Heathland dynamics in biotically disturbed areas: on the role of some features enhancing heath success

It is well accepted that intense anthropogenic activities threaten European heathland communities. The negative impact of high intensities of livestock grazing on heath survival has been demonstrated in many studies, however, it does not seem a straightforward consequence in some cases: in the southern Pyrenees, the existence of large heathland patches in intensively grazed areas suggests that other factors interact and affect heath success. In order to gain insight into the factors affecting heath success, this paper analyses the pattern of occurrence of ericaceous shrubs—particularly Erica vagans—and their effects on soils when compared to herbaceous vegetation, under a complex biotic disturbance regime. Survey plots were established in grassland and heathland communities disturbed by both livestock and by burrowing animals on soils with different levels of basic resources. Relative cover of the species on and off molehills was studied. In heathland plots, ericaceous shrubs were more common on old molehills than off them. This pattern was strongest in the poorest soils. Below established heath shrubs, independently of the initial properties of the soils, some basic resources were more available than in intershrub areas. In grazed ecosystems, where continuous trampling and removal of biomass occurs, the heath canopy provides a site for accumulation of litter and insects, which contribute to the build up of a resource-rich layer. We conclude that successful establishment on ubiquitous molehills and suitable, localised changes in the soil under shrubs are some of the factors that explain the success of ericaceous shrubs in complex biotically disturbed areas.     

Field evidence of plastic growth responses to habitat heterogeneity in the clonal herbRanunculus repens

Stolon internode lengths were measured on plants of the clonal herbRanunculus repens growing in a hay meadow which was subject to disturbance by mole (Talpa europaea) activity. Within the site three habitat types were recognized: closed grassland, the open ground of fresh molehills and the grass-molehill boundary. The lengths of stolon internodes ofR. repens differed significantly in each of the three habitats. The shortest internodes occurred on stolons on the open molchills. The longest occurred in the closed grassland habitat. The type of habitat in which parent ramets were rooted did not significantly influence the length of internodes on their daughter stolons. The length of a stolon internode was determined by its immediate surrounding habitat type. Consecutive internode lengths on a given stolon showed considerable plasticity, shortening significantly as stolons spread onto molehills from surrounding habitats, and increasing significantly as stolons advanced from a molehill into the surrounding closed grassland habitat. These results are consistent with the proposition that under favorable conditions (on the molehills, where resources are expected to be more abundant, and competition absent) internode lengths shorten and the plant forages intensively, whereas under conditions of low resource availability (in the closed grassland, where competition occurs) internode lengths increase, allowingR. repens to forage extensively. Such morphological plasticity may promote more efficient exploitation of resource-rich sites and more rapid vacation of resource-poor sites.     

Changing importance of environmental factors driving secondary succession on molehills

Question: Species composition during secondary succession is influenced by a number of factors, such as soil moisture, disturbance timing and surrounding vegetation. How does the importance of these factors change over the course of succession? Methods: We set up a full‐factorial block design using molehills differing in (a) disturbance timing, (b) soil moisture and (c) composition of surrounding vegetation, and recorded the cover of all species present on the molehills over 3 years. M1ultiple regression analyses on the dissimilarity matrices of community composition and of environmental factors were applied for each of five age classes of molehills to estimate the effect of the single factors at different stages of succession. Results: The timing of disturbance did not significantly affect community composition at any stage of succession. In contrast, the effects of soil moisture and surrounding vegetation changed significantly over time, with moisture being more important at earlier stages of succession and surrounding vegetation at later stages. Conclusion: The importance of environmental factors for species composition change significantly over the course of secondary succession. Instead of aggregating the effects of environmental factors over time, future studies should consider underlying dynamics of recolonization more comprehensively.     

Multi‐nutrient vs. nitrogen‐only effects on carbon sequestration in grassland soils

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi‐nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human‐managed soils. Here, we use a 19‐year old nutrient addition experiment on a permanent grassland to test for nutrient‐induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have ‘constrained’ soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N‐only fertilized soils store, on average, 11 t C ha^?1 more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi‐nutrient additions increase plant productivity, soil C sequestration is increased by N‐only additions. The positive N‐only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N‐only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long‐term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).     

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.     

Is the relation between nutrient supply and biodiversity co‐determined by the type of nutrient limitation?

