Impacts of succession and grass encroachment on heathland Orthoptera

For the conservation of biodiversity, heathlands present important ecosystems throughout Europe. The formerly widespread habitats are nowadays restricted to small and isolated remnants. Without land use heathland vegetation undergoes succession and, in addition, the increasing amount of atmospheric nitrogen deposition has resulted in an encroachment of grasses. In the present study we analysed the effects of succession and grass encroachment on Orthoptera in a coastal heathland on the Baltic island of Hiddensee, Germany. Vegetation, microclimate, soil humidity and Orthoptera were sampled in the five main stages of heathland succession, namely grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, and birch forest. Vegetation and environmental parameters showed strong differences among the successional stages. Orthoptera species richness was highest in transitional stages. The high proportion of grasses offer favourable habitat conditions for graminivorous, chorto- and thamnobiont species. Orthoptera density was highest in grey dunes. Threatened and specialised species were restricted to the young stages grey dunes and dwarf-shrub heath. Hence, in order to maintain a high diversity of Orthoptera in heathlands, maintaining different successional stages is of critical importance and this should be integrated into heathland management practices.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

Response of carabid beetles (Coleoptera: Carabidae) and spiders (Araneae) to coastal heathland succession

We analyzed the impacts of succession and grass encroachment on carabid beetle and spider assemblages in a coastal heathland. Further, indicator species for different successional stages (grey dunes, dwarf-shrub heath, grassy heath, heath with shrubs, birch forest) were identified, and their relations to habitat parameters were analyzed. The study was conducted on the Baltic island of Hiddensee, Germany. Ground-dwelling arthropods were sampled using pitfall traps along a successional gradient containing five stages. Ordination by nonmetric multidimensional scaling (NMDS) and analysis of similarities (ANOSIM) revealed a clear separation of species composition among the successional stages. Both in carabid beetles and spiders, most indicator species were obtained in the youngest stage (grey dunes) and fewest in the intermediate stages (grassy heath, heath with shrubs). Also the proportion of endangered species was highest in grey dunes. Based on our results, conservation management of coastal heathlands should preserve a mosaic of different successional stages with a clear preference on younger stages (grey dunes and dwarf-shrub heath).     

Effects of Montane Heathland Restoration on Leafhopper Assemblages (Insecta: Auchenorrhyncha)

At the beginning of the 20th century, many montane heathlands were abandoned and became subject to natural succession or afforestation by humans. Thus, the formerly large montane heathlands slowly degraded into small and isolated patches. In this study, we evaluate the influence of restoration measures on leafhopper (Auchenorrhyncha) assemblages of montane heathland ecosystems in Central Europe. Our analyses comprised three different site types that were adjacent to each other: (1) montane heathlands, (2) restoration sites, and (3) control sites. Leafhoppers showed a clear response to montane heathland restoration. Thus, after 4–5 years since implementation of restoration measurements restoration sites were characterized by the highest species richness. However, detailed analyses of leafhopper diversity, species composition, and environmental parameters on the three site types revealed that restoration sites were rather similar to control sites and significantly differing from montane heathlands. We conclude that leafhoppers are excellent bioindicators for restoration measurements because they reflected environmental differences between the three site types. Restoration measurements might only be a useful instrument to promote typical montane heathland leafhopper communities in the long run. Colonization by leafhoppers is, however, dependent on many different factors such as leafhopper mobility, vegetation structure, microclimate, and the establishment of ericaceous dwarf shrubs. Practitioners should establish a management regime (grazing and sod‐cutting) that creates a mosaic of different habitat structures and increases typical heathland vegetation, thus, favoring the colonization of typical heathland leafhoppers.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

