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Habitat restoration requires realistic goals. To naturally regenerate European lowland calcareous grassland, whose extent has severely declined, over a century may be required for vegetation to become indistinguishable from that of old calcareous grassland. Progress of natural regeneration can be characterized using member species of the reference vegetation as indicators of favourable site condition. Chronosequence studies have suggested that calcareous-grassland species differ predictably in their ability to colonize ex-arable land, with some usually colonizing early on, and others in later stages. If such patterns are affected by gradually-attenuating establishment limitation, this would have important implications for restoration practice and indication of progress. Particularly, late-colonizing species might be better indicators of favourable site conditions than early colonizers. To explore these aspects, we have reanalysed chronosequence data previously used to investigate causal mechanisms affecting calcareous-grassland restoration progress. We carried out an indicator species analysis to determine which species are indicative of particular stages of natural regeneration. Using correlation analyses, we tested whether species colonization patterns matched those found by previous chronosequence studies that were geographically more limited or relied on more informal approaches to determine species order of colonization. Correlation analyses were also used to test whether order of colonization could be explained by establishment limitation or by dispersal limitation, or by established plant strategies that underlie such limitations. We identified 30 species as indicative of particular stages of natural regeneration, including nine that specifically indicate old calcareous grassland. Correlation results confirmed high congruence with species order of colonization in previous chronosequence studies, and indicated that establishment limitation plays a role in shap

Recent demonstrations of the role of plant–soil biota interactions have challenged the conventional view that vegetation changes are mainly driven by changing abiotic conditions. However, while this concept has been validated under natural conditions, our understanding of the long-term consequences of plant–soil interactions for above-belowground community assembly is restricted to mathematical and conceptual model projections. Here, we demonstrate experimentally that one-time additions of soil biota and plant seeds alter soil-borne nematode and plant community composition in semi-natural grassland for 20 years. Over time, aboveground and belowground community composition became increasingly correlated, suggesting an increasing connectedness of soil biota and plants. We conclude that the initial composition of not only plant communities, but also soil communities has a long-lasting impact on the trajectory of community assembly.

The importance of managing land to optimize carbon sequestration for climate change mitigation is widely recognized, with grasslands being identified as having the potential to sequester additional carbon. However, most soil carbon inventories only consider surface soils, and most large-scale surveys group ecosystems into broad habitats without considering management intensity. Consequently, little is known about the quantity of deep soil carbon and its sensitivity to management. From a nationwide survey of grassland soils to 1 m depth, we show that carbon in grassland soils is vulnerable to management and that these management effects can be detected to considerable depth down the soil profile, albeit at decreasing significance with depth. Carbon concentrations in soil decreased as management intensity increased, but greatest soil carbon stocks (accounting for bulk density differences), were at intermediate levels of management. Our study also highlights the considerable amounts of carbon in subsurface soil below 30 cm, which is missed by standard carbon inventories. We estimate grassland soil carbon in Great Britain to be 2097 Tg C to a depth of 1 m, with ~60% of this carbon being below 30 cm. Total stocks of soil carbon (t ha−1) to 1 m depth were 10.7% greater at intermediate relative to intensive management, which equates to 10.1 t ha−1 in surface soils (0–30 cm), and 13.7 t ha−1 in soils from 30 to 100 cm depth. Our findings highlight the existence of substantial carbon stocks at depth in grassland soils that are sensitive to management. This is of high relevance globally, given the extent of land cover and large stocks of carbon held in temperate managed grasslands. Our findings have implications for the future management of grasslands for carbon storage and climate mitigation, and for global carbon models which do not currently account for changes in soil carbon to depth with management.

Soil biodiversity plays a key role in regulating the processes that underpin the delivery of ecosystem goods and services in terrestrial ecosystems. Agricultural intensification is known to change the diversity of individual groups of soil biota, but less is known about how intensification affects biodiversity of the soil food web as a whole, and whether or not these effects may be generalized across regions. We examined biodiversity in soil food webs from grasslands, extensive and intensive rotations in four agricultural regions across Europe: in Sweden, the UK, the Czech Republic and Greece. Effects of land use intensity were quantified based on structure and diversity among functional groups in the soil food web, as well as on community-weighted mean body mass of soil fauna. We also elucidate land use intensity effects on diversity of taxonomic units within taxonomic groups of soil fauna. We found that between regions soil food web diversity measures were variable, but that increasing land use intensity caused highly consistent responses. In particular, land use intensification reduced the complexity in the soil food webs, as well as the community-weighted mean body mass of soil fauna. In all regions across Europe, species richness of earthworms, Collembolans and oribatid mites was negatively affected by increased land use intensity. The taxonomic distinctness, which is a measure of taxonomic relatedness of species in a community that is independent of species richness, was also reduced by land use intensification. We conclude that intensive agriculture reduces soil biodiversity, making soil food webs less diverse and composed of smaller bodied organisms. Land use intensification results in fewer functional groups of soil biota with fewer and taxonomically more closely related species. We discuss how these changes in soil biodiversity due to land use intensification may threaten the functioning of soil in agricultural production systems. This article is protected