Correlative studies have shown a ‘hump‐backed’ relation between the vegetation N:P ratio and plant species diversity with the highest diversity at balanced N:P ratios (between 10 and 14). We tested the hypothesis that adding growth‐limiting nutrients to mesotrophic grasslands that were in shortage of either N (N:P ratio<10) or P (N:P ratio>14) would lead to an increase of plant diversity. Thereto, we studied the effects of long‐term (11 yr) experimentally increased N and/or P supply on soil nutrient pools, vegetation nutrient dynamics and biodiversity in a riverine grassland in the Netherlands with a low soil N:P ratio (N shortage) and a peat grassland with a high soil N:P ratio (P shortage), respectively. Eleven years of nutrient addition hardly had any effects on the total stocks of C, N and P in the soils of both sites, due to the large size of the soil nutrient pools already present and to the management at both sites (annual hay‐making and ‐removal). However, in the riverine grassland the treatments increased the cycling of the small pool of labile N and P compounds resulting in large increases in annual fluxes of especially N. In the unfertilised controls, species establishments balanced more or less species losses during an 11 year period, thus leading to a dynamic equilibrium of the species pool. However, contrary to our hypothesis, addition of the growth‐limiting nutrient led at both sites to a reduction of species diversity even when total biomass remained below critical levels. Species diversity and species evenness were strongly determined by N mineralisation and to a lesser extent by total soil N and extractable P, respectively. Total aboveground biomass of the vegetation was determined by total soil N. Our study shows that patterns found in correlative studies of the relation between plant diversity and soil and vegetation N:P ratio can not be translated into successful experimental manipulations to enhance biodiversity. The most likely explanation is that colonization limitation occurred in the fertilized plots and that not sufficient diaspores of potentially new species could reach and/or colonize the plots to compensate for the species extinctions as a result of increased nutrient supply.     

Environmentally‐contingent behaviour of invasive plants as drivers or passengers

Invasive plants are often regarded as drivers that actively reduce diversity and alter ecosystem processes such as succession. Alternatively, invaders may be passengers that simply colonize openings produced by anthropogenic disturbance and are present only temporarily. Here we test whether the behaviour of invasive species as drivers or passengers is contingent on disturbance and nutrient availability. We created twelve experimental environments (three levels of annual disturbance × four levels of nitrogen availability) for 18 years in a grassland at the northern edge of the North American Great Plains. Out of 19 invasive species initially present, two perennials (Bromus inermis, a grass, and Cirsium arvense, a forb) acted as drivers, maintaining or increasing dominance, maintaining low species richness, and forming an invader‐ dominated successional sequence. Behaviour as drivers was environmentally contingent: Bromus behaved as a driver only in less disturbed environments, and the tendency of Cirsium to behave as a driver increased significantly with both disturbance and nitrogen availability. Most invasive species (90%), however, consistently behaved as passengers, disappearing or becoming rare. The importance of disturbance and fertility for starting invasions is well‐known, but our study shows that these factors also contribute to the behaviour of some invaders as drivers. The emergence of drivers and invader‐dominated successional sequences suggests that, as rates of invasion, disturbance and eutrophication continue to increase with human activity, invasive species that act as drivers may form low‐diversity communities that persist for decades.     

Soil disturbance favours threatened beetle species in sandy grasslands

Soil disturbance is recognised as an important restoration measure for conserving biodiversity in sandy soils. We used a soil disturbance (ploughing) experiment in a sandy grassland as well as a semi-natural disturbance (slope erosion enhanced by cattle trampling) gradient on a sandy slope to test the soil disturbance effects on the ground-living beetle community. Both experimental disturbance and semi-natural disturbance favoured sandy grassland specialists, but there was no overall effect on beetle richness and abundance. Amara lucida and Harpalus spp. were favoured by disturbance while Calathus melanocephalus was disfavoured. Experimental ploughing significantly increased the proportion of red-listed species in disturbed plots compared to non-disturbed controls. In the semi-natural disturbance gradient we found that the beetle community on the disturbed slope differed from that of the flat areas, and there were tendencies for a higher proportion of red-listed species on the slope. We conclude that increasing the area of bare sand in sandy grasslands can have positive effects on many threatened species. Soil disturbance should thus be included as a regular measure in sandy grasslands under conservation management and as a measure to restore high biodiversity in areas where bare sand is rare.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Dynamics of grazing lawn formation: an experimental test of the role of scale‐dependent processes

