Restoration management of abandoned chalk grassland in the Netherlands

After abandonment of chalk grassland in the Netherlands, Brachypodium pinnatum had become very dominant and this resulted in a dramatic decrease of plant diversity. Restoration of these abandoned sites is important, because of its former high diversity with many nationally endangered species. To restore the chalk grassland vegetation, the impact of the introduction of different experimental cutting regimes was investigated during three years. The thick litter layer and the dominance of Brachypodium was strongly reduced after introduction of all cutting regimes. In all cutting treatments forb phytomass increased considerably, especially in the twice-a-year cutting vegetation, where the forb phytomass became as high as that of Brachypodium. Phanerogamic species diversity was stimulated in all cutting regimes. Especially the number of short-lived forbs. with a persistent seed bank, increased markedly in the twice-a-year cutting treatment. It is concluded that cutting the vegetation twice is adequate to reduce the negative effects of the thick litter layer and the abundant growth of Brachypodium. It creates an appropriate starting point for more detinite management, whether mowing or grazing. However, the restoration of a more complete species assortment strongly depends on the re-invasion possibilities of these species.     

Vertical canopy structure of Dutch chalk grasslands in relation to their management

Analysis of leaf canopy overtopping relationships was carried out using a non-destructive point quadrat method followed by a destructive stratified harvest of the above-ground phytomass in Dutch chalk grasslands with different management: summer sheep grazing and annual autumn mowing. The two methods of analysis are compared: e.g. relative leaf overtopping can be recorded by the point quadrat method but it is obscured in vertical vegetation profiles based on stratified phytomass distribution. However the stratified harvest method describes the relationship between canopy phytomass and light microclimate, recorded by measuring Photosynthetically Active Radiation (PAR) at different heights in the vegetation. Plant growth form during peak standing crop is of greater significance than Raunkiaerian life form in determining structure of chalk grassland vegetation. In annually mown grasslands, the tall graminoid growth form, shown by Brachypodium pinnatum, reduces PAR in the lower canopy and suppresses most other growth forms except those which can reach the higher canopy levels, e.g. clumped herbs such as Origanum vulgare. In contrast, grazing can result in a reduction of dominance from the tall graminoids and reduction of the abundance of taller, grazing-sensitive herbs, e.g. clumped and climbing herbs such as Origanum and Lathyrus pratensis, and an increase in grazing-tolerant species, e.g. smaller rosette herb growth forms, e.g. Leontodon hispidus and shorter rhizomatous or stoloniferous graminoids, e.g. Carex flacca and Briza media. Changes in both the overtopping hierarchy at the peak of the growing season and the intensity of overtopping in the course of a growing season are of conclusive importance in determining the relative abundance of species in the vegetation.     

Sensitivity of seedling recruitment to moss, litter and dominant removal in an oligotrophic wet meadow

The effect of a dominant species, the litter layer, and the moss layer on seedlings and established vegetation was evaluated in two manipulative experiments in an oligotrophic wet meadow (Molinion with some features ofViolion caninae according to phytosociological classification). In the first experiment, the dominant species (Nardus stricta), litter layer, and litter layer together with the moss layer were removed and the response of the established vegetation and seedlings were compared with a control. Results revealed that after one season there was no significant effect of the treatments on established vegetation, but seedling recruitment was significantly affected. The greatest increase of seedling recruitment for many species was observed in the plots with the moss layer removed. In the second experiment, seeds ofSelinum carvifolia were sown into plots subjected to the following treatments: complete vegetation removal, mowing, mowing combined with removal of the moss layer, and an untreated control. The highest number of seedlings was found in plots with the vegetation removed, but the seedling recruitment was nearly as high in plots with the moss layer removed. Low numbers were found in mown plots and the lowest in the untreated control. The results show that seedling recruitment is more sensitive to competition than the established vegetation, at least over one season. This supports the idea of the importance of the regeneration niche for the maintenance of diversity in grassland communities.     

Stimulation of soil nitrification and denitrification by grazing in grasslands: do changes in plant species composition matter?

