Impact of prescribed burning on the nutrient balance of heathlands with particular reference to nitrogen and phosphorus

Question: Can prescribed winter burning compensate atmospheric nutrient loads for dry heathlands? What effects does prescribed burning have on nutrient balances, particularly as regards the limiting nutrients N and P? Location: Lueneburg Heath, NW Germany. Methods: In two burning experiments (in 10/15 year old Calluna‐stands) nutrient balances (for N, Ca, K, Mg, P) were calculated by analysing nutrient inputs (atmospheric deposition, ash deposition), nutrient stores (above‐ground biomass, organic horizon) and nutrient outputs (biomass combustion, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha^‐1.a^‐1 for N and < 0.5 kg.ha^‐1.a^‐1 for P. Nutrient stores in the above‐ground biomass were 95/197 kg.ha^‐1 for N and 5/13 kg.ha^‐1 for P (first/second experiment, respectively). From these stores 90/53% (for N) and 25/14% (for P) were removed by burning. Effects of leaching on nutrient balances were low. In the first two years after burning, leaching rates of N increased by about 4/6 kg.ha^‐1, whereas leaching rates of P did not change significantly. Input/output‐ratios showed that prescribed burning leads to positive nutrient balances for N, Ca and Mg in the long term. For example, the amounts of N removed by prescribed burning are equivalent to ca. five years of atmospheric inputs. Applied in ten‐year cycles, this measure alone cannot prevent N accumulation in the long term. Conclusion: Regarding 10/15 year old Calluna‐heaths, we assume that prescribed burning cannot compensate for atmospheric N inputs, thus making long‐term changes in the nutritional state inevitable. Therefore, prescribed burning should be applied in combination with high‐intensity management measures.     

Seedling emergence after sod cutting in grass heath

Abstract. Experimental plots in grass heath, previously subjected to various sod cutting treatments, contained a seed store of ericoid dwarf shrubs, but seeds of grasses of the former grass heath were not found. The density of emerging dwarf-shrub seedlings in the plots was determined mainly by soil moisture, but with density of seeds stored in the soil and soil acidity as modifying factors. It is concluded that the restoration of a dwarf-shrub heathby sod cutting is relatively successful, as compared to other measures, because the grass seeds become obsolete.     

Effects of nutrient supply,light availability and herbivory on the growth of heather and three competing grass species

Grasses are becoming more abundant in areas in NE Scotland which until recently were dominated by heather (Calluna vulgaris). However, it is not clear if grasses are aggressive competitors which are now able to outcompete the dwarf shrub due to changes in environmental factors (such as grazing pressure and increasing nutrient inputs), or just opportunistic invaders, occupying gaps in the canopy which occur when heather reaches the degenerate stage. Experiments in turves and in field plots were carried out in order to investigate the performance of three grass species, Nardus stricta, Deschampsia cespitosa and Deschampsia flexuosa growing in competition with heather. These three species were selected because they differ in their nutrient requirements, palatability to herbivores and tolerance of shading. The grasses were planted in heather canopies of different structure, either turves of heather of different height and age, or moorland plots with or without heavy grazing by sheep and deer. Fertiliser (NPK) was applied to half the experimental plants. The growth of the grass species and the heather in response to the fertiliser and grazing treatments was measured, together with the light levels penetrating the canopy and received by the grass plants.Results indicated that heather was likely to be outcompeted by grasses only when there are gaps in the canopy, resulting either from heavy grazing or from the heather being in the mature or degenerate phase. Fertiliser enhanced plant growth whereas fencing out herbivores led to strong competition for light as the heather canopy closed. It is concluded that grasses require gaps in the canopy to successfully invade heather moorland, or they tend to be shaded out. Thus better management of heather moorlands to maintain a dense canopy structure may help to preserve heather cover even under increasing nutrient inputs.     

Competition between heather and grasses on Scottish moorlands: Interacting effects of nutrient enrichment and grazing regime

Abstract. Considerable losses and degradation of heathlands (in moorlands and lowlands) have been reported across Europe, with Calluna vulgaris (heather) being replaced by other species, often grasses. Increasing atmospheric nitrogen deposition and overgrazing have been suggested as the driving factors behind this change. This possibility was investigated in a study of the interacting effects of nutrient inputs and grazing on heather and three grass species (Nardus stricta, Deschampsia cespitosa and D. flexuosa) in the field, on a moorland in northeastern Scotland. In addition, the interacting effects of increasing nutrients and Calluna canopy height on N. stricta and D. cespitosa were studied using turves in an outdoor experimental area. In the field, fencing had a larger effect than fertilizer on the growth of all species, except for N. stricta, the species most unpalatable to herbivores. Fencing led to an increase in the height of the Calluna canopy, which may reduce light availability for the grasses. In the turf experiment, the height of the Calluna canopy affected the diameter of the grass tussocks and percentage of green matter (i.e. live leaf material), with plants under the more closed Calluna canopies being smaller. This study suggests that the slow‐growing, evergreen Calluna is a more effective competitor than the faster growing grasses when it has a tall, intact canopy, even at increased levels of nutrient supply. However, overgrazing promotes gap formation in the Calluna canopy, providing an opportunity for grasses to take advantage of increased nutrients. Thus the conservation of heather moorlands requires an understanding of the grazing level which allows Calluna to maintain sufficient canopy structure to outcompete grasses for light.     