Grazing lawns are characteristic for African savanna grasslands, standing out as intensely grazed patches of stoloniferous grazing‐tolerant grass species. Grazing lawn development has been associated with grazing and increased nutrient input by large migratory herds. However, we argue that in systems without mass migrations disturbances, other than direct grazing, drive lawn development. Such disturbances, e.g. termite activity or megaherbivore middens, also increase nutrient input and keep the bunch vegetation down for a prolonged time period. However, field observations show that not all such disturbances lead to grazing lawns. We hypothesize that the initial disturbance has to be of a minimal threshold spatial scale, for grazing intensity to be high enough to induce lawn formation. We experimentally tested this idea in natural tall savanna grassland. We mowed different‐sized plots to simulate initial disturbances of different scales (six times during one year) and applied fertilizer to half of the plots during two years to simulate increased nutrient input by herbivores or termite activity. Allowing grazing by naturally occurring herbivores, we followed the vegetation development over more than three years. Grazing kept bunch grass short in coarser, fertilized plots, while grasses grew out toward their initial height in fine‐scale and unfertilized plots. Moreover, lawn grasses strongly increased in cover in plots with an increased nutrient input but only after coarser scale disturbance. These results support our hypothesis that an increased nutrient input in combination with grazing indeed induces grazing lawn formation, but only above a threshold scale of the initial disturbance. Our results provide an alternative mechanism for the development of grazing lawns in systems that lack mass migrating herds. Moreover, it gives a new spatial dimension to the processes behind grazing lawn development, and hence help to understand how herbivores might create and maintain spatial heterogeneity in grassland systems.     

Effects of Nutrient Manipulations and Grass Removal on Cover,Species Composition,and Invasibility of a Novel Grassland in Colorado

Cover and richness of a 5‐year revegetation effort were studied with ,respect to small‐scale disturbance and nutrient manipulations. The site, originally a relict tallgrass prairie mined for gravel, was replanted to native grasses using a seed mixture of tall‐, mixed‐, and short‐grass species. Following one wet and three relatively dry years, a community emerged, dominated by species common in saline soils not found along the Colorado Front Range. A single species, Alkali sacaton (Sporobolus airoides), composed nearly 50% of relative vegetation cover in control plots exhibiting a negative relationship between cover and richness. Seeded species composed approximately 92% of vegetation cover. The remaining 8% was composed of weeds from nearby areas, seed bank survivors, or mix contaminants. Three years of soil nutrient amendments, which lowered plant‐available nitrogen and phosphorus, significantly increased relative cover of seeded species to 97.5%. Fertilizer additions of phosphate enhanced abundance of introduced annual grasses (Bromus spp.) but did not significantly alter cover in control plots. Unmanipulated 4‐m² plots contained an average of 4.7 planted species and 3.9 nonplanted species during the 5‐year period, whereas plots that received grass herbicide averaged 5.4 nonplanted species. Species richness ranged from an average 6.9 species in low‐nutrient, undisturbed plots to 10.9 species in the relatively high‐nutrient, disturbed plots. The use of stockpiled soils, applied sparingly, in conjunction with a native seed mix containing species uncommon to the preexisting community generated a species‐depauperate, novel plant community that appears resistant to invasion by ruderal species.     

Restoration of Species‐Rich,Nutrient‐Limited Mountain Grassland by Mowing and Fertilization

Mowing and management to reduce nutrient levels have often been successfully used to restore species‐rich grasslands in various parts of Europe. However, such treatments have failed to restore the species‐rich Central European mountain grasslands dominated by Polygonum bistorta. P. bistorta builds an extensive underground rhizome system that monopolizes available nutrients in these nutrient‐poor grasslands, enabling this species to persist at high densities even in the presence of mowing. Therefore, we tested a restoration approach using a factorial combination of fertilization and mowing, as well as a litter removal treatment. The experiment was run over 5 years and species composition response to these treatments was recorded at two spatial scales. Mowing suppressed flowering and cover of P. bistorta and promoted target grassland species and richness. Fertilization prevented nutrient impoverishment and increased height and dominance of the broad‐leaved grasses with which many species‐rich grassland herbs could coexist. The additive effect of the mowing/fertilization treatments was strong enough to act as a driver of P. bistorta suppression and associated community change. The litter removal treatment, however, had little effect on plant composition. The experiment demonstrates that in nutrient‐limited grasslands, increasing nutrient levels in addition to mowing to manage competition for light can be used to control dominants. This contrasts with restoration of systems where after abandonment increased nutrient levels lead to the establishment of tall dominants that suppress other species by competition for light.     

