Floristic and life form diversity in post-pasture successions on a Mediterranean island (Corsica)

The landscapes of the French island of Corsica are the result of thegrazing tradition which has always occupied a pivotal position in the island'seconomy. Furthermore, in the last fifty years, a reduction in the pressure fromhuman activity has lead to changes in plant cover. This article intends tostudythe evolution of dynamic mechanisms during the secondary successions and theeffect of changes in vegetation on plant diversity. Our study was based on asimultaneous study of 375 plots carried out between May and September from 1996to 1998. We studied the floristic composition of each stage both qualitatively(life form, adaptive strategy and seed dispersal mode) and quantitatively(species richness, equitability, Shannon index). The results show a change inspecies and environmental conditions, with a progressive move from heliophilicgrassland and low shrub species with an S or SR strategy, anemochoric and withahigh tolerance for soil quality to a final stage of dense forest composed ofcompetitive (C or CS) zoochoric or barochoric phanerophytes. Moreover, theturning point in Corsica in terms of floristic diversity lies between 20 and 55years after land abandonment. Successions are due to a series of rapidinteractions. Thus, when a site is given reserve or protected area status, weneed to take into account the fact that it will evolve according to thebiological traits of the species. From a research point of view, we need tolookat process diversity and the role of each of the species present during thesuccessions.     

Effects of prescribed burning on plant available nutrients in dry heathland ecosystems

Heathland management is an important tool with which to modify ecosystem impacts caused by atmospheric nutrient deposition. Since changes in nutrient availability as a result of management measures affect the outcomes of heathland succession and species competition, studies on this issue are important from both a nature conservation and management point of view. This study reports the effects of prescribed burning on nutrient availability in dry heathland soils and the nutrient content of the two competing heathland species Calluna vulgaris and Deschampsia flexuosa, with particular reference to N and P. We hypothesise that winter prescribed burning leads to additional N availability, which enhances the importance of P in the context of nutrient limitation in heathland ecosystems. In the nature reserve “Lueneburg Heath” (NW Germany) we examined the availability of nutrients in the humus horizons and in the leachate as well as the relevant C:element ratios in Calluna and Deschampsia before and after a burning experiment. Our results show that prescribed burning resulted in drastically increased NH₄⁺ availability in the O-horizon. We observed only short-term effects (for NO₃⁻, PO₄³⁻, Mg) and insignificant effects on the availability of other nutrients (K, Ca). As a consequence of an increased nutrient availability in the humus horizons and a limited nutrient uptake by plants after burning, leaching increased significantly for N, Ca, K, and Mg after burning treatment. No significant changes were found in the foliar C:N ratios for either species after prescribed burning, although Deschampsia showed an increased deficiency for all the other nutrients, particularly for P, as expressed by increased foliar C:P and N:P ratios. By contrast, the nutrient content of Calluna did not change significantly, suggesting that prescribed burning favours the competitive capacity of Calluna as against Deschampsia. We assume that water shortage as a result of changes in the microclimate was mainly responsible for the deterioration of the nutrient content of Deschampsia. This gives Calluna a competitive advantage, enabling it to out-compete Deschampsia on burned heathlands, with respect to the key factor P-limitation.     

Conservation of Calluno-Vaccinietum heathland in the Belgian Ardennes,an experimental approach

It is now difficult to manage the semi-natural groupings of heathland because of the tendency for succession towards forest stages.Several experiments were conducted in the Hautes Fagnes region (Belgium). After the vegetation was mapped, plots were treated by mowing, burn-beating, and burning.The changing floristic composition was followed from 1972 to 1980 in permanent quadrats. Burn-beating is an excellent method of management for old heath, because it reestablishes the properties of the heath ecosystem.Nomenclature follows de Langhe et al., 1978. Nouvelle flore de la Belgique, du Grand-Duché de Luxembourg, du Nord de la France et des Régions voisines, 2e éd.     

Comparison of the regeneration after burning,cutting and ploughing in a Cistus ladanifer shrubland

The recovery of a Cistus shrubland after experimental treatments (cutting, burning and ploughing) was studied to determine qualitative and quantitative differences in the secondary succession. The study was carried out in three experimental plots of 100 m² each, in which five 1 m² sampling units were fixed and surveyed for five years, estimating cover percentage of each plant species. The data were analysed by means of affinity indexes (Sorensen and Steinhaus Indexes). The results indicated that the cover of the woody species increased progressively, but the degree of recovery depended on the treatment, as well as on the time afterwards. The number of these species remained almost constant throughout the whole period. The herbaceous species diversity was higher in the two first years in all treatments. When treatments were compared, the greatest diversity occurred after cutting. The regeneration of the community occurred by autosuccession in which the diversity decreased when the dominance of Cistus increased, unlike the general trend of the classical theories. The recovery was very quick after all three treatments, which can be interpreted as a great resilience of this ecosystem.     

