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.     

The influence of agricultural land-use intensity on bryophyte species richness

This study is a quantitative approach to the estimation of bryophyte species richness in relation to land-use intensity at three spatial scales in highly cultivated areas. A total of 460 randomly selected habitats and their various substrates within 29 study sites were investigated with regard to their land-use intensity and their bryophyte species richness in an agricultural region of eastern Austria. On bare soils (substrate-scale), low but regular disturbance increases bryophyte diversity in comparison to lower land-use intensity. However, more frequent disturbance (e.g. ploughing more than two times a year) dramatically reduces species richness at these sites, with more than 50% of these sites showing no bryophytes. The production of reproductive units (sporophytes and vegetative units) is highest at an intermediate disturbance regime. On the habitat, as well as on the landscape-scale, there is a significant increase in total bryophyte species number as well as in the number of threatened species with decreasing land-use intensity. This is mainly due to habitat and structural diversity, which increases with decreasing land-use intensity. There are significant correlations between landuse intensity, structural diversity and species richness at the habitat as well as on the landscape scale.     

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).     

Frequent fire promotes diversity and cover of biological soil crusts in a derived temperate grassland

The intermediate disturbance hypothesis (IDH) predicts that species diversity is maximized at moderate disturbance levels. This model is often applied to grassy ecosystems, where disturbance can be important for maintaining vascular plant composition and diversity. However, effects of disturbance type and frequency on cover and diversity of non-vascular plants comprising biological soil crusts are poorly known, despite their potentially important role in ecosystem function. We established replicated disturbance regimes of different type (fire vs. mowing) and frequency (2, 4, 8 yearly and unburnt) in a high-quality, representative Themeda australis–Poa sieberiana derived grassland in south-eastern Australia. Effects on soil crust bryophytes and lichens (hereafter cryptogams) were measured after 12 years. Consistent with expectations under IDH, cryptogam richness and abundance declined under no disturbance, likely due to competitive exclusion by vascular plants as well as high soil turnover by soil invertebrates beneath thick grass. Disturbance type was also significant, with burning enhancing richness and abundance more than mowing. Contrary to expectations, however, cryptogam richness increased most dramatically under our most frequent and recent (2 year) burning regime, even when changes in abundance were accounted for by rarefaction analysis. Thus, from the perspective of cryptogams, 2-year burning was not an adequately severe disturbance regime to reduce diversity, highlighting the difficulty associated with expression of disturbance gradients in the application of IDH. Indeed, significant correlations with grassland structure suggest that cryptogam abundance and diversity in this relatively mesic (600 mm annual rainfall) grassland is maximised by frequent fires that reduce vegetation and litter cover, providing light, open areas and stable soil surfaces for colonisation. This contrasts with detrimental effects of 2-year burning on native perennial grasses, indicating that this proliferation of cryptogams has potentially high functional significance for situations where vegetation cover is depleted, particularly for reducing soil erosion.     

小叶锦鸡儿灌丛化对典型草原群落结构与生态系统功能的影响

草原灌丛化是全球干旱半干旱地区面临的重要生态问题。灌丛化对草原生态系统结构与功能的影响较为复杂, 有待于在更广泛的区域开展研究。该研究在内蒙古锡林郭勒典型草原选择轻度、中度和重度灌丛化草地, 通过群落调查, 结合植物功能性状和土壤理化性质观测, 研究了小叶锦鸡儿(Caragana microphylla)灌丛化对草原群落结构(物种多样性、功能多样性和功能群组成)和生态系统功能(初级生产力、植被和土壤养分库)的影响。结果表明: 1)不同程度灌丛化草地的物种丰富度、功能性状多样性和群落加权性状平均值差异显著, 其中, 中度灌丛化草地的物种多样性和功能多样性较高, 表明一定程度的灌丛化有利于生物多样性维持。2)重度灌丛化草地的地上净初级生产力(ANPP)显著高于轻度和中度灌丛化草地, 其原因主要是随着灌丛化程度加剧, 群落内一/二年生草本植物显著增加, 而多年生禾草和多年生杂类草显著减少。三个灌丛化草地的植被叶片和土壤碳、氮库差异均不显著。3)灌丛化对草原生态系统功能包括ANPP、植被和土壤养分库均没有直接的影响, 而是通过影响功能群组成、土壤理化性质和功能多样性, 间接地影响生态系统功能; 灌丛化导致功能群发生替代和土壤旱碱化是最重要的生物和非生物因素。     

