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.     

Soil properties, grassland management, and landscape diversity drive the assembly of earthworm communities in temperate grasslands

Earthworms are widespread soil organisms that contribute to a wide range of ecosystem services. As such, it is important to improve our knowledge, still scanty, of the factors that drive the assembly of earthworm communities. The aim of the present study was to conjointly evaluate the effects on the assembly of earthworm communities of i) soil properties (texture, organic matter content, and pH), ii) grassland management (grassland age, livestock unit, and type of fertilization), iii) landscape diversity (richness, diversity of surrounding habitats, and grassland plant diversity), and iv) presence of hedgerows. The study was conducted in temperate grasslands of Brittany, France. Earthworms were sampled in 24 grasslands and, in three of these grasslands, they were sampled near a hedgerow or near a ditch (control without a hedgerow). Soil properties explained the larger portion of the variation in the earthworm community parameters compared to grassland management or landscape diversity. The increase in soil organic matter content and pH were the most favorable factors for earthworm abundance and biomass, in particular for endogeic species. Regarding grassland management, the increase in the livestock unit was the most damaging factor for earthworm communities, in particular for the anecic earthworm biomass and endogeic species richness. Surprisingly, landscape diversity negatively affected the total earthworm abundance and epigeic earthworm biomass, but it was related to an increase in the epi-anecic species. At a finer scale, we also demonstrated that the presence of hedgerows surrounding grasslands enhanced earthworm species richness, especially within the epigeic and anecic ecological categories. This study highlights that the earthworm ecological categories respond specifically to environmental filters; further studies need to be conducted to elucidate the factors that drive the assembly of earthworm communities at this ecological category level. We recommend that policymakers should act on landscape management to favor earthworm diversity in order to improve the ecosystem services they drive.     

Grassland management influences spatial patterns of soil microbial communities

Soil micro-organisms play a vital role in grassland ecosystem functioning but little is known about the effects of grassland management on spatial patterns of soil microbial communities. We compared plant species composition with terminal restriction fragment length polymorphism (T-RFLP) fingerprints of soil bacterial and fungal communities in unimproved, restored and improved wet grasslands. We assessed community composition of soil micro-organisms at distances ranging from 0.01 m to 100 m and determined taxa–area relationships from field- to landscape level. We show that land management type influenced bacterial but not fungal community composition. However, extensive grassland management to restore aboveground diversity affected spatial patterns of soil fungi. We found distinct distance–decay and small-scale aggregation of fungal populations in extensively managed grasslands restored from former arable use. There were no clear spatial patterns in bacterial communities at the field-scale. However, at the landscape level there was a moderate increase in bacterial taxa and a strong increase in fungal taxa with the number of sites sampled. Our results suggest that grassland management affects soil microbial communities at multiple scales; the observed small-scale variation may facilitate plant species coexistence and should be taken into account in field studies of soil microbial communities.     

The influence of plant diversity on people’s perception and aesthetic appreciation of grassland vegetation

The conservation of biodiversity critically depends on the value that humans attach to it. Apart from an ecological and economic value, an aesthetic value has often been assigned to biodiversity. However, it is not known whether lay people appreciate the diversity of species and not just certain individual species or nature as a whole. We studied in a series of experiments and field studies people’s perception and appreciation of species diversity. We presented meadow-like arrays of different species richness and evenness but random species composition to lay people and asked them to estimate plant species richness and rank the arrays by attractiveness. The experiments were complemented by two larger-scale field studies using natural meadows. Both in the experiments and the field studies the mean perception of species richness by people increased with true species richness, but was slightly overestimated at low and increasingly underestimated at high diversity levels. Lay people’s aesthetic appreciation of both the experimental grassland arrays and the natural meadows increased with true species richness. Communities consisting of the same number of species were perceived to be more species-rich and were appreciated more when their evenness was high. Our results demonstrate that plant diversity in itself is attractive to humans. The current reduction of the diversity of grasslands due to intensification of management may thus reduce the attractiveness of regions where grasslands are a dominant feature of the landscape. This could have negative consequences for tourism and may add an economic argument for the conservation of biodiversity in grasslands.     

