Effects of removing the litter and humic layers on heathland establishment following plantation removal

We investigated ways of optimising the establishment of Calluna vulgaris-dominated heathland vegetation following removal of pines from afforested heathland using a randomised block experiment. Removal of the litter layer, or both the litter and humic layers, resulted in a higher initial establishment rate of C. vulgaris compared to in the control plots. The effect of adding heathland clippings suggested that the rate of establishment of C. vulgaris was limited by seed availability in plots from which both the litter and humic layers had been removed, but not in those from which just the litter layer had been removed. Removal of both the litter and humic layers did, though, result in a long-term reduction in the depth of the humic layer, a lower loss-on-ignition and lower level of nitrate and total nitrogen compared to in the control plots. The optimal method for establishing heathland vegetation at a site will depend on: (1) whether there is a viable seedbank of C. vulgaris and other desirable heathland species present; (2) the importance of quickly establishing a high cover of desirable heathland species; (3) the likely long-term effects of soil nitrogen levels on the maintenance of C. vulgaris dominance; (4) the practicality of removing and disposing of the litter and humus layers.     

Effects of habitat management on heathland response to atmospheric nitrogen deposition

A field experiment was carried out to investigate the impact of four different management treatments on the response of Calluna-dominated heathland to nitrogen additions of 0 or 30 kg ha⁻¹ yr⁻¹ at a lowland heath in Surrey, England. Post-management Calluna regeneration, growth and canopy development were significantly affected by both the form of management and by nitrogen addition. The effect of nitrogen on shoot growth was lower, in absolute terms, in those plots which had undergone more intensive management treatments. Seedling establishment was higher in plots which received a simulated accidental burn treatment, reflecting the greater proportion of bare ground and the high degree of physical disturbance associated with this treatment. Invasion by Deschampsia seedlings was increased by nitrogen addition, particularly in simulated accidental burn plots. In the long term, the need to maintain a low nutrient environment to favour Calluna dominance, particularly in the face of elevated nitrogen deposition, favours the use of managements which result in the export of a significant proportion of the organic nitrogen stores. However, further study is required to ensure that initial enhancement of seedling invasion by grass and other species, associated with these more intense managements, does not outweigh the long term benefits of associated nutrient removal.     

Long-term heathland restoration on former grassland: The results of a 17-year experiment

European lowland heaths have declined by up to 80% due to land use change and lack of management. There has been considerable research into the restoration of this threatened habitat. However, long-term outcomes of restoration are poorly understood, especially in situations where past agricultural land use imposes severe constraints on community re-assembly. In 1989 a large-scale experiment was established to examine the effectiveness of five treatments to restore heathland on formerly productive grassland: (i) natural regeneration; (ii) herbicide application to facilitate regeneration; (iii) cultivation and application of seed-rich heathland vegetation; (iv) soil removal and incorporation of heathland topsoil; and (v) heathland translocation. After 17 years the pH of the unamended agricultural soil remained significantly higher than that of the adjacent heathland. All treatments showed different trajectories of vegetation change in the long-term. Natural colonisation by heathland species was slow due to seed limitation, resulting in formation of an acid grassland community. Heathland community assembly was not facilitated by destruction of the initial grassland with herbicide. Incorporation of topsoil had an intermediate effect on pH reduction. This may explain the subsequent failure of the plant community to assemble in the anticipated proportions, and the dominance of leguminous scrub species (Ulex spp.). Turf translocation was effective in reducing pH to the required range and restoring the heathland community in the long-term. However, this technique should only be considered as a means of ‘rescue’ when habitat destruction is otherwise unavoidable. The only practical and sustainable means of increasing heathland extent on former farmland is the application of seed-bearing vegetation cut as part of routine management. However, this technique needs refining in order to establish the full range of characteristic heathland species.     