Influence of management on the seed production and seed bank of calcareous fen species

Abstract. The dynamics of the seed production and seed bank of two dominant perennial species of calcareous fens were investigated in response to different mowing frequency, fallow or pasture. The production of fertile shoots responded strongly to management practices. On sites where mowing alternated with fallow, autumn mowing led to an increase of fertile shoot density in Schoenus ferrugineus and a decrease in Molinia caerulea in the subsequent season. Repeated annual mowing after long-term fallow raised the density in Schoenus and inhibited initiation of flowering shoots in Molinia. Seed set and viability were influenced mainly by climatic or other factors. Variation of seed production between sites and in consecutive years at the same site could thus be large. The relationship between total seed production and seed bank was weak, but it improved when germination rates were included in the calculation. As the seed bank was generally smaller than the viable seed production, low survival and a high turnover of seeds in the upper soil layers is assumed for both species. Implications for recruitment strategies are discussed.     

Photosynthesis of ground vegetation in different aged pine forests: Effect of environmental factors predicted with a process‐based model

Question: How do stand age and environmental factors affect the species‐specific photosynthesis of ground vegetation? Location: Five different aged pine forests in Southern Finland. Methods: We measured photosynthesis of common species of ground vegetation during the growing season of 2006. Results: The measured vascular species, especially those with annual leaves, had a clear seasonal cycle in their measured photosynthetic activity (P_maxⁱ). A simple model that uses site‐specific temperature history, soil moisture and recent frost as input data was able to predict the changes in photosynthetic activity in dwarf shrubs with perennial leaves. The P_maxⁱ values of mosses did not have a clear seasonal cycle, but low values occurred after rain‐free periods and high values after precipitation. We modified the model for mosses and included temporary rain events. The model was able to predict most of the large changes in P_maxⁱ of mosses resulting from varying weather events but there was still some uncertainty, which was probably due to difficulties in measuring fluxes over a moss population. Conclusions: Temperature history, recent frosts and soil moisture determine the changes in P_maxⁱ of dwarf shrubs with perennial leaves. The P_maxⁱ of mosses depends mostly on recent precipitation.     

Lignin signatures show the effects of changes in heather and bracken cover on the composition of organic matter in a moorland soil profile

Bracken (Pteridium aquilinum) is aggressively displacing heather (Calluna vulgaris) on many moorlands in Britain. We investigated the use of lignin derivatives to identify the distribution of soil organic matter (SOM) derived from bracken in moorland soil profiles formed under heather. Phenylpropanoids extracted from recently senesced litters, roots and SOM, using alkaline CuO oxidation, showed distinct signatures for bracken and heather, with vanillyl moieties dominating bracken litter extracts and vanillyl and syringyl dominating heather litter extracts. Ratios of vanillyl and syringyl concentrations characterised the SOM derived from heather and bracken better than the concentrations of the individual moieties. The analysis showed up to a depth of 5 cm under pure bracken cover, and at the interface between heather and bracken, the SOM was largely derived from bracken litter but below that depth SOM was apparently derived from heather. The use of these methods to identify the plant origin of SOM not only enables understanding the effects of changing vegetation cover on organic matter dynamics in moorland soils but could also facilitate management techniques in moorland/heathland restoration which involve the removal of comparatively nutrient-rich SOM derived from bracken. Received: 30 November 1997 / Accepted: 11 April 1998     

The effect of increased nutrient availability on vegetation dynamics in wet heathlands

A three year fertilization experiment was conducted in which nitrogen (N series: 20 g N m^–2 yr^–1), phosphorus (P series: 4 g P m^–2 yr^–1) and potassium (K series: 20 g K m^–2 yr^–1) were added to a mixed vegetation of Erica tetralix and Molinia caerulea. At the end of each growing season the percentage cover of each species was determined. At the end of the experiment percentage cover of each species was found to be positively correlated with the harvested biomass. In the unfertilized control series the cover of Erica and Molinia did not change significantly during the experiment. In all fertilized series however, especially in the P series, cover of Erica decreased significantly. The cover of Molinia increased significantly in the P series only.In the fertilized series the biomass of Erica and total biomass per plot did not change significantly compared with the control series. In the P series the biomass of Molinia increased significantly.It is suggested that with increasing phosphorus or nitrogen availability Molinia outcompetes Erica because the former invests more biomass in leaves which in turn permits more carbon to be allocated to the root system, which thereupon leads to a higher nutrient uptake.     