Effects of fire on the vegetation of a lowland heathland in North-western Italy

This study focuses on the effect of fire on lowland heathlands at the extreme southern edge of their European distribution (Vauda Nature Reserve, NW Italy). Forty-nine plots (50 m radius) were surveyed between 1999 and 2006. Each year, fire occurrences were recorded and per cent cover of four vegetation types (grassland, heath, low shrubland, and tall shrubland) was estimated in each plot. Vascular plant species richness was also recorded in 255, 1 m² quadrats. After a fire, grassland vegetation expanded, but then declined rapidly as heath and shrubland recovered: 7 years after a fire, tall shrubland encroached on to more than 40% of the plots, and grassland declined from 50% to 20% cover. Between 1999 and 2006, Betula pendula shrubland greatly expanded, while grassland decreased over most of the Reserve, even where fire frequency was high. Tall shrubland had low plant diversity and was dominated by widespread species of lower conservation value. By contrast, early successional vegetation (grassland and low shrubland) had higher richness and more narrowly distributed species, indication that the development of tall shrubland causes significant species loss in the heathland. Italian lowland heathlands are characterized by high rates of shrubland encroachment that threatens both habitat and species diversity. Burning frequencies of once in 3–6 years seem appropriate in this habitat, but burning alone might not suffice without actions to increase herbivore grazing.     

Spider communities as evaluation tools for wet heathland restoration

Over the last two centuries wet heathlands and associated habitats, such as poor fens and bogs, have suffered extensive fragmentation. Recently, large-scale projects aim to restore these rare habitats throughout Europe. To evaluate post-restoration trajectories of wet heathlands, suitable monitoring tools are urgently needed. Here, we investigated whether spider communities are useful tools for evaluating the restoration success of wet heathlands. Ordination revealed that dissimilarity in spider communities between patches of different age classes resulted mainly from vegetation cover and soil moisture. By using a functional trait-approach, we tested if the time since restoration affects trait distribution of spiders in wet heathlands. Typical wet heathland spider species were less common with increasing vegetation encroachment and lower water content. New patches were inhabited by summer active, eurytopic (non-heathland) spiders, while more typical heathland species were found in middle-aged and old patches. Our results suggest that time-related changes in vegetation structure and moistness of restored wet heathlands are clearly reflected by spider communities. Although mobile spiders quickly recolonize the restored heathlands, it takes time for typical heathland spiders to settle. Restoration measures should prevent the negative effects of a vegetation encroachment and a high density of forested edges and should rehabilitate the hydrological cycle in order to preserve rare heathland spiders. We discuss that accounting for responses of spiders provides additional information to guide wet heathlands restoration.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Rapid response of Orthoptera to restoration of montane heathland

Understanding how to restore threatened ecosystems is of special relevance for nature conservation. The aim of this study was to use Orthoptera as ecological indicators for the effects of montane heathland restoration in Central Europe. We analysed the three following treatments: (i) montane heathlands (MONHEATH) (N = 7), (ii) restoration sites (RESSITE) (N = 3) and (iii) clear-cuts of spruce forests as unprocessed and ungrazed control sites (CONTROL) (N = 3). Vegetation structure and microclimate differed considerably between MONHEATH on the one hand and RESSITE and CONTROL on the other hand. Orthoptera species richness and density did so too. MONHEATH was characterised by a high-growing dense dwarf-shrub and moss layer having a cool microclimate and high soil moisture. In contrast, RESSITE and CONTROL had sparse vegetation and a warm microclimate; Orthoptera species richness and density was highest on these sites. Our study clearly showed that heathland Orthoptera responded rapidly to restoration measures, while Ericaceae dwarf shrubs slowly established. The vast majority of Orthoptera species found on the restoration sites are early and mid-successional species. The colonization of the sites by late-successional Orthoptera species in the future will depend on the further development of the heathland vegetation; that is, if Ericaceae will expand to the sites. We conclude that the realised restoration measures are suitable to promote heathland Orthoptera of early and mid-successional stages. However, the current management of montane heathlands is insufficient and needs to be intensified in order to provide structurally diverse habitats with their characteristic orthopteran assemblages.     

Invasion of Calluna heath by native and non-native conifers: the role of succession, disturbance and allelopathy

