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.     

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.     

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.     

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.     

Influence of soil type and indigenous pathogenic fungi on bean hypocotyl rot caused by Rhizoctonia solani AG4 HGI in Cuba

Calcisol, ferralsol and vertisol soils, representative of different bean production areas of Villa Clara province in Cuba, were selected to determine the impact of soil type on bean hypocotyl rot severity caused by Rhizoctonia solani AG4 HGI (isolate CuVC-Rs7). In inoculated autoclaved soil, hypocotyl rot was most severe in calcisol soil, followed by ferralsol soils and then vertisol soils. In inoculated natural soils, disease severity was lower in vertisol and calcisol soils and higher in ferralsol soil, indicating that biological factors are suppressing or stimulating the pathogenic efficiency of R. solani. Native binucleate Rhizoctonia AGF, Sclerotium rolfsii and R. solani AG 4 HGI were isolated from bean plants grown in natural calcisol, vertisol and ferralsol soils, respectively. Subsequent studies about the interaction between these fungi and R. solani indicated that they were involved in the variability of disease severity caused by R. solani. The addition of R. solani AG4 HGI (isolate CuVC-Rs7) into each autoclaved soil inoculated with binucleate Rhizoctonia or S. rolfsii resulted in a reduction of disease severity caused by this pathogen while in soils inoculated with native R. solani AG4 HGI, disease severity increased. Irrespective of fungal interactions, calcisol was always the most disease conducive soil and vertisol the most disease repressive soil. The mechanisms by which native pathogenic fungi could influence disease severity caused by R. solani are discussed.     

Fraxinus-Glomus-Pisolithus symbiosis: Plant growth and soil aggregation effects

It has been established that arbuscular mycorrhizal (AM) fungi are involved in the conservation of soil structure. However, the effect of ectomycorrhizal (EM) fungi alone or in interaction with AM fungi in soil structure has been much less studied. This experiment evaluated EM and AM fungi effects on soil aggregation and plant growth. Ash plants (Fraxinus uhdei) were grown in pots, and were inoculated with Glomus intraradices and Pisolithus tinctorius separately but also in combination. Our results showed that F. uhdei established a symbiotic association with EM and AM fungi, and that these organisms, when interacting, showed synergistic and additive effects on plant growth compared to singly inoculated treatments. EM and AM fungi prompted changes in root morphology and increased water-stable aggregates. AM fungi affect mainly small-sized macroaggregates, while EM and EM-AM fungi interaction mainly affected aggregates bigger than 0.5 mm diameter. These results suggest that ectomyccorrhizal as well as arbuscular mycorrhizal fungi should be considered in restoration programs with Fraxinus plants.     

Attraction to prey and prey-associated odours by the predatory mite Hypoaspis aculeifer in a soil experimental system

In terms of food searching and trophic interactions among soil organisms volatile signalling plays a crucial role. Still, volatile-based foraging decisions are poorly understood, especially as the dispersal and perception of cues is more restricted within the soil pore space than in simple laboratory approaches where volatiles disperse freely in the air. Focusing on the tritrophic interaction between the soil fungus Trichoderma viride, the fungivorous collembolan Folsomia candida and the predatory mite Hypoaspis aculeifer, we tested the effect of prey and prey-associated volatile signals on the food choice decision of the predatory mite in a laboratory experiment under soil conditions as well as the possible occurrence and relevance of grazing-induced indirect defence signals. The mites showed a clear preference for ungrazed and grazed fungal mycelium. However, we found no evidence of induced indirect defence as ungrazed mycelium was preferred to grazed fungus.