Landscape history and soil properties affect grassland decline and plant species richness in rural landscapes

Past semi-natural grassland extent is thought to have a major influence on contemporary species richness in rural landscapes. The loss of grasslands over the last 300 years was reconstructed for 12 rural landscapes in Sweden, ranging from open modern agricultural landscapes to more forested landscapes. Old maps and aerial photographs from 1950s and today were used to reconstruct landscape patterns in four time-steps to investigate how present plant species richness relates to past grassland extent and decline in old and new grassland habitats. The relative importance of soil properties on the timing of grassland decline was assessed. Plant species occurrence was recorded in managed and abandoned grassland habitats in each landscape.Past and present grassland distribution was a major factor in determining plant species patterns found in grasslands today. All landscapes had an average of 80% grassland 300 years ago. Since then grassland has declined by 90% across all landscapes. Proportion of clay soils influenced the timing of grassland decline, where grasslands in landscapes dominated by clay soils were conversed to crop-fields more than 100 years ago. Grasslands on coarser soils declined later, primarily to forest. Landscapes with more than 10% semi-natural grassland left today had 50% higher species richness in all grasslands, including both abandoned and new grassland. Time since major grassland decline also seems to have an effect on the landscapes’ species richness. The results show that plant species patterns in grasslands at local scales are determined by broader landscape processes which may have occurred many centuries ago.     

Rapid effects of plant species diversity and identity on soil microbial communities in experimental grassland ecosystems

Changes in plant community structure, including the loss of plant diversity may affect soil microbial communities. To test this hypothesis, plant diversity and composition were experimentally varied in grassland plots cultivated with monocultures or mixtures of 2, 3 or 4 species. We tested the effects of monocultures versus mixtures and of plant species composition on culturable soil bacterial activity, number of substrates used and catabolic diversity, microbial biomass N, microbial respiration, and root biomass. These properties were all measured 10 months after seeding the experiment. Soil bacterial activity, number of substrates used and catabolic diversity were measured in the different plant communities using BIOLOG GN and GP microplates, which are redox-based tests measuring capacity of soil culturable bacteria to use a variety of organic substrates. Microbial biomass N, microbial respiration, and root biomass were insensitive to plant diversity. Culturable soil microbial activity, substrates used and diversity declined with declining plant diversity. Their activity, number of substrates used and diversity were significantly higher in plots with 3 and 4 plant species than in monocultures and in plots with 2 species. There was also an effect of plant species composition. Culturable soil microbial activity and diversity was higher in the four-species plant community than in any of the plant monocultures suggesting that the effect of plant diversity could not be explained by the presence of a particular plant species. Our results showed that changes in plant diversity and composition in grassland ecosystems lead to a rapid response of bacterial activity and diversity.     

Small-scale plant species richness and evenness in semi-natural grasslands respond differently to habitat fragmentation

The study explores whether small-scale species diversity, species evenness and species richness in semi-natural grassland communities are similarly associated with present management regime and/or present and historical landscape context (percentage of different land-cover types in the surroundings). Species diversity, evenness and richness were recorded within 441 50 × 50 cm grassland plots in 4.5 × 4.5 km agricultural landscape on Öland, Sweden. Recent and historical land-cover maps (years 2004, 1959, 1938, 1835, and 1800) were used to characterize the present and past landscape context of the sampled vegetation plots. Partial regression and simultaneous autoregressive models were used to explore the relationships between species diversity measures (Shannon diversity, richness and evenness) and different explanatory variables while accounting for spatial autocorrelation in the data. The results indicated that species richness was relatively sensitive to grassland isolation, while the response of species evenness to isolation was characterized by a degree of inertia. Because the richness and evenness components of species diversity may respond differently to habitat fragmentation, we suggest that monitoring projects and empirical studies that focus on changes in biodiversity in semi-natural grasslands should include the assessment of species evenness - as a complement to the assessment of species richness. In addition, our results indicated that the development and persistence of a species-rich and even grassland vegetation was favoured in areas that have historically (in the 19th century) been surrounded by grasslands. Information on landscape history should, whenever possible, be incorporated into the planning of strategies for grassland conservation.     

