Reintroduction of a Rare Plant (Gladiolus imbricatus) Population to a River Floodplain—How Important is Meadow Management?

In seminatural grasslands, the success of reintroduction of locally extinct rare plant species may depend on the ambient management regime. We aimed to study to what extent the success of the restoration of a rare species (Gladiolus imbricatus) depends on management conditions. A management experiment with traditional cutting by scythe and hay removal, mowing (machine cutting and hay removal), mulching (machine cutting without hay removal), spring burning, and unmanaged control, combined with reintroduction of seeds of Gladiolus, was conducted in an Estonian flooded meadow in which the species had become extinct. Seeds were reintroduced in 2003 in all management treatments and populations monitored until 2006. Mulching, mowing, and traditional management resulted in the greatest establishment, whereas the subsequent mortality was not influenced greatly by management regime. The population started to increase in mulching treatment in the third season due to vegetative growth. The results indicate that the establishment of G. imbricatus is primarily seed limited under current conditions, whereas favorable management significantly enhances establishment in a river floodplain meadow. Successful restoration depends on seed addition and proper grassland management—mowing to a height of approximately 15 cm and mulching.     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.     

How do management and restoration needs of mountain grasslands depend on moisture regime? Experimental study from north‐western Slovakia (Western Carpathians)

Question: How does species composition change in traditionally managed meadows after mowing has ceased, and in abandoned meadows after re‐introduction of mowing? Are there differences in the dynamics of dry and moderately wet meadows? Location: Zázrivá‐Ple?ivá (19°11′N, 49°16′E), north‐western Slovakia, western Carpathians. Methods: Pairs of experimental plots (mown and unmown) were established to replicate each combination of dry/wet and traditionally managed/abandoned meadows. Changes in species composition were studied over 5 years. The data on changes in species composition was analysed by constrained and unconstrained ordinations, and visualized using Principal Response Curves. Results: Species composition of newly abandoned wet grasslands was changing towards the corresponding long‐abandoned plots even in the first year of abandonment. Similarly, newly established restoration mowing in abandoned dry grasslands rapidly shifted the stand species composition towards that of traditionally managed ones. Nevertheless, 4 year after reintroduction of mowing, the species composition of the restored plots was still far from the target composition. The effect of mowing in abandoned wet grasslands and abandonment in dry grasslands was much less pronounced and slower. Conclusions: Moisture regime is a very important factor determining the management needs of various grassland types. Wet grasslands are much more sensitive to abandonment, with a rapid degradation rate and limited possibilities for restoration, which can be extremely slow. Even in the dry grasslands, that quickly responded to restoration mowing, restoration is a long‐term process.     

Plant species diversity in an old, traditionally managed hay meadow compared to abandoned hay meadows in southwest Norway

A chronosequence, representing a successional series, was used for the comparison of a hay meadow site managed in an old traditional way for at least a hundred years, and hay meadow sites abandoned for about 10, 20 and 30 years, respectively. Old traditional management included grazing early and late in the growing season, mowing in August and light or no fertilizing. The tree cover was the most important factor deciding the composition of vegetation. Time since abandonment was not completely correlated to tree cover, as some plots had a dense canopy and others were situated in the open. The total species number decreased with number of years since abandonment in plots > 0.001 m²and <100 m². The highest species number in 1 m² plots was recorded in the managed site, with 38 species of phanerogams. Fourtyeight % of the indicators of traditional management present in the managed site was recorded in the site which had been abandoned for 30 years. Frequency — log area curves made it possible to group species according to persistence in the sward. As a result, a group of functional indicators of rare hay meadows in the region was distinguished.     

Evaluating alternative mowing regimes for conservation management of Central European mesic hay meadows: A field experiment

Conservation status of hay meadows highly depends on their management. The main goal of this study was to assess the efficiency of different mowing regimes in maintenance of plant species richness and diversity of mesic hay meadows. The field experiment was carried out on a species rich, mesic hay meadow in Western Hungary. We evaluated the effects of four alternative types of management on the plant community after 7 years of continuous treatment: (1) mowing twice a year, typical traditional management, (2) mowing once a year in May, most practised currently by local farmers, (3) mowing once a year in September, often proposed for conservation management and (4) abandonment of mowing. Traditional mowing resulted in significantly higher number and higher diversity of vascular plant species than other mowing regimes. Mowing twice a year was the only efficient way to control the spread of the invasive Solidago gigantea, and mowing in September was more successful in it than mowing in May. We conclude that the traditional mowing regime is the most suitable to maintain botanical diversity of mesic hay meadows; however, other regimes should also be considered if certain priority species are targeted by conservation.     