Evaluating patterns of biodiversity in managed grasslands using spatial turnover metrics

Market and policy incentives that encourage agricultural intensification, such as incentives for bioenergy, may contribute to biodiversity decline when they encourage a large-scale conversion of native and semi-natural ecosystems to production fields. In order to appreciate the impact of these incentives on biodiversity, it is imperative to better understand how native and semi-natural ecosystems contribute to plant diversity and composition. We studied the five most common types of managed grasslands in Northeastern Kansas, a region undergoing agricultural intensification. We analyzed plant community data recorded at three spatial scales in 98 managed grassland sites, and examined patterns of diversity and composition of plant species and functional groups; and spatial turnover of species and functional groups measured at different spatial scales. We found reduced soil quality and plant diversity as well as a lack of scale-dependent community patterns on sites that were historically cultivated. Forage management practices (haying or grazing) altered plant diversity and composition in native grassland remnants but not in non-native grasslands. We identified several opportunities where changes to existing management practices could benefit both conservation and bioenergy production objectives. Recommendations to conserve biodiversity include increasing the use of hay management or other biomass collection on native grassland remnants and improving the Conservation Reserve Program by increasing enrollment, adding more native species to seed mixes, and incorporating a periodic biomass collection. We also found that using measures of spatial turnover in community composition added important insights in understanding the effects of management decisions on biodiversity.     

Vegetation development of sown grass margins in arable fields under a countrywide agri-environment scheme

Farmers were paid to establish 6 m wide sown grass strips in arable field margins under the Countryside Stewardship Scheme in England, UK. One hundred and sixteen sites in eight regions were surveyed to determine if grass margins had been established successfully and the extent to which they were colonised by forbs. Sown margins had more grass and fewer weed species than naturally regenerated sites. Grass margins contrasted with normally cropped sites, having greater species richness of grasses, forbs and perennials and more bird, butterfly larva and bumblebee foodplants. Mesotrophic grassland forbs were scarce in margins established from basic grass seed mixtures but significantly more abundant if included in the seed mixture. Annuals were more prevalent in sites up to 2 years old but species composition was not related to age in sites over 2 years old. Variation partitioning showed that overall species composition was related to seed mixture type, region and soil properties but there was little overlap in the variation explained by these environmental variable sets. Habitat context and management practices did not explain any variation in species composition. Perennial grassy vegetation was established successfully using basic grass seed mixes but only competitive species colonised subsequently. Diverse seed mixtures containing mesotrophic grassland forbs merit support in agri-environment schemes because they do enhance the botanical diversity of sown grass margins.     

Grazing as a management tool in dune grasslands: Evidence of soil and scale dependence of the effect of large herbivores on plant diversity

In nature management, the introduction of large herbivores into human-influenced grasslands is thought to be effective to maintain or enhance plant diversity. In order to test the validity of this assumption, we studied the effect of grazing by large herbivores on plant species richness and community heterogeneity across a soil acidity gradient at different spatial scales in dry coastal dune grasslands in western Belgium and north-western France. The effect of grazing on plant richness varied with scale and soil acidity. Grazing had a predominantly positive effect on plant species richness in all habitats at the small scale (0.25 × 0.25 m). However, at site scale (8 × 8 m) it had only positive effects in grasslands with higher soil pH (6-7.4). Similarly, grazing resulted in a homogenization of grassland vegetation at lower pH, while heterogeneity increased with grazing on soil with higher pH. In general, grazing increased the number of rare species, independent of soil pH. The results confirm that the impact of grazing on plant diversity depends on the scale considered and that the effects further depend on soil acidity which was correlated to biomass production at the given soil pH range in this study. Although grazing seems an appropriate management tool to maintain and even enhance plant biodiversity under many circumstances, it may negatively affect plant species richness, where soil resources limit plant biomass production.     