Stimulation of nitrification and denitrification by long term (from years to decades) grazing has commonly been reported in different grassland ecosystems. However, grazing generally induces important changes in plant species composition, and whether changes in nitrification and denitrification are primarily due to changes in vegetation composition has never been tested. We compared soil nitrification- and denitrification-enzyme activities (NEA and DEA, respectively) between semi-natural grassland sites experiencing intensive (IG) and light (LG) grazing/mowing regimes for 13 years. Mean NEA and DEA (i.e. observed from random soil sampling) were higher in IG than LG sites. The NEA/DEA ratio was higher in IG than LG sites, indicating a higher stimulation of nitrification. Marked changes in plant species composition were observed in response to the grazing/mowing regime. In particular, the specific phytomass volume of Elymus repens was lower in IG than LG sites, whereas the specific volume of Lolium perenne was higher in IG than LG sites. In contrast, the specific volume of Holcus lanatus, Poa trivialis and Arrhenatherum elatius were not significantly different between treatments. Soils sampled beneath grass tussocks of the last three species exhibited higher DEA, NEA and NEA/DEA ratio in IG than LG sites. For a given grazing regime, plant species did not affect significantly soil DEA, NEA and NEA/DEA ratio. The modification of plant species composition is thus not the primary factor driving changes in nitrification and denitrification in semi-natural grassland ecosystems experiencing long term intensive grazing. Factors such as trampling, N returned in animal excreta, and/or modification of N uptake and C exudation by frequently defoliated plants could be responsible for the enhanced microbial activities.     

Species composition and above ground phytomass in chalk grassland with different management

During the last decades chalk grasslands lost their agricultural importance in the greater part of their distribution area in W Europe. Due to their botanical richness a number of chalk grassland sites were established as Nature Reserve. As a consequence of the semi-natural character of these grasslands an appropriate management is necessary to maintain or to re-create this vegetation including a great number of rare and endangered species.This paper deals with the results of three different management practices, viz. mowing in autumn, sheep grazing and abandoning, of a medium term (8–11 yr) permanent plot experiment. Sheep grazing was considered the best management since it resulted in the highest number of species, phanerogams as well as bryophytes, and the highest number of characteristic chalk grassland species. Abandoning resulted in a decrease in species number and a dominance of a few species only.The changes in species number are related to the above ground biomass. Under the canopy in the abandoned plot light intensity and the Red/Far-red ratio are very low, which partly explains the decrease in species number as such conditions are not favourable to seedling emergence and survival.Plant nomenclature follows Heukels & Van Ooststroom (1975) for phanerogams, Margadant & During (1976) and van der Wijk et al. (1969) for bryophytes, and Westhoff & den Held (1969) for syntaxa.Presented at the Symposium on Vegetation Dynamics, organized by the International Society for Vegetation Science at Montpellier, 15–20 September 1980. I am grateful to Dr. H. J. During for useful comments.     

Effects of vegetation management by mowing on ground-dwelling arthropods

Species-rich grasslands are rare in the Netherlands and need consistent vegetation management to retain their characteristic biodiversity. Roadside verges are important refuges for grassland plants since the mowing management no longer aims at traffic safety only but also strives for botanical diversity. Although arthropods are highly abundant in roadside verges, the effect of different mowing practices on this group is largely unknown. During 4 years, we studied ground beetles, weevils, ants and ground-dwelling spiders with pitfall traps in experimental plots in roadside verges with five different mowing treatments: (i) no management, (ii) and (iii) mowing once a year with and without hay removal, (iv) and (v) mowing twice a year with and without hay removal. This was done in a plant productivity gradient; the experiment was repeated in low-, medium- and high-productive verges. In the low-productive site, the effect of management on the arthropods only existed in a higher abundance in plots mown twice per year with hay removal. In the medium- and high-productive sites, mowing twice a year with hay removal resulted not only in highest abundances but also in highest arthropod species richness. Mowing twice without hay removal and mowing once with removal showed intermediate values, while mowing once per year without removal and particularly the absence of management resulted in low diversity and low abundance. To promote ground-dwelling arthropods in medium-to-high-productive grassland verges, we recommend a management of mowing twice a year with the removal of hay. It is reasoned that some form of rotational management, aiming at leaving some vegetation refuges intact after mowing events, may further promote arthropod survival. However, caution should be taken that these refuges are not too large, as overall suitability for ground-dwelling arthropod decreases rapidly in such patches. Out of several studied vegetation characteristics, the number of flowering plant species (medium-productive verge) and total flower abundance (high-productive verge) appeared to represent suitable, and easily monitored, proxies that significantly mirror arthropod diversity.     

