Nutrient supply in undrained and drained Calthion meadows

Abstract. Plant species-rich Calthion meadows on mesotrophic fen peat soil extensively cut for hay are among the endangered semi-natural vegetation types in northwestern Europe. They are often badly affected by lowering the groundwater table (drainage) and fertilization. In a comparative study of an undrained site with a Calthion meadow and an adjacent drained site, availability of N, P and K was biologically assessed under field conditions (for two years) as well as in a greenhouse (for 18 weeks) by measuring shoot responsiveness. Also, experimental wetting of intact turf samples taken from both sites was applied in order to study the interaction between nutrient supply and anaerobic soil conditions. It was concluded that the above-ground phytomass yield in the undrained site was restricted by a major shortage of N-supply and a moderate shortage of K-supply by the fen peat soil. The above-ground phytomass yield of the drained site was only reduced by a strongly limited supply of K by the soil. The extent of K-deficiency was larger for the drained site. No P-deficiency was observed in any of the drained or undrained sites. Rewetting turf samples, taken from the drained site, did not change above-ground phytomass yields, suggesting that nutrient supplies were not affected by rewetting. Leaching has likely resulted in a strong reduction of K-supply in the drained site. It is assumed that a shortage in K-supply from the peat soil may have become an important environmental constraint for characteristic plant species of Calthion meadows. This may hamper the development of this meadow type on drained peat soils after rewetting by groundwater discharge.     

Nutrient limitations in wet, drained and rewetted fen meadows: evaluation of methods and results

Restoration of wet grassland communities on peat soils involves management of nutrient supply and hydrology. The concept of nutrient limitation was discussed as well as its interaction with drainage and rewetting of severely drained peat soils. Different methods of assessing nutrient limitation were compared and the type and extent of nutrient limitation were determined for several wet grassland communities. It was concluded that a full-factorial field fertilisation experiment is the most preferable method. Plant tissue analyses and soil chemical analyses were considered less suitable, although they may provide helpful additional information. Fertilisation experiments in the laboratory using sods or using test plants appear to be the proper means to study mechanisms or processes, but have a restricted predictive value for field situations. Generalising the results, it seems that many relativily undisturbed grassland plant communities on peaty soils are characterised by N limitation. Phosphate limitation for vegetation on peat soils is mainly observed in specific circumstances such as extreme calcium richness, high concentrations of Fe or as a result of drainage or long-term hay cropping. The latter two may also cause K limitation. Rewetting is regarded as a prerequisite in restoring wet grassland communities. Further restoration measures to influence nutrient availability depend on aims of the management and the individual site conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nutrient limitation and nutrient‐driven shifts in plant species composition in a species‐rich fen meadow

Question: We studied the development and persistence of the effects of nutrient pulses on biomass production and species composition in a fen meadow. Location: Nature reserve, central Netherlands, 5 m a.s.l. Methods: Single pulse fertilization with N and P in a factorial design on an undrained central and a drained margin site in a species‐rich fen meadow (Cirsio dissecti‐Molinietum). Biomass production and species composition were monitored during four years. Results: At the central site, N addition boosted biomass production, but only during one year. The species composition was not changed. P fertilization increased the biomass production and changed the species composition from a vegetation dominated by Carex panicea to a grassland community with abundant Holcus lanatus, but not before the second year. At the margin site, P fertilization changed the species composition in a similar way, but biomass production was not increased. N fertilization had no effect. At both sites the P induced shift in species composition persisted for four years although the P effect declined during the experiment. Conclusions: The biomass responses show that N was limiting in the central site. Another nutrient, besides N and P (probably K) must have been limiting in the marginal site. The fast decline of the N effect on biomass is ascribed to increased denitrification and biomass removal. The delay in the P effect on biomass and species composition and the persistence of the P effect on species composition are ascribed to fast immobilisation and subsequent slow release of fertilizer P in the peat soil. Recurrence of the P pulses is expected to cause permanent changes in species composition.     