Do They Ever Come Back? Responses of Leafhopper Communities to Extensification of Land Use

We studied leafhopper communities in meadows subject to progressive extensification of land use, particularly (i) delay of the first cut, (ii) cessation of fertilising and (iii) reduction of cutting events. Within a gradient from conventionally used high-productivity meadows (as control) through our extensified plots to extensively managed wet hay meadows (as control), we found an increasing species number correlated with extensification of land use. However, a separate analysis of generalists and specialists showed that the latter group increased significantly whereas generalists did not respond at all. Even after 12 years of extensification there was only little evidence for the recovery or recolonisation of former hay meadow insect communities. Instead the increase in species numbers was rather due to immigration of more xerophilous or mesophilous species. We conclude that leafhoppers principally respond positively to extensification of land use, but that restoration of former moisture conditions is necessary in order to achieve a full recovery of original hay meadow communities. Finally we propose a model extensification ecogram for meadow leafhoppers which can be used as a predictive tool for extensification and as an indicator of restoration progress and success.     

Diaspore and gap availability are limiting species richness in wet meadows

Species pool theory claims that diaspore and microsite availability limit species richness in plant communities. Wet meadows (Calthion) and litter meadows (Molinion, Caricion davallianae) belonging to the most species-rich meadows in the foothills of the Alps have suffered from a strong decrease since the 1970s. Restoration efforts including nutrient impoverishment and rewetting management after drainage and fertilization did not result in the re-establishment of the former species richness although the abiotic filter would have been appropriate for the re-colonization of many locally extinct species. In our experiment at four sites in the largest fen of Southwest-Germany we tested if the restoration success was seed- and gap-limited. We applied sowing and hay spreading (hay seed) as treatments to make seeds available and harrowing to increase gap availability. Sowing seeds or hay seed of species of the former meadow types increased species richness immediately. The proportion of re-established species was higher when additional harrowing was applied. Species richness could be increased not only in vascular plants but also in bryophytes when hay spreading was applied. The strongest re-development towards the target communities (defined through the abiotic filter and the species richness before drainage and fertilization) took place on those sites where hay spreading and harrowing were applied. Sowing seeds and hay seed were traditional techniques to establish e.g. litter meadows, both techniques have been applied for centuries. Even harrowing was described as early as the 19th century to increase the chance of establishing certain species. Therefore, the “application of the knowledge coming from the species pool theory” (although not named during this time) has been common practice since at least the 19th century.     

Restoring riparian meadows currently dominated by Artemisia using alternative state concepts ‐ the establishment component

Abstract. We evaluated the potential for restoring riparian grass and sedge meadows currently dominated by Artemisia tridentata var. tridentata with an experiment in which we burned sites with low, intermediate, and high water tables, i.e., dry, intermediate, and wet sites. To define the alternative states and thresholds for these ecosystems, we examined burning and water table effects on both abiotic variables and establishment of grasses adapted to relatively high (Poa se‐cunda ssp. juncifolia), intermediate (Leymus triticoides), or low (L. cinereus) water tables. Wet sites had lower soil temperatures and higher soil water contents than dry sites. Burning increased soil temperatures on all sites. Undershrub microsites on control plots had the lowest temperatures, while former undershrub microsites on burn plots had the highest temperatures. Surface soil water was low on burn plots early in the growing season due to desiccation, but higher at deeper depths after active plant growth began. Emergence was generally greater on wet sites, but survival was microsite‐ and species‐specific. Undershrub microsites on control plots facilitated emergence and first‐year survival, but seedlings that survived initially harsh conditions on burn plots had similar numbers alive at the end. In general, favorable environments and establishment of species adapted to mesic conditions indicate that wet sites represent an alternative state of the naturally occurring dry meadow ecosystem type, and can be restored to grass and sedge meadows. Harsh environments and lack of establishment of species adapted to mesic conditions indicate that dry sites have crossed a threshold and may represent a new ecosystem type. Understory vegetation and seed banks on dry sites have been depleted, and restoration will require burning and reseeding with species adapted to more xeric conditions.     