The potential for lowland heath regeneration following plantation removal

Historical records show that large areas of lowland heath have been planted with coniferous woodland. Such sites will increasingly be targeted for heathland restoration in order to achieve the objectives set by current conservation policy. Therefore information on management to achieve effective restoration is a priority for research. In this study we examined the size, composition and distribution of the seed bank beneath conifer plantations of different ages at two important heathland sites in Britain. Changes in the physical and chemical properties of heathland soil under the plantations were also described. There were significant, exponential declines in the mean density of viable heather seeds with plantation age at both sites. However, relatively large seed banks survived for 40 years and under some circumstances for more than 70 years, suggesting that restoration on first rotation sites is unlikely to be seed limited. There were small, but significant differences in the soil nutrient status under the plantations compared to the adjacent heaths. It was concluded that the main constraint on heathland regeneration is likely to be the burial of the seed bank by a thick layer of conifer leaf litter. The mechanical removal of this layer has been shown to be practically feasible.     

Seasonal variations and effects of nutrient applications on N and P and microbial biomass under two temperate heathland plants

Eutrofication is a threat against nutrient-poor habitats as increased amounts of nutrients in ecosystems may cause changes in the vegetation. Nitrogen (N) deposition leads to conversion of Calluna heathlands into graminoid dominated heath, but low availability of P may hinder or slow down this process.In this study the soil properties under two dominant heathland plants, the dwarf shrub Calluna vulgaris and the grass Deschampsia flexuosa, were investigated, with focus on nutrient content in the organic top soil and soil microbes during the main growing season and effects of nutrient amendments. The concentration of inorganic and dissolved organic N was significantly higher under D. flexuosa than C. vulgaris all though there were the same amounts of total N in the soil below the two species. N and P amendment enhanced available N and P in the soil, but added nutrients had little direct effects on microbes. The microbial biomass on the other hand was positively related to soil water content in fertilized plots indicating that this was due to an indirect effect of enhanced nutrient availability. Microbial N and P pools were respectively 1000 and 100 times higher than the pool of inorganic N and P, and microbes therefore may play an important role in regulating plant nutrient supply. Judged from responses of inorganic and microbial N and P concentrations to added N and P, N seemed to limit C. vulgaris and soil microbes below while P seemed to limit D. flexuosa and soil microbes below this species. There were lower rates of net nitrification, net ammonification and DOC and DON production rates during winter in the soil under C. vulgaris than below D. flexuosa, although all these rates were equal under the two species on an annual basis. This indicates that these microbial processes were taking place during winter but were affected by exudates from C. vulgaris.     

Impact of sod-cutting and choppering on nutrient budgets of dry heathlands

Heathlands are endangered by both atmospheric nutrient deposition and natural succession. High-intensity management measures are considered necessary, as low-intensity measures (e.g. mowing, prescribed burning) are not able to compensate for atmospheric nutrient loads. Choppering (i.e. the near-complete removal of the O-layer) has several advantages over sod-cutting, including less waste material, faster vegetation recovery and lower costs. This raises the question addressed in this study as to the extent to which choppering and sod-cutting affect nutrient budgets in dry heathlands.We compared the quantities of N, Ca, K, Mg, and P removed by choppering and sod-cutting in the Lueneburg Heath (NW Germany). Nutrient balances were calculated by analysing atmospheric inputs, elevated leaching rates following management, and output due to the removal of above-ground biomass and humus horizons.Nutrient loss was particularly high after removal of O- and A-horizons. In contrast, increased leaching after management was of minor importance for nutrient budgets. Although considerably more nutrients were removed by sod-cutting than by choppering (e.g. N: 1712/1008 kg ha⁻¹), nutrient output by choppering was still sufficient to compensate for 60.7 years of net N-input. Choppering was able to remove more N per volume unit than sod-cutting due to higher N-contents in the organic layer than in the A-horizon. For this reason, choppering is more economical than sod-cutting and, thus, should be considered the preferable method at sites not dominated by Molinia caerulea. A combination of high-intensity measures with prescribed burning would appear to be suitable as this would ensure more selective removal of N.     