Sustaining recovered grasslands is not likely without proper management: vegetation changes after cessation of mowing

Grasslands recovered by sowing low diversity seed mixtures of local provenance are usually managed by mowing. Besides restoration success only a few studies have focused on the direct effects of post-restoration mowing on recovered grassland vegetation. In this study we followed vegetation changes in 13 successfully recovered grasslands in 5 × 5-m-sized sites with continuous and ceased mowing at Hortobágy National Park, East-Hungary. We asked the following questions: (i) What are the effects of cessation of mowing on the vegetation structure and diversity of recovered grasslands? (ii) What are the effects of cessation of mowing on the abundance of sown grasses, target and undesirable species? (iii) Is yearly mowing an appropriate management tool for the maintenance of recovered grasslands? Our results showed that the cessation of mowing caused litter accumulation, while diversity, total vegetation cover and the cover of sown grasses decreased compared to the mown sites. The cover of undesirable perennial species was significantly higher in unmown sites than in mown ones. The species composition of mown sites remained more similar to near-natural grasslands than the unmown ones. Our results suggest that without regular post-restoration mowing the favourable status of recovered grasslands can rapidly decline due to litter accumulation and by the expansion of undesirable species, even in the short-run. We also stress that while yearly mowing is enough to maintain grasslands recovered by low-diversity seed sowing, it cannot be considered to be enough to recover target vegetation composition.     

Species diversity and primary productivity inMiscanthus sinensis grasslands

In an old field grassland dominated byMiscanthus sinensis Anderss. the community structures, phytomass, dominance ofM. sinensis and species diversity were measured. Species and life form composition of the stand were characterized by higher percentages of therophytes, woody and shrubby species, liana and alien species. From May so September in 1982, all the aboveground parts were harvested from each of the four quadrats (2 m×2 m) once a month. Seasonal peak of aboveground phytomass, in September, was 1027 g d.w.m⁻² to whichM. sinensis contributed as much as 96.5%. With the progress of the growing season,M. sinensis became increasingly important both in stand phytomass and in dominance, whereas species diversity based on the dry weight contributions of constituent species decreased. Our analysis of these seasonal trends showed that the diversity was largely a function of dominance of the most important species, rather than that of stand phytomass or productivity. The simultaneous measurements of 20 quadrats in late August 1983, also supported the above conclusion.     

Effectiveness of mowing for the flora diversity preservation: a case study of steppe-like grassland in Croatia (NATURA 2000 site)

Intensive human activities have resulted in a critical reduction of grasslands in Croatia, one of the richest Europian countries in terms of biodiversity. Sub-Pannonic steppic grasslands are very rare, only few occur in the continental part of Croatia and most of them are protected as NATURA 2000 sites. We studied vascular flora of the Sub-Pannonic stepic grassland with the aim to examine the changes in plant communities 15 years from the mowing application. Results showed that hand mowing once per year at the end of vegetation season increased the species diversity as well as the number of medium and low growth taxa. The critically endangered, nearly threatened and vulnerable species remained preserved. The occurrence of trees, shrubs and invasive species as a consequence of succession and anthropogenic influences could be important threat for steppe-like grassland flora. From a conservation point of view infrequent mowing regime could be an effective management tool, although it needs to be adapted to regional and local circumstances for maintaining high biodiversity of steppe-like grasslands and valuable plant species.     

