The occurrence of former heathland vegetation in the coastal areas of the south-east Baltic sea,in particular Lithuania: a review

Previously published pollen diagrams from the coastal area of the south-east Baltic Sea including parts of the Polish and Lithuanian peninsulas, Mierzeja Wilana (Frische Nehrung) and Kuri Nerija (Courland Spit) respectively, document the former presence of heathland vegetation and an associated burning practice. This type of human influence can be dated back to ca. 1200 B.P. at the latest, and was maintained until the 18th century. It is suggested that this former Calluna-dominated vegetation belonged to the known west to north-west European heathland vegetation. Several dwarf shrub taxa, such as Genista spp. and Erica tetralix, normally connected to heathland vegetation, may have had a former wider distribution.     

Transition from heathland to grassland initiated by the heather beetle

This paper discusses the transition from heather-dominated vegetation to one dominated by grass. The dominating species involved are Calluna vulgaris and Deschampsia flexuosa. The study covered three years, 1980–82, and was done at the Hoorneboeg heathland in the Netherlands. Several factors account for this transition: damage caused by the chrysomelid heather beetle Lochmaea suturalis Thomson, soil type, and interaction between beetle damage and Deschampsia when already present underneath or in the immediate vicinity of a Calluna canopy. There was a time-lag between beetle damage and grass establishment or expansion. In fact, this transition concerns a type of vegetation succession and can be treated as such. This succession therefore involves several causative factors external to the system Calluna-Deschampsia. The successional process is conceived as a stochastic one defined by transition probabilities which depend on a time component, a space component and a logistic response component. All these components are discussed in this paper, but the discussion of yet another component, the crowding of Deschampsia itself, will be treated separately.     

Effects of a heather beetle attack on soil moisture and water balance at a Danish heathland

The effects of reclamation and high atmospheric nitrogen deposition have for long threatened the existence of heathlands in Denmark. A high nitrogen input increases the frequency and intensity of heather beetle attacks. However, any indirect effects of these attacks on the soil water balance are seldom investigated. In autumn 1994 a 2000-year old Danish inland heath was struck by a severe heather beetle attack and the effects on the soil moisture and the water balance were studied. Soil water content, gross precipitation and throughfall were measured continuously from 1993 to 1998 at the heath. The first signs of the attack on the water balance were seen in the dry summer of 1995 when the soil water content was relatively high. Four years after the beetle attack, new heather plants covered the area again and during summer the soil water seemed to be depleted to the same degree as before the beetle attack. In the years after the beetle attack a high coefficient of variation between individual soil moisture measurements was seen. It is proposed that the inhomogeneous wetting was caused by heterogeneous throughfall, water-repellent soil and break down of the structure of the organic top-horizons due to the beetle attack. The effect of the beetle attack was examined using a simple water balance model. Model simulations showed that evapotranspiration was reduced by respectively, 14, 29 and 5% in the three years following the beetle attack. From 1993 to 1998 percolation was on average 62% of precipitation with very little variation from year to year. Evapotranspiration was on average 38%, but in the years affected most by the beetle attack transpiration was relatively low whereas evaporation from soil was increased. A comparison between the modelled and measured throughfall, as well as percolation estimated by the chloride mass balance method, showed that the water balance parameters were estimated well in the two years which were most affected by the beetle attack, i.e. 1995 and 1996.     

The heathers of Europe and adjacent areas

McCLINTOCK, D., 1989. The heathers of Europe and adjacent areas. Andromeda polifolia, Bruckenthalia spiculifolia, Calluna vulgaris , two species of Daboecia , 19 of Erica with five subspecies and seven natural hybrids are discussed. Comment is chiefly given on E. andevalensis, E. maderensis and the group currently denominated: E. anthura, E. manipuliflora, E. multiflora and E. vagans. Two new combinations are made.     