Long-term effects of drainage and hay-removal on nutrient dynamics and limitation in the Biebrza mires,Poland

To provide a reference for wetlands elsewhere we analysed soil nutrients and the vegetation of floodplains and fens in the relatively undisturbed Biebrza-valley, Poland. Additionally, by studying sites along a water-table gradient, and by comparing pairs of mown and unmown sites, we aimed with exploring long-term effects of drainage and annual hay-removal on nutrient availabilities and vegetation response. In undrained fens and floodplains, N mineralization went slowly (0–30 kg N ha⁻¹ year⁻¹) but it increased strongly with decreasing water table (up to 120 kg N ha⁻¹ year⁻¹). Soil N, P and K pools were small in the undisturbed mires. Drainage had caused a shift from fen to meadow species and the disappearance of bryophytes. Biomass of vascular plants increased with increasing N mineralization and soil P. Annual hay-removal tended to have reduced N mineralization and soil K pools, but it had increased soil P. Moreover, N concentrations in vascular plants were not affected, but P and K concentrations and therefore N:P and N:K ratios tended to be changed. Annual hay-removal had induced a shift from P to K limitation in the severely drained fen, and from P to N limitation in the floodplain. The low nutrient availabilities and productivity of the undisturbed Biebrza mires illustrate the vulnerability of such mires to eutrophication in Poland and elsewhere. In nutrient-enriched areas, hay removal may prevent productivity increase of the vegetation, but also may severely alter N:P:K stoichiometry, induce K-limitation at drained sites, and alter vegetation structure and composition.     

Restoration experiments in middle European wet terrestrial ecosystems: an overview

Most wetlands in the central European lowlands have been severely altered by cultivation. As a consequence they no longer fulfil their function as habitats for specialised species and communities, nor as retention areas for water and solid materials. Therefore, a number of renaturation experiments are in progress, which intend to develop and test strategies and measures to improve this defect in landscape diversity. For this purpose experiments on re-wetting, nutrient depletion of eutrophicated areas and re-establishment of typical wetland plant species and phytocoenoses have been performed. An ecological development concept defining the aims and describing their feasibility precedes such experiments. Preliminary results indicate that the reconstruction of the former state (regeneration) is impossible within reasonable time spans. In drained raised bogs overgrown with heather, as well as in those which have been industrially exploited, the primary aim must always be to restore efficient peat formation as far as possible; as a rule one succeeds only with well-growing and nutrient-demanding fen and transitional bog species. In cultivated fens the aim is to reduce peat loss. As a first step this is accomplished by converting arable fields and sown meadows into permanent grassland, if possible with reduced fertilization and low mowing or grazing frequencies, and accompanied by rewetting during winter. Many experiments have sought to impoverish eutrophicated fen soils and introduce typical fen species by sowing or planting, so well tested techniques are available. However, the total prevention of peat loss is only possible by permanent rewetting throughout the year, so that peat accumulation can start again. Only in this way could fens regain their former function as sinks in landscape processes.     

Establishment of Carex stricta Lam. seedlings in experimental wetlands with implications for restoration

The loss of Carex dominated meadows due to agricultural drainage in the previously glaciated midcontinent of North America has been extensive. The lack of natural Carex recruitment in wetland restorations and the failures of revegetation attempts underscore the need for information on the establishment requirements of wetland sedges. In this study, seedlings of Carex stricta Lam. were planted in three experimental wetlands in east-central Minnesota, USA to investigate the biotic and abiotic environmental limitations to establishment. Seedlings were planted along an elevational water depth gradient to assess the effects of water depth and water level fluctuation on seedling survival and growth. A different water level fluctuation regime was assigned to each of the experimental wetlands to assess seedling tolerance for seasonal water level changes. The effects of seedling planting density and the presence or absence of non-sedge colonizers on seedling survival and growth were also studied. The experiment was followed for three growing seasons. The results of this study indicate that C. stricta seedlings were sensitive to the timing and duration of inundation during the first growing season. Once established, plants tolerated a broad range of seasonal drying and flooding conditions. Seedling and juvenile growth was slowed by non-sedge colonizers during the first two growing seasons, but by the third growing season, C. stricta was able to out-grow all annual and perennial weeds, except the aggressive perennial, Phalaris arundinacea L. The rapid growth of C. stricta plants, once established, indicates that the use of seedlings is a successful method for (re) introducing this tussock sedge into wetland restorations under a variety of environmental conditions. Comparison with other studies performed under similar conditions suggests that planting of seedlings is a more appropriate method of establishing this species than the use of transplanted rhizomes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Vegetation heterogeneity and ditches create spatial variability in methane fluxes from peatlands drained for forestry