The coastal heathlands of North-western Europe are treeless anthropogenic landscape of high conservation value, now threatened by encroachment by both native (Pinus sylvestris L.) and non-native (Picea sitchensis (Bong) Carr.) conifers. This study wants to gain insight into the different external drivers which interact with the natural successional processes to determine the future development of the heathland landscape. We tested how seedling emergence and establishment is related to competitive effects from the standing vegetation. Heathlands are nutrient-stressed habitats and we tested the seedling emergence and establishment along a nutrient gradient using mean Ellenberg values. In addition we tested if phytotoxic substances are affecting the seedling emergence and establishment related to different successional stages. Study species differed in seed size and we expected different colonization rates and competitive effects from the heathland vegetation. A factorial experiment was carried out in three different successional stages to isolate the effects of competition versus phytotoxins on seedling emergence and establishment by combining vegetation cutting and charcoal addition treatments. Both conifers were able to invade all successional stages, but invasibility decreased during succession. This was largely attributable to competitive effects from the standing vegetation, although phytotoxic effects were also detected, especially in late-successional stages. Seedling emergence and establishment was higher in the larger-seeded P. sylvestris than in P. sitchensis. These results present a management challenge as fire is an important component of the traditional management of the coastal heathlands, while at the same time increasing invasibility. Managers at sites susceptible to invasion should ensure that resources for tree seedling control are available before fire or other management-related disturbances are applied.     

Fine‐scale microhabitat selection for dense vegetation in a heathland rodent,Rattus lutreolus: Insights from intraspecific and temporal patterns

Abstract Vegetation is a dynamic habitat component and successional changes in vegetation structure can lead to concomitant changes in the communities of animals living in a particular area. Heathland rodents are a classic example, with vegetation at different ages post fire being dominated by different species. While broad associations are often demonstrated between the distribution and abundance of species and vegetation structure, the causal relationships are poorly understood. Studies of temporal and sex‐ or age‐specific patterns can provide strong insights into the processes underling patterns of habitat selection. In an attempt to better understand the mechanistic links between rodent successional patterns and vegetation structure in heathlands, we conducted a detailed study of microhabitat use by the swamp rat, Rattus lutreolus, in a native heathland in south‐eastern Australia. Rattus lutreolus typically occurs in late‐succession heath and is frequently associated with high vegetation density. Our assessment of vegetation at trapping stations, and also along trails used by the animals (using the spool‐and‐line tracking technique), revealed strong selection by the rats for dense vegetation by both day and night. The spool‐and‐line tracking approach revealed distinct intraspecific and temporal patterns. During the day, females foraged in vegetation of much higher density than did juveniles, with males behaving intermediately. During the night, however, all animals selected dense vegetation irrespective of sex or age, although the mean density of vegetation selected during the night was lower than it was during the day. These patterns were independent of daily maximum and minimum air temperature and were therefore unlikely to be related to microclimate. We propose instead that high vegetation density acts as a source of protection from predators, allowing R. lutreolus to forage safely both by day and by night.     

Vkgetation analysis along a successional gradient from heath to oak forest

At Hjelm Hede in Denmark where an oak shrub is invading into a heathland the successional process has been studied on poor sandy outwashed plain. The study used a transect through a successional gradient from open heath to oak shrub where vegetation, major topsoil parameters and tree characteristics were analysed. The study enables us with spatial variation of key parameters to combine numerous detailed studies from the same site into an overall picture of changing vegetation and soil parameters reflecting the succession, which has been documented previously. The data were analysed by the use of Detrended Correspondence Analysis (DCA). Along the gradient from heath to forest the thick mor layer of the heathland soil decomposed, the CM ratio decreased from 25 to 15, pH increased and ammonium became available for the plants during a time span of approximately 120 years. The vegetation on the open heath was typical Danish inland heath vegetation dominated by Calluna vulgaris, Empetrum nigrum and Deschampsiaflexuosa. Under the young oak shrub the vegetation was dominated by Deschampsia Jexuosa, Trientalis europaea , and Maianthemum bijiolium. In the oldest and most forest like parts of the shrub a vegetation dominated by Holcus mollis, Anemone nemomsa and Stellaria holostea was found. Woody chamaephytes were replaced first by hemicryptophytes and later by geophytes. The vegetation's Ellenberg values indicated an increase in nitrogen availability and a decrease in acidity and light availability through secondary succession.     

A comparative study of the seedbanks of heathland and successional habitats in Dorset, Southern England

1 Many areas of lowland heaths are being lost due to invasion by Betula spp., Pinus sylvestris, Pteridium aquilinum, Rhododendron ponticum and Ulex europaeus. One of the factors influencing the success of restoration of heathland on such sites will be the content of their viable seedbanks.
2 Ten heathland areas in the Poole Basin area of Dorset, where succession to one or more of the above species had occurred were studied. The viable seedbanks of the successional sites were compared with those of nearby heathland using Canonical Discriminant Analysis.
3 The seedbanks of all the successional stages were significantly different from the seedbank of the heath.
4 The seedbanks from the Pinus sylvestris and Pteridium aquilinum successional stages contained significantly lower numbers of heathland species than did the heathland seedbank, although few non heathland species were present.
5 The seedbanks from the Betula spp., Rhododendron ponticum and Ulex europaeus successional sites contained both significantly lower numbers of heathland species and significantly higher numbers of non heathland species than the heathland seedbank.
6 The results are discussed in relation to the restoration of heathland on successional sites and the use of the seedbank as a source of propagales for the establishment of heathland species.     