Disturbance persistence in managed grasslands: shifts in aboveground community structure and the weed seed bank

The length of time and form in which disturbances persist in systems depends on the intensity and frequency of disturbance and on the abilities of resident species to recover from such events. In grazed grasslands, trampling by large mammalian herbivores can periodically facilitate weed establishment by exposing patches of bare ground but whether an intense soil disturbance event results in a temporary increase in weed abundance or a persistent weed problem remains unclear. In May 2002, cattle trampling following heavy rain caused severe damage to nine-month old, rotationally grazed, cool-season pastures (Midwest USA). In September 2002, we compared the aboveground composition of paddocks (i.e., fenced pasture sections) that were heavily disturbed to those that received no damage. Relative to undisturbed paddocks, forage species relative cover was 17% lower in disturbed paddocks, and weed species and bare ground relative cover was 61% and 100% higher, respectively. By September 2004, paddock types did not differ in all aboveground community components. However, the abundance and species richness of weed seeds in the soil seed bank averaged respectively 82% and 30% higher in disturbed paddocks between 2003 and 2004. These findings indicate that a spatially extensive, intense soil disturbance event may soon become undetectable in components of aboveground pasture structure but can persist as an augmented weed seed bank. Because of high weed seed bank longevity, disturbances to formerly disturbed pastures would likely result in higher weed recruitment, with more species represented, than in those which lack previous disturbance. Disturbance history may thus be a useful predictor of weed community composition following subsequent disturbance. Based on empirical data supporting this proposition, we recommend that grassland managers explicitly incorporate disturbance history into dynamic management planning and do not rely exclusively on aboveground characters to evaluate the invasion status or colonization potential of an area by undesirable plants. We emphasize that the ecological legacies of past soil disturbance events cannot only influence the contemporary patterns and processes of grasslands, but importantly, affect their compositional trajectories following subsequent perturbation.     

Species richness and species turnover in a successional heathland

Abstract. Changes in species richness and species turnover during secondary succession following experimental disturbance were studied in eight permanent plots in a species‐rich dry heathland in the southern part of the Czech Republic. The treatments applied were sod‐cutting, burning, cutting of above‐ground biomass, and control. The plots were sampled annually between 1992 and 2000; species richness was analysed at three scales, 25 cm × 25 cm, 1 m × 1 m, and 3 m × 3 m. Disturbances resulted in increased species richness. The highest vascular plant richness was attained during the secondary succession after sod‐cutting, where species established on exposed bare ground. Less severe disturbances by burning and cutting also caused a slight increase in the number of vascular plant species. For bryophytes and lichens, the highest increase in the number of species was also found in the sod‐cut plots, where all cryptogams were removed by the disturbance. At the scale of 3 m × 3 m, species richness of both vascular plants and cryptogams peaked in 1995–1996, i.e. 3–4 yrs after the disturbance, and slowly decreased or slightly fluctuated without any trend thereafter. At smaller scales it either peaked later or constantly increased over the entire observation period of 9 yrs. Species mobility, expressed as species accumulation over time, was lower than reported from grasslands. Rates of species turnover, calculated as Jaccard dissimilarity between pairs of consecutive years, corresponded across different scales. This implies that successional dry heathlands have a higher small‐scale mobility than heathlands which are apparently stable at larger scale.     