Reinforced Traditional Management is Needed to Save a Declining Meadow Species. A Demographic Analysis

The changes in agricultural practices during the last century have led to a drastic decrease in the number of traditionally managed hay meadows. Also, traditional management practices are often applied more cursorily in the remaining meadows. In combination with an increase in aerial anthropogenic nitrogen deposition, this has led to a loss of biodiversity. To investigate whether the current management is sufficient for maintaining viable populations of a typical meadow plant, Succisa pratensis, we experimentally reinforced the raking and mowing parts of the traditional management over four years in a two-by-two factorial experiment in three traditionally managed wooded hay meadows on the Baltic island of Gotland, Sweden. We found decreased litter and hay production in two of the three studied meadows as a result of our treatments. Plant sizes and asymptotic population growth rates (??) of S. pratensis increased, particularly in plots receiving the combined raking and mowing treatment. Stochastic long-term population growth rates (?? _s ) increased under the reinforced management: projected population sizes 50?years into the future showed a three-fold increase in raked plots and a 17-fold increase in plots that were both raked and mown. Because we found positive responses even in these seemingly well-managed meadows we conclude that it is essential that management is carried out more thoroughly to ensure viable population sizes. Our conclusion applies to most semi-natural grasslands receiving anthropogenic nitrogen, or where traditional management practices are less rigorously applied. We also suggest using biomass estimation instead of vegetation height as a measure of management strength.     

How can we protect rare hemiparasitic plants? Early-flowering taxa ofEuphrasia andRhinanthus on the Baltic island of Gotland

Euphrasia stricta var.suecica, E. stricta var.tenuis, andRhinanthus serotinus subsp.vernalis are three endangered, hemiparasitic annual herbs found in traditionally managed hay meadows on Gotland, Sweden. We have studied — experimentally and in the field — how some features in the present and traditional management cycle affect their fitness. We set up permanent 50 × 50-cm plots in a coastal hay meadow where bothE. stricta var.suecica andRhinanthus are still quite common. The survival, growth and fecundity of cohorts ofEuphrasia andRhinanthus were followed throughout the growing season for three years. The length of the growing season was vital forEuphrasia andRhinanthus performance. In cooler summers, the percentage ofEuphrasia individuals that produced mature seeds was decreased by 20% at the time of mowing. This indicates the need to individually adjust the time of mowing, as was formerly done, to the actual phenological development in order to maintain healthy populations. It is also important to consider the effect of time of mowing on total species richness, asEuphrasia more successfully established in 10 × 10-cm squares with high species richness. Also, the time in spring when hemiparasite growth started was crucial. Cohorts of bothEuphrasia stricta var.suecica andRhinanthus that connected to hosts and started growing early in the season had a strong advantage over later cohorts by having fitness values four to eight times higher. From this we conclude that the traditional practice of spring raking is important for the long-term persistence of these hemiparasites, since raking promotes an earlier onset of both host and hemiparasite growth. Another important issue is the traditional, but nowadays often neglected practice of letting the hay dry in the meadow after mowing. Hay that was left to dry in the meadow contributed significantly moreRhinanthus seed to the meadow than hay that was taken away immediately after mowing. We also found that seedling emergence and subsequent growth ofEuphrasia stricta var.suecica was greatly enhanced by a second hay cut in September. A second cutting mimics some of the positive effects of the traditional practice of aftermath grazing, which is nowadays often abandoned.     

Impact of management on biodiversity-biomass relations in Estonian flooded meadows

A negative species richness–productivity relationship is often described in grasslands at smaller scales. We aimed to study the effect of management on this relationship. In particular, we addressed the relative importance of biomass cutting, hay removal and nutrient impoverishment on species richness and growth form structure. We conducted fieldwork in flooded meadows in Alam-Pedja Nature Reserve, central Estonia. We sampled vegetation in managed and abandoned stands of two types of alluvial meadows, sedge and tall forb meadow. Aboveground biomass and litter were harvested, weighed and analysed for major plant nutrients by near infrared reflectance spectroscopy. Three groups of general additive models were developed and compared, addressing the impact of (i) productivity, (ii) nutrients and (iii) management regime on species richness. Management—mowing and hay removal—reduced the amount of litter but not aboveground biomass. Management led to a decrease in nitrogen in the biomass and enhanced species richness, particularly in the tall forb meadow. The strongest determinant of species richness was the amount of litter, exhibiting a hump-shaped relationship with richness. The effect of nitrogen supply was significant, but explained less variation. Management increased the proportion of sedges in the sedge meadow and of small herbs in the tall forb meadow. We conclude that litter removal is the most important management means to support biodiversity. On highly productive sites, reducing nutrients via hay removal is of secondary importance within a timeframe of 10 years.     