The importance of traditional meadow management techniques for conservation of vascular plant species richness in Norway

Using information from a regional survey of vascular plants of 130 sites in western Norway, a selection of sites based on a heuristic iterative complementarity-based nature reserve selection procedure was performed. The results indicate that conservation of traditionally managed hay meadows is of major importance as they contributed 60.1% of all native species recorded; afforested grasslands (deciduous woodlands < 70 years old) contributed 26.8%, whereas artificially fertilized hay meadows and intensively cultivated grasslands taken together contributed 13.1% of the species. The species composition of the meadows was significantly nested. Thus, if you conserve the most species-rich meadows, you also conserve most of the species in the less species-rich meadows. Nestedness in meadows was significantly correlated with within-meadow habitat diversity and soil pH. The most species-rich meadows were traditional meadows, characterized by high habitat diversity and high soil pH. These meadows will support nearly all species including habitat specialists and regionally rare species, whilst artificially fertilized hay meadows only support the generalist subset, i.e. common species. Area was not significantly correlated with nestedness suggesting that it is more important to cover many habitats than to preserve large traditional meadows just because they are large.     

Contrasting responses of plant and insect diversity to variation in grazing intensity

The effects of grazing intensity on plant and insect diversity were examined in four different types of grassland (intensively and extensively cattle-grazed pastures, short-term and long-term ungrazed grassland; 24 study sites). Vegetation complexity (plant species richness, vegetation height, vegetation heterogeneity) was significantly higher on ungrazed grasslands compared to pastures but did not differ between intensively and extensively grazed pastures. However, insect species richness was higher on extensively than on intensively grazed pastures, established by suction sampling of four insect taxa (Auchenorrhyncha, Heteroptera, Coleoptera, Hymenoptera Parasitica). This may be due to intensive grazing disrupting plant-insect associations as predicted by a “trophic-level” hypothesis. Local persistence and small-scale recolonization of insects on plants appeared to be difficult in the highly disturbed environment of intensive grazing. Insect diversity increased across the four treatments in the following order: intensively grazed<extensively grazed<short-term ungrazed<long-term ungrazed. The major predictor variable of differences in species diversity was found to be vegetation height. Predator-prey ratios within the investigated insect groups were not affected by grazing intensity.     

Soil fauna feeding activity in temperate grassland soils increases with legume and grass species richness

Edaphic fauna contributes to important ecosystem functions in grassland soils such as decomposition and nutrient mineralization. Since this functional role is likely to be altered by global change and associated shifts in plant communities, a thorough understanding of large scale drivers on below-ground processes independent of regional differences in soil type or climate is essential. We investigated the relationship between abiotic (soil properties, management practices) and biotic (plant functional group composition, vegetation characteristics, soil fauna abundance) predictors and feeding activity of soil fauna after accounting for sample year and study region. Our study was carried out over a period of two consecutive years in 92 agricultural grasslands in three regions of Germany, spanning a latitudinal gradient of more than 500 km. A structural equation model suggests that feeding activity of soil fauna as measured by the bait-lamina test was positively related to legume and grass species richness in both years. Most probably, a diverse vegetation promotes feeding activity of soil fauna via alterations of both microclimate and resource availability. Feeding activity of soil fauna also increased with earthworm biomass via a pathway over Collembola abundance. The effect of earthworms on the feeding activity in soil may be attributed to their important role as ecosystem engineers. As no additional effects of agricultural management such as fertilization, livestock density or number of cuts on bait consumption were observed, our results suggest that the positive effect of legume and grass species richness on the feeding activity in soil fauna is a general one that will not be overruled by regional differences in management or environmental conditions. We thus suggest that agri-environment schemes aiming at the protection of belowground activity and associated ecosystem functions in temperate grasslands may generally focus on maintaining plant diversity, especially with regard to the potential effects of climate change on future vegetation structure.     

Effects of insularisation on plant species richness in the prairie-forest ecotone

Data on plants from five groups of remnant prairies and forests in the prairie-forest ecotone of the midwestern United States show that: (1) Archipelagos of small sites tend to contain more species than do single large ones of equal total area. (2) No species are excluded from small sites. (3) Small sites tend to have surprisingly many species, and large sites surprisingly few, relative to a random colonisation model. (4) ‘Rare’ species (those that occur in only one site) are found more often in small sites than a random colonisation model would predict. (5) There is no evidence for any of these sites that species number is ‘relaxing’, but all these sites have been remnants for only a short time. Result (1) is consistent with results from similar studies on other taxa and suggests that there is no automatic reason to believe that species richness is maximised by single large refuges.     

