Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Predicting bryophyte diversity in grassland and eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the environmental factors associated with bryophyte diversity in remnants in a fragmented, agricultural landscape. Location Eighty‐two remnants of tussock grassland, eucalypt woodland and eucalypt forest in the subhumid Midlands region of Tasmania, Australia. Methods Remnants were surveyed for bryophytes and predictor variables, such as vascular plant cover, climate, and topography. Management histories for each remnant were compiled using both site observation and landowner surveys. Bryophyte cover, richness, and composition were related to the independent variables using simple correlation and general linear models. Results We found weak relationships between the dependent variables and the fragmentation variables (remnant area, remnant perimeter to area ratio, distance to nearest remnant, distance to nearest larger remnant, and remnant age). Instead, climatic variables were important in predicting bryophytes, in particular those affecting humidity (minimum temperature of the coldest month, precipitation). Despite extensive sheep grazing in this landscape, grazing was not correlated with bryophyte diversity. Bryophyte diversity was not explained by vascular plant richness and was only weakly explained by composition, but was predicted by the cover of vascular plants. There was greater bryophyte cover and richness and different composition where the cover of native vascular plants was lower. Main conclusions The implications of our results are that all remnants, regardless of area, age and isolation, appear to be valuable for bryophyte conservation in this highly altered landscape. Our results also suggest that the cover of the vascular plant community, rather than its diversity, holds promise as a guide to bryophyte diversity. Bryophyte composition was similar between sites and a focus on the most species‐rich sites may be the best conservation strategy in this ecosystem.     

Bryophyte biomass and species richness on the Park Grass Experiment,Rothamsted, UK

The relationships between bryophyte biomass and species richness and soil pH, nutrient applications and vascular plant biomass and species richness were analyzed for the Park Grass Experiment (Rothamsted, UK). The study examined the abundance of bryophytes in relation to long-term fertilizer and lime application and to fertilizer treatments recently being ceased on some plots. The probability of bryophytes being present on a plot increased with increasing soil pH, and on plots at soil pH 3.3–4.5, the lowest values in this experiment, there were virtually no mosses present. Total bryophyte biomass decreased with increasing vascular plant biomass and vascular plant richness. Both bryophyte biomass and species richness showed a curvilinear response to soil pH. Bryophyte biomass was markedly increased on plots where nitrogen (N) fertilization had recently been ceased. The abundance of the common bryophyte species showed individualistic responses to treatments. N had a negative effect on the abundance of Brachythecium rutabulum. Increasing soil pH, and the application of phosphorus (P) and potassium (K) fertilizer together, had a positive effect on Eurhynchium praelongum. This species was also negatively affected by N, but tolerated larger amounts of it (100–150 kg ha^–1 N) than B. rutabulum. An ephemeral moss, Bryum subapiculatum, had a unimodal response to soil pH but showed no response to N, P, K or other explanatory variables.     

Vascular Plants Facilitated Bryophytes in a Grassland Experiment

In grassland communities vascular plants and bryophytes form two distinct layers. In order to understand the factors responsible for plant community structure, more information about interactions between these plant groups is needed. Often negative correlations between vascular plant and bryophyte covers have been reported, suggesting competition. Here we tested experimentally whether different grassland vascular plant species (Trifolium pratense, Festuca pratensis, Prunella vulgaris) had different influences on the cover of two bryophyte species (Rhytidiadelphus squarrosus, Brachythecium rutabulum). In a two-year garden pot experiment one bryophyte species and one vascular plant species were planted per pot. Bryophytes were planted at a constant density, vascular plants in four densities. The cover of both bryophyte species increased with increasing vascular plant cover, showing the facilitative effect of vascular plants through creating better microclimate, e.g., optimising temperature. Bryophyte responses to vascular plant species were species-specific. Festuca had significantly positive effects on both bryophyte species in the second year, and Trifolium on Brachythecium in both years, whereas Prunella had no significant effect on bryophytes. The facilitative effect of vascular plants was stronger at the second experimental year. In summary, the biotic effects between bryophytes and grassland vascular plants are species-specific and positive interactions are prevailing at low vascular plant densities.     