Effects of N-deposition on ion trapping by B-horizons of Danish heathlands

Coastal and inland heaths constitute a dominating fraction of the relatively few semi-natural ecosystems in Denmark. At the inland heath, the soil forming factors have been active with the same vegetation type during the last 2000 years, whereas at the coastal heath, these soil forming factors have only been active over a couple of hundred years, due to the activity of windblown sand. Effects on ion-balance and soil processes of nitrogen deposition on a young dune heath and an old inland heath have been investigated. Different levels of NH₄NO₃have been applied to the two heathlands, and rain, throughfall water and soil water chemistry below the experimental plots have been analysed. Results show that the ability to retain added, as well as deposited, nitrogen requires the presence and the integrity of a humifyed H-subhorizon. When the mor-layer for some reason breaks down, the nutrients will be retained by the B-horizons as a kind of back-up, and most probably be reused by the new vegetation. One year after destructive heather beetles attack on the inland heath, the mor-layer started to decompose. A large translocation of organic compounds, complexed associated cations and nitrogen-species, started from the mor-layer to the top of the B-horizon where they precipitate. The podzolic B-horizons can be seen as nutrient trap to the mor-layer. This revised version was published online in June 2006 with corrections to the Cover Date.     

Re-establishment of dominance by dwarf shrubs on grass heaths

Experimental fields were established at three sites on different soil types in Dutch heathlands, where grass species have become dominant, in order to find methods for re-establishment of an ericoid dwarf-shrub heath. Treatments included mowing, mowing and cutting for hay, ploughing, milling, sod cutting and burning.Re-establishment of ericoid dwarf shrubs was only observed after creation of gaps or patches of open soil. The results after nine years for two of the sites and six years for the other site indicate that sod cutting is the most suitable method provided that the mineral top soil is not removed or disturbed. More profound removal of the sod including mineral soil causes a delay in re-establishment of ericoid dwarf shrubs and an increase of grasses.Factors that may explain different responses to gap creation including availability of seeds, soil moisture, soil fertility and plant survival strategies are briefly discussed.Abbreviations Calluna = Calluna vulgaris - Deschampsia = Deschampsia flexuosa - Erica = Erica tetralix - Molinia = Molinia caerulea - Sarothammus = Sarothammus scoparius     

Effects of habitat fragmentation and soil quality on reproduction in two heathland Genista species

Habitat fragmentation decreases plant population size and increases population isolation, as well as altering patterns of plant–animal interactions, all of which may reduce plant fitness. Here, we studied effects of habitat fragmentation (in terms of population size and isolation) and soil quality on the reproduction of two rare legume species, Genista anglica (13 populations) and Genista pilosa (14 populations), confined to remnants of acidic and nutrient‐poor Calluna heathlands. Single individuals of the Genista plants are impossible to distinguish; population size was therefore estimated according to the area occupied (referred to as population size hereafter). We collected soil samples in all heathland sites to determine content of soil water, C, N, P, Ca, K and Mg. In both species values of soil pH and C/N ratio, as well as content of soil P and base cations, reflected the highly acidic and nutrient‐poor environment of the heathlands. Population sizes were unrelated to soil quality. Although the two Genista species are similar in morphology and ecology, effects of explanatory variables on reproduction were largely inconsistent across species. In G. anglica, population size had a positive impact on all reproductive variables except germination rate, which, in contrast, was the only variable affected positively by population size in G. pilosa. In both species, mean total reproductive output, calculated as the product of total seed mass per shoot and total germination, increased with increasing water content and decreased with increasing P. In G. anglica, we found positive effects of the C/N ratio on all reproductive variables except mean single and total seed mass per shoot. In summary, in both species reproductive success per shoot decreased with increasing soil nutrient availability in the heathland sites. The infestation of two large populations of G. pilosa with the pre‐dispersal, seed‐predating weevil Apion compactum had no significant effect on reproduction of the populations.     

Effects of nitrogen addition on the stress sensitivity of Calluna vulgaris

The effects of low levels of nitrogen addition (7·7 and 15·4 kg N ha⁻¹ yr⁻¹) on plant sensitivity to biotic and abiotic stress were studied at a lowland heath in the south of England that has received N treatments since 1989. Larval growth rates and adult weights of heather beetles were found to be significantly higher when insects were reared on plants that had received additional N, with implications for insect survival and reproductive success. Electrolyte leakage measurements failed to reveal any significant impact of N addition on plant sensitivity to frost episodes in early winter. In April, however, there was some evidence of slightly decreased frost hardiness in plants receiving additional N. Accelerated spring bud burst also suggested earlier physiological activity in N-treated plots. The rate of water loss from excised shoots of Calluna vulgaris (L.) Hull was significantly faster in plants receiving additional N, although no difference in plant water potential was measured in the field after a prolonged dry spell. Whilst experimental N addition had only a small effect on plant sensitivity to abiotic stress, the relationship between enhanced deposition and increased insect performance was clear, with the potential for substantially increased insect damage at deposition rates around the critical load contributing to the formation of gaps in the Calluna canopy.