective measure for fen meadow restoration, provided that groundwater seepage can be re-established. There are also several biotic limitations to fen meadow restoration, due to limited propagule availability of target species and the legacy of the former vegetation in form of its soil seed bank and high seed production by unwanted species. Under the present environmental conditions, the re-development of fen meadows on degraded fens will result in species compositions different from those observed in the past and such restoration may require considerable time and effort. [Copyright &y& Elsevier]

Abstract: Clonal multiplication is a predominant type of reproduction in wetland species. However, both wetlands and plant organs of clonal growth are diverse, thus due to different stress factors operating in various wetlands preponderance of plants with specific clonal growth organs (CGOs) can be expected. To test this hypothesis the CGO spectra of wetland communities of the Netherlands were analysed, including a bog, a fen, heathland, a floodplain, river beds, fresh water pools, open salt water and a salt marsh. Moreover, it was evaluated whether different CGOs are characterised by different functional traits (shoot cyclicity, persistence of connections between ramets, number of offspring produced per year and lateral spread per year) in wetland species. Data on types of CGO, i.e., epigeogenous and hypogeogenous rhizomes, fragments and budding plants, stolons, tubers and bulbs, root-splitters, root-sprouters and special adaptations (turions) as well as their functional traits, were taken from the CLO-PLA 3 database. CGO spectra of wetland communities were analysed using two methods: comparison of observed vs. expected CGO spectra based on the presence/absence data and multivariate analysis (CCA) for inter-community differences considering species frequency. Moreover, relationships between CGOs and their functional traits were tested using multidimensional contingency tables. Apart from 26% of non-clonal species, the majority of wetland species was rhizomatous (51%). Other types of CGO were represented in less than 10% of species and root-derived CGOs were underrepresented (<2%) in comparison with terrestrial habitats. Among communities, fresh water pools and open salt water hosted higher proportion of species with fragments (∼10%) and turions (∼30%). Multivariate analysis divided wetland communities along the disturbance and hydric (water) gradients. Highly disturbed communities (salt marshes) were characterised by non-clonal species and species with root-derived CGOs. Aquatic communities (fresh water pools and open salt water) hosted species with the ability to spread by fragmentation and turions, contrary to permanently wet communities (bog and wet heathland) with the prevalence of species with epigeogenous rhizomes. It was also confirmed that the CGOs of wetland species differed in their traits. The most important functional trait characterising individual CGOs in the wetland flora was the degree of lateral spread (explained variability: 53%) followed by duration of persistence of connections between ramets (explained variability: 74%), which is in accordance with earlier distinguished strategies of clonal growth: integrator/splitter and spreading/non-spreading clones. [Copyright &y& Elsevier]

Techniques such as rewetting, topsoil removal, diaspore transfer or combinations of these are increasingly applied in fen meadow and flood meadow restoration in Western Europe. In this paper, we present a quantitative assessment of the effectiveness of the commonly used meadow restoration methods. We use the change in ‘saturation index’ to evaluate the degree of success. The index reflects the completeness of restored communities in comparison to regional target communities. Meadow restoration has limited success in most cases, with an average increase in species richness below 10% of the regional species pool. Restoration success was partly determinated by the starting situation. The more species-rich the starting situation, the higher the saturation index after restoration but, at the same time, the smaller the increase in the number of target species due to restoration. Top soil removal and diaspore transfer were found to contribute most to restoration success. A combination of top soil removal and diaspore transfer and a combination of all three techniques appeared to be the most effective measure and resulted in an increase in the saturation index of up to 16%. Rewetting alone had no measurable effect on restoration success. [Copyright &y& Elsevier]

