Soil Disturbance as a Grassland Restoration Measure—Effects on Plant Species Composition and Plant Functional Traits

Soil disturbance is recognized as an important driver of biodiversity in dry grasslands, and can therefore be implemented as a restoration measure. However, because community re-assembly following disturbance includes stochastic processes, a focus only on species richness or establishment success of particular species will not inform on how plant communities respond ecologically to disturbance. We therefore evaluated vegetation development following disturbance by quantifying species richness, species composition and functional trait composition. Degraded calcareous sandy grassland was subjected to experimental disturbance treatments (ploughing or rotavation), and the vegetation was surveyed during four subsequent years of succession. Treated plots were compared with control plots representing untreated grassland, as well as nearby plots characterized by plant communities representing the restoration target.Species richness and functional diversity both increased in response to soil disturbance, and rotavation, but not ploughing, had a persistent positive effect on the occurrence of specialist species of calcareous sandy grassland. However, no type of soil disturbance caused the plant species composition to develop towards the target vegetation. The disturbance had an immediate and large impact on the vegetation, but the vegetation developed rapidly back towards the control sites. Plant functional composition analysis indicated that the treatments created habitats different both from control sites and target sites. Community-weighted mean Ellenberg indicator values suggested that the observed plant community response was at least partially due to an increase in nitrogen and water availability following disturbance. This study shows that a mild type of disturbance, such as rotavation, may be most successful in promoting specialist species in calcareous sandy grassland, but that further treatments are needed to reduce nutrient availability. We conclude that a functional trait based analysis provides additional information of the vegetation response and the abiotic conditions created, complementing the information from the species composition.     

Acidification of sandy grasslands – consequences for plant diversity

Questions: (1) Does soil acidification in calcareous sandy grasslands lead to loss of plant diversity? (2) What is the relationship between the soil content of lime and the plant availability of mineral nitrogen (N) and phosphorus (P) in sandy grasslands? Location: Sandy glaciofluvial deposits in south‐eastern Sweden covered by xeric sand calcareous grasslands (EU habitat directive 6120). Methods: Soil and vegetation were investigated in most of the xeric sand calcareous grasslands in the Scania region (136 sample plots distributed over four or five major areas and about 25 different sites). Environmental variables were recorded at each plot, and soil samples were analysed for exchangeable P and N, as well as limestone content and pH. Data were analysed with regression analysis and canonical correspondence analysis. Results: Plant species richness was highest on weakly acid to slightly alkaline soil; a number of nationally red‐listed species showed a similar pattern. Plant species diversity and number of red‐listed species increased with slope. Where the topsoil had been acidified, limestone was rarely present above a depth of 30 cm. The presence of limestone restricts the availability of soil P, placing a major constraint on primary productivity in sandy soils. Conclusions: Acidification of sandy grasslands leads to reduced abundance of desirable species, although the overall effect is rather weak between pH 5 and pH 9. Slopes are important for high diversity in sandy grasslands. Calcareous soils cannot be restored through shallow ploughing, but deep perturbation could increase the limestone content of the topsoil and favour of target species.     

Pastoral Practices to Reverse Shrub Encroachment of Sub-Alpine Grasslands: Dung Beetles (Coleoptera,Scarabaeoidea) Respond More Quickly Than Vegetation

In recent decades, pastoral abandonment has produced profound ecological changes in the Alps. In particular, the reduction in grazing has led to extensive shrub encroachment of semi-natural grasslands, which may represent a threat to open habitat biodiversity. To reverse shrub encroachment, we assessed short-term effects of two different pastoral practices on vegetation and dung beetles (Coleoptera, Scarabaeoidea). Strategic placement of mineral mix supplements (MMS) and arrangement of temporary night camp areas (TNCA) for cattle were carried out during summer 2011 in the Val Troncea Natural Park, north-western Italian Alps. In 2012, one year after treatment, a reduction in shrub cover and an increase in bare ground cover around MMS sites was detected. A more intense effect was detected within TNCA through increases in forage pastoral value, and in the cover and height of the herbaceous layer. Immediately after treatment, changes in dung beetle diversity (total abundance, species richness, Shannon diversity, taxonomic and functional diversity) showed a limited disturbance effect caused by high cattle density. In contrast, dung beetle diversity significantly increased one year later both at MMS and TNCA sites, with a stronger effect within TNCA. Multivariate Regression Trees and associated Indicator Value analyses showed that some ecologically relevant dung beetle species preferred areas deprived of shrub vegetation. Our main conclusions are: i) TNCA are more effective than MMS in terms of changes to vegetation and dung beetles, ii) dung beetles respond more quickly than vegetation to pastoral practices, and iii) the main driver of the rapid response by dung beetles is the removal of shrubs. The resulting increase in dung beetle abundance and diversity, which are largely responsible for grassland ecosystem functioning, may have a positive effect on meso-eutrophic grassland restoration. Shrub encroachment in the Alps may therefore be reversed, and restoration of grassland enhanced, by using appropriate pastoral practices.     

