Effects of plant diversity, plant productivity and habitat parameters on arthropod abundance in montane European grasslands

Arthropod abundance has been hypothesized to be correlated with plant diversity but the results of previous studies have been equivocal. In contrast, plant productivity, vegetation structure, abiotic site conditions, and the physical disturbance of habitats, are factors that interact with plant diversity, and that have been shown to influence arthropod abundance. We studied the combined effect of plant species diversity, productivity and site characteristics on arthropod abundance in 71 managed grasslands in central Germany using multivariate statistics. For each site we determined plant species cover, plant community biomass (productivity), macro- and micronutrients in the soil, and characterized the location of sites with respect to orographic parameters as well as the current and historic management regimes. Arthropods were sampled using a suction sampler and classified a priori into functional groups (FGs). We found that arthropod abundance was not correlated with plant species richness, effective diversity or Camargo's evenness, even when influences of environmental variables were taken into account. In contrast, plant community composition was highly correlated with arthropod abundances. Plant community productivity influenced arthropod abundance but explained only a small proportion of the variance. The abundances of the different arthropod FGs were influenced differentially by agricultural management, soil characteristics, vegetation structure and by interactions between different FGs of arthropods. Herbivores, carnivores and detritivores reacted differently to variation in environmental variables in a manner consistent with their feeding mode. Our results show that in natural grassland systems arthropod abundance is not a simple function of plant species richness, and they emphasize the important role of plant community composition for the abundance patterns of the arthropod assemblages.     

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.     

Bird assemblages in fragmented agricultural landscapes: the role of small brigalow remnants and adjoining land uses

Agricultural intensification typically leads to changes in bird diversity and community composition, with fewer species and foraging guilds present in more intensively managed parts of the landscape. In this study, we compare bird communities in small (2–32 ha) brigalow (Acacia harpophylla) remnants with those in adjacent uncultivated grassland, previously cultivated grassland and current cropland, to determine the contribution of different land uses to bird diversity in the agricultural landscape. Twenty remnant brigalow patches and adjacent agricultural (‘matrix’) areas in southern inland Queensland, Australia were sampled for bird composition and habitat characteristics. The richness, abundance and diversity of birds were all significantly higher in brigalow remnants than in the adjacent matrix of cropping and grassland. Within the matrix, species richness and diversity were higher in uncultivated grasslands than in current cultivation or previously cultivated grasslands. Forty-four percent of bird species were recorded only in brigalow remnants and 78% of species were recorded in brigalow and at least one other land management category. Despite high levels of landscape fragmentation and modification, small patches of remnant brigalow vegetation provide important habitat for a unique and diverse assemblage of native birds. The less intensively managed components of the agricultural matrix also support diverse bird assemblages and thus, may be important for local and regional biodiversity conservation.     

Soil phosphorus availability determines the contribution of small,individual grassland remnants to the conservation of landscape-scale biodiversity

Questions

Small, remnant habitats embedded in degraded, human-dominated landscapes are generally not a priority in conservation, despite their potential role in supporting landscape-scale biodiversity. To warrant their inclusion in conservation management and policy, we question under which conditions they may exhibit the largest conservation value.

Location

Nine landscapes spread across the counties of Stockholm and Södermanland, Sweden.

Methods

Per landscape, plant communities were surveyed in 6 and 12 1 × 1 m² plots across large, intact semi-natural grasslands and small remnant grasslands, respectively. These two contrasting grassland types served as a model system. A topsoil sample was taken in each plot to determine habitat quality in terms of soil pH, plant-available P, and C:N ratio. We used a joint species distribution model to analyse the extent to which grassland type and habitat quality define and predict resident community diversity and composition, including whether they support grassland specialists.

Results

At the landscape scale, the combined remnant grasslands sustained diverse plant communities which did include a significant subset of habitat specialists. Yet, the contribution of individual remnants clearly varied with local-scale habitat quality; soil phosphorus availability lowered plot-level species richness, mostly by constraining the occurrence of grassland specialists. Semi-natural grassland communities were comparatively insensitive to variation in soil phosphorus availability.