Abstract: Current knowledge about the spatial variation of atmospheric nitrogen deposition on a local scale is limited, especially for vegetation with a low canopy. We measured nitrogen deposition on artificial vegetation at variable distances of local nitrogen emitting sources in three nature reserves in the Netherlands, differing in the intensity of agricultural practices in the surroundings. In the nature reserve located in the most intensive agricultural region nitrogen deposition decreased with increasing distance to the local farms, until at a distance of 1500m from the local nitrogen emitting sources the background level of 15kgNha−1 yr−1 was reached. No such trend was observed in the other two reserves. Interception was considerably lower than in woodlands and hence affected areas were larger. The results are discussed in relation to the prospects for the conservation or restoration of endangered vegetation types of nutrient-poor soil conditions. [Copyright &y& Elsevier]

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Many endangered plant species remain absent in rewetted, previously drained fens. We performed a 3-year introduction experiment with endangered fen species (9 Carex - and 6 bryophyte species) in 4 hydrologically restored fens to investigate which factors hamper establishment and survival. Carex species were introduced as adults and seedlings, mosses as gametophytes. Introductions were done on (initially) bare soil, which allowed us to exclude excessive competition for light during the first year. First year survival of the transplants was high in all fens (mean survival = 96%), indicating that there were no direct abiotic constraints on establishment. However, survival analysis revealed that a decrease in relative light intensity (RLI) at the soil surface during consecutive years (indicating an increase in biotic competition for light) drove high mortality rates in most species. As a result, overall final survival was lowest in the two most productive (low light) fens (mean survival = 38%), while most transplants persisted in the two less productive (high light) fens (mean survival = 79%). Taller and faster-growing Carex species were able to outgrow light limitation near the soil surface, and thus had a higher overall survivability than smaller and slower-growing species. Light limitation also drove the loss of 5 out of 6 bryophyte species. We conclude that both dispersal limitation and asymmetric competition for light may explain the lack and loss of small and endangered plant species in rewetted fens. A minimum empirical threshold of c. 30% relative light intensity near the soil surface is required for successful introduction. [ABSTRACT FROM AUTHOR]

Global land-use intensification and drainage has altered the biogeochemical properties of many peatlands, and concomitant eutrophication has led to a loss of low-competitive fen species. We investigated the hypothesis that removal of a degraded and eutrophied top peat layer, thereby exposing an underlying peat layer, can improve conditions for rich fen restoration. We studied the long-term (3–18 years) effects of past topsoil removal in six rich fens in Western Europe by comparing topsoil removal plots with (untouched) control plots. Overall, topsoil removal plots were characterized by lower bulk densities and soil nutrient pools of P and KCl-extractable NH 4 + , while organic matter contents and soil C:N ratios were higher. Pore water concentrations of NO 3 − and NH 4 + also decreased in the topsoil removal plots, while concentrations of base cations (Ca 2+ , Mg 2+ , Na + , K + ) and HCO 3 − increased. Furthermore, lower nutrient levels appeared to restrict herb biomass production in the topsoil removal plots, so that optimized light conditions led to the establishment of light-demanding target species and a significant increase in bryophyte cover. Multivariate analysis revealed that most variation in vegetation assembly was due to higher groundwater levels in the topsoil removal plots, closely followed by a higher relative light intensity (RLI) at surface level, lower pore water nutrient (NH 4 + ) concentrations, and higher concentrations of base cations. We conclude that topsoil removal can be an effective mechanism to “reset” a degraded peatland to its initial state of nutrient limitation, base saturation and high availability of light, thereby improving the conservation prospects of endangered rich fen communities. [ABSTRACT FROM AUTHOR]