Ecological response hides behind the species abundance distribution: Community response to low‐intensity disturbance in managed grasslands

Land‐use and management are disturbance factors that have diverse effects on community composition and structure. In traditional rural grasslands, such as meadows and pastures, low‐intensity management is maintained to enhance biodiversity. Maintenance of road verges, in turn, creates habitat, which may complement traditional rural grasslands. To evaluate the effect of low‐intensity disturbance on insect communities, we characterized species abundance distributions (SAD) for Carabidae, Formicidae, and Heteroptera in three grassland types, which differed in management: meadows, pastures, and road verges. The shape of SAD was estimated with three parameters: abundance decay rate, dominance, and rarity. We compared the SAD shape among the grassland types and tested the effect of environmental heterogeneity (plant species richness) and disturbance intensity (trampling in pastures) on SADs. The shape of SADs did not differ among the grassland types but among the taxonomic groups instead. Abundance decay rate and dominance were larger for Formicidae, and rarity smaller, than for Carabidae and Heteroptera. For Carabidae and window‐trapped Heteroptera, rarity increased with increasing plant species richness. For Formicidae, dominance increased with trampling intensity in pastures. Although the SAD shape remained largely unchanged, the identity of the dominant species tended to vary within and among grassland types. Our study shows that for a given taxonomic group, the SAD shape is similar across habitat types with low‐intensity disturbances resulting from different management. This suggests that SADs respond primarily to the intensity of disturbance and thus could be best used in monitoring communities across strong disturbance and environmental gradients. Because taxonomic groups can inherently have different SADs, taxon‐specific SADs for undisturbed communities must be empirically documented before the SAD shape can be used as an indicator of environmental change. Because the identity of the dominant species changes from management type to another, the SAD shape alone is not an adequate monitoring tool.     

Nutrient limitations in wet, drained and rewetted fen meadows: evaluation of methods and results

Restoration of wet grassland communities on peat soils involves management of nutrient supply and hydrology. The concept of nutrient limitation was discussed as well as its interaction with drainage and rewetting of severely drained peat soils. Different methods of assessing nutrient limitation were compared and the type and extent of nutrient limitation were determined for several wet grassland communities. It was concluded that a full-factorial field fertilisation experiment is the most preferable method. Plant tissue analyses and soil chemical analyses were considered less suitable, although they may provide helpful additional information. Fertilisation experiments in the laboratory using sods or using test plants appear to be the proper means to study mechanisms or processes, but have a restricted predictive value for field situations. Generalising the results, it seems that many relativily undisturbed grassland plant communities on peaty soils are characterised by N limitation. Phosphate limitation for vegetation on peat soils is mainly observed in specific circumstances such as extreme calcium richness, high concentrations of Fe or as a result of drainage or long-term hay cropping. The latter two may also cause K limitation. Rewetting is regarded as a prerequisite in restoring wet grassland communities. Further restoration measures to influence nutrient availability depend on aims of the management and the individual site conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Species–soil associations, disturbance, and nutrient cycling in an Australian tropical rainforest