Differential effects of ammonium and nitrate on three heathland species

Arnica montana and Cirsium dissectum, typical of species-rich heathlands and acidic grasslands, have declined rapidly in The Netherlands in recent years. Field surveys suggest that the decline is caused by soil acidification as a result of enhanced atmospheric N and S deposition. Therefore, the survival, growth and development of these species were studied in a water culture experiment, using nutrient solutions which differed both in mineral nitrogen form and in ammonium concentration. For comparison, the performance of a third, acid tolerant species, Calluna vulgaris, was studied. The results showed that both Arnica and Cirsium performed better using nitrate than when using ammonium as a sole nitrogen source, whereas ammonium toxicity became apparent when ammonium concentrations were raised above 100 µM. Ammonium toxicity was expressed by an increase in mortality of Arnica plants with increasing ammonium concentrations and by a reduction of biomass in Arnica and Cirsium. Furthermore, cation concentrations in both roots and shoots decreased when ammonium was supplied as a nitrogen source. In contrast, Calluna showed optimal development when using ammonium as a sole nitrogen source. In this species, only root biomass was negatively affected by high ammonium concentrations. The ecological implications of these preferences are discussed in relation to soil acidification.     

Leaf-nutrient accumulation and turnover at three stages of succession from heathland to forest

Abstract. Accumulation of nutrients in leaves of the dominating species of three ecosystems, characterizing the secondary succession from Genisto-Callunetum heathland through Leucobryo-Pinetum birch-pine woodland to mature Querco-Fagetum oak-beech forest, as well as nutrient turnover within these ecosystems was studied. The objective of the study was to establish potential variations in quantity and quality of nutrient supply to the plants with respect to succession dynamics. The results show very low leaf nutrient concentrations of all species investigated, coinciding with low nutrient availability in the soil. However, the nutrient content of leaves and leaf litter of Quercus petraea and Fagus sylvatica, which dominate the late succession stages, and in Betulapéndula are higher than in the photosynthetic organs (leaves and young shoots) of Calluna vulgaris and Pinus sylvestris. The combination of the higher nutrient content of the leaves and an increasing leaf-litter production during succession results in an increased nutrient turnover via leaf-litter fall. However, due to the high leaf biomass, the storage of nutrients in the leaf biomass is highest within the birch-pine woodland. From this, it may be assumed that the low demand and the low loss of nutrients via leaf-litter fall are favourable for Pinus at the early stages of forest succession on poor sandy soils. In contrast, Quercus and Fagus are provided with better growth conditions at the later stages of succession resulting from the accumulation of plant-available nutrients in the ecosystem by Pinus sylvestris, combined with a higher nutrient turnover as compared with the heathland.     

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.     

Seed-shedding in heather (Calluna vulgaris (L.) Hull

Abstract

Background: Accumulating evidence, mainly from small-scale experiments, suggests that species diversity helps stabilise ecological communities; however, this relationship needs testing at larger scales in a wider range of natural communities.

Aims: In experimentally burnt ericaceous shrub stands, we aimed to determine whether more diverse stands had more stable vegetation cover.

Methods: Using two prescribed fire management experiments – one on heathland and one in forest – at scales of 100–700 m², we compared pre-disturbance vegetation characteristics (measures of diversity, traits and composition) with resistance (degree of perturbation on disturbance), resilience (here defined as post-disturbance recovery rate) and stability (the inverse of temporal variability).

Results: Responses to disturbance were usually best explained by a measure of dominance: Simpson's index. High dominance was associated with high resilience, but low resistance and low stability. Within the forest, the shrub community had lower dominance and higher stability than it did on heathland. Dominance and diversity were strongly linked as opposite poles of a principal axis of vegetation variation; therefore, more diverse vegetation was more stable.

Conclusions: Ericaceous shrub cover is known to support important ecosystem services, such as soil protection, carbon sequestration, flood prevention and livestock grazing. Our results suggest that high shrub diversity would support stable provision of these services. Land managers who prioritise this aim in British heathlands and pinewoods should consider a wider range of management approaches than those of typical current practice.     

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.