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Although temporal heterogeneity is a well-accepted driver of biodiversity, effects of interannual variation in land-use intensity (LUI) have not been addressed yet. Additionally, responses to land use can differ greatly among different organisms; therefore, overall effects of land-use on total local biodiversity are hardly known. To test for effects of LUI (quantified as the combined intensity of fertilization, grazing, and mowing) and interannual variation in LUI (SD in LUI across time), we introduce a unique measure of whole-ecosystem biodiversity, multidiversity. This synthesizes individual diversity measures across up to 49 taxonomic groups of plants, animals, fungi, and bacteria from 150 grasslands. Multidiversity declined with increasing LUI among grasslands, particularly for rarer species and aboveground organisms, whereas common species and belowground groups were less sensitive. However, a high level of interannual variation in LUI increased overall multidiversity at low LUI and was even more beneficial for rarer species because it slowed the rate at which the multidiversity of rare species declined with increasing LUI. In more intensively managed grasslands, the diversity of rarer species was, on average, 18% of the maximum diversity across all grasslands when LUI was static over time but increased to 31% of the maximum when LUI changed maximally over time. In addition to decreasing overall LUI, we suggest varying LUI across years as a complementary strategy to promote biodiversity conservation.
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.
Land use is increasingly recognized as a major driver of biodiversity and ecosystem functioning in many current research projects. In grasslands, land use is often classified by categorical descriptors such as pastures versus meadows or fertilized versus unfertilized sites. However, to account for the quantitative variation of multiple land-use types in heterogeneous landscapes, a quantitative, continuous index of land-use intensity (LUI) is desirable. Here we define such a compound, additive LUI index for managed grasslands including meadows and pastures. The LUI index summarizes the standardized intensity of three components of land use, namely fertilization, mowing, and livestock grazing at each site. We examined the performance of the LUI index to predict selected response variables on up to 150 grassland sites in the Biodiversity Exploratories in three regions in Germany(Alb, Hainich, Schorlheide). We tested the average Ellenberg nitrogen indicator values of the plant community, nitrogen and phosphorus concentration in the aboveground plant biomass, plant-available phosphorus concentration in the top soil, and soil C/N ratio, and the first principle component of these five response variables.
The LUI index significantly predicted the principal component of all five response variables, as well as some of the individual responses. Moreover, vascular plant diversity decreased significantly with LUI in two regions (Alb and Hainich).
Inter-annual changes in management practice were pronounced from 2006 to 2008, particularly due to variation in grazing intensity. This rendered the selection of the appropriate reference year(s) an important decision for analyses of land-use effects, whereas details in the standardization of the index were of minor importance. We also tested several alternative calculations of a LUI index, but all are strongly linearly correlated to the proposed index.
The proposed LUI index reduces the complexity of agricultural practices to a single dimension and may serve as a baseline to test how different groups of organisms and processes respond to land use. In combination with more detailed analyses, this index may help to unravel whether and how land-use intensities, associated disturbance levels or other local or regional influences drive ecological processes.
Bryophytes constitute an important and permanent component of the grassland flora and diversity in Europe. As most bryophyte species are sensitive to habitat change, their diversity is likely to decline following land-use intensification. Most previous studies on bryophyte diversity focused on specific habitats of high bryophyte diversity, such as bogs, montane grasslands, or calcareous dry grasslands. In contrast, mesic grasslands are rarely studied, although they are the most common grassland habitat in Europe. They are secondary vegetation, maintained by agricultural use and thus, are influenced by different forms of land use. We studied bryophyte species richness in three regions in Germany, in 707 plots of 16 m(2) representing different land-use types and environmental conditions. Our study is one of the few to inspect the relationships between bryophyte richness and land use across contrasting regions and using a high number of replicates. Among the managed grasslands, pastures harboured 2.5 times more bryophyte species than meadows and mown pastures. Similarly, bryophyte cover was about twice as high in fallows and pastures than in meadows and mown pastures. Among the pastures, bryophyte species richness was about three times higher in sheep grazed plots than in the ones grazed by cattle or horses. In general, bryophyte species richness and cover was more than 50% lower in fertilized than in unfertilized plots. Moreover, the amount of suitable substrates was linked to bryophyte diversity. Species richness of bryophytes growing on stones increased with stone cover, and the one of bryophytes growing on bark and deadwood increased with larger values of woody plant species and deadwood cover. Our findings highlight the importance of low-intensity land use and high structural heterogeneity for bryophyte conservation. They also caution against an intensification of traditionally managed pastures. In the light of our results, we recommend to maintain low-intensity sheep grazing on sites with low productivity, such as slopes on shallow soils.