Conclusions

The combined habitat amount and the significant number of habitat specialists sustained by remnant grasslands with high habitat quality, shows they can represent a valuable resource to support landscape-scale biodiversity conservation. This offers no wildcard to neglect the continued biotic and abiotic threats on semi-natural grassland plant diversity such as chronic and accumulating P eutrophication, discontinuation of management or poor matrix permeability, as semi-natural grasslands harbour the majority of habitat specialists, while sourcing surrounding remnant grassland communities.

     

An experimental test of the effect of plant functional group diversity on arthropod diversity

Characteristics used to categorize plant species into functional groups for their effects on ecosystem functioning may also be relevant to higher trophic levels. In addition, plant and consumer diversity should be positively related because more diverse plant communities offer a greater variety of resources for the consumers. Thus, the functional group composition and richness of a plant community may affect the composition and diversity of the herbivores and even higher trophic levels associated with that community. We tested this hypothesis by sampling arthropods with a vacuum sampler (34 531 individuals of 494 species) from an experiment in which we manipulated plant functional group richness and composition. Plant manipulations included all combinations of three functional groups (forbs, C₃ graminoids, and C₄ graminoids) removed zero, one, or two at a time from grassland plots at Cedar Creek Natural History Area, MN. Although total arthropod species richness was unrelated to plant functional group richness or composition, the species richness of some arthropod orders was affected by plant functional group composition. Two plant characteristics explained most of the effects of plant functional groups on arthropod species richness. Nutritional quality, a characteristic related to ecosystem functioning, and taxonomic diversity, a characteristic not used to designate plant functional groups, seemed to affect arthropod species richness both directly and indirectly. Thus, plant functional groups designated for their effects on ecosystem processes will only be partially relevant to consumer diversity and abundance.     

Floristic diversity of meadow steppes in the Western Siberian Plain: effects of abiotic site conditions,management and landscape structure

Temperate grasslands have suffered from severe habitat loss and degradation worldwide. In Russia, vast areas of forest-steppe grasslands have been converted to cropland during Soviet times, whilst remaining grasslands were often intensively grazed. Contrastingly, the collapse of the Soviet Union have resulted in a massive reduction in livestock numbers and cessation of management. Albeit relatively large natural grassland areas remained in the Western Siberian Plain, their present condition is poorly studied. We analysed plant species composition, functional structure and richness of grassland communities and tested for the effect of local factors (management, abiotic site conditions) and landscape factors (patch size, proportion of land cover types) on diversity patterns. Abiotic site conditions, mainly soil moisture and salinity, differentiated distinct community types. Overall, species richness was highest in meadow steppe communities with lower soil moisture and salinity. Grazing intensity and litter accumulation due to cessation of management were significant negative related to species richness and shaped the functional structure. At the landscape scale, diversity in meadow steppe grasslands was higher in forest-grassland mosaics and in small remnants isolated in a matrix of cropland. Our findings highlight that meadow steppes suffered massively under the historical habitat loss and high grazing pressure. Small species-rich remnants are evidence of the former extent of meadow steppe habitats in agricultural landscape, but are likely threatened by an extinction debt. Low intense, irregular mowing maintained species-rich meadow steppe in forest- grassland mosaics, but currently such practices are declining.     

How do sheep affect plant communities and arthropod populations in temperate grasslands?

Grasslands being used in sheep farming systems are managed under a variety of agricultural production, recreational and conservational objectives. Although sheep grazing is rarely considered the best method for delivering conservation objectives in seminatural temperate grasslands, the literature does not provide unequivocal evidence on the impact of sheep grazing on pasture biodiversity. Our aim was therefore to review evidence of the impacts of stocking rate, grazing period and soil fertility on plant communities and arthropod populations in both mesotrophic grasslands typical of agriculturally improved areas and in native plant communities. We therefore conducted a literature search of articles published up to the end of the year 2010 using ‘sheep’ and ‘grazing’ as keywords, together with variables describing grassland management, plant community structure or arthropod taxa. The filtering process led to the selection of 48 articles, with 42 included in the stocking rate dataset, 9 in the grazing period dataset and 10 in the soil fertility dataset. The meta-analysis did not reveal any significant trends for plant species richness or plant community evenness along a wide stocking rate gradient. However, we found frequent shifts in functional groups or plant species abundance that could be explained by the functional properties of the plants in the community. The meta-analysis confirmed that increasing soil fertility decreased plant species richness. Despite the very limited dataset, plant species richness was significantly greater in autumn-grazed pastures than in ungrazed areas, which suggests that choosing an appropriate grazing period would be a promising option for preserving biodiversity in sheep farming systems. Qualitative review indicated that low grazing intensity had positive effects on Orthoptera, Hemiptera (especially phytophagous Auchenorrhyncha) and, despite a diverse range of feeding strategies, for the species richness of Coleoptera. Lepidoptera, which were favoured by more abundant flowering plants, also benefited from low grazing intensities. Spider abundance and species richness were higher in ungrazed than in grazed pastures. In contrast, there are insufficient published studies to draw any firm conclusions on the benefits of late grazing or stopping fertilization on insect diversity, and no grounds for including any of this information in decision support tools at this stage.     