Resource availability and disturbance are important factors that shape the composition, structure, and functioning of ecosystems. We investigated the effects of soil fertility and disturbance on plant–soil interactions and nutrient cycling in a diverse tropical rainforest. Our goal was to determine how common soil specialisation is among species and how plant–soil interactions affect ecosystem functioning in the presence of disturbance. Most species (59%) showed significant fidelity to either fertile (basalt) or infertile (schist) soils. Obligate schist specialists (six species) contributed 39 and 37% to total stand-level basal area and aboveground net primary productivity, respectively. High nutrient use efficiency of schist specialists reduced the rates of within-stand nutrient cycling through the production of nutrient-poor plant tissues and litter. Although forests on schist soils had higher basal area and similar rates of productivity to forests on basalt, uptake of Mg, K, P, and N were markedly less on schist than on basalt, particularly after a cyclone disturbance. Stands on schist soils were also less affected by the cyclone and, as a result, contributed less (ca. 50%) Mg, K, P, and N inputs to the forest floor (via litterfall) than stands on basalt soils. System “openness” (i.e. the risk of nutrient loss) from cyclone-affected basalt forests was minimised by high rates of uptake following disturbance and large effective cation exchange capacities of soils. Soil–plant-disturbance interactions are likely to engender different fitness-enhancing strategies on fertile and infertile soils, possibly leading to the development and/or maintenance of diversity in rainforests.     

Tailoring restoration interventions to the grassland‐savanna‐forest complex in central Brazil

Defining the reference system for restoration projects in regions characterized by complex vegetation mosaics is challenging. Here we use the Cerrado region of Brazil as an example of the importance of clearly defining multiple natural and anthropogenically altered states in grassland‐savanna‐forest mosaics. We define three main, natural vegetation types–grassland, savanna, and scleromorphic (cerradão) forest–to (1) distinguish between original and degraded states and (2) set appropriate targets for and guide restoration. We contend that the differences in Cerrado vegetation composition originally were driven by soil conditions and secondarily by fire frequency and precipitation patterns that differ from the core to the edge of the Cerrado region. Grasslands are found on the shallowest, least fertile soils and/or in waterlogged soils; scleromorphic forests are generally located on deeper, more fertile soils; and savannas occupy an intermediate position. In recent decades, this biophysical template has been overlain by a range of human land‐use intensities that strongly affect resilience, resulting in alternative anthropogenic states. For example, areas that were originally scleromorphic forest are likely to regenerate naturally following low‐ or medium‐intensity land use due to extensive resprouting of woody plants, whereas grassland restoration requires reintroduction of grass and forb species that do not tolerate soil disturbance and exotic grass competition. Planting trees into historic grasslands results in inappropriate restoration targets and often restoration failure. Correctly identifying original vegetation types is critical to most effectively allocate scarce restoration funding.     

A preliminary test of catabolic nutrients in explanation of the puzzling treelessness of grassland in mesic Australia

The puzzling exclusion of native trees from grassland under mesic climates can hypothetically be explained by nutritional regimes that make grasses competitively superior to trees. One hypothesis holds that where nutrient concentrations in soils allow catabolism to be as rapid as anabolism at the scale of the whole plant, photosynthates are respired instead of being allocated to stems, resulting in dominance of herbaceous over woody growth‐forms. This matching of catabolic and anabolic rates would depend on the ratios of catabolic to anabolic nutrient elements in soils at a particular site. We consequently investigated elemental concentrations of soils near boundaries between treeless grassland and eucalypt woodland in New South Wales, Australia. Soils were analysed for Be, Na, Mg, Al, Si, K, Ca, Ti, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, As, Se, Sr, Mo, Cd, Ba and Pb. Based on our preliminary classification of nutrients as mainly catabolic (e.g. Cu) or mainly anabolic (e.g. Mn), the results showed that, relative to adjacent woodland, the soils under treeless grassland were naturally enriched in anabolic nutrients, but even more so in catabolic nutrients. Furthermore, the greater concentration of Mo – which can induce deficiency of the catabolic nutrient Cu – in soils under woodland than in soils under treeless grassland suggests that catabolic dystrophy allows eucalypt seedlings to prevail in competition with grasses. The results support the theory that where ratios of catabolic to anabolic nutrients are sufficient grasses will achieve competitive superiority over woody seedlings. Our study indicates that soil nutrient concentrations may partly explain the incidence of natural patches of treeless grassland in Australian landscapes otherwise dominated by woody plants.     