Fire history and vegetation recovery in two raised bogs at the Baltic Sea

Questions: What were the bog fire patterns and frequencies in two boreal peatlands during the last 5000 years? What is the nature and time‐scale of post‐fire vegetation successions? Were fire events related to climate? Location: Männikjärve bog, central east Estonia; Kontolanrahka bog, southwest Finland. Methods: Macroscopic charcoal, plant macrofossils and radiocarbon dating were examined. Redundancy analysis was used in the assessments. Results: During the last 5000 years, both of the above peatlands have experienced several fire events. A typical pre‐fire vegetation community consisted of dry hummock Sphagnum spp., often accompanied by Calluna vulgaris. Only the most severe occasional fires resulted in a dramatic change in the vegetation composition. In these cases, a wet shift occurred, where the pre‐fire hummock community was replaced by a wet hollow community. Calluna vulgaris was found to be a key species in both pre‐ and post‐fire vegetation dynamics. The recovery time of dry microtopes following severe combustion and the subsequent hydrological change could take up to 350 years. Even after a long‐lasting wet phase, the post‐fire disturbance succession led towards a dry hummock community. Conclusions: Fire succession appeared to be cyclic, starting as and developing towards a dry hummock community. Fires have been a regular phenomenon in boreal bogs, even in regions with rather low human impact. The fire history records did not indicate any direct link to the regional long‐term climate.     

Nutrient leaching in dry heathland ecosystems: effects of atmospheric deposition and management

Atmospheric nutrient deposition has contributed to widespread changes in sensitive seminatural ecosystems throughout Europe. For an understanding of underlying processes it is important to quantify input–output flows in relation to ongoing atmospheric inputs and current management strategies. In this study we quantified losses of N, P, Ca, Mg, and K via leaching in heathland ecosystems (Lüneburger Heide, NW Germany) as a function of current deposition rates and different management measures (mowing, prescribed burning, choppering, sod-cutting) which aim to prevent shrub and tree encroachment. Leaching was only moderately related to atmospheric input rates, indicating that leaching was mostly affected by internal turnover processes. Leaching significantly increased for most of the nutrients after the application of management measures, particularly in the choppered and sod-cut plots. However, atmospheric nutrient inputs exceeded leaching outputs for most of the nutrients, even in the plots subjected to management. Despite high deposition rates (20–25 kg N ha⁻¹ year⁻¹), retention of atmospheric N input ranged between 74% and 92% in the control plots. In the treated plots, N retention decreased to 59–80%. However, in the study area mean N leaching in the controls has almost doubled since 1980 and currently amounts to 3.7 kg ha⁻¹ year⁻¹, indicating an early stage of N saturation. Our study provides evidence that leaching did not compensate for atmospheric nutrient deposition, particularly as regards N. Management, thus, will be an indispensable tool for the maintenance of the low-nutrient status as a prerequisite for the long-term preservation of heathland ecosystems.     

Fate of airborne nitrogen in heathland ecosystems: a 15N tracer study

In the present study, we analyze the fate of airborne nitrogen in heathland ecosystems (NW Germany) by means of a ¹⁵N tracer experiment. Our objective was to quantify N sequestration and N allocation patterns in an ecosystem that is naturally limited by N, but that has been exposed to airborne N inputs exceeding critical loads for more than 3 decades. We hypothesized that the system has a tendency towards N saturation, which should be indicated by low N sequestration and high N leaching. We analyzed ¹⁵N partitioning (aboveground biomass and soil horizons) and investigated ¹⁵N leaching over 2 years following a ¹⁵N tracer pulse addition. ¹⁵N tracer recovery was 90% and 76% in the first and second year, respectively. Contrary to our expectations, more than 99% of the tracer recovered was sequestered in the biomass and soil, while leaching losses were <0.05% after 2 years. Mosses were the most important short‐term sink for ¹⁵N (64% recovery in the first year), followed by the organic layer. In the second year, the moss layer developed from a sink to a source (23% losses), and soil compartments were the most important sink (gains of 11.2% in the second year). Low ¹⁵N recovery in the current year's shoots of Calluna vulgaris (<2%) indicated minor availability of ¹⁵N tracer sequestered in the organic layer. N partitioning patterns showed that the investigated heaths still have conservative N cycling, even after several decades of high N loads. This finding is mainly attributable to the high immobilization capacities for N of podzols in soil compartments. In the long term, the podzol A‐ and B‐horizons in particular may immobilize considerable amounts of incoming N. Since N compounds of these horizons are not readily bio‐available, podzols have a high potential to withdraw airborne N from the system's N cycle.     