Edge effect in different groups of organisms: vascular plant, bryophyte and fungi species richness across a forest-grassland border

Six transects were established on the edges of two deciduous forests near Krosno (Carpathian Foothills) to compare the species richness pattern of vascular plants, bryophytes and fungi. The transects had the shape of a cross with one arm 10 m along the forest edge and the other across the edge, 50 m into the forest interior and 12 m into the grassland. They consisted of 2×2 quadrats. The strongest edge effects were recorded for bryophyte, shrub and tree species richness, the weakest for herbaceous plant species richness. Overall vascular plant species richness and herbaceous species richness were higher in grassland than in the forest and peaked in grassland, 3 m from the forest edge. The shrub species richness was highest 1 m from the edge (in the forest) and the tree species richness 3 m from edge. Bryophyte species richness had roughly the same level across the grassland and within the first several meters of the forest, except for the 2 m zone on the edge itself where species richnes was as low as in the forest interior. Fungi species richness was low in the grassland and on the forest edge and rose dramaticaly a few meters from the edge, ramaining at the same level within the forest. The species composition across the forest-grassland border was analysed using detrended correspondence analysis. It revealed that in the case of bryophytes the increase in species richness did not correspond to a change in species composition, such as might have been caused by a general increase in bryophyte density.     

高海拔幼林地表苔藓组成与结构——皆伐与造林实践的影响评估

 对壤塘县二林场4个不同时间皆伐和造林后形成的幼林地, 开展了苔藓植物和维管植物多样性调查, 采用方差分析法对苔藓植物特征进 行差异性检验, 对苔藓植物结构特征和环境因子作偏相关性分析。结果表明: 1)幼林地4个阶段的发展进程中苔藓物种组成变化显著。优势种相 似, 为喜光耐旱的侧蒴藓物种; 一些耐阴喜湿的种类种群扩大成为林下局部生境优势物种; 同时由于干扰造成地表裸露后产生耐旱土生或石生 性广泛分布的藓类。2)随着幼林地植被恢复进程, 苔藓物种丰富度(22～37)和多样性指数(0.37～1.08)呈增长趋势。结构特征在4个幼林地表现 出较强的差异性。3)乔木与灌木层盖度与苔藓植物的发育呈正相关, 灌木层盖度的发育明显促进了苔藓植物丰富度的增加, 而基于喜光先锋的 禾草为主的草本发育程度很高, 与凋落物生物量一起抑制了苔藓的发育。随幼林地发展, 乔、灌、凋落物发育性质及其相互作用发生动态变化 。因此加速幼林地乔、灌层片的发育对于促进地表苔藓植物多样性的恢复与层片发育具有重要作用。     

Is vascular plant species diversity a predictor of bryophyte species diversity in Mediterranean forests?

This study aimed to (i) investigate the congruence among the species composition and diversity of bryophytes and vascular plants in forests; (ii) test if site prioritization for conservation aims by the maximization of the pooled number of vascular plant species is effective to maximize the pooled number of bryophyte species. The study was performed in six forests in Tuscany, Italy. Four-hundred and twenty vascular plant species (61 of which were woody) and 128 bryophyte species were recorded in 109 plots. Despite the good predictive value of the compositional patterns of both woody plants and total vascular with respect to the compositional pattern of bryophytes, the species richness of the latter was only marginally related to the species richness of the former two. Bryophyte rare species were not spatially related to rare plant species and neither coincided with the sites of highest plant species richness. The species accumulation curves of bryophytes behaved differently with respect to those of woody plants or total vascular plants. Reserve selection analysis based on the maximization of the pooled species richness of either woody plants or total vascular plants were not effective in maximizing the pooled species richness of bryophytes. This study indicates that species diversity of vascular plants is not likely to be a good indicator of the bryophyte species diversity in Mediterranean forests.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