Landscape-scale crop diversity interacts with local management to determine ground beetle diversity

Crop diversification and maintenance of semi-natural habitats (grasslands and field boundaries) are suggested to enhance farmland biodiversity, but the relative importance of these factors remains poorly known. We evaluated how crop diversity and availability of semi-natural grasslands at a landscape-scale interacted with local farming management (three management types from low to high intensity: ley < winter wheat < sugar beet) in their effect on ground beetle assemblages in southern Sweden. Ground beetle diversity increased with crop diversity either independently of local management (Simpson species diversity), or only in the less intensively managed habitats (rarefied species richness). While ground beetle diversity in leys tended to increase with field boundary length, no such relationship was observed in winter wheat or sugar beet fields. In contrast, the landscape proportions of leys and semi-natural grasslands did not affect ground beetle species richness and diversity. We conclude that (a) semi-natural grasslands and leys may not function as source habitats at a landscape-scale if they comprise a low proportion of the total land-use, while (b) increasing crop diversity is correlated to ground beetle richness and diversity in agricultural landscapes dominated by arable land. The beneficial effect of landscape-scale crop diversification on farmland biodiversity may depend on the general level of agricultural intensity of a region.     

Close association between grasshopper and plant communities in suburban secondary grasslands and the indicator value of grasshoppers for conservation

Semi-natural grasslands in Japan have decreased due to management abandonment and urbanization over the last 100 years, but they remain in suburban areas in addition to rural areas. Because suburban grasslands have various land-use histories and disturbance regimes, plant and herbivorous insect communities are likely to differ among grassland types. To identify grasslands with high conservation value, we conducted a comprehensive survey of grasshoppers and plants in 150 grasslands with 5 grassland types differing in land-use history and current management in northern Chiba prefecture, Japan. We then analyzed the association of the distributions of grasshopper and plant species compositions. Our results showed that grasshoppers were classified into habitat specialists and generalists. Three out of four habitat specialists were almost exclusively found in semi-natural grasslands and vacant lots, while habitat generalists were commonly observed at the cropland margins. This habitat specialist–generalist distribution gradient corresponded well to that found in plant communities, which was probably due to current disturbance regimes. We suggest that vacant lots as well as semi-natural grasslands have high conservation value for grassland organisms of various taxa in suburban areas, and grasshoppers are candidate indicator species for monitoring grassland environments.

     

Recreating Grasslands in Swedish Rural Landscapes – Effects of Seed Sowing and Management History

Recent loss of plant species richness in Swedish semi-natural grasslands has led to an increase in grassland recreation and restoration. To increase the establishment of declining species favoured by grazing and to re-establish original species richness, seed sowing has been discussed as a conservation tool. In this study, I examined to what extent seed sowing in former arable fields increases species richness and generates a species composition typical of semi-natural grasslands. Six grassland species favoured by grazing (target species) and six generalist species favoured by ceased grazing, were studied in a seed-addition experiment. Four different seed densities were used on four different grassland categories, two grazed former arable fields, one continuously grazed grassland and one abandoned grassland. Target and generalist species emerged in all grassland categories, but seedling emergence was higher in the grazed than in the abandoned grassland. Target species had higher emergence in the two grasslands with the longest grazing continuity. Seedling emergence and frequency of established plants of each target species were positively associated. The largest fraction of seeds germinated at an intermediate sowing density, 20–50 seeds/dm², suggesting that aggregation of seeds positively affects emergence up to a certain threshold. In conclusion, artificial seed sowing may induce the recreation of typical grassland communities on former arable fields, which may be an important contribution to increase the total grassland area and species richness in the landscape.     