Conservation of isolated Atlantic heathlands in the Mediterranean region: effects of land-use changes in the Montseny biosphere reserve (Spain)

In the Mediterranean region, cycles of controlled burning combined with continuous grazing appear to have been an effective tool for maintaining isolated Calluna vulgaris heathlands in the form in which they occur in many places in the Atlantic region. Changes in land use and management of the mosaic of extensively exploited heathland and associated grassland over recent decades, such as bringing land into cultivation followed by its abandonment and the prohibition of fires has resulted in a process of transformation into new shrub communities with lower biodiversity. In the Mediterranean region, these changes are similar to those described in the Atlantic area, but encroachment occurs faster and could lead ultimately to afforestation by Mediterranean woodland.In a study area of 300 ha of heathland in the Spanish Mediterranean basin (specifically, in the Montseny Natural Park and Biosphere Reserve), comparison of present and former vegetation showed that shrub cover increased from 15% in 1967 to 32% in 2000. Broom (Cytisus scoparius) was the main invasive species in abandoned crop fields, whereas Mediterranean holm oak forest (Quercus ilex) increased by 18%. The surface area of fernlands doubled and C. vulgaris heathlands decreased from 35% to just 9% during the same period. Intermixed grasslands also decreased moderately and progressively from 4% to 3%.It seems probable that cycles of fires are more important in terms of shrub control and biodiversity conservation than continuous grazing alone, even at a high rate of stocking (four small ruminants per hectare per year). This encroachment process throws into relief the role that isolated habitats can play as a monitor of land use changes.     

The establishment of heathland vegetation on ex-arable land: the response of Calluna vulgaris to soil acidification

The UK Biodiversity Action Plan has identified the creation of lowland heathland as an important objective. Heathland restoration studies have identified soil pH, elevated soil nutrients and large weed seed banks as major problems in the restoration of heathland vegetation on ex-arable land. Heathland vegetation is usually found on nutrient-poor acidic soils. Creating acidic soil conditions on ex-arable sites thus may produce a suitable environment for the establishment of heath vegetation. Soil acidification by the addition of sulphur has been shown to reduce the soil pH and the availability of nutrients in arable soils. A series of experiments was established to investigate the effects of soil acidification using sulphur on the establishment of Calluna vulgaris and the development of weed vegetation. The application of sulphur at 0.24 kg m⁻² to an arable soil was found to increase the survival rate of C. vulgaris cuttings planted in it. The mechanism of increased C. vulgaris survival appeared to be by sulphur application significantly reducing the cover of arable weeds arising from the soil seed bank. Higher rates of sulphur application (0.36 and 0.48 kg m⁻²) resulted in the death of many C. vulgaris plants. However C. vulgaris seedlings were able to establish successfully on these ex-arable soils within 18-24 months following the addition of these levels of sulphur. The application of sulphur appears to offer a practical solution to heathland creation on ex-arable land. However, it may be necessary to provide an interval of between 18 and 24 months between the application of sulphur and the addition of C. vulgaris plants or seeds for the successful establishment of heathland vegetation.     

The nature of eurytopy in polyploid earthworm species in relation to their use in biological soil amelioration

Summary Many of the most widely distributed species of Lumbricidae are polyploid. So far polyploid forms have been found to be distributed more widely than diploid ones. Taking Eisenia nordenskioldi as an example, it was shown that polyploids are able to occupy new ecological niches within the area of the diploid form. Ecological differences observed in representatives of different chromosomal races provide polyploids with the chance to coexist in one biocurrent, occupying different ecological niches.Dedicated to the late Prof. Dr. M.S. Ghilarov     

HeathMod: a model of the impact of seasonal grazing by sheep on upland heaths dominated by Calluna vulgaris (heather)