Diversity and floristic patterns of mediterranean grasslands: the relative influence of environmental and land management factors

Managed grasslands are normally ecosystems with high species diversity. They are associated with traditional extensive livestock grazing. Land-use changes, in particular gradual abandonment or grazing intensification, lead to major changes in floristic patterns of these grasslands, some included in priority habitats of the EU Habitats Directive. In order to analyse these patterns of change, Mediterranean grasslands of eastern areas of mainland Portugal were studied aiming to: (1) establish ecological gradients underlying their floristic patterns; (2) examine the relative importance of both environmental and land-use factors on their floristic composition; (3) assess how floristic composition and species richness are affected by land-use factors. Vegetation sampling was carried out from 2008 to 2009 following phytosociological procedures. Canonical Correspondence Analysis was applied. Variation partitioning was used to assess the relative influence of land-use variables. Richness, legumes cover, endemic species and bryophytes cover were compared in four land-management regimes: unmanaged; extensive grazing by sheep; extensive grazing by cattle and sheep; frequent soil tillage. Significant differences were found in floristic and diversity patterns, revealing the importance of grazing as a management tool for maintaining or improving floristic diversity. From our findings slight extensive grazing can contribute to both maintenance of species diversity and to the increase in legumes cover and thus to palatability and economic value of studied grasslands. Interactions between herbivores and grasslands, considering also their influences on soil properties, have an important role in the equilibrium states which promote biodiversity and conservation of the services of these ecosystems.     

Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences

In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species‐richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co‐existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation. In contrast to the diversifying effects of local disturbances, productivity‐related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro‐evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats. During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co‐existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.     

A long-term experimental test of the dynamic equilibrium model of species diversity

The dynamic equilibrium model of species diversity predicts that ecosystem productivity interacts with disturbance to determine how many species coexist. However, a robust test of this model requires manipulations of productivity and disturbance over a sufficient timescale to allow competitive exclusion, and such long-term experimental tests of this hypothesis are rare. Here we use long-term (27 years), large-scale (8 × 50-m plots), factorial manipulations of soil resource availability and sheep grazing intensity (disturbance) in grasslands to test the dynamic equilibrium model. As predicted by the model, increased productivity not only reduced plant species richness, but also moderated the effects of grazing intensity, shifting them from negative to neutral with increasing productivity. Reductions in species richness with productivity were associated with dominance by faster growing (i.e. high specific leaf area) and taller plants. Conversely, grazing favoured shorter plants and this effect became stronger with greater productivity, consistent with the view that grazing can lead to weaker asymmetric competition for light. Our study shows that the dynamic equilibrium model can help to explain changes in plant species richness following long-term increases in soil resource availability and grazing pressure, two fundamental drivers of change in grasslands worldwide.     

Impact of different‐sized herbivores on recruitment opportunities for subordinate herbs in grasslands

Abstract. Potential effects of herbivores on plant species diversity depend on herbivore size, species and density. In this study we examine the effect of different‐sized herbivores (cattle and rabbits) on recruitment of subordinate herbs in grasslands. We show that in a grazed floodplain, grassland plant species richness is mainly determined by the presence of many species of subordinate herbs. These herbs experience high colonization and extinction rates. We conclude that the creation of colonization opportunities for subordinate herbs plays a crucial role in maintaining plant species richness in productive grasslands. We found that cattle disperse large amounts of seeds via their dung, over ten times more than rabbits. Rabbits create more and on average larger bare soil patches than cattle. In a field experiment artificial disturbances improved germination success tremendously for four tested herb species. We found that bare soil is the best regeneration site, while cattle dung gave a too strong nutrient stimulus, resulting in tall vegetation and therefore light limitation. These results can be confirmed with results from field monitoring plots where plant species richness was positively related to the occurrence of bare soil patches. Therefore both large and small herbivores have a major impact on dispersal and colonization, but for different reasons. Cattle are identified as most important for seed dispersal whereas rabbits have a main effect as creators of disturbances. These results emphasize the importance of distinguishing between herbivore species in assessing their (potential) effects.     