The effects of pH and disturbance on the bryophyte flora in calcareous sandy grasslands

The unique flora and fauna of calcareous sandy grasslands is threatened by acidification, eutrofication and changed land use. Restoration experiments in such grasslands in southern Sweden, including deep and shallow soil perturbation, allowed us to examine the effects of pH and disturbance on the bryophyte flora. We also studied natural pH and disturbance gradients in order to compare the natural pH and disturbance variation with the responses of experimental manipulation. We found that increased pH due to soil perturbation resulted in increased species richness. However, in naturally disturbed areas, pH seemed to have a lower effect, and instead, the location of the site and possibly the site history had a larger influence on species richness. We also found that some of the species that were common at naturally high pH (mainly acrocarps) were favoured by experimental pH manipulation. Our results pointed out Syntrichia ruraliformis as a sensitive pH indicator, being almost exclusively restricted to pH values above 7. The species can also serve as an indicator of sand steppe vegetation.     

环境因子和干扰强度对高寒草甸植物多样性空间分异的影响

采用样带与样地结合的方法在三江源自然保护区的核心区沿海拔梯度在阴坡、阳坡分别进行草本植被调查,通过因子分析和偏相关分析研究丰富度指数、多样性指数与环境梯度（包括海拔梯度、裸斑面积、坡度、土壤总碳、总氮含量、土壤pH值、土壤总可溶性盐含量）和干扰强度（鼠类干扰强度、放牧强度）之间的关系。研究结果表明：杂类草丰富度指数（DMa杂）与总物种丰富度指数（DMa总）极显著相关（P<0.01）;阳坡DMa杂和DMa总均呈现“中海拔膨胀”现象,阴坡DMa杂和DMa总与海拔梯度呈正相关,莎草科和禾本科的丰富度指数（DMa莎和DMa禾）随海拔升高并无明显规律;通过主成分分析,及偏相关分析,第一主成分（裸斑面积、鼠类干扰和放牧强度）与除莎草科Margalef丰富度指数、禾本科Simpson指数和禾本科Pielou均匀度指数外的其他草地多样性指数均显著相关,是影响阳坡草地植物多样性的主要因子,土壤总碳、总氮含量对阳坡禾本科类群的多样性指数和均匀度指数有极显著影响,土壤pH值、TDS含量和坡度对阳坡莎草科类群的丰富度有显著影响;海拔梯度、土壤总碳、总氮以及pH值对阴坡草本植物群落的多样性影响较大。研究结论认为,植物群落生物多样性的空间分异特征是地理环境、土壤环境以及干扰强度等因素综合作用的结果。无干扰或干扰较弱时,物种多样性主要受土壤环境状况所影响;而在强干扰存在条件下,干扰强度对物种丰富度和多样性的影响比环境因子更显著;遏制高寒草甸植物多样性降低应首先控制放牧及鼠类等强干扰活动。     

Mechanical soil disturbance as a determinant of arbuscular mycorrhizal fungal communities in semi-natural grassland

While the effect of disturbance on overall abundance and community composition of arbuscular mycorrhizal (AM) fungi has been researched in agricultural fields, less is known about the impact in semi-natural grasslands. We sampled two AM plant species, Festuca brevipila and Plantago lanceolata, from an ongoing grassland restoration experiment that contained replicated plowed and control plots. The AM fungal community in roots was determined using nested PCR and LSU rDNA primers. We identified 38 phylotypes within the Glomeromycota, of which 29 belonged to Glomus A, six to Glomus B, and three to Diversisporaceae. Only three phylotypes were closely related to known morphospecies. Soil disturbance significantly reduced phylotype richness and changed the AM fungal community composition. Most phylotypes, even closely related ones, showed little or no overlap in their distribution and occurred in either the control or disturbed plots. We found no evidence of host preference in this system, except for one phylotype that preferentially seemed to colonize Festuca. Our results show that disturbance imposed a stronger structuring force for AM fungal communities than did host plants in this semi-natural grassland.     