The dwarf shrub heather Calluna vulgaris is an important component of the UK landscape, with value for amenity, nature conservation and animal production. A model, called “HeathMod”, is described, which investigates the impact of grazing on upland heather. It combines an empirical model of production with a simple process-based model of grazing impacts. Validation from short-term studies in the literature indicated that the model predicts grazing impacts well, although more studies of grazing impacts are required to confirm the model’s accuracy. Analysis of the model allowed derivation of a simple general formula for predicting sustainable levels of grazing, given knowledge of local site productivity. Predicted levels of sustainable grazing were lower than previous estimates, mainly because the model predicts the long-term impacts of sustained grazing. Application of the model is most appropriate in areas where heather is in mixed-age stands, because there is inadequate information available to model interactions between grazing and degeneration in even-aged stands of heather.     

Five years of simulated atmospheric nitrogen deposition have only subtle effects on the fate of newly synthesized carbon in Calluna vulgaris and Eriophorum vaginatum

To understand the implications of atmospheric nitrogen deposition on carbon turnover in peatlands, we conducted a ¹³C pulse labeling experiment on Calluna vulgaris and Eriophorum vaginatum already receiving long-term (5 years) amendments of 56 kg N ha⁻¹ y⁻¹ as ammonium or nitrate. We examined shoot tissue retention, net ecosystem respiration returns of the ¹³C pulse, and soil porewater DOC content under the two species. ¹³C fixation in Eriophorum leaves was enhanced with nitrogen addition and doubled with nitrate supply. This newly fixed C appeared to be relocated below-ground faster with nitrogen fertilization as respiration returns were unaffected by N inputs. By contrast, increases in ¹³C fixation were not observed in Calluna. Instead, net ecosystem respiration rates over Calluna increased with N fertilization. There was no significant label incorporation into DOC, suggesting a conservative strategy of peatland vegetation regarding allocation of C through root exudation. Greater concentrations of total DOC were identified with nitrate addition in Calluna. Given the long-term nature of the experiment and the high N inputs, the overall impacts of nitrogen amendments on the fate of recently synthesized C in Eriophorum and Calluna in this ombrotrophic peatland were surprisingly more moderate than originally hypothesized. This may be due to N being effectively retained within the bryophyte layer, thus limiting, and delaying the onset of, below-ground effects.     

Soil feedback effects to the foredune grass Ammophila arenaria by endoparasitic root-feeding nematodes and whole soil communities

In coastal foredunes, the grass Ammophila arenaria develops a soil community that contributes to die-back and replacement by later successional plant species. Root-feeding nematodes and pathogenic soil microorganisms are involved in this negative feedback. Regular burial by wind-blown beach sand results in vigorous growth of A. arenaria, probably because of enabling a temporary escape from negative soil feedback. Here, we examine the role of root-feeding nematodes as compared to the whole soil community in causing negative feedback to A. arenaria. We performed a 3-year sand burial experiment in the field and every year we determined the feedback of different soil communities to plant growth in growth chamber bioassays.In the field, we established A. arenaria in tubes with beach sand, added three endoparasitic root-feeding nematode species (Meloidogyne maritima, Heterodera arenaria and Pratylenchus penetrans) or root zone soil to the plants, and created series of ceased and continued sand burial. During three subsequent years, plant biomass was measured and numbers of nematodes were counted. Every year, bioassays were performed with the field soils and biomass of seed-grown A. arenaria plants was measured to determine the strength of feedback of the established soil communities to the plant.In the field, addition of root zone soil had a negative effect on biomass of buried plants. In the bioassays, addition of root zone soil also reduced the biomass of newly planted seedlings, however, only in the case when the field plants had not been buried with beach sand. Addition of the three endoparasitic root-feeding nematodes did not influence plant biomass in the field and in the bioassays. Our results strongly suggest that the negative feedback to A. arenaria is not due to the combination of the three endoparasitic nematodes, but to other components in the soil community, or their interactions with the nematodes.     