Secondary succession of terrestrial isopod,centipede, and millipede communities in grasslands under restoration

The aim of this study was to investigate the impact of restoration management on the composition of a macro-invertebrate community in a formerly, nutrient-poor grassland. Four grassland plots were selected that were last fertilised 7, 11, 24 or 29 years before sampling in 1996. In the same plots it was observed that nutrient impoverishment as a restoration tool resulted in a decrease in primary production and a directional shift in vegetation composition after cessation of fertiliser application. Terrestrial isopods, millipedes, and centipedes were sampled with pitfall traps in the four plots. The directional shift observed in vegetation composition before this study was not accompanied by a directional change in macro-invertebrate composition. Both the field poorest in nutrients and the one richest in nutrients showed the lowest density and species richness, while the species composition was similar across intermediate succession stages. By far the most specimens and species were caught in the field that had not received fertilisers for 24 years. Succession theory could only partly explain the observed results. Canonical correspondence analysis of the data revealed that only a small part of the pattern could be explained by the nutrient status of the grasslands. The C accumulation due to secondary succession of plants was hypothesised to influence the densities and diversity of macro-invertebrate communities in these grasslands.     

The importance of the gravel excavation industry for the conservation of grassland butterflies

Conservation biology often relies on the protection of (semi)natural habitat remnants. However, the ever increasing human population is taking over natural resources and habitats. Here, contrary to most other studies, we ask how human-associated severe changes in the environment can be used to enrich local biodiversity. We tested if industrial activity (gravel excavation) leads to the creation of habitats that support grassland butterflies and how these areas add to the richness of local species when compared to typical semi-natural habitats (grasslands). We also identified key factors affecting the richness, abundance, diversity and commonness of butterfly species to provide practical recommendations. Species richness, diversity index and the occurrence of rare species were higher in gravel-pit shores than in grasslands. The richness of butterfly species and their abundance were positively affected by the richness of plant species, shrub density and age of the gravel-pit but negatively by the cover of water reservoirs in the surrounding area and the isolation of gravel-pits from grasslands. Butterfly diversity was positively influenced by the richness of plant species and proximity of human settlement but negatively by area of the shore and isolation. Our study is the first one to show the high value of gravel-pits for the conservation of butterflies. We recommend the inclusion of gravel-pits in a system of ecological networks and management of their surroundings to improve the colonization rate of rare species. We suggests that directing interest to the possible positive effects of industrial development on biodiversity may support conservation efforts.     

Remnant grassland habitats as source communities for plant diversification in agricultural landscapes

Lately there has been a shift in Sweden from grazing species-rich semi-natural grasslands towards grazing ex-arable fields in the modern agricultural landscape. These fields normally contain a fraction of the plant species richness compared to semi-natural grasslands. However, small remnant habitats have been suggested as important for plant species diversity and conservation as they may function as refugia for grassland specialists in fragmented and highly modified agricultural landscapes. In this study, we examined whether plant communities on small remnant habitats, i.e. midfield islets, can function as sources for grassland species to disperse out into surrounding grazed ex-fields (former arable fields). We examined species richness and grassland specialists (species favoured by grazing) and their ability to colonize fields after 5 and 11 years of grazing. The fields that had been grazed for a shorter time were fairly species-poor with few grassland specialists. A longer period of grazing had a positive effect on total and small-scale species diversity in both islets and fields. Species composition became more similar with time, and the number of grassland specialists in both habitats increased. We found that grassland specialists dispersed step-wise into the fields, and the number of grassland specialists decreased with distance from the source. Our study suggests that remnant habitats, such as midfield islets, do function as a source community for grassland specialists and enhance diversification of grassland species when grazing is introduced. For long-term conservation of plant species, incorporating small refugia into larger grazing complexes may thus enhance species richness.     

Are spiders reacting to local or landscape scale effects in Hungarian pastures?

Declines of West-European farmland biodiversity have been associated with intensive agricultural practices, Central and Eastern European grasslands still harbour a diverse and unique arthropod community. However, our understanding of the effects of farmland management both at local and landscape levels is rather limited there. A paired field approach was used to compare extensively (0.5 cows/ha) and intensively (>1 cows/ha) grazed pastures in 42 fields in three distinct biogeographic regions within Hungary. Spiders belonging to the hunting and web-building communities were sampled using funnel traps. We found no management effect either on richness and abundance or on species composition, which shows that both forms of grazing management at the intensity levels studied support valuable spider fauna. At the local scale plant and litter cover were the two most important variables that significantly affected the communities overall, meaning both the hunting and to some degree the web-building communities. No significant landscape effects were found in the analyses on spider richness and abundance, but community structure was affected by two landscape level factors (grassland patch density and grassland percentage). We suggest that to protect the valuable spider and other fauna of these regions, the recently launched national agri-environmental program should be further supported and enlarged to maintain and reintroduce the traditional grazing management on these semi-natural grasslands.     