Changes in local species richness of farmland birds in relation to land-use changes and landscape structure

It has been suggested that an increase in the area of low-intensity land-use on arable land (e.g. set-aside fields and short-rotation coppice), and high or increased farmland habitat heterogeneity, may halt or reverse the observed population decline of farmland birds. We tested these hypotheses by undertaking farmland bird censuses during two contrasting periods of agricultural policies and land-use (i.e. 1994 and 2004) in a farmland region covering a gradient of forest- to farmland-dominated landscapes in Sweden. Local species richness (i.e. at 3 hectare sites) declined significantly between 1994 and 2004. Local species richness was positively related to habitat heterogeneity in both years of study whereas temporal change in species richness was not. Local change in species richness was positively associated with a change in the proportion of non-rotational set aside and short-rotation coppice (i.e. low-intensity land-use forms), but also to changes in the amount of spring-sown crops. However, the effect of low-intensity land-use was significantly dependent on the amount of forest in the surrounding landscape. An increase in low-intensity land-use was linked to an increase (or less marked decrease) in species richness at sites located in open farmland surroundings but to a decrease in richness at sites located in forest surroundings. This interaction between amount of forest and low-intensity land-use could be interpreted as a “rare habitat effect”, where an increase in a farmland habitat only positively affects biodiversity when it was originally uncommon (i.e. open farmland areas). Our results suggest that conservation measures of farmland biodiversity have to be put in a landscape context.     

The influence of land-use practices and economics on plant species richness in meadows

Thirty-one meadows were investigated within five sites representing various farming styles found in Austrian cultural landscapes. The meadows were analysed regarding (a) biodiversity (vascular plant and bryophyte species richness), (b) land-use practices (fertiliser input, mowing intensity, the use of silage), and (c) economic aspects (variable costs, profit margin and subsidies per ha). There were significant negative correlations between plant species richness and mowing intensity and intensity of fertiliser application. Bryophytes were good indicators of low nutrient regimes, having high species richness at low fertiliser input. Vascular plants showed highest species richness at an intermediate nitrogen supply. The total plant species richness decreased with increasing nitrogen supply. Intensive silage production was also negatively correlated with plant diversity. Species with a very narrow ecological niche of soil moisture and nutrients declined, whereas species adapted to wider ecological conditions increased. Profit margin and variable costs correlated negatively with plant species richness, with meadows that offered low or no profit margins showing highest species richness. There was no significant relationship between species richness and the amount of subsidies invested at the study meadows. Estimated costs of maintaining a species are shown. It is concluded that if plant species richness are to be maintained in these meadows, farmers have to receive increased financial incentives through agro-environmental subsidies for appropriate meadow management, and these have to be linked to clearly defined measures.     

Changes in Collembola richness and diversity along a gradient of land-use intensity: A pan European study

Changes in Collembola richness and diversity along a land-use intensity gradient were studied in eight European countries (Portugal, Spain, France, Switzerland, Hungary, UK, Ireland and Finland). In each country a set of six 1 km² land-use units (LUUs) were selected forming a gradient ranging from natural forest to agricultural dominated landscapes, passing through mixed-use ones. In addition to data on Collembola, detailed information regarding landscape diversity and structure was collected for each LUU. A total of 47,774 individuals were identified from 281 species. Collembola reacted not only to changes in the diversity of the landscape, but also to the composition of that diversity and the area occupied by each land-use type at each LUU. Although species richness patterns were not concordant among the different countries, the total number of species per LUU (landscape richness) was generally higher in natural forests and mixed-used landscapes, and lower in agricultural dominated landscapes. Moreover, high richness and diversity of Collembola at each LUU were associated with a diverse landscape structure, both in terms of number of patches and patch richness. Despite this comparable species richness between mixed-use landscapes and those dominated by natural forests, average species richness on forested areas (local richness) decreased along the gradient, showing that forest patches on mixed-use landscapes support a lower richness than in landscapes dominated by forest. This aspect is important when addressing the role of native forests in structuring biodiversity in disturbed and fragmented landscapes. Although a diverse landscape can support a high biodiversity, the results suggest that intensive fragmentation should be avoided with the risk of collapsing local species richness with the consequent result for regional biodiversity.     