Re-establishment of suppressiveness to soil- and air-borne diseases by re-inoculation of soil microbial communities

The aim of this study was to investigate the potentials and limitations in restoring soil suppressiveness in disturbed soils. Soils from three sites in UK and Switzerland (STC, REC, THE) differing in their level of suppressiveness to soil-borne and air-borne diseases were γ-irradiated and this soil matrix was re-inoculated with 1% (w/w) of either parent native soil or native soil from the other sites (‘soil inoculum’). Suppressiveness to air-borne and soil-borne diseases was quantified by means of the host-pathogen systems Lepidium sativum (cress)-Pythium ultimum, an oomycete causing root rot and seedling damping-off, and Arabidopsis thaliana-Hyaloperonospora parasitica, an oomycete causing downy mildew. Soil microbial biomass, activity and community structure, as determined by phospholipid fatty acid (PLFA) profiles, were measured in native, γ-irradiated, and re-inoculated soils. Both, L. sativum and A. thaliana were highly susceptible to the pathogens if grown on γ-irradiated soils. Re-inoculation completely restored suppressiveness of soils to the foliar pathogen H. parasitica, independently of soil matrix or soil inoculum, whereas suppressiveness to P. ultimum depended on the soil matrix and, to a lesser extent, on the soil inoculum. However, the soil with the highest inherent suppressiveness did not reach the initial level of suppressiveness after re-inoculation. In addition, native microbial populations as defined by microbial biomass, activity and community structure, could not be fully restored in re-inoculated soils. As for suppressiveness to P. ultimum, the soil matrix, rather than the source of soil inoculum was identified as the key factor for re-establishing the microbial community structure. Our data show that soils do not or only slowly fully recover from sterilisation by γ-irradiation, indicating that agricultural soil management practices such as soil fumigation or heat treatments frequently used in vegetable cropping should be avoided.     

Moorland restoration aids the reassembly of associated phytophagous insects

A change in the vegetation of UK moorland from Calluna vulgaris-dominated heath to Molinia caerulea- or Nardus stricta-dominated grassland is frequently deemed undesirable on economic and biodiversity grounds. Moorland vegetation restoration is an increasingly practiced management option. However there has been little research on the effectiveness of such management for aiding the restoration of other aspects of moorland biodiversity such as associated insect populations.We sampled Hemiptera and Lepidoptera assemblages from restored, degraded and C. vulgaris-dominated (target) moorland at eight restoration sites in England and Scotland. Vegetation assemblages and additional environmental variables were sampled at the same locations.Vegetation restoration aided the reassembly of moorland insects to a variable extent. Hemiptera restoration success was greatest at mechanically restored sites dominated by C. vulgaris. In addition to treatment, latitude best explained variation in the Hemiptera assemblage. A range of plant species, all either dwarf shrubs or graminoids, acted as further determinants. Lepidoptera restoration success was correlated most closely with the success of restoring the full diversity of the vegetation assemblage and was highest at sites restored by grazing control.The results demonstrate that vegetation restoration brings wider biodiversity benefits and that the form these take may depend on the restoration method used and consequent characteristics of the resultant vegetation assemblage.     

野葛及百喜草对红壤坡地水土保持及土壤改良的效应

试验研究红壤坡地野葛水土保持和土壤改良效应结果表明,横峰野葛和百喜草可有效增大土壤持水量、土壤孔隙度和降低土壤容重,提高土壤pH值、有机质和N、P、K含量,减少土壤侵蚀量和径流量,水土保持和改良土壤效果明显,且野葛综合效果优于百喜草。     

The effect of turf cutting on plant and arbuscular mycorrhizal spore recolonisation: Implications for heathland restoration