Soil inoculation from plant species-rich into species-poor grasslands may enable the establishment of self-facilitating networks between microbes and vegetation, thereby steering ecosystem development. We conducted a three-year experiment that covered a wide range of post-agricultural grasslands to determine how succession is affected by the interactive effects of edaphic properties, grass layer removal and hay or soil fragment transfer from a late-successional donor grassland. Soil inoculation generally impacted community assembly of vegetation and fungi, but not of prokaryotes. Effects were strongest when preceded by removal of the grass layer, indicating the importance of priority effects and dispersal limitation. Inoculation enabled the establishment of putative rhizosphere-associated fungal taxa, particularly from the families Helotiaceae, Glomeraceae and Archaeorhizomycetaceae. Nonetheless, effect sizes were mostly small, as was overall resemblance of the receptor grasslands to the donor. Fungal communities were primarily shaped by environmental filters and only reached a high resemblance to the donor in nutrient-poor sites. Shifts in the vegetation were strongest in those grasslands where the mycobiome more closely resembled that of the donor. Soil inoculation generally facilitates colonization by target plant and fungal communities, where establishment success of the former can be predicted by the latter, but the final outcome of succession is environmentally determined. • Soil inoculation impacts community assembly of vegetation and fungi in grasslands. • Inoculation enables the establishment of putative rhizosphere-associated fungal taxa. • Local environmental filters determine the final outcome of succession. [ABSTRACT FROM AUTHOR]

Abstract: The northwest European heathland landscape with its characteristic communities of nutrient-poor and acidic soils has a high nature value, because of its locally high biodiversity and the distinct site conditions. In order to conserve and restore the heathlands, numerous rehabilitation projects have been performed, although with varying success. This is partly due to the fact that the key biogeochemical processes distinguishing the various vegetation types within the heathlands are not known in detail. Therefore, we performed a statistical survey on the main communities and their soil characteristics. In addition, we analyzed the data for key factors determining biodiversity in the heathland landscape. Data from previous studies and surveys was used to compile a dataset of 267 vegetation relevés (classified as EUNIS habitat types) with extensive soil measurements (22 parameters). A canonical discriminant analysis revealed that soil acidity explained most of the differences between the habitat types, while soil moisture content and soil fertility were less important. Acidity-related factors as Al3+, Al/Ca-ratio and pH were also strongly correlated to plant diversity in the majority of the habitat types, respectively, the species-rich Nardus grasslands, the Rhynchosporion communities and the species-rich Molinia meadows. In the dry heaths and over the total heathland landscape, plant diversity was negatively correlated with soil -concentrations. Only in wet heath, nutrient availability, in this case P, was the primary factor in explaining plant diversity. This study presents ranges for all major soil parameters for the studied well-developed heathland habitat types, thereby providing clear guidelines for conservation and restoration. [Copyright &y& Elsevier]

Abstract: Structural patterns of tall stands (“tussock”) and short stands (“lawn”) are observed in grazed vegetation throughout the world. Such structural vegetation diversity influences plant and animal diversity. A possible mechanism for the creation and preservation of such patterns is a positive feedback between grazing and plant palatability. Although some theoretical studies have addressed this point in a non-spatial setting, the spatial consequences of this feedback mechanism on the stability and spatial characteristics of vegetation structure patterns have not been studied. We addressed this issue by analyzing a spatially explicit individual-based plant-grazer simulation model, based on published empirical relations and the assumption of optimal foraging. In the model, the selection by the grazer of short stands (that have a higher energy content and are more palatable) is affected by traveling costs and the spatial organization of swards. Nevertheless, the most selected biomass in this type of short stands was the optimal biomass predicted by cropping and digestion constraints. As a result of the optimal foraging strategy, the grazers displayed Lévy-flight traveling behavior during the simulations with characteristic exponent μ ≈2. Patterns of short and tall stands created by grazing were preserved for at least a decade. Even in seasonal habitat, the spatial organization of the patterns remained relatively constant, despite fluctuations in the area of short stands. Heterogeneity of initial vegetation increased heterogeneity of the grazing-induced pattern, but did not affect its stability. The area of short stands that was preserved by grazing scaled with the herbivore mass to the power 0.4 and with the carrying capacity of the vegetation to the power −0.75. Patterns of tall and short stands can be created and perpetuated by optimally grazing ruminants, irrespective of possible underlying soil patterns. The simulations generate predictions for the stability and spatial characteristics of such structural vegetation patterns. [Copyright &y& Elsevier]