小叶锦鸡儿灌丛化对典型草原群落结构与生态系统功能的影响

草原灌丛化是全球干旱半干旱地区面临的重要生态问题。灌丛化对草原生态系统结构与功能的影响较为复杂, 有待于在更广泛的区域开展研究。该研究在内蒙古锡林郭勒典型草原选择轻度、中度和重度灌丛化草地, 通过群落调查, 结合植物功能性状和土壤理化性质观测, 研究了小叶锦鸡儿(Caragana microphylla)灌丛化对草原群落结构(物种多样性、功能多样性和功能群组成)和生态系统功能(初级生产力、植被和土壤养分库)的影响。结果表明: 1)不同程度灌丛化草地的物种丰富度、功能性状多样性和群落加权性状平均值差异显著, 其中, 中度灌丛化草地的物种多样性和功能多样性较高, 表明一定程度的灌丛化有利于生物多样性维持。2)重度灌丛化草地的地上净初级生产力(ANPP)显著高于轻度和中度灌丛化草地, 其原因主要是随着灌丛化程度加剧, 群落内一/二年生草本植物显著增加, 而多年生禾草和多年生杂类草显著减少。三个灌丛化草地的植被叶片和土壤碳、氮库差异均不显著。3)灌丛化对草原生态系统功能包括ANPP、植被和土壤养分库均没有直接的影响, 而是通过影响功能群组成、土壤理化性质和功能多样性, 间接地影响生态系统功能; 灌丛化导致功能群发生替代和土壤旱碱化是最重要的生物和非生物因素。     

Montane heathland rejuvenation by choppering—Effects on vascular plant and arthropod assemblages

Land-use changes and atmospheric nitrogen deposition have negatively affected heathland biota. Active habitat management is one possible way of counteracting the biodiversity loss associated with these habitat alterations. However, management practices for lowland heathlands often have been transferred to montane heathlands, irrespective of the differences in environmental conditions or assemblage composition. The objective of this study was to evaluate the effects of so-called choppering for the rejuvenation of montane heathland. Choppering involves chaffing and removing the largest part of the organic layer down to the mineral soil. In this study, we compared montane heathlands that were rejuvenated through the application of choppering (CHOPPER) to old-growth montane heathlands (CONTROL). Thirteen years after the rejuvenation measures had been conducted, the environmental conditions between CHOPPER and CONTROL still differed. CHOPPER was characterised by shorter vegetation (herbs/grasses and dwarf shrubs), more bare soil, less litter and higher temperatures. Although, the vascular plants and all studied arthropod groups were affected by the environmental changes, their responses were somewhat different. CHOPPER had a unique assemblage of each taxonomic group that included at least a few heathland species that mainly occurred in this treatment. However, choppering was most beneficial for vascular plants, grasshoppers and carabid beetles. As shown for lowland heathlands, choppering is also a suitable management measure for montane heathland to rejuvenate vegetation with its characteristic arthropod fauna. Due to the intact seed banks and Ericaceae root systems with their mycorrhizas, in combination with the availability of bare soil, heathland vegetation can rapidly regenerate after choppering. The keystone structures that explain the high relevance of CHOPPER, especially for vascular plants, grasshoppers and carabid beetles, are low-growing vegetation and bare soil, which result in light and warm microclimatic conditions. Based on the results of our study, we recommend choppering as a regular management measure to rejuvenate montane heathland.     