Quality of remnant indigenous grassland linkages for adult butterflies (Lepidoptera) in an afforested African landscape

Retention of interconnected, remnant grassland linkages is proposed here to reduce the adverse effects of alien pine afforestation in Afromontane grasslands. Adult butterflies were sampled at 38 grassland sites, representing increasing levels of disturbance both within the afforested area and outside it. Butterfly species richness and abundance in the lesser disturbed grassland remnants within the afforested area were similar to those of the surrounding natural grasslands. In contrast, butterfly species richness, but not necessarily abundance, decreased significantly in the highly disturbed sites, both in the grassland linkages and outside. Although some highly disturbed sites were relatively rich in species, most were visited by geographically widespread and vagile species. In contrast, wide, relatively undisturbed grassland linkages, as well as grasslands outside, were important for localised, sedentary and local endemic butterfly species. Nectar plants, especially the alien Verbena bonariensis, were the most significant variable explaining local butterfly distribution. In addition, tall grasses, hills, topographical landmarks, thermoregulatory sites, shelter and water features were also vital for particular species. It did not matter how deep the grassland linkages were situated inside the afforested area, as long as they were made up of good habitat. Retention of wide, quality grassland linkages are a way forward to maximise biodiversity alongside agroforestry.     

The importance of disturbance for the conservation of a low-competitive herb in mesotrophic grasslands

For the protection or re-establishment of species-rich grasslands, the limiting factors controlling species richness have increasingly become of scientific interest. This study aims to analyze the role of disturbance for the occurrence of a low-competitive herb, Centaurium erythraea Rafn, in mesotrophic, lowland grasslands in NW Germany. We sampled a total of 38 plots with presence of C. erythraea and 24 control (random) plots in semi-natural grasslands. As a proxy for disturbance, we estimated the cover of bare ground and Ellenberg flooding indicator species and measured the distance between the plot and the nearest path. Moreover, we counted the number of C. erythraea individuals within each plot. In the GLM analyses the distance from path was the only predictor; both the presence of the species and the number of individuals decreased with the distance from path. The grasslands at the path edges had the highest disturbance intensity. Here horse riding, military-vehicle traffic and regular sod-cutting directly create bare ground. In general, disturbance creating bare ground seems to be the key factor enabling germination and growth of C. erythraea in mesotrophic grasslands and other low-competitive short-lived species. Disturbance enhances the expression of the seed bank, favours the development of the shade-avoiding and low-growing rosettes and oppresses tall-growing competitors. Therefore we suggest grazing as the best management method.     

不同母质温带草地植物群落多样性对人为干扰的响应

本研究以赛罕乌拉保护区2种不同土壤母质(黄土和沙母质)与3种利用方式(围栏打草、季节性放牧和自由放牧)草地为对象,基于群落组成调查数据,计算人为干扰指数与物种多样性指数,研究草地植物群落物种多样性与草地退化程度间的关系。结果表明: 土壤母质与人为利用方式的差异使草地处于不同的退化状态,黄土与沙母质类型草地退化程度均随人为利用强度的增加而上升,且当人为利用方式相同时,黄土母质草地人为干扰指数(均值1.21)比沙母质草地(均值1.48)低。各样地物种多样性指数总体随土壤母质的砂质化和人为利用强度的增加而下降,其中Margalef丰富度指数为1.57～4.27,Shannon多样性指数为1.16～2.39,Simpson优势度指数为0.76～0.87,Pielou均匀度指数为0.71～0.80。随着人为干扰指数增加,丰富度指数、多样性指数和优势度指数均下降,而均匀度指数有上升趋势。过度放牧对2种土壤母质草地均会造成严重威胁,黄土与沙母质草地最适宜的利用方式分别为围封打草和季节性放牧。在今后生物多样性保护的实施过程中,需兼顾草地不同土壤母质与人为利用方式的影响,针对不同土壤母质条件规划不同的草地利用方式,实现因地制宜的草地恢复与管理。     

Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape

The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strá?ovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16?m². The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.     

How do subordinate and dominant species in semi-natural mountain grasslands relate to productivity and land-use change?