Bryophyte Species Richness and Composition along an Altitudinal Gradient in Gongga Mountain,China

An investigation of terrestrial bryophyte species diversity and community structure along an altitudinal gradient from 2,001 to 4,221 m a.s.l. in Gongga Mountain in Sichuan, China was carried out in June 2010. Factors which might affect bryophyte species composition and diversity, including climate, elevation, slope, depth of litter, vegetation type, soil pH and soil Eh, were examined to understand the altitudinal feature of bryophyte distribution. A total of 14 representative elevations were chosen along an altitudinal gradient, with study sites at each elevation chosen according to habitat type (forests, grasslands) and accessibility. At each elevation, three 100 m × 2 m transects that are 50 m apart were set along the contour line, and three 50 cm × 50 cm quadrats were set along each transect at an interval of 30 m. Species diversity, cover, biomass, and thickness of terrestrial bryophytes were examined. A total of 165 species, including 42 liverworts and 123 mosses, are recorded in Gongga mountain. Ground bryophyte species richness does not show any clear elevation trend. The terrestrial bryophyte cover increases with elevation. The terrestrial bryophyte biomass and thickness display a clear humped relationship with the elevation, with the maximum around 3,758 m. At this altitude, biomass is 700.3 g m⁻² and the maximum thickness is 8 cm. Bryophyte distribution is primarily associated with the depth of litter, the air temperature and the precipitation. Further studies are necessary to include other epiphytes types and vascular vegetation in a larger altitudinal range.     

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).     

Endozoochory by slugs can increase bryophyte establishment and species richness

Herbivory can affect plant community composition and diversity by removing biomass and reducing light competition. Herbivory may particularly benefit low growing species such as bryophytes, which are frequently limited by light competition. Gastropods are important herbivores of seed plants and cryptogams, furthermore, they can disperse propagules such as seeds and spores via endozoochory. However, whether gastropod herbivory can reduce the dominance of vascular plants and thereby promote the germination and establishment of endozoochorously dispersed bryophyte spores has never been tested experimentally. Moreover, it is unclear whether these possible interacting effects can influence bryophyte species richness. Here, we tested for endozoochorous spore dispersal by slugs, in combination with sowing of vascular plants, in a fully factorial common garden experiment. Enclosures contained either slugs previously fed with bryophyte sporophytes, control slugs, or no slugs. After 21 days the bryophyte cover was on average 2.8 times higher (3.9% versus 1.4%) and after eight months the bryophyte species richness 2.6 times higher (5.8 versus 2.2) in enclosures containing slugs previously fed with bryophyte sporophytes than in the other treatments. Furthermore, after eight months high vascular plant cover reduced bryophyte diversity. On average enclosures without seed sowing harboured 1.6 times more bryophyte species than the ones with seed sowing (4.2 versus 2.6), indicating competitive effects of vascular plants on bryophytes. Our findings suggest that slugs are important dispersal vectors for bryophytes and that they can increase bryophyte populations and maintain bryophyte diversity by reducing the dominance of vascular plants.     

Bryophyte and Lichen Diversity Under Simulated Environmental Change Compared with Observed Variation in Unmanipulated Alpine Tundra

Effects of simulated environmental change on bryophyte and lichen species richness and diversity in alpine tundra were investigated in a 5-year experiment at Latnjajaure, northern Sweden. The experiment had a factorial design including fertilisation and temperature enhancement in one meadow and one heath plant community. Responses in species richness, biodiversity, and species composition of bryophytes and lichens to experimental treatments were compared to the observed variation in six naturally occurring plant communities. The combination of fertilisation and enhanced temperature resulted in a species impoverishment, for bryophytes in the bryophyte-dominated community, and for lichens in the lichen-dominated communities, but the species composition stayed within the observed natural variation. During the course of the study, no species new to the investigated mid-alpine landscape were recorded, but that scenario is realistic within a decade when comparing with the processes seen in vascular plants.     