Ecological Succession of a Relict Central European Peat Bog and Variability of Its Insect Biodiversity

The isolated habitat of the ervené Blato bog (South Bohemia, Czech Republic) and its relict insect fauna have been the subject of long-term monitoring. The species composition and abundance of Lepidoptera (light traps) and Coleoptera (pitfall traps) were monitored for 4 years (1994–1997) simultaneously on two sites – in the edaphic climax pine forest and in wetland successional habitats. The method of statistical evaluation by RDA and CCA ordination, representing the habitat preference of species of Coleoptera (Carabidae only) and Lepidoptera (all nocturnal phototactic taxa) between the edaphic climax forest and succession stages, was used. All categories of the peatland taxa (tyrphobiontic, tyrphophilous and tyrphoneutral species) were analysed. Ten highly stenotopic tyrphobiontic species and 23 tyrphophilous species of Lepidoptera (out of 487) were most characteristic of the bog habitat. Only two tyrphophilous carabid species (out of 20) were characteristic of the bog. The most important relict species (tyrphobionts) of Lepidoptera are most diverse and abundant within the successional habitats and in the open wet forest. The relict fauna of the closed climax pine forest is much less diverse and composed mostly of abundant tyrphophilous and tyrphoneutral forest species. Preservation or restoration of sufficiently constant hydrological conditions, which prevents formation of the closed forest, is the basic management for habitat conservation of all relict tyrphobiontic species of the ervené Blato bog and similar peat land habitat islands. The peat bog is a unified complex system of specific diverse and relict taxa. The most specific taxa are tyrphobiontic Lepidoptera, but a number of other vulnerable tyrphophilous and tyrphoneutral insects are associated with the peat bog as well.     

The role of ant nest-mounds in maintaining small-scale patchiness in dry grasslands in Central Germany

Nest-mounds of Lasius flavus, Lasius alienus and Formica rufibarbis are mainly on north-facing slopes with a markedly patchy distribution at Gimritz, northwest of Halle (Saale). Nest-mound soils had higher sodium and potassium levels but lower phosphorus and nitrogen levels than soils away from mounds. Moisture content was lower but soil pH was significantly higher in nest-mound soils. However, there was no difference in the relative abundance of plants of alkaline or acid soils growing on mounds compared to plants off mounds and no significant difference between the numbers of nitrophilous plants on and off mounds. Fewer plant species (35 spp.) in total occurred on mounds than off mounds (42 spp.). Plant species richness and total plant cover was significantly less on the nest-mounds than off the mounds. Similarly, the mean number of grass (1.39 spp dm−2) and forb species (1.13 spp dm−2) on mounds was lower than the mean number of grass (1.99 spp dm−2) and forb species (1.91 spp dm−2) off mounds. Some shrubs and forbs had higher occurrence and cover on the mounds, although only in Calluna vulgaris, Thymus serpyllum and Cerastium arvense was this difference particularly marked. Ant nest-mounds may favour plant species that cannot compete with tall grasses on unmanaged, set-aside meadows. This revised version was published online in November 2006 with corrections to the Cover Date.     

The influence of ammonium nitrate on the root growth and ericoid mycorrhizal colonization of Calluna vulgaris (L.) Hull from a Danish heathland

Conversion of European heathlands to grassland has been reported as a response to increased nutrient availability, especially of nitrogen; a direct effect upon mycorrhizal colonization has been proposed as an likely explanation.This hypothesis was tested in a random block experiment with four blocks and four replicates on a Danish inland heath, Hjelm Hede. Ammonium nitrate was applied (0, 35, 50 and 70 kg N ha^–1 year^–1) to a stand of Calluna vulgaris (L.) Hull four times annually for 2 years. Calluna roots were sampled on four occasions in the 2nd year of the nitrogen treatment. The extent of ericoid mycorrhizal colonization was determined by direct observation of the roots using a line-intersection method. The nitrogen content of the current-year shoots of Calluna increased when they were treated with nitrogen. Nitrogen fertilization had no significant effects on ericoid mycorrhizal colonization of Calluna nor on root biomass. The seasonal variation in mycorrhizal colonization of the Calluna roots was highly significant. The spatial variability of mycorrhizal colonization, both in replicated plots and in the two contrasted soil horizons – the mor layer and the bleached sand – within the plots, were considerable. I conclude that heather decline under enhanced nitrogen input is unlikely to be caused by a direct impact on the ericoid mycorrhizae of Calluna. Received: 1 December 1998 / Accepted: 7 December 1999     