Not only the butterflies: managing ants on road verges to benefit Phengaris (Maculinea) butterflies

Obligate myrmecophilic butterfly species, such as Phengaris (Maculinea) teleius and P. nausithous, have narrow habitat requirements. Living as a caterpillar in the nests of the ant species Myrmica scabrinodis and M. rubra, respectively, they can only survive on sites with both host ants and the host plant Great Burnet Sanguisorba officinalis. After having been reintroduced into a nature reserve in the Netherlands in 1990, both butterfly species expanded their distribution to linear landscape elements such as road verges and ditch edges outside this reserve. As additional habitat of both butterfly species, vegetation management of these landscape elements became important. Our results show that a management beneficial for Phengaris butterflies should aim to increase the nest density of Myrmica species, at the same time reducing the density of nests of the competitor Lasius niger or at least keeping them at a low density. Unfavourable grassland management under which L. niger thrives, includes not mowing or flail-cutting the grass, or depositing dredgings along the side of the ditch. Management favourable for the two Myrmica species differs, demanding some flexibility if both species are to benefit. M. scabrinodis is best supported with early mowing of the road verge vegetation or late mowing in the nature reserve, both of which result in an open vegetation and warm microclimate. In contrast, the nest sites of M. rubra should be left undisturbed during the summer, and mown in late autumn. Mowing of butterfly habitat should be avoided between mid-June and mid-September as this would remove the flowerheads of the Sanguisorba plants, on which the butterflies lay their eggs.     

禁牧条件下不同类型草地群落结构特征

利用幂乘方法则模型探讨了鄂尔多斯3种不同类型草地在禁牧情况下的群落结构特征,包括物种组成、物种多样性、生物量和空间分布规律。结果表明:幂乘方法则在解析鄂尔多斯不同类型草地的空间异质性时具有很好的吻合性;羊草(Leymuschinensis)草地、芨芨草(Achnatherum splendens)草地和油蒿(Artemisia ordosica)灌丛草地均比随机分布呈现了较强的空间异质性,群落整体的空间异质性指数表现为油蒿灌丛草地>羊草草地>芨芨草草地;群落整体的物种多样性指数为油蒿灌丛草地>羊草草地>芨芨草草地;L-样方(50cm×50cm)内的平均物种数和物种多样性指数均表现为羊草草地极显著地高于油蒿灌丛草地,油蒿灌丛草地又极显著地高于芨芨草草地(P<0.001);L-样方内的平均生物量表现为油蒿灌丛草地极显著地高于芨芨草草地(P<0.001),而羊草草地与油蒿灌丛草地以及芨芨草草地之间没有显著性差异;这3种类型的草地其L-样方内的平均生物量和物种多样性指数都随着群落整体空间异质性指数的增大而增大。     

Soil feedback does not explain mowing effects on vegetation structure in a semi-natural grassland

Due to its ability to create aboveground conditions that favour plant diversity, mowing is often used to preserve the high conservation value of semi-natural species-rich grasslands. However, mowing can also affect belowground conditions. By decreasing plant carbon supply to soil, mowing can suppress the activity of soil decomposers, diminish plant nutrient availability and thus create a feedback on plant growth. In this study, we first documented the effects of three-year mowing on plant community structure in a species-rich grassland. We found that mowing decreased the total areal cover of woody plants and increased the total cover of leguminous forbs. At the species level, mowing further increased the cover of two non-leguminous forbs, Prunella vulgaris and Sagina procumbens. Mowing did not affect the species number, diversity or evenness of the plant community. To study whether any of these effects could be explained by mowing-induced changes in the soil, and particularly by reduced nutrient availability, we then collected soil from different treatment plots and monitored the growth of nine plant species in these soils in a greenhouse. Plant growth did not differ between soils collected from mowed and unmowed plots, suggesting that our mowing regimes did not impose such changes in soil decomposer activity and nutrient supply that would feedback on plant growth. Moreover, each of the nine species responded equally to the different nutrient availability in different parts of the grassland, which indicates that even if mowing had reduced plant nutrient supply, this would not have led to changes in plant community structure. It appears that those changes in aboveground vegetation that we recorded after three years of mowing were purely due to the aboveground effects, such as frequent cutting of woody plants and enhanced light availability for low-growing forbs.     