Disturbance–diversity models: what do they really predict and how are they tested?

The intermediate disturbance hypothesis (IDH) and the dynamic equilibrium model (DEM) are influential theories in ecology. The IDH predicts large species numbers at intermediate levels of disturbance and the DEM predicts that the effect of disturbance depends on the level of productivity. However, various indices of diversity are considered more commonly than the predicted number of species in tests of the hypotheses. This issue reaches beyond the scientific community as the predictions of the IDH and the DEM are used in the management of national parks and reserves. In order to compare responses with disturbance among measures of biodiversity, we used two different approaches of mathematical modelling and conducted an extensive meta-analysis. Two-thirds of the surveyed studies present different results for different diversity measures. Accordingly, the meta-analysis showed a narrow range of negative quadratic regression components for richness, but not evenness. Also, the two models support the IDH and the DEM, respectively, when biodiversity is measured as species richness, but predict evenness to increase with increasing disturbance, for all levels of productivity. Consequently, studies that use compound indices of diversity should present logical arguments, a priori, to why a specific index of diversity should peak in response to disturbance.     

Long‐term development of species richness in a central European beech (Fagus sylvatica) forest affected by windthrow—Support for the intermediate disturbance hypothesis?

On the basis of long‐term surveys of permanent plots and traps, we examined the communities of saproxylic beetles, fungi, herbs, and trees on an untreated 22 ha large beech forest windthrow and asked whether the results lend support to the intermediate disturbance hypothesis (IDH). We studied species richness and the similarity of community composition. Additionally, we grouped species by their frequency trend over time to successional model types to examine whether, corresponding to the IDH, the diversity of these groups explained peak richness at intermediate intervals after the disturbance. In line with the IDH, species richness showed a hump‐backed temporal course for alpha and gamma diversity. We found evidence for a linear succession directly after the disturbance. This, however, did not continue, and in all species groups, a partial recovery of the initial community was observed. In the case of fungi, herbs, and trees, but not for saproxylic beetles, alpha diversity was driven by the diversity of the successional model types. Our results underline that the mechanisms driving species richness after disturbances are more complex than the IDH suggests and that these mechanisms vary with species group. We assumed that, besides competition, legacy effects, facilitation, habitat heterogeneity, and random saturation of the species pool are important. In case of trees and herbs, we found indications for strong legacy and competition effects. For fungi and beetles, substrate heterogeneity and microclimate were assumed to be important. We concluded that disturbances contribute to increasing species richness not only by reducing the effectiveness of competitors but also by increasing the amount and diversity of resources, as well as their rate of change over time.     

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.     

武汉市城区苔藓植物多样性和分布及与环境因子的关系

对武汉市7个中心城区和部分远郊城区共26个样点73个样地的苔藓群落进行调查,采集苔藓植物样本共431份。经鉴定调查区内共有苔藓植物20科35属91种,其中狭叶小羽藓（Haplocladium angustifolium（Hampe et C.Muell.）Broth.）和钝叶绢藓（Entodon obtusatus Broth.）等为优势种,分布于其中的24个样点。多样性指数分析结果显示,位于青龙山国家森林公园的α多样性Patrick和Shannon-Wiener指数最高,说明该样点苔藓植物群落的复杂程度高,群落所含信息量大。Patrick指数与环境因子的Pearson相关性分析结果显示,土壤含水率和与主要干道距离是显著影响地面生苔藓植物多样性的环境因子,而显著影响树附生苔藓植物多样性的环境因子仅与距主要干道距离相关。用典范对应分析法（CCA）研究26个样点中苔藓植物的盖度与主要环境因子的关系,结果发现人为干扰程度、草本盖度、与城市主要干道的距离等对地面生和树附生苔藓植物的分布都有显著影响。其中地面生苔藓植物的分布受人为干扰和草本盖度影响最为显著;而树附生苔藓植物的分布受附生树干胸径及与主要干道距离影响最为显著。研究结果表明武汉市区苔藓植物的多样性和分布受人为干扰较大,树附生苔藓对汽车尾气等因素更为敏感。     