This study focuses on the restoration of chalk grasslands over a six-year period and tests the efficacy of two management practices, hay spreading and soil disturbance, in promoting this process for phytophagous beetles. Restoration success for the beetles, measured as similarity to target species-rich chalk grassland, was not found to be influenced by either management practice. In contrast, restoration success for the plants did increase in response to hay spreading management. While the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonise, few beetle species colonised as soon as their host plants became established. Morphological characteristics and feeding habits of 27 phytophagous beetle species were therefore tested to identify factors that limited their colonisation and persistence. The lag time between host plant establishment and colonisation was greatest for flightless beetles. Beetles with foliage-feeding larvae both colonised at slower rates than seed-, stem- or root-feeding species, and persisted within the swards for shorter periods. While the use of hay spreading may benefit plant communities during chalk grassland restoration it did not directly benefit phytophagous beetles. Without techniques for overcoming colonisation limitation for invertebrate taxa short-term success of restoration may be limited to the plants only.

1. Declines in area and quality of species-rich mesotrophic and calcareous grasslands have occurred all across Europe. While the European Union has promoted schemes to restore these grasslands, the emphasis for management has remained largely focused on plants. Here we focus on restoration of the phytophagous beetles of these grasslands. Although local management, particularly that which promotes the establishment of host plants, is key to restoration success, dispersal limitation is also likely to be an important limiting factor during the restoration of phytophagous beetle assemblages. 2. Using a 3-year multi-site experiment, we investigated how restoration success of phytophagous beetles was affected by hay-spreading management (intended to introduce target plant species), success in restoration of the plant communities and the landscape context within which restoration was attempted. 3. Restoration success of the plants was greatest where green hay spreading had been used to introduce seeds into restoration sites. Beetle restoration success increased over time, although hay-spreading had no direct effect. However, restoration success of the beetles was positively correlated with restoration success of the plants. 4. Overall restoration success of the phytophagous beetles was positively correlated with the proportion of species-rich grassland in the landscape, as was the restoration success of the polyphagous beetles. Restoration success for beetles capable of flight and those showing oligophagous host plant specialism were also positively correlated with connectivity to species-rich grasslands. There was no indication that beetles not capable of flight showed greater dependence on landscape scale factors than flying species. 5.Synthesis and applications. Increasing the similarity of the plant community at restoration sites to target species-rich grasslands will promote restoration success for the phytophagous beetles. However, landscape context is also important

Genetic differentiation among plant populations and adaptation to local environmental conditions are well documented. However, few studies have examined the potential contribution of plant antagonists, such as insect herbivores and pathogens, to the pattern of local adaptation.Here, a reciprocal transplant experiment was set up at three sites across Europe using two common plant species, Holcus lanatus and Plantago lanceolata. The amount of damage by the main above-ground plant antagonists was measured: a rust fungus infecting Holcus and a specialist beetle feeding on Plantago, both in low-density monoculture plots and in competition with interspecific neighbours. Strong genetic differentiation among provenances in the amount of damage by antagonists in both species was found. Local provenances of Holcus had significantly higher amounts of rust infection than foreign provenances, whereas local provenances of Plantago were significantly less damaged by the specialist beetle than the foreign provenances. The presence of surrounding vegetation affected the amount of damage but had little influence on the ranking of plant provenances. The opposite pattern of population differentiation in resistance to local antagonists in the two species suggests that it will be difficult to predict the consequences of plant translocations for interactions with organisms of higher trophic levels.

Root herbivores can have a positive or negative effect on the abundance and/or performance of foliar phytophages. In addition, abiotic factors such as drought can either strengthen or weaken this effect, depending on the system under investigation. One explanation for these varying responses lies in differences in the physiological response of host plants to drought and root herbivores. Here, the impacts of root phytophages on a leaf-mining species feeding on annual and perennial plant species (four Sonchus species) were compared. The responses of plants and leaf-miners to drought and root herbivore treatments were not related to whether the host plant was an annual or perennial. However, where root feeders did affect foliar phytophage performance, this occurred only under a drought treatment, demonstrating the potential for climate change to alter the outcome of plant-mediated interactions.