Assessing the impact of land abandonment, nitrogen enrichment and fairy-ring fungi on plant diversity of Mediterranean grasslands

Semi-natural calcareous grasslands are of great conservation interest because of their high species richness, but they are threatened by land abandonment and nitrogen eutrophication. These plant communities evolved as a result of a long history of human activity, which generated and maintained these habitats by extensive grazing and mowing. Calcareous grasslands are listed as a priority for conservation in the EC Habitats Directive. However, the effects of different management regimes, nitrogen enrichment, and soil-borne pathogens on plant species diversity are less clear for grasslands of the Mediterranean Basin, compared to meadows in Northern and Central Europe. In this study, we assessed the impact of land abandonment, nitrogen enrichment, and fairy-ring fungi on species diversity in semi-natural grasslands found in the Mediterranean Basin by comparing the available literature with findings from recent studies carried out in Central Italy. In a series of field experiments, the cutting of abandoned grassland consistently reduced the living biomass of the dominant perennial grasses, such as Brachypodium rupestre and Bromus erectus, and promoted a rapid increase in species richness and diversity by allowing the establishment of rare species. There was a similar, but less effective, restoration of species diversity and composition in mowed grassland after litter removal. We also show that nitrogen enrichment at levels comparable to atmospheric deposition depresses species diversity, which also hampers the positive effects of litter removal. Our findings are consistent with previous results achieved in Northern and Central Europe, which however, mainly focused on grasslands with intermediate to high primary productivity levels. The limited availability of data from low-productivity, drought-prone Mediterranean grasslands requires further studies to assess the impact of land abandonment and nitrogen eutrophication in such ecosystems. Finally, we discuss the role of fairy-ring fungi in the maintenance of plant diversity in species-rich grassland. We show that fairy-ring fungi (e.g. Agaricus campestris) critically affect the spatial distribution and diversity of coexisting plant species. By killing the dominant perennial herbs, these radially growing plant pathogens produce empty niches for rare, short-lived species, thus affecting the vegetation pattern. Overall, our results are of interest for environmental managers, as they provide guidelines for the restoration of abandoned areas and the conservation of these species-rich habitats.     

Can grassland plant communities be preserved on road and railway verges?

Abstract. Semi‐natural grasslands are a threatened biotope type in many countries. Typical grassland plant species are adapted to continuous grazing or mowing that keeps the environment open. With the decline in grassland area, these species are ever more reliant on alternative habitats such as road verges. To find out whether plant communities comparable to those of semi‐natural grasslands can be maintained on modern road and railway sides, the vegetation of 92 grasslands and 90 road and railway verges was studied. These biotope types were compared with each other according to their community structure, total number of species, number of grassland species and restricted‐range diversity. Further study of the vegetation of road and railway verges was carried out in order to identify the treatments and environments which are most likely to support diverse plant communities. The species number and the restricted‐range diversity proved to be higher next to roads and railways than on grasslands. Grassland species were, however, most abundant on grasslands. Furthermore, the community structure of these biotope types was totally divergent. In their present state, road and railway verges are not a substitute for semi‐natural grasslands. Nevertheless, the occurrence of grassland species in verges may be enhanced by a suitable mowing regime, by giving up the use of de‐icing salt and herbicides and by allowing natural establishment of vegetation on the verges.     