The effects of windthrow on plant species richness in a Central European beech forest

The effects of soil disturbance caused by the uprooting of a single or a few canopy trees on species richness and composition of vascular plant species and bryophytes were examined in a temperate beech forest (Fagus sylvatica) in northern Germany. We recorded the vegetation in 57 pairs of disturbed and adjacent undisturbed plots and established a chronosequence of mound ages to study the effect of time since microsite formation on plant species richness and composition. We found significant differences in plant species richness and composition between disturbed and adjacent undisturbed plots. Species richness of both vascular plants and bryophytes was higher in the disturbed than in the undisturbed plots, but these differences were more pronounced for bryophytes. We suggest that three main factors are responsible for this differential response. The availability of microsites on the forest floor that are suitable for the recruitment of bryophytes is lower than for vascular plants. Establishment of bryophytes in disturbed microsites is favoured by a greater abundance of propagules in the close vicinity and in the soil of the disturbed microsites, as well as by a greater variety of regeneration strategies in bryophytes than in vascular plants. Time since mound formation was a major factor determining plant species richness and composition. A significant decrease in the mean number of species was found from young mounds to intermediate and old mounds. However, differences were observed between vascular plants and bryophytes in the course of changes through time in species richness and composition. A large number of exclusive and infrequent vascular plant species was observed on young mounds, among them several disturbance specialists. We suggest that the establishment of many vascular plant species was infrequent and short-lived due to unfavourable light conditions and a low abundance of propagules. By contrast, the development of a litter layer was the main reason for the decreased mean number of bryophytes on old mounds. Our study supports the view that groups of species differing in important life history traits exhibit different responses to soil disturbance.     

Evaluating the role of fire disturbance in structuring small reptile communities in temperate forests

Fire is an integral disturbance shaping forest community dynamics over large scales. However, understanding the relationship between fire induced habitat disturbance and biodiversity remain equivocal. Ecological theories including the intermediate disturbance hypothesis (IDH) and the habitat accommodation model (HAM) offer predictive frameworks that could explain faunal responses to fire disturbances. We used an 80 year post-fire chronosequence to investigate small reptile community responses to fires in temperate forests across 74 sites. First, we evaluated if changes in species richness, abundance and evenness post-fire followed trends of prior predictions, including the IDH. Second, using competing models of fine scale habitat elements we evaluated the specific ways which fire influenced small reptiles. Third, we evaluated support for the HAM by examining compositional changes of reptile community post-fire. Relative abundance was positively correlated to age post-fire while richness and evenness showed no associations. The abundance trend was as expected based on the prior prediction of sustained population increase post-disturbance, but the trend for richness contradicted the prediction of highest diversity at intermediate levels of disturbance (according to IDH). Abundance changes were driven mainly by changes in overstorey, ground layer, and shelter, while richness and evenness did not associate with any vegetation parameter. Community composition was not strongly correlated to age since fire, thus support for the HAM was weak. Overall, in this ecosystem, frequent fire disturbances can be detrimental to small reptiles. Future studies utilizing approaches based on species traits could enhance our understanding of biodiversity patterns post-disturbance.     

Management impacts on vegetation dynamics of hyper‐eutrophicated fields at Berlin,Germany