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.     

Hay strewing, brush harvesting of seed and soil disturbance as tools for the enhancement of botanical diversity in grasslands

Since the middle of the last century agricultural intensification within Europe has led to a drastic decline in the extent of botanically diverse grasslands. Whilst measures to enhance the diversity of agriculturally-improved grasslands are in place, success has often been limited. One of the primary factors limiting success is the paucity of sources of propagules of desirable species in the surrounding landscape. The restoration of two contrasting grassland types (lowland hay meadow and chalk grassland) was examined using a replicated block experiment to assess the effectiveness of two methods of seed application (hay strewing and brush harvesting) and two methods of pre-treatment disturbance (power harrowing and turf stripping). The resulting changes in botanical composition were monitored for 4 years. Seed addition by both methods resulted in significant temporal trends in plant species composition and increases in plant species richness, which were further enhanced by disturbance. Power harrowing increased the effectiveness of the seed addition treatments at the lowland hay meadow site. At the chalk grassland site a more severe disturbance created by turf stripping was used and shown to be preferable. Whilst both hay strewing and brush harvesting increased plant species richness, hay strewing was more effective at creating a sward similar to that of the donor site. Soil disturbance and seed application rate at the recipient site and timing of the hay cut at the donor site are all factors to be considered prior to the commencement of restoration management.     

Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment

Calcareous grasslands, most of which are man made and therefore depend on some kind of human interference for their maintenance, are among the most species-rich communities on Earth at a small scale. For many centuries, most of these grasslands have been used as extensive pasture. However, after 1900, and particularly from 1940 onwards, livestock grazing has declined throughout Europe leading to the abandonment of low intensity grasslands over large areas. To conserve the remaining grasslands or to restore recently abandoned grasslands, better insights about the effects of grassland management on above and belowground species diversity are needed. Here, we describe the results of an 11-year experiment to investigate the role of grassland management (grazing, mowing and abandonment) in determining species composition and diversity both in the aboveground vegetation and the seed bank of a calcareous grassland in Belgium. Species diversity declined by about 60% 11 years after abandonment, from 29 species m⁻² to as few as 12 species m⁻². Plots that were grazed remained constant in species richness, whereas mown sites lost about 20% of their original species. Abandoned plots were largely dominated by a few grass species, in particular Festuca rubra. Concomitant with changes in the aboveground vegetation, both the number of species found in the seed bank and seed density (number of seeds m⁻²) had changed significantly 11 years after abandonment. Species diversity and seed density were significantly lower in abandoned plots than in grazed or mown plots. We conclude that abandonment of calcareous grasslands may lead to rapid decline of plant species diversity both in the aboveground vegetation and in the seed bank. As a result, seed banks probably have a limited role to play in the restoration of recently abandoned grasslands.     

Are higher taxa adequate surrogates for species-level assemblage patterns and species richness in stream organisms?