Flood events overrule fertiliser effects on biomass production and species richness in riverine grasslands

Question: Do severe winter flood events lift the nutrient limitation of biomass production in a river floodplain? How does this affect plant species richness? How long do the effects last? Location: Floodplain grassland on calcareous sandy loam near river Rhine in The Netherlands. Methods: Plots were fertilised with four treatments (control, N, P, N+P) for 21 years; plant species composition, vegetation biomass and tissue nutrient concentrations were determined every year between 1985 and 2005. Results: Fertilisation with N generally increased biomass production and reduced species richness, but these effects varied over time. During the first four years of the experiment, biomass production appeared to be co‐limited by N and P, while N fertilisation dramatically reduced plant species richness; these effects became weaker subsequently. Following two extreme winter floods in 1993–94 and 1994–95 and a drought in spring 1996, the effects of fertilisation disappeared between 1998 and 2001 and then appeared again. Flooding caused an overall reduction in species richness (from c. 24 to 15 species m^‐2) and an increase in biomass production, which were only partly reversed after ten years. Conclusions: Long time series are necessary to understand vegetation dynamics and nutrient limitation in river floodplains, since they are influenced by occasional flood and drought events, whose effects may persist for more than ten years. A future increase in flooding frequency might be detrimental to species richness in floodplain grasslands.     

The physiological basis of bryophyte production

In the main features of their carbon metabolism and physiological responses, bryophytes behave as normal C_:) plants. However, their small size and frequent poikilohydric habit have important effects on the context in which these characteristics are expressed, and on their environmental physiology. Many are tolerant of drying out to low water contents (c. 5–10%, of dry weight). Photosynthesis declines rapidly with water loss, and resumes with greater or lesser delay on remoistening. The rate and completeness of recovery depend on the intensity and duration of desiccation, and on drought-hardening (perhaps largely related to protection of cell components from oxidative damage) which lakes place as the bryophyte dries. Most bryophytes, including species of well-illuminated habitats, function in effect as shade plants, with low chlorophyll a/b ratios, and become light-saturated at relatively low irradiance. Boundary-layer resistance is critically important in determining water loss from bryophytes in many situations. The time for which a poikilohydric species can photosynthesize after rain is determined by storage capacity and rate of water loss, both strongly influenced by growth-form. In sheltered habitats with extensive bryophyte cover water loss is largely determined by radiation balance, and may be very slow in deeply shaded places. Bryophyte growth-forms must represent an adaptive balance between water economy and needs for light capture and carbon and mineral nutrient acquisition.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Epigeic bryophytes do not improve bioindication by Ellenberg values in mountain forests

Based on a stratified random sample of 93 vegetation plots and coincident measurements of ecological conditions in mountain forests of the Bavarian Alps, the degree to which species composition and Ellenberg indicator values derived thereof were related to measured environmental variables was assessed for vascular understorey plants and epigeic bryophytes. According to Mantel tests vascular composition contained ca. 30% more ecological information than bryophyte composition. When expressed as average Ellenberg or Düll values, vascular plant-based values reflected 60% more of measured variables than bryophyte-based values. The differences remained after rarefaction of the vascular matrix to the gamma diversity of bryophytes, showing that indication is not a function of indicator richness. Analysing vascular plants and bryophytes combined yielded very similar, or even slightly less stringent relationships with the environment than using vascular plants only.Bivariate relationships of indicator values with corresponding ecological measurements confirmed the specific potential of the values to estimate ecological factors from both plant groups, but vascular plants performed better for all factors. Bryophyte indication was particularly poor for light, temperature and base saturation. Bryophyte-based indicator values did not significantly predict the residuals of measured ecological variables against vascular plant-based Ellenberg values.For the study region, it is concluded that indicator values of vascular forest understorey should be used without consideration of Düll's indicator values for epigeic bryopyhtes. There appears to be potential to improve bioindication by recalibrating indicator values of epigeic bryophytes based on ecological measurements and vascular plant indicator values.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Species richness of vascular plants,bryophytes, and lichens along an altitudinal gradient in western Norway