Response of rhizosphere microbial communities to plant succession along a grassland chronosequence in a semiarid area

Journal of Soils and Sediments - Changes in microbial communities during natural succession in semiarid areas have been widely studied but their association with plant and soil properties remains...     

Fifty years of change in Central European grassland vegetation: Large losses in species richness and animal-pollinated plants

There is growing concern that biodiversity loss in European agricultural landscapes is having negative effects on functional trait diversity. Long-term studies examining vegetation changes from the period before agricultural industrialisation are however rare. Here, we ask how management intensification and increased nutrient input initiated in the 1950/1960s have altered grassland plant community composition, species diversity and functional trait composition using comprehensive datasets from five floodplain regions (plus one protected reference region) in northern Germany. Sites with available historical relevés and vegetation maps (1950/1960s, 1990s) were resampled in 2008 to facilitate the analysis of a period spanning four to five decades.Plant community composition changed tremendously in all study regions during the 50 year period, which was related to increasing Ellenberg indicator values for nutrient availability. Species richness at the plot-level fell by 30–50% over the period, and losses in functional diversity were equally large. A non-formal comparison with the results from the protected reference study region indicates that the changes may mostly be attributable to local nutrient input rather than to supra-regional climate change. Our results indicate a consistent trend towards much more species-poor communities dominated by mow-tolerant, N-demanding competitive grasses, whereas species with more ruderal strategies, species flowering early in the season and, in particular, insect-pollinated herbs have all decreased. The substantial loss of nectar-producing grassland herbs is likely to have negative effects on the abundance of pollinating insects, with consequences for the grassland animal communities. This highlights the growing need for adequate grassland management schemes with low N input to preserve high-nature-value grassland.     

Belowground responses by AM fungi and animal trophic groups to repeated defoliation in an experimental grassland community

We tested a hypothesis that the effects of defoliation on plants and soil organisms vary with the number of successive defoliations. We established a 23-week greenhouse experiment using replicated grassland microcosms that were composed of three plant species, Trifolium repens, Plantago lanceolata and Phleum pratense, growing together in grassland soil with a diverse soil community. The experiment consisted of two treatment factors-defoliation and harvest time-in a fully factorial design. The defoliation treatment had two levels, i.e. no trimming and trimming of plants every 2 weeks, and the harvest time five levels, i.e. harvests after 1-3, 5 and 7 trimmings. Shoot production (trimmed plus harvested shoot mass), harvested shoot and root mass and root N and C concentrations increased with time but were reduced by defoliation. Colonization rates of arbuscular mycorrhizal (AM) fungi decreased with time in T. repens roots but were enhanced by defoliation, whereas AM colonization rates in P. pratense roots were not affected by harvest time or defoliation. The abundance of bacterivorous and fungivorous nematodes decreased and that of herbivorous and predatory nematodes increased with time, while the abundance of omnivorous nematodes and detritivorous enchytraeids varied in time without a linear trend. Defoliation had no effect on fungivores and predators but increased the abundance of bacterivores. Defoliation also increased the abundance of herbivores, omnivores and detritivores after 2 trimmings and that of omnivores and detritivores after 5 trimmings, but had a negative effect on omnivores after 3 trimmings and on herbivores after 7 trimmings. Among nematode genera, some deviation from the trophic group responses existed: for instance, defoliation reduced the abundance of bacterivorous Acrobeloides spp. and did not affect the abundance of herbivorous Filenchus spp. and Paratylenchus spp. Our results show that the effects of defoliation on plants, AM fungi and some soil animal trophic groups may remain constant all the way through several defoliations, whereas other animal trophic groups may have different and even opposite responses to defoliation depending on the length of the defoliation period before monitoring. This shows how separate studies with defoliation periods of different length can produce contradictory results of the effects of defoliation on the abundance of soil animals.     