Vegetation mapping and nature conservation: a case study in Terceira Island (Azores)

In order to emphasize the importance of vegetation mapping for nature conservation purposes a case study in Terceira island (Azores) is presented, in which the importance of the natural vegetation of the eastern slope of Santa Bárbara volcano (which is part of the Site of Community Importance of Santa Bárbara–Pico Alto) is evaluated through the elaboration of its vegetation map. Fourteen (14) different natural vegetation types were identified: grasslands (1 type), peat bogs (2 types), scrubs (2), forests (5), successional vegetation (3) and vegetation of rocky slopes (1). All communities are protected under the Habitat and Species Directive (EC/92/43) and most of them are endemic to the Azores Islands. This fact, together with the significant number of Azorean endemic taxa (18), Macaronesian endemic taxa (5) and species protected under the Habitat and Species Directive (7), gives this area an important conservation value that justifies future protection actions. Vegetation mapping is an important tool for the characterization, evaluation and implementation of managing plans of natural areas of the Azores islands. The use of a floristic-based classification, supported by multivariate analysis and structural data, is an efficient methodology for the construction of these maps. The data collected comprise an important set of information about the distribution and abundance of natural vegetation types and endemic and rare species. This information was not available until now and is indispensable for the elaboration of management plans of Special Zones for Conservation that will be part of the NATURA 2000 network.     

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 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.     

Short-term study of effects of fertilisation and cutting treatments on the vegetation dynamics of mountain heathlands in Spain

The influence of management and nutrient availability on the vegetation dynamics of heathlands characterised by Calluna vulgaris and Erica tetralix were studied in three mountain sites in Northern Spain. A total of 90 plots (1 m² each) received different combinations of cutting and twice the estimated background atmospheric deposition of nitrogen (56 kg ha⁻¹ yr⁻¹). One of the two dominant ericaceous species was selectively cut by hand at ground level and their regeneration compared in the presence or absence of the other. The results after 2 years showed significant effects of the fertiliser on the vegetation cover, mainly by favouring perennial herbaceous species. There were less noteworthy effects on the number of flowers and on the annual growth of the ericaceous species. It is concluded that, in the short term, increased nutrients alone, at twice the estimated current atmospheric deposition for the area, will not alter significantly the composition of the mountain heathlands. However, once the stands reach the mature phase, the capacity of the community to regenerate after a severe disturbance diminishes. A drastic impact, such as cutting may not result in re-growth of the same shrub species but in replacement by herbaceous species, which will also benefit from the increased nutrients.     

Isolation and characterization of microsatellite loci from Lochmaea suturalis, the heather beetle (Coleoptera: Chrysomelidae), a pest in Europe and a biocontrol agent in New Zealand

The heather beetle Lochmaea suturalis which is native to northwest Europe has been released as a biocontrol agent for heather in New Zealand. We have isolated and optimized eight microsatellite loci from New Zealand beetles. These loci provide markers with high polymorphism ranging from four to 20 alleles per locus. Observed heterozygosity averaged 0.631 per locus. These results suggest the markers are useful for population studies that will contribute to assessment of L. suturalis as a biocontrol agent.     

Habitat associations of the Small Red Damselfly (<Emphasis Type="Italic">Ceriagrion tenellum)</Emphasis> (De Villiers) in heathland in southern England (Zygoptera: Coenagrionidae)

The Small Red Damselfly (Ceriagrion tenellum) (De Villiers) (Odonata: Coenagrionidae: Ceriagrion) is classed as vulnerable (Shirt, British Red Data Book, Nature Conservancy Council, Peterborough, UK, 1987) throughout the UK, and is included in certain Local Biodiversity Action Plans (LBAPs) in the south. A large proportion of any Biodiversity Action Plan is concerned with the requirement of conservation and management programmes. In order to guide them, information about the habitat preferences of the species concerned is vital. Detailed habitat information was collected to include a variety of physical parameters particularly vegetation, both in-channel and bankside. The species was found to be primarily associated with in-channel emergent broad-leaved plants, bankside grasses and rushes, and shallow, narrow channels with dark organic substrate. The consequences of these findings are discussed in relation to the conservation and management of C. tenellum.