Species richness patterns of ground-dwelling vascular plants, bryophytes, and lichens were compared along an altitudinal gradient (310–1135 m a.s.l.), in western Norway. Total species richness peaked at intermediate altitudes, vascular plant species richness peaked immediately above the forest limit (at 600–700 m a.s.l.), bryophyte species richness had no statistically significant trend, whereas lichen richness increased from the lowest point and up to the forest limit, with no trend above. It is proposed that the pattern in vascular plant species richness is enhanced by an ecotone effect. Bryophyte species richness responds to local scale factors whereas the lichen species richness may be responding to the shading from the forest trees.     

Impact of climate change on alpine vegetation of mountain summits in Norway

Climate change is affecting the composition and functioning of ecosystems across the globe. Mountain ecosystems are particularly sensitive to climate warming since their biota is generally limited by low temperatures. Cryptogams such as lichens and bryophytes are important for the biodiversity and functioning of these ecosystems, but have not often been incorporated in vegetation resurvey studies. Hence, we lack a good understanding of how vascular plants, lichens and bryophytes respond interactively to climate warming in alpine communities. Here we quantified long-term changes in species richness, cover, composition and thermophilization (i.e. the increasing dominance of warm-adapted species) of vascular plants, lichens and bryophytes on four summits at Dovrefjell, Norway. These summits are situated along an elevational gradient from the low alpine to high alpine zone and were surveyed for all species in 2001, 2008 and 2015. During the 15-year period, a decline in lichen richness and increase in bryophyte richness was detected, whereas no change in vascular plant richness was found. Dwarf-shrub abundance progressively increased at the expense of lichens, and thermophilization was most pronounced for vascular plants, but occurred only on the lowest summits and northern aspects. Lichens showed less thermophilization and, for the bryophytes, no significant thermophilization was found. Although recent climate change may have primarily caused the observed changes in vegetation, combined effects with non-climatic factors (e.g. grazing and trampling) are likely important as well. At a larger scale, alpine vegetation shifts could have a profound impact on biosphere functioning with feedbacks to the global climate.     

Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands

Aims and Methods Disturbance is supposed to play an important role for biodiversity and ecosystem stability as described by the intermediate disturbance hypothesis (IDH), which predicts highest species richness at intermediate levels of disturbances. In this study, we tested the effects of artificial soil disturbances on diversity of annual and perennial vascular plants and bryophytes in a field experiment in 86 agricultural grasslands differing in land use in two regions of Germany. On each grassland, we implemented four treatments: three treatments differing in application time of soil disturbances and one control. One year after experimental disturbance, we recorded vegetation and measured biomass productivity and bare ground. We analysed the disturbance response taking effects of region and land-use-accompanied disturbance regimes into account.Important findings Region and land-use type strongly determined plant species richness. Experimental disturbances had small positive effects on the species richness of annuals, but none on perennials or bryophytes. Bare ground was positively related to species richness of bryophytes. However, exceeding the creation of 12% bare ground further disturbance had a detrimental effect on bryophyte species richness, which corresponds to the IDH. As biomass productivity was unaffected by disturbance our results indicate that the disturbance effect on species richness of annuals was not due to decreased overall productivity, but rather due to short-term lowered inter- and intraspecific competition at the newly created microsites. Generally, our results highlight the importance of soil disturbances for species richness of annual plants and bryophytes in agricultural grasslands. However, most grasslands were disturbed naturally or by land-use practices and our additional experimental soil disturbances only had a small short-term effect. Overall, total plant diversity in grasslands seemed to be more limited by the availability of propagules rather than by suitable microsites for germination. Thus, nature conservation efforts to increase grassland diversity should focus on overcoming propagule limitation, for instance by additional sowing of seeds, while the creation of additional open patches by disturbance might only be appropriate where natural disturbances are scarce.