Plant species richness does not attenuate responses of soil microbial and nematode communities to a flood event

Human activities are causing climatic changes and alter the composition and biodiversity of ecosystems. Climate change has been and will be increasing the frequency and severity of extreme climate events and natural disasters like floods in many ecosystems. Therefore, it is important to investigate the effects of disturbances on ecosystems and identify potential stabilizing features of ecological communities. In this study, soil microbial and nematode communities were investigated in a grassland biodiversity experiment after a natural flood to investigate if plant diversity is able to attenuate or reinforce the magnitude of effects of the disturbance on soil food webs. In addition to community analyses of soil microorganisms and nematodes, the stability indices proportional resilience, proportional recovery, and proportional resistance were calculated. Generally, soil microbial biomass decreased significantly due to the flood with the strongest reduction in gram-negative bacteria, while gram-positive bacteria were less affected by flooding. Fungal biomass increased significantly three months after the flood compared to few days before the flood, reflecting elevated availability of dead plant biomass in response to the flood. Similar to the soil microbial community, nematode community structure changed considerably due to the flood by favoring colonizers (in the broadest sense r-strategists; c–p 1, 2 nematodes), particularly so at high plant diversity. None of the soil microbial community stability indices and few of the nematode stability indices were significantly affected by plant diversity, indicating limited potential of plant diversity to buffer soil food webs against flooding disturbance. However, plant diversity destabilized colonizer populations, while persister populations (in the broadest sense K-strategists; c–p 4 nematodes) were stabilized, suggesting that plant diversity can stabilize and destabilize populations depending on the ecology of the focal taxa. The present study shows that changes in plant diversity and subsequent alterations in resource availability may significantly modify the compositional shifts of soil food webs in response to disturbances.     

Evenness and plant species identity affect earthworm diversity and community structure in grassland soils

Diversity of plant species and the species composition (identity) are known to influence below-ground diversity. In this paper we examine the effects of plant species diversity (richness and evenness), rates of nitrogen application and planting density, on earthworm community structure in grassland. The study was carried out at three sites in Ireland using a Simplex experimental design to define the compositions of the experimental plant communities used.A negative relationship was detected between diversity (evenness) of plant species and diversity of earthworms in the soils. However, plant species identity also affected the structure of the earthworm assemblage. In particular, the legume, Trifolium repens had a strong effect but this was conditional on the rate of nitrogen application. No earthworm species favoured communities dominated by slow growing grasses (Phleum pratense and Dactylis glomerata) (P = 0.02).Higher N inputs reduced earthworm abundance and biomass under T. repens. Earthworm richness, was negatively influenced by elevated amounts of N inputs. No effect of planting density was detected but this factor also did not affect plant biomass production.     

Effect of cutting frequency on species richness and dry matter yield of permanent grassland after grazing cessation

The effect of grazing cessation management has only rarely been investigated. In this study, the influence of cutting regime in the absence of fertilization was examined during 2005–2011 on a semi-natural grassland in Rapotin (Czech Republic) which had been grazed for more than 30 years before the start of the trial. This was conducted as a randomized plot design with five levels of treatment: 4 C – intensive (four cuts per year); 3 C – medium intensive (three cuts per year); 2 C – low intensive (two cuts per year); 1 C – extensive (one cut per year); 0 C – abandoned (control; without any management). Species richness was significantly lower in treatments 1 C and 0 C than in other treatments. Management and year had similar and significant influence on species composition and explained 23% of its variability. Dry matter (DM) yields were found to be lower in three-cut than in two-cut treatments (5.56 t ha^?1 and 6.22 t ha^?1, respectively). In the case of grazing cessation, a suitable compromise of three cuts per year can be recommended to guarantee maintenance of species richness, botanical composition, and appropriate grass forage yield under similar site conditions.     