Effects of changes in farming of subalpine meadows on a grassland bird, the whinchat ( Saxicola rubetra)

The increasing intensity of farming of meadows is supposed to be the major cause for strong declines of many meadow breeding birds. The whinchat, Saxicola rubetra, a Palaearctic migratory bird, is an indicator species of open grassland farmed at a low intensity. Originally widespread throughout Switzerland, it is now restricted to mountain and subalpine grassland. We document the changes in meadow cultivation in subalpine farmland from 1988 to 2002, and the breeding performance and density of the whinchat. We explored the impact of habitat degradation on the population dynamics of this meadow bird. The cultivation of hay meadows changed markedly within the 15 years: the onset of mowing was shifted forward by about 20 days, and farmers applied new techniques such as silage and irrigation. This shift was more pronounced in the favourable farmland in the valley bottom (Pradellas) than on the slopes (Vnà). The percentage of successful whinchat broods, ranging from 5% to 78% in different years, strongly depended on mowing date on both sites. In spite of earlier mowing, birds did not change their time schedule of breeding. Breeding success in Pradellas was too low to compensate for mortality (sink population), but because of immigration the number of breeding pairs did not decrease untill 2000. Population size may therefore be a misleading indicator of local population viability. Based on the breeding schedule of whinchats at different altitudes, we recommend mowing dates in agreement with the reproductive cycle of ground nesting meadow birds.     

Artificial Dispersal as a Restoration Tool in Meadows: Sowing or Planting?

Habitat fragmentation and the abandonment of former agricultural practices have led to extremely low dispersal rates for plant species growing in traditionally managed hay meadows in Sweden. Historically, seed dispersal between populations was maintained by hay movement, grazing animals, and farmers sharing their equipment. Because these means of dispersal typically are no longer occurring, artificial dispersal using seeds and plug-plants is tested here as a restoration tool. In this study, we chose two perennial herbs commonly occurring in meadows as test species, viz, Hypochoeris maculata L. (Asteraceae) and Succisa pratensis Moench. (Dipsacaceae). We found that plug-plant transplants were twice as effective as seed sowing for both species. The seed collection site was found to be important for seed-based establishment and survival; consequently, the choice of donor meadow is important when acquiring seeds used for restoration. We also found that survival of plants introduced as seeds was generally lower at sites harboring species favored by nitrogen as well as at sites in later successional phases. Both methods of introducing meadow species worked well, even though long-term establishment may well be more successful with the plug-plant method due to higher plug-plant establishment 2 years after introduction in the field.     

Seed bank diversity in mesic grasslands in relation to vegetation type,management and site conditions

Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow‐pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above‐ground vegetation, edaphic factors and land‐use history? 3. Are there differences in C‐S‐R plant strategy types and seed longevity under different management regimes? Location: Lahn‐Dill Highlands in central‐western Germany. Methods: Above‐ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above‐ground vegetation, site conditions and land‐use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above‐ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R‐strategy compared to hay meadows. In contrast, in hay meadows and meadow‐pastures proportions of S‐strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re‐establish from the seed bank after disappearance from above‐ground vegetation.     

Disturbance by mowing affects clonal diversity: the genetic structure of Ranunculus ficaria (Ranunculuaceae) in meadows and forests

To study the impact of disturbance by mowing on clonal variation, we compared the genetic structure of Ranunculus ficaria (Ranunculaceae) in meadows and forests located in southeast Germany. We applied random amplified polymorphic DNA (RAPD) analysis to investigate the clonal and genetic diversity and analysed a total of 117 samples from three study plots in each habitat type. Polymerase chain reaction with six primers resulted in 57 fragments. Clonal diversity differed clearly between the two analysed habitat types and was significantly higher in the study plots from meadows than in those from forests. The mean percentage of distinguishable genotypes (PD) was 0.80 in meadow plots and 0.36 in forest plots, and the detected genets were smaller in meadow plots than in forest plots. Mean genetic diversity measured as percentage of polymorphic bands, Shannon’s information index and Nei’s gene diversity was also higher in meadows (44.4, 0.22 and 0.14) than in forests (25.1, 0.09 and 0.05). The higher level of clonal diversity in meadow plots is most likely due to the effects of disturbance by mowing, which increases the dispersal of bulbils and promotes the establishment of new plants in meadows compared to forests.     

Is regular mowing the most appropriate and cost-effective management maintaining diversity and biomass of target forbs in mountain hay meadows?