Drainage of peatlands for forestry starts a succession of ground vegetation in which mire species are gradually replaced by forest species. Some mire plant communities vanish quickly following the water-level drawdown; some may prevail longer in the moister patches of peatland. Drainage ditches, as a new kind of surface, introduce another component of spatial variation in drained peatlands. These variations were hypothesized to affect methane (CH₄) fluxes from drained peatlands. Methane fluxes from different plant communities and unvegetated surfaces, including ditches, were measured at the drained part of Lakkasuo mire, Central Finland. The fluxes were found to be related to peatland site type, plant community, water-table position and soil temperature. At nutrient-rich fen sites fluxes between plant communities differed only a little: almost all plots acted as CH₄ sinks (−0.9 to −0.4 mg CH₄ m⁻² d⁻¹), with the exception of Eriophorum angustifolium Honck. communities, which emitted 0.9 g CH₄ m⁻² d⁻¹. At nutrient-poor bog site the differences between plant communities were clearer. The highest emissions were measured from Eriophorum vaginatum L. communities (29.7 mg CH₄ m⁻² d⁻¹), with a decreasing trend to Sphagna (10.0 mg CH₄ m⁻² d⁻¹) and forest moss communities (2.6 mg CH₄ m⁻² d⁻¹). CH₄ emissions from different kinds of ditches were highly variable, and extremely high emissions (summertime averages 182–600 mg CH₄ m⁻² d⁻¹) were measured from continuously water-covered ditches at the drained fen. Variability in the emissions was caused by differences in the origin and movement of water in the ditches, as well as differences in vegetation communities in the ditches. While drainage on average greatly decreases CH₄ emissions from peatlands, a great spatial variability in fluxes is emerged. Emissions from ditches constantly covered with water, may in some cases have a great impact on the overall CH₄ emissions from drained peatlands.     

Groundwater chemistry and vegetation of gradients from rich fen to poor fen in the Naardermeer (the Netherlands)

In the eastern part of the Naardermeer peatlands (the Netherlands) a regional calcium-rich groundwater flow discharges (here often called the seepage area), whereas in the western part infiltration takes place. The ecological consequence of this hydrological pattern is reflected by the pattern in reedland communities. In the seepage area, which is characterized by Thelypteris-reedlands including many rare and endangered species (Caricion davallianae, Calthion palustris), there is a complex gradient of water types. The lime potential in the peat soil is clearly influenced by the hydrological gradient. In the ombrotrophic (poor fen) part of the gradient (containing species of Caricion curto-nigrae) the lime potentials are low and the groundwater contains low amounts of dissolved ions. The rare and endangered species (Caricion davallianae) are restricted to a small area with high lime potentials which is nourished by regional calcium-rich groundwater. High lime potentials were also measured in eutrophic reedlands influenced by brackish groundwater. Several species which generally occur in wet meadows even show a preference for this brackish environment.In a part of the seepage area succession from rich fen to poor fen and Alnus wood has taken place over a period of 40 years. This development has been caused by the diminishing amount of fresh seepage due to a lowering of the water levels in the surrounding area.The characteristics of poikilotrophic zones (contact zones between water flows) are discussed in relation to their significance for the preservation of endangered marsh species.Abbreviations EC₂₅= Electrical conductivity measured at 25 °C     