The significance of habitat continuity and current management on the compositional and functional diversity of grasslands in the uplands of Lower Saxony,Germany

There is a growing concern that land use intensification is having negative effects on semi-natural grasslands and that it leads to a general loss of biodiversity among all types of formerly extensively managed grasslands of poor to medium nutrient richness. Since the 1950s, many Central European uplands have been subject to an increase in grassland cover as a result of changes in land use practices. Using such a landscape in Lower Saxony, Germany, as a model region, we assessed environmental factors that control grassland diversity, including plant community composition, species richness and pollination trait composition. In 2007, 189 vegetation sampling sites were randomly distributed among grasslands covering some 394 ha within a 2500 ha study area. Plant communities were classified using TWINSPAN and the effects of environmental factors (soil, topography, current management and habitat continuity) were analysed by canonical correspondence analysis and regression analysis reducing for the effects of spatial autocorrelation by using principal coordinates of neighbour matrices.We found a wide range of six species-poor (<15 plant spp.) to extremely species-rich (>27 spp.) grassland types under mesic to dry site conditions, including sown, Cynosurion, Arrhenatherion and semi-natural grasslands. Grassland community composition was best explained by soil factors and species richness and pollination type composition by combined effects of current management and habitat continuity. During the 1950/60s, the extent of grassland area within the studied landscape rapidly increased to more than double its previous extent, and in 2007, grasslands comprised 16%. Natura 2000 grassland types comprised 1% of the surveyed site and medium-rich, high-nature-value grasslands a further 5%. While the number of wind-pollinated plant species was equal among all grassland types, there was a parallel decline in insect-pollinated plants and overall median species richness in the grassland communities along a gradient of increasing land use intensity (mowing, nutrient supply). Moreover, insect-pollinated plants occurring in intensively managed grasslands were found to additionally have the ability for self-pollination. Species-rich grasslands – including semi-natural grasslands and a semi-improved, species-rich Arrhenatherion community – occurred exclusively on old sites (with >100 years of habitat continuity) that had been used for traditional sheep grazing (environmental contracting). Medium-rich Arrhenatherion grasslands were established primarily on less productive, formerly arable fields (<30 years). We conclude that conservation efforts should focus on extant species-rich grassland types and should aim to implement traditional land use practices such as sheep grazing. Additional restoration efforts should focus on establishing new grasslands on less productive sites in the proximate surroundings of species-rich grasslands to facilitate seed dispersal, but nitrogen deposition should be buffered where appropriate. These measures would enhance the interaction between nature reserves and agricultural grasslands and thus improve the ecological quality of grasslands at the landscape scale.     

Bottom–up and top–down forces structuring consumer communities in an experimental grassland