Short-term effects of nitrogen enrichment,litter removal and cutting on a Mediterranean grassland

Semi-natural grassland communities are of great interest in conservation because of their high species richness. These communities are being threatened by both land abandonment and nitrogen eutrophication, and their continued existence will depend upon correct management. However, there is a distinct lack of studies of the ecological mechanisms that regulate species diversity and productivity in Mediterranean grasslands. We have conducted a 3-year field experiment in a species-poor grassland in central Italy to investigate the effects of nitrogen fertilization coupled with removal of plant litter and artificial cutting on species diversity and community productivity. Vegetation cutting reduced living biomass but increased species diversity. In fact, cutting had positive effects on the cover of almost all of the annual and biennial species, while it had a negative effect on the dominant perennial grasses Brachypodium rupestre and Dactylis glomerata. Litter removal had similar effects to cutting, although it was far less effective in increasing species diversity. In contrast, nitrogen enrichment strongly increased the living biomass while maintaining very low species diversity. Our results have indicated that semi-natural Mediterranean grasslands need specific management regimes for maintenance and restoration of species diversity. In the management of these grasslands, attention should be paid to the potential threat from nitrogen enrichment, especially when coupled with land abandonment.     

Community Associations of Chalk Grassland Leafhoppers (Hemiptera: Auchenorrhyncha): Conclusions for Habitat Conservation

Due to their confinement to specific hostplants or restricted habitat types, Auchenorrhyncha have the potential to make suitable biological indicators to measure the quality of chalk grassland under different management practices for nature conservation. The Auchenorrhyncha data from a study designed to identify the factors influencing the invertebrate diversity of chalk grasslands in southern England was used to evaluate the potential use of this group of insects as biological indicators. Between 1998 and 2002 altogether 81 chalk grassland sites were sampled. Vegetation structure and composition were recorded, and Auchenorrhyncha were sampled at each site on three occasions in each of two seasons using a ‘Vortis’ suction sampler. Auchenorrhyncha assemblages were then linked to the different grassland plant communities occurring on chalk soils according to the British National Vegetation Classification (NVC). Altogether 96 Auchenorrhyncha species were recorded during the study. Using data on the frequency and dominance of species, as is commonly done for plant communities, it was possible to identify the preferential and differential species of distinct Auchenorrhyncha assemblages. Significant differences between the Auchenorrhyncha assemblages associated with the various chalk grassland plant communities of the NVC were observed down to a level of sub-communities. We conclude that data on Auchenorrhyncha assemblages can provide valuable information for the setting of conservation management priorities, where data on floristic composition alone may not be sufficient, providing additional information on aspects of vegetation structure and condition.     

Quick Target Vegetation Recovery after Restorative Shrub Removal and Mowing in a Calcareous Fen

Removal of shrubs and trees is an important management and restoration practice to promote openness and light‐dependent vegetation in fens, especially as tree cover is increasing in previously open wetlands. The effects of woody vegetation removal on target species have been poorly documented in wetlands up to now. In this study, I investigated the effect of tree and shrub removal (especially of Juniperus communis) on the target vegetation in a partly overgrown and degraded grazed rich fen after 6 years. I also tested whether additional intensified management by mowing could promote initial recovery. Shrub removal resulted in a rapid recovery of species‐rich fen vegetation such that after 6 years brown moss cover more than tripled and target species richness doubled and became similar to values of a reference area in a favorable conservation status. Additional mowing resulted in a much higher abundance of the target rich fen vascular plants. The reasons for the success at this site may be the proximity to well‐developed rich fen vegetation, presence of cattle that dispersed diaspores, and presence of bare, colonizable substrate. Thus, it may be more beneficial to restore and expand already existing sites in a partly favorable status than to restore severely deteriorated sites. Extensive management by woody vegetation removal may be an alternative method to maintain high conservation values of open mires and other wetlands, where grazing or mowing is not necessary or feasible to meet future needs in response to overgrowth caused by global warming.     