Basis for restoration of saltcedar (Tamarix spp., Tamaricaceae) invaded sites through an adaptive management approach

Biological invasions are considered one of the most serious drivers of global biodiversity degradation in the face of ecological restoration. Saltcedar (Tamarix spp.) is an aggressive invader in arid environments of the United States, Mexico, Australia, and Argentina, causing profound alteration of riparian habitats, the composition and structure of natural communities, and ecosystem functioning. Given the severity of the reported invasion processes, and considering that Tamarix’s responses to the wide range of existing control techniques are still poorly explored, the objectives of this investigation were to assess the implementation of different control techniques based on active adaptive management and to define indicators to measure the effectiveness of both the techniques and the recovery of the system, as a first action in ecological restoration projects of invaded sites. The experiment was carried out in the Llancanelo Wetland Provincial Reserve and Ramsar site, where 540 ha of marsh environment were replaced by monospecific saltcedar forests. Thirteen treatments were proposed combining control techniques (mechanical and chemical), saltcedar population situation (forest, shrubland, resprouts), and times of the year. Simultaneously, an assisted revegetation experiment was performed on four of the controlled plots. Finally, the costs associated with each treatment were estimated. The most effective control techniques were mechanical extraction and root burning for areas with mature, low-density forests, and cutting and herbicide application for shrublands and resprouts in late summer-early spring. The cutting and shading technique is recommended in places where the herbicide application can put the ecosystem integrity at risk. Assisted revegetation only provided 2% coverage in the intervened plots, a non-significant value considering that natural recovery presented up to 35% coverage and a low percentage of invasive alien species. In spite of this, the species composition reported in the monitoring suggests adjusting revegetation techniques to assist native species recovery.     

Fossil insects and ecosystem dynamics in wetlands: implications for biodiversity and conservation

We review the uses of fossil insects, particularly Coleoptera (beetles) and Chironomidae (non-biting midges) from ancient deposits to inform the study of wetland ecosystems and their ecological and restoration processes. In particular, we focus on two contrasting ecosystems, drawing upon research undertaken by us on British raised mire peats and shallow lake systems, one an essentially terrestrial ecosystem, the other aquatic, but in which wetland insects play an important and integral part. The study of raised mire peats suggests that faunal stability is a characteristic of these wetland systems, over what appear to be extensive periods of time (up to several millennia), whilst studies of shallow lake ecosystems over recent timescales indicates that faunal instability appears to be more common, usually driven by increasing eutrophication. Drawing upon a series of fossil Coleoptera records spanning several thousand years from Hatfield Moors, south Yorkshire, we reconstruct in some detail the mire’s ontogeny and fluctuations in site hydrology and vegetation cover, illustrating the intimate association between substrate, topography and peat development. A comparison between fossil and modern beetle populations indicates that the faunal characteristics of this mire and its adjacent neighbour, Thorne Moors, become established during the early phases of peat development, including its rare endemics, and that the faunal biodiversity on the sites today is dictated by complex site histories. The over-riding characteristic of these faunas is of stability over several thousand years, which has important implications for the restoration of degraded sites, especially those where refugial areas are limited. In contrast, analyses of fossil Chironomidae from shallow lakes allow researchers to track changes in limnological status and while attempts have been made to reconstruct changes in nutrient levels quantitatively, the chironomids respond indirectly to such changes, typically mediated through complex ecosystem dynamics such as changes in fish and/or macrophyte communities. These changes are illustrated via historic chironomid stratigraphies and diversity indices from a range of shallow lakes located across Britain: Slapton Ley, Frensham Great Pond, Fleet Pond, Kyre Pool and Barnes Loch. These sites have shown varying degrees of eutrophication over recent timescales which tends to be associated with a decline in chironomid diversity. While complex functional processes exist within these ecosystems, our evidence suggests that one of the key drivers in the loss of shallow lake chironomid diversity appears to be the loss of aquatic macrophytes. Overall, while chironomids do show a clear response to altered nutrient regimes, multi-proxy reconstructions are recommended for a clear interpretation of past change. We conclude that if we are to have a better understanding of biota at the ecosystem level we need to know more of the complex interactions between different insect groups as well as with other animal and plant communities. A palaeoecological approach is thus crucial in order to assess the role of insect groups in ecosystem processes, both in the recent past and over long time scales, and is essential for wetland managers and conservation organisations involved in long term management and restoration of wetland systems     