Synthesis The interplay between bottom‐up and top‐down effects is certainly a general manifestation of any changes in both species abundances and diversity. Summary variables, such as species numbers, diversity indices or lumped species abundances provide too limited information about highly complex ecosystems. In contrast, species by species analyses of ecological communities comprising hundreds of species are inevitably only snapshot‐like and lack generality in explaining processes within communities. Our synthesis, based on species matrices of functional groups of all trophic levels, simplifies community complexity to a manageable degree while retaining full species‐specific information. Taking into account plant species richness, plant biomass, soil properties and relevant spatial scales, we decompose variance of abundance in consumer functional groups to determine the direction and the magnitude of community controlling processes. After decades of intensive research, the relative importance of top–down and bottom–up control for structuring ecological communities is still a particularly disputed issue among ecologists. In our study, we determine the relative role of bottom–up and top–down forces in structuring the composition of 13 arthropod functional groups (FG) comprising different trophic consumer levels. Based on species‐specific plant biomass and arthropod abundance data from 50 plots of a grassland biodiversity experiment, we quantified the proportions of bottom–up and top–down forces on consumer FG composition while taking into account direct and indirect effects of plant diversity, functional diversity, community biomass, soil properties and spatial arrangement of these plots. Variance partitioning using partial redundancy analysis explained 21–44% of total variation in arthropod functional group composition. Plant‐mediated bottom–up forces accounted for the major part of the explainable variation within the composition of all FGs. Predator‐mediated top–down forces, however, were much weaker, yet influenced the majority of consumer FGs. Plant functional group composition, notably legume composition, had the most important impact on virtually all consumer FGs. Compared to plant species richness and plant functional group richness, plant community biomass explained a much higher proportion of variation in consumer community composition.     

Soil pH and phosphorus drive species composition and richness in semi-natural heathlands and grasslands unaffected by twentieth-century agricultural intensification

Background: Increased soil phosphorus (P) caused by agricultural intensification has been associated with decreased plant species richness (SR) in central Europe. How plant communities and soil P gradients are related in unimproved open habitats remains unclear.

Aims: The aim of this article was to characterise the relationship between soil chemical parameters and plant species composition and richness in unimproved open habitats.

Methods: The influence of soil chemical parameters (pH, P, K, Mg) on species composition was assessed, using data from 40 heathland and 54 grassland plots, by non-metric multidimensional scaling and permutational multivariate analysis of variance. The relationship between soil chemical parameters and SR was tested by linear mixed effects models.

Results: A direct relationship between heathland community composition and pH was observed, explaining 10% of variation in species composition, while P, Mg and pH together explained 17% of variation in grassland composition. In heathlands, SR increased with increasing pH, whereas in grasslands, SR decreased with increasing soil P.

Conclusions: Soil chemical parameters were substantially related to plant community composition and richness. In an area spared from a century of agricultural intensification, reduced pH appeared to constrain SR in heathlands, while even slight P increases (<10 mg kg^?1) depressed plant SR in semi-natural grasslands.     

Sod cutting and soil biota effects on seedling performance

Sod cutting (i.e. top soil removal) is a restoration management option for enhancing seedling establishment and for lowering the nutrient concentration in eutrophicated soils of nutrient-poor species-rich grasslands. Removal of the upper soil changes not only abiotic soil properties but may also affect the resident soil community. We investigated the effects of sod cutting on the establishment and performance of two endangered plant species (Cirsium dissectum and Succisa pratensis) while simultaneously manipulating the interaction between seedlings and soil biota. In intact grassland and sod-cut areas at two localities, seedlings were grown in plastic tubes. Half of the tubes had a filter that excluded roots but allowed entry of fungal hyphae and soil microorganisms. The other tubes were closed (i.e. no contact with the surrounding soil). In a greenhouse experiment we studied the effect of soil solutions (with or without fungal tissue) from three grasslands and three sod-cut areas on seedling growth. Sod cutting had a positive net effect on seedling growth for S. pratensis. Access to (mycorrhizal) fungi and other soil biota resulted in a negative impact on seedling growth of both plant species, both in grassland and sod-cut areas. The greenhouse experiment confirmed that the soil biota in these meadows reduced seedling growth. Although sod cutting did not mitigate negative plant-soil feedback, it enhanced seedling growth, presumably by decreasing competition for light. Sod cutting is therefore very useful when seedling establishment needs to be stimulated.     

Direct and indirect effects of land‐use intensification on ant communities in temperate grasslands