Abstract. This paper reports on vegetation dynamics on terrestrial, temperate grassland sites at the upper range of the productivity scale, i.e. on abandoned sewage fields (fields once used for waste water disposal) at Berlin‐Blankenfelde, Germany. I studied regeneration and the influence of different management practices (removal of top soil and mowing in late summer). Changes in species composition and cover were followed on permanent plots of 2m × 2m size through five years of vegetation development. At the outset of the experiment the abandoned fields were dominated by dense Urtica dioica /Elymus repens stands. Species richness was 7 species/ 4m², and it remained low on unmanaged plots during the time of observation (7.6 species/plot in year 5). Removal of 20 cm of top soil caused a severe decline of Urtica and a large increase in species richness (21 species in year 1 after disturbance). Mowing was slightly higher compared with unmown plots on both initially excavated and unexcavated plots. Total cover was always near 100 % (except immediately after top soil removal). Colonization of bare soil was very rapid and in late summer of the first year after disturbance cover already increased towards 100%. On all plots the vegetation was mostly dominated by perennial herbs and grasses. Winter season gaps are occupied by Galium aparine, a large‐seeded annual scrambling climber. Monocarpic perennials behaved as winter annuals in most cases. Woody species were inhibited by dense above‐ground biomass and litter cover. The paper questions whether succession on abandoned sewage fields may proceed towards a woodland stage and advises how vegetation of such hyper‐eutrophicated sites can be managed towards higher diversity.     

Short-term invasibility patterns in burnt and unburnt experimental Mediterranean grassland communities of varying diversities

This paper reports the findings of a short-term natural invasibility field study in constructed Mediterranean herbaceous communities of varying diversities, under a fire treatment. Three components of invasibility, i.e. species richness, density and biomass of invaders, have been monitored in burnt and unburnt experimental plots with resident diversity ranging from monocultures to 18-species mixtures. In general, species richness, density and biomass of invaders decreased significantly with the increase of resident species richness. Furthermore, the density and biomass of invading species were significantly influenced by the species composition of resident communities. Although aboveground biomass, leaf area index, canopy height and percent bare ground of the resident communities explained a significant part of the variation in the success of invading species, these covariates did not fully explain the effects of resident species richness. Fire mainly influenced invasibility via soil nutrient levels. The effect of fire on observed invasibility patterns seems to be less important than the effects of resident species richness. Our results demonstrate the importance of species richness and composition in controlling the initial stages of plant invasions in Mediterranean grasslands but that there was a lack of interaction with the effects of fire disturbance.     

Community phylogenetic diversity and abiotic site characteristics influence abundance of the invasive plant Rhamnus cathartica L.

Aims Theory predicts that the success of introduced species is related to the diversity of native species through trait-based processes. Abiotic site characteristics may also affect a site's susceptibility to invasion. We quantified resident plant species richness, phylogenetic diversity and several abiotic site characteristics for 24 oak forests in Minnesota, USA, to assess their impact on the abundance of a widespread, introduced terrestrial plant species, common buckthorn (Rhamnus cathartica L.). Specifically, we asked (1) whether resident species richness and phylogenetic diversity affected the abundance of R. cathartica and (2) what site characteristics explained the overall abundance of R. cathartica .Methods Our survey included 24 oak-dominated stands in Minnesota's deciduous forests. In each stand, we identified all species in 16 plots. We also measured a series of environmental site characteristics, including canopy openness (a proxy for light availability), percent bare soil, soil pH, percent sand, an index of propagule availability, duff layer thickness (a proxy for earthworm activity), an index of insolation and slope. For all species present in at least one site, we estimated a community phylogeny. We combined all site-level characteristics, including phylogenetic diversity of the resident plant species, in a multiple regression model to examine site level drivers of community invasibility.Important findings Results indicate that sites with higher overall plant phylogenetic diversity harbor less R. cathartica, even though native species richness was not significantly related to R. cathartica abundance. Regression analyses indicated that, in addition to resident species phylogenetic diversity, the most important predictors of R. cathartica abundance were canopy openness and the amount of bare soil, both positively related to the abundance of the invader. By combining the effects of abiotic site characteristics and resident species phylogenetic diversity in a model that predicted the abundance of R. cathartica, we were able to simultaneously account for a wide range of factors that might influence invasibility. Overall, our results suggest that management strategies aimed at reducing disturbances that lead to increased bare soil and light levels may be more successful if they also maximize phylogenetic diversity of the resident plant community.