Arbuscular mycorrhiza of introduced and native grasses colonizing zinc wastes: implications for restoration practices

The analysis of mycorrhizal status of grasses introduced under conventional phytoremediation programmes (including covering of the waste area with a layer of uncontaminated soil and watering) and of grasses that spontaneously established in later successional stages, was carried out on the slopes of the 1- to 30-year-old tailings of ZG Trzebionka Mining Company located near Chrzanów (southern Poland). Most grasses were mycorrhizal and the parameters of mycorrhization were higher on the older parts of the waste. Grasses selected for restoration practices were well developed only if the waste was covered with a layer of soil and continuous irrigation took place, both being expensive practices. The field experiment showed that the introduction of standard inoculum containing mycorrhizal fungi indeed improved the development of grasses during early stages, but still was not effective enough due to high erosion of the substratum. A slight improvement was observed when AgroHydroGel was used to lock moisture in the soil, but the plants did not survive for longer time periods. On the contrary, much better results were obtained when vegetatively multiplied grasses selected from specimens originating from natural succession were used to stabilize the vegetation on the bare industrial waste.     

Nutrient status and retention in pristine and disturbed wetlands in Uganda: management implications

Wetlands in Uganda experience different forms of human pressure ranging from drainage for agriculture and industrial development to over harvesting of wetland products. In order to develop sustainable management tools for wetland ecosystems in Uganda and the Lake Victoria Region, water quality analyses were carried out in a rural undisturbed (pristine) wetland (Nabugabo wetland in Masaka) and two urban wetlands that are experiencing human and urban development pressure (the Nakivubo wetland in Kampala and Kirinya wetland in Jinja). The former wetland forms the main inflow into Lake Nabugabo while the other two border the northern shore of Lake Victoria, Uganda. Nabugabo wetland buffers Lake Nabugabo against surface runoff from the catchment, while Nakivubo and Kirinya wetlands provides a water treatment function for wastewater from Kampala City and Jinja town respectively, in addition to buffering Lake Victoria against surface runoff. Water quality was assessed in all the wetland sites, and in addition nutrient content and storage was investigated in the main plant species (papyrus, Phragmites, Miscanthidium and cocoyam) in Nakivubo and Kirinya wetlands. A pilot experiment was also carried out to assess the wastewater treatment potential of both the papyrus vegetation and an important agricultural crop Colocasia esculenta (cocoyam). Low electrical conductivity, ammonium–nitrogen and ortho-phosphate concentrations were recorded at the inflow into Nabugabo wetland (41.5 μS/cm; 0.91 mg/l and 0.42 mg/l respectively) compared to the Nakivubo and Kirinya wetlands (335 μS/cm; 31.68 mg/l and 2.83 mg/l and 502 μS/cm; 10 mg/l and 1.87 mg/l respectively). The papyrus vegetation had higher biomass in Nakivubo and Kirinya wetlands (6.7 kg DW m⁻²; 7.2 kg DW m⁻² respectively), followed by Phragmites (6.5, 6.7), cocoyams (6.4, 6.6) and Miscanthidium (4.0, 4.2). The papyrus vegetation also exhibited a higher wastewater treatment potential than the agricultural crop (cocoyam) during the pilot experiment (maximum removal degree of ammonium–nitrogen being 95% and 67% for papyrus and yams). It was concluded that urbanisation pressure reduces natural wetland functioning either through the discharge of wastewater effluent or the degradation of natural wetland vegetation. It is recommended that wetland vegetation be restored to enhance wetland ecosystem functioning and for wetlands that are not yet under agricultural pressure, efforts should be made to halt any future encroachment.     