Land‐use intensification is a major driver of local species extinction and homogenization. Temperate grasslands, managed at low intensities over centuries harbored a high species diversity, which is increasingly threatened by the management intensification over the last decades. This includes key taxa like ants. However, the underlying mechanisms leading to a decrease in ant abundance and species richness as well as changes in functional community composition are not well understood. We sampled ants on 110 grassland plots in three regions in Germany. The sampled grasslands are used as meadows or pastures, being mown, grazed or fertilized at different intensities. We analyzed the effect of the different aspects of land use on ant species richness, functional trait spaces, and community composition by using a multimodel inference approach and structural equation models. Overall, we found 31 ant species belonging to 8 genera, mostly open habitat specialists. Ant species richness, functional trait space of communities, and abundance of nests decreased with increasing land‐use intensity. The land‐use practice most harmful to ants was mowing, followed by heavy grazing by cattle. Fertilization did not strongly affect ant species richness. Grazing by sheep increased the ant species richness. The effect of mowing differed between species and was strongly negative for Formica species while Myrmica and common Lasius species were less affected. Rare species occurred mainly in plots managed at low intensity. Our results show that mowing less often or later in the season would retain a higher ant species richness—similarly to most other grassland taxa. The transformation from (sheep) pastures to intensively managed meadows and especially mowing directly affects ants via the destruction of nests and indirectly via loss of grassland heterogeneity (reduced plant species richness) and increased soil moisture by shading of fast‐growing plant species.     

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

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

Habitat type plays a greater role than livestock grazing in structuring shrubsteppe plant–pollinator communities

Livestock grazing is a widespread grassland disturbance and can negatively impact biodiversity. Pollinators constitute a vital component of grassland ecosystems, but the impact of grazing on pollinator diversity has seldom been evaluated in North America. We assessed vegetation structure, and pollinator and flowering plant abundance, richness, diversity, and community composition in four pairs of spring-grazed/ungrazed sites in south-central British Columbia, Canada. We also investigated whether pollinator or floral communities differed between the two threatened shrubsteppe habitat types we sampled—antelope-brush and big sagebrush shrubsteppe. Pan-trapping surveys captured 5907 bees, flies, beetles, wasps and butterflies constituting 253 species. We found that the percent cover of shrubs and bare soil increased with grazing, while the height of grasses and forbs decreased. In contrast, pollinator and flowering plant abundance, richness, diversity, and community composition were not significantly affected by grazing. Flowering plant and pollinator community composition did differ significantly between shrubsteppe habitats. Our results indicate that grasslands in North America, when managed responsibly, can maintain pollinator and flowering plant diversity under grazing pressure. The continued effort of land managers to balance ecological integrity and economic viability will be important for the conservation of grassland pollination systems.     

Successful restoration of abandoned terraced vineyards and grasslands in Southern Switzerland

Extensively managed semi-natural grasslands represent species-rich habitats and therefore play a key role for the maintenance of biodiversity in agricultural areas. In marginal and poorly accessible areas, the traditional management of grassland is frequently abandoned, which leads to the spread of forest. In Southern Switzerland, terraced vineyards (a special grassland type) and terraced grasslands are part of the cultural heritage and local biodiversity hotspots. Yet, many of them are overgrown by forest. In the past years, several abandoned terraced vineyards and grasslands have been restored by removing the forest, rebuilding the walls and re-introducing the traditional management. We examined restoration success by assessing plant species richness, diversity and species composition in both the aboveground vegetation and soil seed bank in (1) restored, (2) abandoned for 25–50 years, and (3) permanently used areas of six terraced vineyards and six terraced grasslands. Plant species richness and diversity were reduced and species composition altered in the aboveground vegetation of abandoned vineyards and grasslands compared to the permanently used and restored ones. However, species richness, Shannon-diversity and species composition of the aboveground vegetation did not differ between restored and permanently used areas, indicating a successful restoration of the vegetation 10–15 years after restoration. In abandoned vineyards, species richness of plants emerging from the soil seed bank was slightly higher than in permanently used and restored vineyards. No difference in seedling species richness was found between abandoned, permanently used and restored terraced grasslands. Our results showed that the soil seed bank played a minor role for the re-establishment of the above-ground vegetation. We assume that the large species pool in the surroundings and the presence of dispersal vectors are essential for the successful passive restoration of abandoned grassland in this region.     