Conservation of grassland biodiversity is a key issue in the EU agro-biodiversity policy. We assessed the effects of yearly mowing on target forb biomass in years with contrasting precipitation (2006–2007) in mountain fen and dry-mesophilous hay meadows in NE-Hungary. We hypothesised: (i) Species richness and biomass of target forbs is higher in mown than in abandoned stands. (ii) Mowing has more an effect on the biomass of target forbs, graminoids and litter than precipitation.     

Seedling recruitment in flood‐meadow species: The effects of gaps,litter and vegetation matrix

Question: Is the failure of establishment of rare flood‐meadow species in habitat restoration primarily due to seed or microsite limitation? How do sown species respond to artificially created gaps and added litter at two neighbouring sites with similar physical conditions but contrasting vegetation matrix (young arable fallow field vs species‐poor meadow sward)? Location: Upper Rhine valley, southwestern Germany, 85 m a.s.l . Methods: Seeds of six typical flood‐meadow species were sown in four treatment combinations of the factors gap creation and litter addition. Seedling recruitment was monitored for three years. Results: Five of the six species established successfully at both sites largely irrespective of treatments, indicating seed limitation. Only in the small‐seeded Arabis nemorensis, which was revealed to be strictly gap‐dependent at the meadow site, could an obvious microsite limitation be shown. The non‐significance of gap treatments in all other species at the relatively high productive meadow site is probably due to biomass removal by mowing in early summer. Only at the extremes of the seed size spectrum did the results meet predictions of plant ecological theory, such as the strict gap dependence of small‐seeded species in closed swards or the positive to neutral response of large‐seeded species to litter layers. Conclusions: Species identity was revealed to be the major factor influencing differences in recruitment. Due to the lack of a general trend in the response towards treatments the results support conceptual models that describe the interplay of facilitation and interference as a highly dynamic equilibrium, driven by variable abiotic and biotic marginal conditions.     

How is Regeneration of Plants after Mowing Affected by Shoot Size in Two Species-Rich Meadows with Different Water Supply?

Mowing a meadow is an example of an equalizing process that reduces differences among species by removing aboveground biomass approximately 5 cm above ground. This regular disturbance that affects all plants prevents competitive exclusion of small species and thus allows coexistence of numerous species differing in shoot size. In this paper we search for the mechanism behind this by comparing the shoot biomass of 41 common species in dry and wet species-rich meadows in mown and recently abandoned plots in June (before mowing) and in October. We asked the following questions: i) Do the plants differ in proportion of biomass lost by mowing? ii) Are the mown plants able to compensate for biomass lost by mowing? iii) Is the compensatory ability of mown plants related to their size? iv) Is the compensatory ability of plants related to severity of disturbance (removed biomass)? v) Does water availability in meadows affect these features? Our results revealed that the earlier explanation of equalization of meadow plants after mowing due to the proportionally larger biomass loss in larger plants than small plants does not represent the entire mechanism. Even when larger plants in the wet meadow lost more biomass, the proportion of lost biomass was not dependent on plant size, and compensation ability (growth of mown in comparison with unmown plants) was not related to the lost biomass in this meadow type. On the contrary, the observed pattern could be explained by different compensation abilities of small versus tall plants. In addition, according to our expectations, the compensation for lost biomass in the wet meadow was higher than in the dry one.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

Nutrient status,disturbance and competition: an experimental test of relationships in a wet meadow

Abstract. In many species-rich hay meadows in Central Europe, the traditional extensive (low input, low output) management is no longer economical and meadows are either fertilized or abandoned. Both these practices lead to changes in species composition and usually to a loss in species diversity. The response of a species-rich meadow plant community to fertilization, mowing and removal of dominant species was studied in a manipulative factorial experiment over four years. Both species richness and seedling recruitment were positively influenced by mowing and to a lesser extent by removal of the dominant species, Molinia caerulea, and were negatively influenced by fertilization. Fertilization caused an immediate increase in community biomass. Response to removal of the dominant species was delayed by one season, continued over the whole period, and by the fourth year the biomass reached a similar value as in the plots with Molinia present. Changes in species composition were evaluated by RDA for repeated measurements. The best and only significant predictor of species response to fertilization was plant height. This shows that with increased nutrient availability, nutrient limitation weakens and competition for light becomes the decisive factor. Competition for light appears to be more asymmetric than competition for nutrients, and consequently, it is more likely to drive inferior species to extinction. This is, together with seedling recruitment limitations, the most important cause of a decrease in species richness under high nutrient levels.