Changes in agricultural practices of semi-natural mountain grasslands are expected to modify plant community structure and shift dominance patterns. Using vegetation surveys of 11 sites in semi-natural grasslands of the Swiss Jura and Swiss and French Alps, we determined the relative contribution of dominant, subordinate and transient plant species in grazed and abandoned communities and observed their changes along a gradient of productivity and in response to abandonment of pasturing.The results confirm the humpbacked diversity–productivity relationship in semi-natural grassland, which is due to the increase of subordinate species number at intermediate productivity levels. Grazed communities, at the lower or higher end of the species diversity gradient, suffered higher species loss after grazing abandonment. Species loss after abandonment of pasturing was mainly due to a higher reduction in the number of subordinate species, as a consequence of the increasing proportion of dominant species.When plant biodiversity maintenance is the aim, our results have direct implications for the way grasslands should be managed. Indeed, while intensification and abandonment have been accelerated since few decades, our findings in this multi-site analysis confirm the importance of maintaining intermediate levels of pasturing to preserve biodiversity.     

Size-related deterioration of semi-natural grassland fragments in Sweden

Abstract. One of the most dramatic landscape changes during the 20th century in Sweden, like in most of Europe, has been the reduction and fragmentation of semi-natural grasslands. Using a set of remnant semi-natural grasslands, chosen to be as similar as possible, but differing in size, we have examined whether size of remnant fragments of traditionally managed semi-natural grasslands in Sweden is related to patterns of species richness and composition. We focused on edge-to-interior relationships, since we expected that a possible impact from invasive habitat generalists would be manifested in a gradient from the edge of fragments to their interior. We found no relationship between size of grassland fragments and (a) overall species richness, (b) species richness at different spatial scales, and (c) abundance of some typical invader species or species characteristic of semi-natural grasslands. However, the results indicated that larger grasslands have a comparatively larger number of species in the edges, whereas the opposite pattern was found in smaller grasslands. The similarity in species composition between the edge and the interior of the pastures also increased with grassland size. Thus, even though the overall species richness is still unaffected by reduction in grassland fragment size, the edges of smaller grasslands show signs of degradation, i.e. reduction in species richness and a decreased similarity to the grassland interior. We suggest that these kinds of effects may be early signs of fragmentation effects that in the future will result in species loss even if the present distribution of semi-natural grasslands is maintained.     

Can re-establishment of cattle grazing restore bryophyte diversity in abandoned mesic semi-natural grasslands?

Changes of agricultural practices have led to decline of semi-natural habitats sustained by traditional animal husbandry in many European regions. The abandonment of semi-natural pastures leads to increase of vascular plant biomass and subsequent decline of weak competitors such as bryophytes. Re-establishing traditional animal husbandry may potentially restore biodiversity but the success of such measures remains insufficiently known. In this study, we asked if re-establishing cattle grazing on previously abandoned grasslands will restore their bryophyte communities. The effect of cattle grazing on bryophyte communities of mesic semi-natural grasslands was studied in south-western Finland in a comparison of (i) continuously grazed pastures, (ii) previously abandoned pastures where grazing was re-established during 1990s, and (iii) abandoned pastures, where grazing had ceased during late 1960s to early 1980s. The average cover, species richness, species density and species diversity of bryophytes were significantly higher in the continuously grazed than in the abandoned grasslands. Ordination analyses revealed clear differences also in community structure between the management classes. Re-established grasslands were ecologically heterogeneous and situated in between the continuously grazed and abandoned grasslands in all characteristics, indicating variable effect of the restoration measure. Seventeen bryophyte species were recognized as significant indicators of the three grassland classes, four of which could be used as indicators of valuable grassland habitats. Although there was variation in the consequences of re-introduction of grazing, the results give evidence of positive effect of grazing on regaining bryophyte diversity of abandoned grasslands.     

Woody elements benefit bird diversity to a larger extent than semi-natural grasslands in cereal-dominated landscapes