Importance of molehill disturbances for invasion by Bunias orientalis in meadows and pastures

Small-scale soil disturbances by fossorial animals can change physical and biotic conditions in disturbed patches and influence spatial and temporal dynamics, and the composition of plant communities. They create regeneration niches and colonization openings for native plants and, according to the intermediate disturbance hypothesis, they are expected to increase plant community diversity. However, it also has been reported that increased disturbance resource availability and decreased competition with native species may result in the invasion of communities by alien plant species, as predicted by the fluctuating resources theory of invasibility. In this study, we investigated the importance of European mole disturbances for the invasion of semi-natural fresh meadows and pastures by the alien plant, Bunias orientalis, which has mainly spread throughout Central Europe on anthropogenically disturbed sites. We hypothesized that the invader, being particularly well adapted to anthropogenic disturbances, enters into dense vegetation of meadows and pastures mainly on mole mounds. To assess the seedling recruitment of B. orientalis in relation to disturbance, we counted the number of seedlings that emerged on molehills and control plots in meadows and pastures. The establishment of juvenile (0–1 year) rosette plants on and off molehills was surveyed on 5 × 5 m plots. In accordance with our hypothesis, mole disturbances were found to serve as a gateway for B. orientalis by which the invader may colonize semi-natural grasslands. The seedlings of the species emerged almost solely on molehills and the young rosettes were established predominantly on mole mounds. Although the seedling density did not differ significantly between the meadows and pastures, the number of established plants in the pastures was considerably higher. We suggest that the invasion by B. orientalis in pastures may be facilitated by vegetative regeneration following root fragmentation by sheep pasturing.     

水淹干扰对羊草草地地上生物量影响的初步研究

为在一定程度上揭示水淹干扰后草地净初级生产力变化的机制,对松嫩平原羊草草地水淹干扰梯度上的地上生物量进行了测定和比较,并对经历水淹干扰后土壤水分及主要养分(N、P)的变化以及植被物种组成的变化作了比较分析。结果表明,轻度和重度水淹干扰样带的地上生物量显著高于未受水淹干扰的对照样带,分别高出对照样带89．54％和113.45％,表明水淹干扰消除了对草地生产力起限制作用的因素,使草地净初级生产力有了大幅度提高,水淹干扰首先改变了土壤的水分状况,而水分状况是限制草地生产力的最主要的因素,尤其是在干旱年份,消除了干旱对草地生产力的限制;土壤水分的增加导致土壤养分(N、P等)的有效性显著增加,消除了原来土壤养分匮乏对草地生产力造成的限制;在水淹干扰作用下,群落的物种组成由相对低矮的物种组合趋向于向形态高大的、具有更高生产力潜力的物种组合转变。     

Land use intensity,rather than plant species richness,affects the leaching risk of multiple nutrients from permanent grasslands

The intensification of land use constitutes one of the main drivers of global change and alters nutrient fluxes on all spatial scales, causing landscape‐level eutrophication and contamination of natural resources. Changes in soil nutrient concentrations are thus indicative for crucial environmental issues associated with intensive land use. We measured concentrations of NO₃–N, NH₄–N, P, K, Mg, and Ca using 1,326 ion‐exchange resin bags buried in 20 cm depth beneath the main root zone in 150 temperate grasslands. Nutrient concentrations were related to land use intensity, that is, fertilization, mowing, grazing intensities, and plant diversity by structural equation modeling. Furthermore, we assessed the response of soil nutrients to mechanical sward disturbance and subsequent reseeding, a common practice for grassland renewal. Land use intensity, especially fertilization, significantly increased the concentrations of NO₃–N, NH₄–N, K, P, and also Mg. Besides fertilization (and tightly correlated mowing) intensity, grazing strongly increased NO₃–N and K concentrations. Plant species richness decreased P and NO₃–N concentrations in soil when grassland productivity of the actual year was statistically taken into account, but not when long‐term averages of productivity were used. Thus, we assume that, in the actual study year, a distinct drought period might have caused the observed decoupling of productivity from fertilization and soil nutrients. Breaking up the grassland sward drastically increased NO₃–N concentrations (+146%) but reduced NH₄–N, P, and K concentrations, unbalancing soil nutrient stoichiometry and boosting the risk of N leaching. Reseeding the sward after disturbance did not have a short‐term effect on nutrient concentrations. We conclude that renewal of permanent grassland should be avoided as far as possible and future grassland management has to strongly rise the effectiveness of fertilization. Additionally, grassland management might have to increasingly taking care of periods of drought, in which nutrient additions might not increase plant growth but potentially only facilitate leaching.