Linking Above- and Belowground Responses to 16 Years of Fertilization,Mowing, and Removal of the Dominant Species in a Temperate Grassland

Species-rich oligotrophic meadows are affected by a wide range of management interventions that influence their functioning and capacity to deliver ecosystem services, but long-term studies on the above- and belowground adaptations to different management tools are still scarce. We focused on the interactive effects of NPK fertilization, mowing, and removal of the initially dominant species (Molinia caerulea) on plant, soil, and microbial responses in wet oligotrophic grassland in a 16-year full-factorial manipulative experiment. Changes in vegetation composition, soil pH, and nutrient availability were accompanied by altered microbial phospholipid fatty acid (PLFA) composition, whereas treatment effects on soil microbial biomass and carbon (C) mineralization were mainly related to changes in soil organic matter (SOM) content and nutrient availability. Fertilization decreased plant species richness aboveground and lowered SOM storage and microbial activity belowground. Mowing preserved high plant diversity and led to more efficient recycling of N within the grassland, whereas Molinia removal significantly affected only plant community composition. Mowing combined with fertilization maintained high species richness only in the short term. Belowground, mowing reduced N leaching from the fertilized system but did not prevent SOM depletion, soil acidification, and concomitant adverse effects on soil microbes. We conclude that annual mowing is the appropriate type of extensive management for oligotrophic species-rich meadows, but the concomitant nutrient depletion should not be compensated for by regular NPK fertilization due to its adverse effects on soil quality.     

Long-term population dynamics of Dactylorhiza incarnata (L.) Soó after abandonment and re-introduction of mowing

The orchid Dactylorhiza incarnata (L.) Soó is a highly polymorphic species listed as endangered in many regional red lists of Central Europe and Scandinavia. The dramatic decline of its populations during recent decades has been caused by the loss and degradation of their natural and semi-natural habitats (fens and wet meadows, respectively) as a result of secondary succession following intensification or abandonment of traditional land use.In this study, we analysed the effects of abandonment and re-introduction of mowing on the long-term (28 years) population dynamics of Dactylorhiza incarnata at Lake Barsbek in northern Germany. In this area, to preserve a remaining population of D. incarnata, an annual mowing regime was re-established in 1981 on site M-1 of the investigated plots after a period of abandonment. Annual mowing was introduced on a second site, M-2, in 1987. Two- to three-year mowing was introduced on a third site, M-3. Site A, abandoned since 1970, was used as a reference. On each of these sites, flowering individuals were counted once a year. Population structure and accompanying vegetation were recorded simultaneously. In 2006, light measurements were carried out in the mowed areas.The D. incarnata population at M-1 increased exponentially during the first 10 years after re-introduction of mowing. Pronounced decreases in the number of flowering individuals were recorded in 1997 and 2003. Population dynamics at M-2 generally resembled the temporal development at M-1. D. incarnata disappeared on site A during the investigation period, while vegetation height and litter layer increased by 60 and 100%, respectively. D. incarnata was able to withstand reduced light availability to a certain extent by increasing its vertical growth (shade avoidance). It is concluded that the maintenance of D. incarnata populations in Central Europe requires continuation or re-establishment of wet meadow management. On previously abandoned sites, an initially higher mowing frequency is recommended. Management intensity can be reduced after phytomass production of the vegetation has declined.     

Fine-scale spatial distribution of leaves and shoots of two chalk grassland perennials

The fine-scale spatial distribution of leaves and shoots of Brachypodium pinnatum and Carex flacca, two rhizomatous graminoids, was investigated in two chalk grasslands in South Limburg (The Netherlands). The objective was to examine whether leaves and shoots of Brachypodium, a dominant species, had a regular distribution on a small scale, as has been suggested for other clonal species that form high-density stands. Patterns were compared to Carex, which is never found to be as abundant as Brachypodium.The number of shoots and leaf contacts were counted in small quadrats, grouped in a grid. Using Moran's I analysis for autocorrelation, it appeared that leaves and shoots of both species were arranged in clumps, and that these clumps were randomly distributed across the soil surface. Shoot clumps in Carex were smaller in diameter and not as pronounced as those in Brachypodium.In most cases, patterns of leaves and shoots were positively correlated, indicating that leaves were predominantly positioned above and around the groups of quadrats where the shoots were attached. However, in dense stands of Brachypodium the positions of leaf clumps were not correlated to those of shoot clumps. This is a result of the tall growth form of this species and its high shoot densities, and it is suggested that this will be a characteristic of any species that dominates a dense stand.