Increasing landscape complexity can mitigate negative effects of agricultural intensification on biodiversity by offering resources complementary to those provided in arable fields. In particular, grazed semi-natural grasslands and woody elements support farmland birds, but little is known about their relative effects on bird diversity and community composition. In addition, the relative importance of local habitat versus landscape composition remains unclear. We investigated how the presence of semi-natural grasslands, the number of woody elements and the composition of the wider agricultural landscape affect bird species richness, true diversity (exponential Shannon diversity) and species composition. Bird communities were surveyed four times on 16 paired transects of 250 m each with 8 transects placed between a crop field and a semi-natural grassland and 8 transects between two crop fields with no semi-natural grasslands in the vicinity. The number of woody elements around transects was selected as an important predictor in all models, having a positive effect on species richness and true diversity, while the local presence of semi-natural grasslands was not selected in the best models. However, species richness and true diversity increased with increasing cover of ley and semi-natural grasslands, whereas species composition was modified by the coverage of winter wheat at the landscape scale. Furthermore, bird species richness, true diversity and species composition differed between sampling dates. As bird diversity benefited from woody elements, rather than from the local presence of semi-natural grasslands as such, it is important to maintain woody structures in farmland. However, the positive effect of grassland at the landscape scale highlights the importance of habitat variability at multiple scales. Because species richness and true diversity were affected by different landscape components compared to species composition, a mosaic of land-use types is needed to achieve multiple conservation goals across agricultural landscapes.     

Grassland management for meadow birds in the Netherlands is unfavourable to pollinators

Agricultural intensification and loss of semi-natural grassland have contributed to biodiversity decline, including pollinator species, in pastures around the world. To reverse the decline, agri-environmental schemes have been implemented, varying widely in effectiveness. In addition, many countries, including the Netherlands, have established nature reserves in which semi-natural grasslands are restored and are often managed for specific groups of species, e.g. meadow birds or plants. The effects of such measures on insect biodiversity are not well known but recent reports on the dramatic decline of insect biomass in nature reserves have put even more attention to the impact of land use and management on biodiversity. This study compares pollinator abundance and species richness in three common semi-natural grassland management types in the Netherlands: (1) hay meadows, (2) herb-rich grasslands and (3) meadow bird grasslands. Pollinator abundance and species richness were assessed in eleven study areas, each with all three management types present. Standardized transects, insect sampling within a standard 20 min time frame and plot-based flower surveys were used in spring and summer to assess the relationships between management regime, floral abundance and diversity and pollinator communities. The results show that meadow bird grasslands have lower pollinator abundance and diversity and a less unique pollinator assemblage than both other types. Moreover, flower abundance has a positive effect on pollinator abundance and flower diversity has a positive effect on pollinator species richness. These results indicate that meadow-bird grasslands are a comparatively unfavourable habitat for bees, hoverflies and butterflies, which may be explained by a lack of flowers as well as unsuitable mowing practices. Measures benefitting both insectivorous birds and flower-visiting insects, such as rotational mowing, could remediate this imbalance.     

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.     

Complex vegetation responses to soil disturbances in mountain grassland

We studied vegetation responses to disturbances originated by ants and voles in subalpine grasslands in the Eastern Pyrenees. We compared the effects of these small-scale disturbances with those of a large-scale disturbance caused by ploughing. We wanted to know if these soil disturbances promoted species richness through the existence of a specific guild of plants colonizing these areas, and if this guild was the same for all soil disturbances, independently of their extent. In general, grassland vegetation seemed to recover relatively quickly from soil-displacement disturbances, and the effects could be scaled up in time and space in terms of species richness and composition. Vole mound composition was similar to that in the surrounding grassland, suggesting that mounds were rapidly colonized by the neighbouring vegetation. Vegetation composition differed between the grassland and the ant mounds. Grasses and erect dicots coped well with repeated disturbance, while rosette-forming species and sedges were very sensitive to it. Landscape processes could be important to understanding recolonization. Species from xeric grasslands were found in mesic grasslands when disturbed by ploughing and on the tops of active ant mounds. Furrows in mesic grasslands recovered well, but decades after disturbance showed long persistence of some xeric species and increased species richness compared to terraces, while xeric grasslands showed decreased richness. This suggests that, because of those disturbances, within-habitat diversity was increased, although landscape diversity was not. However, specific disturbances showed idiosyncratic effects, which could enhance the species richness globally. In ant-affected areas, the grassland itself showed the highest plant species richness, partially associated to the presence of some species with elaiosomes not, or only rarely, found in adjacent grasslands without ant mounds. Therefore, soil disturbances occurring at different spatial scales contributed to complexity in vegetation patterns in addition to abiotic factors and grazing. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Nomenclature of the species follows Tutin et al. (1964–1980) and Bolòs et al. (1993).