Combining field incubation with nitrogen-15 labelling to examine nitrogen transformations in low to high intensity grassland management systems

 The ¹⁵N isotope dilution method was combined with a field incubation technique to provide simultaneous measurements of gross and net rates of N turnover in three long-term swards: unfertilized (Z) or receiving N either from N fixation as clover (C), or as 200 kg fertilizer N ha^–1 year^–1 (F). Uniform N enrichment of soil microplots was achieved with a multi-point soil injector to measure mineralization/immobilization turnover and nitrification over a 4-day incubation. Net rates of mineralization ranged between 0.6 and 2.9 μg N g^–1 day^–1 and in all three treatments were approximately half the gross rates. Nitrification rates (gross) were between 1.0 and 1.6 μg N g^–1 day^–1. In the F treatment, the turnover of NH₄ ⁺-N and NO₃ ^–-N pools was on a 2- and 4-day cycle, respectively, whereas in the N-limited treatments (C and Z) turnover rates were faster, with the NO₃ ^–-N pools turning over twice as fast as the NH₄ ⁺-N pools. Therefore, available N was recycled more efficiently in the C and Z treatments, whereas in the F treatment a higher N pool size was maintained which would be more vulnerable to leakage. A large proportion of the added ¹⁵N was recovered in the soil microbial biomass (SMB), which represented a 4–5 times larger sink for N than the plant biomass. Although the C treatment had a significantly lower SMB than the grass-only treatments, there were no differences in microbial activity. Gross rates of nitrification increased along the gradient of N input intensity (i.e. Z<C<F), and the addition of a nitrification inhibitor (C₂H₂) tended to increase microbial immobilization, but did not influence plant N uptake. In this study, the value of combining different techniques to verify net rates was demonstrated and the improved methodology for ¹⁵N labelling of soil enabled measurements to be obtained from relatively undisturbed soil under natural field conditions. Received: 25 May 1999     

Effect of density and species richness of soil mesofauna on nutrient mineralisation and plant growth

The aim of this study was to test the relative importance of changes in density and species richness of soil mesofauna as determinants of nutrient mineralisation and plant growth. The experiment was carried out using microcosms containing a mixture of plant litter and soil in which seedlings of Lolium perenne were planted, and a range of combinations of levels of density and species richness of microarthropods added. Over the duration of the experiment, nutrient release, measured as concentrations of NO₃ ^--N and total N in leachates, increased significantly with increasing microarthropod density, but decreased with increasing species richness. Leachate concentrations of NH₄ ⁺-N, dissolved organic N and C (DON and DOC) were not affected by the faunal treatments. Soil respiration, a measure of microbial activity, decreased with increasing density of microarthropods, whereas microbial biomass was not affected by microarthropods. Increasing density of soil animals had a negative effect on the shoot biomass of L. perenne while the effect of species richness was positive. Neither the species richness nor density of soil microarthropods was found to significantly influence root biomass. We conclude that variations in animal density had a greater influence on soil nutrient mineralisation processes than did species richness. Possible reasons for these opposing effects of animal density and diversity on soil N mobilization are discussed.     

Landscape changes caused by high altitude ski-pistes affect bird species richness and distribution in the Alps

There is a paucity of research on the wider landscape-level effects of ski-piste construction on alpine fauna. In this study, the response of alpine bird communities to the landscape changes induced by the construction of ski-pistes was investigated in the western Italian Alps. The aims were: (i) to test the hypothesis that ski-pistes have a detrimental effect on alpine grassland bird communities at a landscape-scale; and, (ii) to model local probability distributions of bird species according to different scenarios of ski-piste restoration and ski-piste proliferation above the treeline. Species richness and presence/absence data from point counts were analyzed in relation to GIS-derived landscape variables based on a 16 ha hexagon grid. Predictive variables were selected through the LASSO model selection procedure, and logistic regression was used to estimate the probability of occurrence of each species in each hexagon. Grassland species richness, and probability of occurrence of water pipit, wheatear and black redstart, significantly decreased with increasing extent of ski-piste edge. Length of ski-piste edge was in the set of best models considering a large range of habitat and landscape predictors, and are therefore clearly features that exert a strong negative effect on high alpine grassland bird communities. Predictions of species occurrence were made by applying the models to different scenarios of habitat change. These showed predicted detrimental impacts of a relatively small 10% increase in ski-piste extent, but also that grassland restoration on existing ski-pistes could result in significantly increased occurrence rates of alpine grassland species. This study suggests that ski-pistes are perceived by birds as detrimental features of the alpine landscape. To minimize their impact, new, environmentally–friendly ways of constructing pistes should be developed, which could include habitat restoration and management to obtain a level of grass cover such that edges of ski-pistes are no longer perceived by birds.