We examined the utility of using genera and families as surrogates of species-level assemblage patterns. We assessed cross-taxonomic level congruence, as well as taxonomic structure-environment and taxon richness-environment relationships in stream macroinvertebrate and diatom data surveyed across Finland. A number of interesting and important findings emerged from our analyses. For both macroinvertebrates and diatoms, species richness was strongly correlated with genus and family richness, and similarities in assemblage structure between species-, genus- and family-level matrices were also significant and relatively strong. For macroinvertebrates, the key environmental variables for assemblage structure and taxon richness remained the same irrespective of taxonomic level. For diatoms, however, the environmental variables related to variation in diatom taxon richness varied between taxonomic data sets, while determinants of assemblage structure remained largely the same. For macroinvertebrates, relatively similar amounts of variation in assemblage structure and taxon richness for species-, genus-, and family-level data were accounted for by the environmental variables, shared environmental and spatial variables, spatial variables, and total variation explained. By contrast, for diatoms, these components of variability differed somewhat between taxonomic levels. Our results suggest that, for macroinvertebrates, even family-level data could be used in rapid biodiversity assessments, where the objective is to examine biodiversity patterns and rank sites according to their conservation value in our study region. Families might also perform well in this task in other regions that are characterized by low faunal species diversity, but the situation is likely to be different for regions with higher species diversity. Although weaker than the relationships in macroinvertebrate data, species-level data for diatoms also showed rather similar patterns to genus- and family-level data. However, due to the low number of diatom families, it is likely that families do not provide enough variability in biodiversity patterns. Thus, for diatoms, genus-level data could perhaps be more efficient than family-level data in correlating with species-level data and portraying assemblage patterns for conservation purposes.     

The influence of field shape, area and surrounding landscape on plant species richness in grazed ex-fields

Over the past 100 years species-rich semi-natural grasslands have decreased dramatically in Western Europe, where former arable fields (ex-fields) are used instead as pasture. The disappearance of semi-natural grasslands has caused a threat to the biodiversity in agricultural landscapes. Many typical grassland plants are dispersal limited, thus grazed ex-fields can be used to investigate if species spontaneously colonise these new grassland habitats. We examined the relationship between surrounding landscape, field area, shape, distance between edge and centre, and plant species diversity in ex-fields that had been grazed for 15-18 years. The results showed that there were 35% more plant species in fields surrounded by commercial forestry production compared to those surrounded by open agricultural landscape. Area and shape did not influence species richness, although there was increasing number of species in the centre with decreasing distance from the edge. Twenty-five percent of the species where typical grassland species, and ex-fields surrounded by forest had 91% more grassland species compared to those in the open landscape. It is possible to increase grassland plant occurrences by grazing ex-fields surrounded by forest or other grassland remnant habitats, particularly in landscapes where grazed semi-natural grasslands are scarce.     

东北平原典型玉米种植区农业景观植物多样性研究——以昌图县为例

非耕作生境是农业景观中的重要组成部分,为生物提供食物和栖息地,对生物多样性保护具有重要意义。本文以辽宁省昌图县为例,选取了20个非耕作生境比例为0~50%的网格(1 km?1 km)为样区,采用典型样地法对调查区域内农业景观中的林地、田间路、草地、沟渠、果园5类主要非耕作生境中的植物群落进行调查,探讨不同非耕作生境类型和比例中植物多样性及其与景观异质性之间的关系。研究结果表明,随着非耕作生境比例增加,植物物种多样性和均匀度先增加后减小,在20%~30%时达到峰值;非耕作生境比例在0~40%范围内时,植物物种丰富度同样呈现先增加后减小,且在20%~30%时达到峰值;非耕作生境比例高于40%时,植物物种丰富度再次出现峰值且高于20%~30%。随着非耕作生境比例的增加,农业景观香浓多样性、香浓均匀度指数和景观斑块形状复杂程度逐渐增加,蔓延度恰好相反。不同生境中重要值最高的均为禾本科植物,但植物种不同;优势种优势度为果园沟渠草地田间路林地。果园、草地和沟渠的物种多样性和均匀度指数显著高于林地和田间路,丰富度指数为草地林地沟渠果园田间路,且草地、果园和田间路的不同调查区域间植物群落指数存在较大的变异性。草地、果园、沟渠、田间路和林地中景观异质性与物种多样性相关性顺次降低。研究结论表明,东北平原典型玉米种植区农业景观中,最适宜非耕作生境比例为20%~30%;非耕作生境中林地、沟渠对物种多样性维持、湿生植物多样性保护具有积极作用;不同生境干扰类型和方式不同,景观异质性与植物物种多样性相关性程度也不同。在今后农业景观生物多样性保护中应综合考虑非耕作生境的类型和比例,干扰的类型和方式等多种因素对生物多样性、生态服务及农业产量的影响。     