Changes in heathland vegetation under goat grazing: effects of breed and stocking rate

Questions: How are heathland vegetation dynamics affected by different goat grazing management? Location: Cantabrian heathlands in Illano, Asturias, northern Spain. Methods: During 4 years, vegetation dynamics (structural composition, canopy height and floristic diversity) were studied under three goat grazing treatments with three replicates: high stocking rate (11.7 goats ha^?1) with a local Celtiberic breed, and high (15 goats ha^?1) and low (6.7 goats ha^?1) stocking rates with a commercial Cashmere breed. Results: The relative cover of woody plants, particularly heather species, decreased more while herbaceous cover increased more under local Celtiberic than under Cashmere breed grazing. Within Cashmere treatments, the cover and height of live shrubs decreased more and the herbaceous cover increased more under high than under low stocking rate. Redundancy analysis showed a significant effect of treatment × year interaction on floristic composition. Greater species richness was recorded under local goat grazing, but Shannon diversity index fell in the fourth year on these plots because of dominance by two grass species. Conclusions: Local Celtiberic goat grazing at such a high stocking rate (11.7 goats ha^?1) hinders the development of sustainable systems on these heathlands, both in environmental and productive terms, owing to the limitations in soil fertility. Nevertheless, Celtiberic goats could be useful for controlling excessive shrub encroachment and reducing fire hazard. Cashmere goat grazing at high stocking rate promoted the highest Shannon diversity by generating a better balance between woody and herbaceous plants, while shrub dominance was not altered under the low stocking rate.     

Soil seed bank dynamics in alpine wetland succession on the Tibetan Plateau

The primary goal was to address several questions with regard to how soil seed banks change in a successional series. How does the composition of the viable seed bank change, and how does the relationship of the soil seed bank and vegetation change with succession? Can the seed bank be regarded as a potential as a source of seeds for wetland restoration? We collected soil seed bank samples and sampled the vegetation in four different successional stages and used the NMDS (nonmetric multidimensional scaling) to evaluate the relationship of species composition between the seed banks and vegetation. The difference of seed density and species richness in different habitats and soil depths also was compared. Viable seeds of half (37) the species in the early-successional stage were found in all the successional stages. Similarity between seed bank and vegetation increased with succession. Both seed density and species richness in the seed bank increased with successional age and decreased with soil depth. The majority of species from the early-successional stage produced long-lived seeds. Seed density and species richness increased with succession, mainly as a result of increasing seed production, and hypotheses predicting decreasing density of buried seeds and species richness were not confirmed. Seed banks play a minor role in contributing to the regeneration of vegetation, and managers cannot rely on soil-stored seed banks for restoration of wetlands.     

Structural changes of vegetation and its association with microclimate in a successional gradient of low thorn forest in northeastern Mexico

Understanding how microclimate and vegetation are associated during secondary succession is of primary importance for plant conservation in the face of the increasing land cover modification. However, these patterns are still unstudied for many plant communities. This study aimed to evaluate the structure (species richness, Shannon's diversity index, Simpson´s dominance index, abundance of each species, average height of species, species cover (%), species composition, and indicator values) of a low thorn forest fragment and to analyze its relation with microclimate along a successional gradient. Four stages of succession were delimited by the analysis of Landsat images, in the state of Tamaulipas, northeast Mexico. Statistical models incorporated species richness, diversity indices, abundance, height, and cover, as variables for searching differences between stages, or to evaluate microclimate associations. A total of 70 species, 54 genera, and 27 families were determined. Height of tree layer was the most important variable for discrimination of the successional stages. Conserved areas differed floristically from other stages, associated mainly with the lowest values of wind speed originated by tree layer characteristics. A significant association between species and microclimate was found, being wind speed and relative humidity the most important variables. Some species, due to their high importance values and their patterns of association with microclimate, may be considered as key taxa for low thorn forest, which is a threatened semitropical community in northeast Mexico. Conserved and late successional areas account for climatic regulation of this plant community, and the importance of these forest patches may be considered when establishing biodiversity protection areas.

     

Problems in heathland and grassland dynamics

A review of factors governing vegetation change in heathlands and grasslands is presented, with emphasis on soil factors. Climate, microclimate, fire, fauna, grazing by vertebrates, grazing by invertebrates, disease, dispersal, establishment, competition, allelopathy, stabilization, soil formation and podzolization are discussed with emphasis on inconsistencies and lacks in our present knowledge. Examples are mainly from NW European Calluna heath and related woodlands.Nomenclature follows Clapham, Tutin & Warburg (1962) for vascular plants; species not included in this work have authorities cited in the text.     

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.