Patterns of invasion of an urban remnant of a species-rich grassland in southeastern Australia by non-native plant species

Abstract. The invasion by non-native plant species of an urban remnant of a species-rich Themeda triandra grassland in southeastern Australia was quantified and related to abiotic influences. Richness and cover of non-native species were highest at the edges of the remnant and declined to relatively uniform levels within the remnant. Native species richness and cover were lowest at the edge adjoining a roadside but then showed little relation to distance from edge. Roadside edge quadrats were floristically distinct from most other quadrats when ordinated by Detrended Correspondence Analysis. Soil phosphorus was significantly higher at the roadside edge but did not vary within the remnant itself. All other abiotic factors measured (NH₄, NO₃, S, pH and % organic carbon) showed little variation across the remnant. Non-native species richness and cover were strongly correlated with soil phosphorus levels. Native species were negatively correlated with soil phosphorus levels. Canonical Correspondence Analysis identified the perennial non-native grasses of high biomass as species most dependent on high soil nutrient levels. Such species may be resource-limited in undisturbed soils. Three classes of non-native plants have invaded this species-rich grassland: (1) generalist species (> 50 % frequency), mostly therophytes with non-specialized habitat or germination requirements; (2) resource-limited species comprising perennial species of high biomass that are dependent on nutrient increases and/or soil disturbances before they can invade the community and; (3) species of intermediate frequency (1–30 %), of low to high biomass potential, that appear to have non-specialized habitat requirements but are currently limited by seed dispersal, seedling establishment or the current site management. Native species richness and cover are most negatively affected by increases in non-native cover. Declines are largely evident once the non-native cover exceeds 40 %. Widespread, generalist non-native species are numerous in intact sites and will have to be considered a permanent part of the flora of remnant grasslands. Management must aim to minimize increases in cover of any non-native species or the disturbances that favour the establishment of competitive non-native grasses if the native grassland flora is to be conserved in small, fragmented remnants.     

Species diversity of remnant calcareous grasslands in south eastern Germany depends on litter cover and landscape structure

Species diversity depends on, often interfering, multiple ecological drivers. Comprehensive approaches are hence needed to understand the mechanisms determining species diversity. In this study, we analysed the impact of vegetation structure, soil properties and fragmentation on the plant species diversity of remnant calcareous grasslands, therefore, in a comparative approach.We determined plant species diversity of 18 calcareous grasslands in south eastern Germany including all species and grassland specialists separately. Furthermore, we analysed the spatial structure of the grasslands as a result of fragmentation during the last 150 years (habitat area, distance to the nearest calcareous grassland and connectivity in 1830 and 2013). We also collected data concerning the vegetation structure (height of the vegetation, cover of bare soil, grass and litter) and the soil properties (content of phosphorous and potassium, ratio of carbon and nitrogen) of the grassland patches. Data were analysed using Bayesian multiple regressions.We observed a habitat loss of nearly 80% and increasing isolation between grasslands since 1830. In the Bayesian multiple regressions the species diversity of the studied grasslands depended negatively on cover of litter and to a lower degree on the distance to the nearest calcareous grassland in 2013, whereas soil properties had no significant impact.Our study supports the observation that vegetation structure, which strongly depends on land use, is often more important for the species richness of calcareous grasslands than fragmentation or soil properties. Even small and isolated grasslands may, therefore, contribute significantly to the conservation of species diversity, when they are still grazed.     

Earthworms promote greater richness and abundance in the emergence of plant species across a grassland-forest ecotone

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.     

Role of Soil Organisms in the Maintenance of Species-Rich Seminatural Grasslands through Mowing

To preserve species-rich grasslands, management practices such as mowing are often required. Mowing is known to promote aboveground conditions that help to maintain plant species richness, but whether belowground effects are important as well is not known. We hypothesized that if mowing decreases belowground carbon transfer by reducing root mass, this will reduce the abundance and activity of soil decomposers and lead to diminished nutrient availability in soil. In grasslands, this would provide a means to mitigate the negative effects of nitrogen enrichment on plant species richness. We established experimental plots on grassland with one-third of plots growing untouched, one-third mowed once a summer, and one-third mowed twice a summer for three growing seasons. Root and soil attributes were measured at each plot 1 month after each mowing. Mowing decreased root mass but had few effects on belowground biota and soil NH₄-N, NO₃-N, and PO₄-P concentrations. Mowing did not affect root N concentrations. Our results show that despite reducing root mass, mowing may have few effects on those soil biota that control nutrient supply and, as a result, have no clear-cut effects on nutrient availability in soil. This suggests that the effects of mowing on soil decomposers and soil nutrient availability may not provide an effective means to mitigate N enrichment and enhance plant species richness in grasslands in a timescale of a few years. Whether such effects could, however, be achieved with longer lasting mowing remains open.     

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

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