Old cultural traditions, in addition to land use and topography, are shaping plant diversity of grasslands in the Alps

Socio-economically motivated land use changes are a major threat for species diversity of grasslands throughout the world. Here, we comprehensively explore how plant species diversity of grasslands in the species-rich cultural landscape of the Swiss Alps depends on recent land use changes, and, neglected in previous studies, on old cultural traditions. We studied diversity in 216 grassland parcels at three altitudinal levels in 12 villages of three cultural traditions (Romanic, Germanic, and Walser). In valleys of Romanic villages more different parcel types tended to occur than in those of Germanic and Walser villages, suggesting that socio-economic differences among cultural traditions still play a role in shaping landscape diversity. Moreover, at the village level, higher man-made landscape diversity was associated with higher plant species richness. All observed changes in land use reduced the farmers’ workload. Plant species richness was lower in fertilized than in unfertilized parcels and in abandoned compared with used parcels. Grazing slightly reduced species richness compared with mowing among unfertilized parcels, while in fertilized parcels it had a positive influence. The highest species diversity was found in mown unfertilized subalpine grasslands. Nevertheless, moderate grazing of former meadows can be a valuable alternative to abandonment. We conclude that the ongoing changes in land use reduce plant species richness within parcels and at the landscape level. To preserve plant species diversity at the landscape level a high diversity of land use types has to be maintained.     

The response of plant and insect assemblages to the loss of Calluna vulgaris from upland vegetation

Across large areas of upland Britain Molinia caerulea and Nardus stricta are replacing Calluna vulgaris as the dominant plant species of the vegetation assemblage. Associated changes in the composition of the whole plant assemblage and a phytophagous insect group, the Hemiptera, were investigated. Vegetation and hemipteran assemblages were sampled at six moorland sites in northern England and Scotland. Sampling at each site was carried out in Calluna vulgaris-dominated areas and grass-dominated areas that had previously been dominated by C. vulgaris. Vegetation assemblages of heath and grass sites differed significantly even when species by which sites were selected for sampling (C. vulgaris, M. caerulea and N. stricta) were removed from the analysis with the highest species richness in dry grassland sites. Hemipteran assemblages differed significantly between heath and grass sites with greater species richness also being in grass sites. Plant species composition was the most important explanatory variable of hemipteran assemblages with temperature, latitude, soil loss on ignition and vegetation height also explaining a significant proportion of the variation. Overall a range of vegetation composition and structural variables explained 40.8% of variation in the Hemiptera species data with ranges of soil and biogeographic variables explaining an additional 8.7%.This research demonstrates that the replacement of one dominant plant species by another is associated with a change in other aspects of moorland biodiversity. In particular the loss of C. vulgaris actually leads to an increase in the species richness of Hemiptera and plants although the number of specialist species declines.     

No evidence of a plant extinction debt in highly fragmented calcareous grasslands in Belgium

Long transient times in response to decreasing habitat area and increasing isolation may cause the present plant species distribution to reflect the historical rather than the present landscape configuration, resulting in a so-called extinction debt. To investigate how plant species richness is shaped by both present and historical landscape configuration and local environmental conditions in fragmented calcareous grasslands, 64 sites in Southern Belgium were surveyed for their total species richness and environmental characteristics. Species were subdivided in specialist and generalist groups and the former were clustered into emergent groups (EGs) according to 16 relevant life-history traits. Four specialist emergent groups were derived: orchids, rosette species, annuals and half-rosette species. Both specialist and generalist species richness increased with present fragment area, while present fragment connectivity only affected the number of specialist species. This trend was maintained for most EGs, although habitat area was mostly more important than connectivity. All species groups responded to an insolation gradient, except EGs of rosette and annual species. Differential response to fragmentation among emergent groups could be interpreted in terms of dispersal and persistence traits. No relation between species richness and historical area or connectivity was found, making an extinction debt unlikely. Hence, present-day variation in plant species diversity seems to be no legacy of past landscape configuration. Our results clearly indicate that plant species may be quickly lost in response to new fragmentation events. To restore plant diversity, management should focus both on mitigating landscape fragmentation and restoring habitat quality.