Soil properties of an Antarctic inland site: implications for ecosystem development

Inland Antarctic nunataks typically have simple physically weathered soils and limited ecosystem complexity. In this paper we present quantitative measurements of soil physical and chemical properties at one Antarctic nunatak. We measured pH, grain size, field capacity, soil organic carbon, phosphate, nitrate, ammonium and the cations magnesium, calcium and potassium along two transects. The data obtained indicated that very low levels of nutrients were present/available to biota, and that liquid water was absent, at least from the surface depths of soil, except during periods of active snow melt. Consequently, biological activity is severely limited. We conclude that, due to the climatic and microclimatic conditions at this location, the development of biological communities and soils is maintained in an extremely simple but still apparently stable ‘quasi climax’ state. Increased soil development and biological complexity can be expected if the contemporary rapid regional warming in the Antarctic Peninsula region continues.     

Zostera marina population genetics in Barnegat Bay, New Jersey, and implications for grass bed restoration

Within Barnegat Bay, New Jersey, Zostera marina populations have declined by 62% over the last 20 years, and restoration efforts have met with mixed success. We have completed a microsatellite-based genetic investigation of eight populations of Z. marina within Barnegat Bay to determine whether the genetic stock origins of the plants used in management projects may affect restoration success. Additionally, we assessed the genetic diversity of Z. marina in Barnegat Bay to better understand its population structure. Clonal diversity ranged from 0.70 to 0.95 for the populations studied. Individually, Barnegat Bay populations are not genetically diverse, and there is also little divergence among populations. The Atlantic populations had mean Hobs values (0.20–0.34) that were far lower than the Hexp values (0.69–0.83). Also, the F _IS values in all of the eastern populations indicate a surfeit of homozygotes over heterozygotes, suggesting a low degree of outcrossing in the Barnegat Bay populations. Six of the ten populations studied (Ham Island, Manahawkin Bay, Shelter Island, Marsh Elder, Harvey Cedar Sedge, and Long Island) show evidence of historical bottlenecks. Mean estimated F _ST values would suggest that most alleles are undergoing moderate genetic differentiation, with values that range from 0.06 to 0.13. Oyster Creek and Sedge Island demonstrate the largest estimated effective population sizes and may be the most appropriate populations for use in future eelgrass restoration projects.     

Single gene restoration of cytoplasmic male sterility in wheat and its implications in the breeding of restorer lines

Summary Cultivars of T. aestivum crossed onto two lines with male sterility induced by the cytoplasm of T. timopheevi gave a high level of restoration in the F₁ generation. The ratio of fertile to sterile plants segregating in the F₂ generation was consistent with that expected for a single dominant restorer gene. The possible association between this gene and mildew resistance or some other desirable character derived from CI 12633, a common ancestor of the cultivars used, is discussed.     

Post-fire and post-quarry rehabilitation successions in Mediterranean-like ecosystems: Implications for ecological restoration

Resilience against sudden changes in the environment is a very desirable trait in plants used for ecosystem restoration. Mediterranean-like vegetation exhibits particularly strong fire resilience. There are two main functional groups of fire-prone species among Mediterranean-like vegetation: seeders and resprouters. Our aims were to describe how the theory of succession after fire relates to rehabilitation and to use this knowledge to improve the results of rehabilitation attempts in Mediterranean-like ecosystems. Eight post-fire (PF) sites, 14 post-rehabilitated (PR) quarry sites and two woodland sites were sampled. Detrended Canonical Correspondence Analysis (DCCA) showed that PF and PR successions were quite different. Both displayed an increasing abundance of resprouters over time, but seeder density increased throughout PR succession and decreased during PF succession. Nine species were common to both successions in all studied stages. The results showed that until 15-21 years of succession, the post-rehabilitation sites had not become as resilient to fires as sites populated by indigenous vegetation due to the lack of a seeder seed bank. However, after 21 years of PR succession, the exponentially increasing seeder population may allow for seed bank formation and thus eventually improve the fire resilience of the site.