During the last century changing patterns of land use in the UK have led to the large scale loss and degradation of species-rich chalk grasslands, a habitat of conservation importance across Europe. In particular, the conversion of chalk grasslands into arable land has represented a major threat to this habitat type. The re-creation of chalk grasslands on ex-arable land therefore has the potential to both increase the area of this threatened habitat, and to provide links between existing remnant fragments. We present a four year study investigating the effects of three different methods of introducing plant species during grassland re-creation: 1) a simple grass seed mix, intended to suppress the establishment of pernicious weeds; 2) spreading of hay; 3) sowing of brush harvested seeds. The latter two methods introduce local provenance seeds collected from species-rich chalk grassland. Using a replicated block design we investigate the efficacy of these management practices in promoting the re-creation of both plant and phytophagous beetle assemblages. We compare re-creation success relative to a target chalk grassland and contrast differences in the responses between these two trophic levels. The introduction of local provenance seeds by hay spreading and brush harvesting is shown to be beneficial during re-creation. However, if used in combination with simple grass-only seed mixtures the establishment of some chalk grassland species may be reduced. We discuss the relevance of these findings for the re-creation of plant and phytophagous beetle assemblages of species-rich grasslands on ex-arable land. We also consider the implications of differences in establishment rates between these two taxa for the long term success of habitat creation.

Summer droughts are predicted to increase in severity and frequency in the United Kingdom, due to climate change. Few studies have addressed the impacts of drought on interactions between species, and the majority have focussed on increases in CO2 concentration and changes in temperature. Here, the effect of experimental summer drought on the strength of the plant-mediated interaction between leaf-mining Stephensia brunnichella larvae and root-chewing Agriotes larvae was investigated. Agriotes larvae reduced the abundance and performance of S. brunnichella feeding on a mutual host plant, Clinopodium vulgare, as well as the rate of parasitism of the leaf-miner. The interaction did not, however, occur on plants subjected to a severe drought treatment, which were reduced in size. Changes to summer rainfall, due to climate change, may therefore reduce the occurrence of plant-mediated interactions between insect herbivores.

Soil invertebrate communities are likely to be highly vulnerable to low soil moisture, caused by a reduction in summer rainfall which is predicted for some regions under current climate change scenarios. However, the effects of changes in summer rainfall on soil invertebrate assemblages have rarely been tested experimentally. In this study, samples were taken in 2003 and 2004 from a long-running field experiment, to investigate the impact of 10 years of experimental summer drought and increased summer rainfall manipulations on the soil fauna of a calcareous grassland. Summer drought altered the soil invertebrate assemblage in the autumn, immediately following treatment application, but by the following spring treatment effects were no longer apparent. The two most common root herbivore species responded differently to the summer rainfall manipulations. Larvae of the dominant root-chewing species, Agriotes lineatus, were more numerous under enhanced rainfall in both the spring and autumn. In contrast, abundance of the Coccoidea Lecanopsis formicarum was unaffected by the rainfall manipulations. The responses of root herbivores to an increased incidence of summer droughts are therefore likely to vary, depending on their feeding strategy and life history.

Evolutionary theory suggests that divergent natural selection in heterogeneous environments can result in locally adapted plant genotypes. To understand local adaptation it is important to study the ecological factors responsible for divergent selection. At a continental scale, variation in climate can be important while at a local scale soil properties could also play a role. We designed an experiment aimed to disentangle the role of climate and (abiotic and biotic) soil properties in local adaptation of two common plant species. A grass (Holcus lanatus) and a legume (Lotus corniculatus), as well as their local soils, were reciprocally transplanted between three sites across an Atlantic–Continental gradient in Europe and grown in common gardens in either their home soil or foreign soils. Growth and reproductive traits were measured over two growing seasons. In both species, we found significant environmental and genetic effects on most of the growth and reproductive traits and a significant interaction between the two environmental effects of soil and climate. The grass species showed significant home site advantage in most of the fitness components, which indicated adaptation to climate. We found no indication that the grass was adapted to local soil conditions. The legume showed a significant home soil advantage for number of fruits only and thus a weak indication of adaptation to soil and no adaptation to climate. Our results show that the importance of climate and soil factors as drivers of local adaptation is species-dependent. This could be related to differences in interactions between plant species and soil biota.

The impact of climate change on phytophages is difficult to predict, due in part to variation between species in their responses to factors such as drought stress. Here, the hypothesis that several species within the leaf-mining feeding guild will respond in a consistent way to changes in rainfall patterns is tested, using a manipulative field experiment. Summer drought, enhanced summer rainfall, and control treatments were imposed on a calcareous grassland community, and the responses of five leaf-mining species were assessed. One leaf-mining species was more abundant under enhanced rainfall, one was more abundant under drought, and the other three species showed no consistent response to the rainfall treatments. Higher parasitism levels under drought may partly explain the response of one species (Stephensia brunnichella) to the treatments. These results show that generalisations relating to drought stress impacts cannot be drawn at the feeding guild level for leaf-mining insects.