In two natural heathland vegetations, we analysed the effect of turf cutting on spore numbers of arbuscular mycorrhizal fungi (AMF). Next to this, we performed a controlled factorial experiment to examine the role of AMF for germination and establishment of Arnica montana in both turf cut and non-turf cut situations. AMF spore numbers decreased with soil depth, and, along with the topsoil, almost all AMF spores were removed with the removal of the acidified and/or eutrophied organic layer. Recolonisation of AMF spore numbers after turf cutting was slow: spore numbers of approximately 60-95 spores g⁻¹ dry soil were found two and a half years after turf cutting, corresponding with 55-70% of AMF spore numbers found in natural field populations of A. montana. Since AMF colonisation increased establishment and biomass, and decreased mortality of A. montana, it was suggested that lack of AMF after turf removal might complicate the establishment of this herbaceous species. Removal of organic material as a management measure should therefore carefully be applied, taking in consideration the low recolonisation rates of AMF as this can markedly effect the success of restoration.     

A field assessment of the role of selective herbicides in the restoration of British moorland dominated by Molinia

Upland heather moors are high priority biotopes for conservation in Britain, but are under threat from a number of sources. One such threat is a change from a vegetation dominated by dwarf shrubs, mainly Calluna vulgaris, to a grassland dominated by Molinia caerulea. Restoration of a dwarf shrub community has proved problematic with studies showing that herbicide use may be necessary to control Molinia. Previously the non-selective herbicide, glyphosate, has been recommended, but this herbicide may damage Calluna and other moorland species severely. Here, we tested a range of selective herbicides (graminicides) for use in an herbicide-based conservation strategy under field conditions. Two selective herbicides, propaquizafop and quizalofop-ethyl, gave a short-term check to Molinia and another, cycloxydim, provided a reduction for at least 1 year, but this effect disappeared after 3 years. Damage to Calluna was less than that caused by glyphosate, and the selective herbicides had little effect on other moorland species present. Our findings suggest that selective herbicides have a potential role in reducing Molinia during moorland restoration, although follow-up applications, and appropriate moorland management, will also be needed.     

Biological soil amelioration as the basis of sustainable agriculture and forestry

Before the discovery of plant nutrients, soil amelioration and fertilization was by force of circumstance biological, i.e. done with substances taken from nature. After the discovery of plant nutrients, the chemical concept of fertility displaced the biological one. The defects of chemical cultivation — inefficiency, deterioration in product quality, diminishing productivity of soils and negative effects on the environment — have created an urgent need for the study of fertility as a result of the activity of the biocycles of the ecosystem. With the aid of the advances of modern science, we can understand the defects and deficiencies of the chemical concept of fertility and return the biological concept of fertility to its proper place. This in turn is a good starting point for productive research and developmental work towards an ecologically sustainable agriculture and forestry.     

Are meadows with high ecological and patrimonial value endangered by heathland invasion in the French central Pyrenees?

Due to their high ecological and patrimonial value Festuca eskia and Nardus stricta meadows and Rhododendron ferrugineum heathlands need to be locally preserved in the Pyrenees. This could pose dilemmas to conservation managers if heathlands invade and replace the meadows. However, studies showing that the heathlands do actually invade meadows are lacking in the Pyrenees and the invasion hypothesis therefore remains to be proved. So, vegetation pattern changes in a particularly vulnerable site from the central Pyrenees was investigated by two means: (i) interpretation of stereo pairs of aerial photographs and (ii) analysis of the demography of Rhododendron populations in ten 150 m² experimental plots. The main results from photo interpretation indicated that, during the period between 1954 and 1995, 15% of a 17 ha meadow area shifted to heathland. This corresponded with a 592 m² year⁻¹ colonization rate and to an overall front progression of heathland over meadows of 3-60 m. Demographic data showed that meadow colonization by heathland passed through successive phases but appeared to be a rather slow process that can take as long as 120-320 years. During the process the cover rate over the 150 m² plots varied between 1 and 0.4 m² year⁻¹. Heathland extension does not seem to have been favored by the decrease in grazing pressure. Despite this, the progression of heathlands over meadows seems endangering the meadows. Finally we propose practices locally to limit the progression of Rhododendron.