Bryophytes constitute an important and permanent component of the grassland flora and diversity in Europe. As most bryophyte species are sensitive to habitat change, their diversity is likely to decline following land-use intensification. Most previous studies on bryophyte diversity focused on specific habitats of high bryophyte diversity, such as bogs, montane grasslands, or calcareous dry grasslands. In contrast, mesic grasslands are rarely studied, although they are the most common grassland habitat in Europe. They are secondary vegetation, maintained by agricultural use and thus, are influenced by different forms of land use. We studied bryophyte species richness in three regions in Germany, in 707 plots of 16 m2 representing different land-use types and environmental conditions. Our study is one of the few to inspect the relationships between bryophyte richness and land use across contrasting regions and using a high number of replicates.Among the managed grasslands, pastures harboured 2.5 times more bryophyte species than mead-ows and mown pastures. Similarly, bryophyte cover was about twice as high in fallows and pastures than in meadows and mown pastures. Among the pastures, bryophyte species richness was about three times higher in sheep grazed plots than in the ones grazed by cattle or horses. In general, bryophyte species richness and cover was more than 50% lower in fertilized than in unfertilized plots. Moreover, the amount of suitable substrates was linked to bryophyte diversity. Species richness of bryophytes growing on stones increased with stone cover, and the one of bryophytes growing on bark and deadwood increased with larger values of woody plant species and deadwood cover. Our findings highlight the importance of low-intensity land use and high structural heterogeneity for bryophyte conservation. They also caution against an intensification of traditionally managed pastures. In the light of our results, we recommend to maintain low-intensity sheep grazing on sites with low productivity, such as slopes on shallow soils.
Assessing diversity is among the major tasks in ecology and conservation science. In ecological and conservation studies, epiphytic cryptogams are usually sampled up to accessible heights in forests. Thus, their diversity, especially of canopy specialists, likely is underestimated. If the proportion of those species differs among forest types, plot-based diversity assessments are biased and may result in misleading conservation recommendations. We sampled bryophytes and lichens in 30 forest plots of 20 m x 20 m in three German regions, considering all substrates, and including epiphytic litter fall. First, the sampling of epiphytic species was restricted to the lower 2 m of trees and shrubs. Then, on one representative tree per plot, we additionally recorded epiphytic species in the crown, using tree climbing techniques. Per tree, on average 54% of lichen and 20% of bryophyte species were overlooked if the crown was not been included. After sampling all substrates per plot, including the bark of all shrubs and trees, still 38% of the lichen and 4% of the bryophyte species were overlooked if the tree crown of the sampled tree was not included. The number of overlooked lichen species varied strongly among regions. Furthermore, the number of overlooked bryophyte and lichen species per plot was higher in European beech than in coniferous stands and increased with increasing diameter at breast height of the sampled tree. Thus, our results indicate a bias of comparative studies which might have led to misleading conservation recommendations of plot-based diversity assessments.
There is a wealth of smaller-scale studies on the effects of forest management on plant diversity. However, studies comparing plant species diversity in forests with different management types and intensity, extending over different regions and forest stages, and including detailed information on site conditions are missing. We studied vascular plants on 1500 20 m x 20 m forest plots in three regions of Germany (Schwabische Alb, Hainich-Dun, Schorfheide-Chorin). In all regions, our study plots comprised different management types (unmanaged, selection cutting, deciduous and coniferous age-class forests, which resulted from clear cutting or shelterwood logging), various stand ages, site conditions, and levels of management-related disturbances. We analyzed how overall richness and richness of different plant functional groups (trees, shrubs, herbs, herbaceous species typically growing in forests and herbaceous light-demanding species) responded to the different management types. On average, plant species richness was 13% higher in age-class than in unmanaged forests, and did not differ between deciduous age-class and selection forests. In age-class forests of the Schwabische Alb and Hainich-Dun, coniferous stands had higher species richness than deciduous stands. Among age-class forests, older stands with large quantities of standing biomass were slightly poorer in shrub and light-demanding herb species than younger stands. Among deciduous forests, the richness of herbaceous forest species was generally lower in unmanaged than in managed forests, and it was even 20% lower in unmanaged than in selection forests in Hainich-Dun. Overall, these findings show that disturbances by management generally increase plant species richness. This suggests that total plant species richness is not suited as an indicator for the conservation status of forests, but rather indicates disturbances.
Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have consi-dered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community-weighted means (CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition-related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf-economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters.
Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have considered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community-weighted means (CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition-related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf-economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters.