The study of chalk grassland in Northern France: An historical review

The study of chalk grassland in Northern France has evolved through three main phases. During the period from the early 19th century to the First World War, the general flora of Northern France was studied in some detail, but there was little ecological understanding of plant communities. Some workers, however, did attempt to group chalk grassland species ecologically, and the primitive concepts of plant sociology unconsciously developed. Between the wars, with the growth of plant sociology, chalk grassland studies became more elaborate. Specific associations were now described and the wider aspects of chalk grassland phytogeography considered. The post-war period has seen a great increase in botanical activity in Northern France. There have been many regional chalk grassland studies, all of which, in varying degrees, have employed the techniques of plant sociology. It may at last be possible to synthesise these studies, and to describe the chalk grassland associations of Northern France.     

The study of chalk grassland in Northern France An historical review

The study of chalk grassland in Northern France has evolved through three main phases. During the period from the early 19th century to the First World War, the general flora of Northern France was studied in some detail, but there was little ecological understanding of plant communities. Some workers, however, did attempt to group chalk grassland species ecologically, and the primitive concepts of plant sociology unconsciously developed. Between the wars, with the growth of plant sociology, chalk grassland studies became more elaborate. Specific associations were now described and the wider aspects of chalk grassland phytogeography considered. The post-war period has seen a great increase in botanical activity in Northern France. There have been many regional chalk grassland studies, all of which, in varying degrees, have employed the techniques of plant sociology. It may at last be possible to synthesise these studies, and to describe the chalk grassland associations of Northern France.     

Sampling soil wood charcoals at a high spatial resolution: a new methodology to investigate the origin of grassland plant communities

Questions: (i) Can sampling of soil wood charcoals at high spatial resolution produce new evidence concerning the presence of chalk grassland before or during the Neolithic, Bronze and Iron Ages? (ii) Are there correlations between vegetation history and archeological data during these periods at this particular site? Location: The chalk hillsides of Saint‐Adrien in the lower Seine Valley, Upper Normandy, northwest France. Methods: The study was carried out at a high spatial resolution in chalk grassland using soil wood charcoal analysis, in which charcoals found in the soil were identified and dated in an area of several hundred square meters. Results: Late‐successional woody species (Fagus sylvatica, Quercus sp.) were still present in the study site in an area inconsistent with the existence of large chalk grassland herbaceous plant communities (several hectares) in the Neolithic (6500–3800 BP) and Bronze Age (3800–2700 BP). Conclusions: The presence of late‐successional woody species on the studied hillside suggests that fires in the Neolithic were linked to forest clearance for pastoral activities, as already demonstrated for similar ecosystems in eastern France and Germany. Nevertheless, our methodology clearly demonstrates that palaecological studies need to take into account the spatial organisation of plant communities as a complementary element to validate their potential existence in former times.     

Community Associations of Chalk Grassland Leafhoppers (Hemiptera: Auchenorrhyncha): Conclusions for Habitat Conservation

Due to their confinement to specific hostplants or restricted habitat types, Auchenorrhyncha have the potential to make suitable biological indicators to measure the quality of chalk grassland under different management practices for nature conservation. The Auchenorrhyncha data from a study designed to identify the factors influencing the invertebrate diversity of chalk grasslands in southern England was used to evaluate the potential use of this group of insects as biological indicators. Between 1998 and 2002 altogether 81 chalk grassland sites were sampled. Vegetation structure and composition were recorded, and Auchenorrhyncha were sampled at each site on three occasions in each of two seasons using a ‘Vortis’ suction sampler. Auchenorrhyncha assemblages were then linked to the different grassland plant communities occurring on chalk soils according to the British National Vegetation Classification (NVC). Altogether 96 Auchenorrhyncha species were recorded during the study. Using data on the frequency and dominance of species, as is commonly done for plant communities, it was possible to identify the preferential and differential species of distinct Auchenorrhyncha assemblages. Significant differences between the Auchenorrhyncha assemblages associated with the various chalk grassland plant communities of the NVC were observed down to a level of sub-communities. We conclude that data on Auchenorrhyncha assemblages can provide valuable information for the setting of conservation management priorities, where data on floristic composition alone may not be sufficient, providing additional information on aspects of vegetation structure and condition.     

The Restoration of Phytophagous Beetles in Species‐Rich Chalk Grasslands

This study focuses on the restoration of chalk grasslands over a 6‐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. Although the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonized, few beetle species colonized 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 colonization and persistence. The lag time between host plant establishment and colonization was greatest for flightless beetles. Beetles with foliage‐feeding larvae both colonized at slower rates than seed‐, stem‐, or root‐feeding species and persisted within the swards for shorter periods. Although the use of hay spreading may benefit plant communities during chalk grassland restoration, it did not directly benefit phytophagous beetles. Without techniques for overcoming colonization limitation for invertebrate taxa, short‐term success of restoration may be limited to the plants only.     

Problems, Approaches, and Results in Restoration of Dutch Calcareous Grassland During the Last 30 Years

This paper is based on research of the restoration of species‐rich calcareous grasslands in The Netherlands, over the last 30 years. Chalk grassland is a semi‐natural vegetation with a high density of species at a small scale. This type of vegetation was once widespread in Western Europe as common grazing land, mainly for flocks of sheep for which the main function was dung production. In some regions of Central Europe, these grasslands were also used for hay production. The dung was used to maintain arable field production at a reasonable level. In the chalk district in the southernmost part of The Netherlands some 25 sites of this vegetation, varying in area from 0.05–4.5 ha, are still present. Chalk grassland completely lost its significance for modern agricultural production after the wide application of artificial fertilizer following World War II. This grassland has a high conservation value both for plants and animal species, of which a large number of species are exclusively restricted to this biotope. When conservation activities started at a large scale in the early 1960s, three different types of restoration activities could be distinguished: (1) restoration of fertilized sites; (2) restoration of abandoned grasslands; and (3) recreation of chalk grassland on former arable fields. The main aim of the restoration attempt is to create and/or improve sustainable conditions for both plant and animal species characteristic of the chalk grassland ecosystem. In the process of restoration, several phases of different activities can be distinguished: (1) pre‐restoration phase, during which information of the land use history is collected and, based on these data, clear restoration goals are established; (2) initial restoration phase, during which effects of former, non‐conservational land use has to be undone in order to stimulate germination and establishment of target species originating from soil seed bank and species pool; (3) consolidation phase, including the introduction and continuation of a regular management system for sustainable conservation; and (4) long‐term conservation strategy, including measures to prevent disturbance from the outside and genetic erosion and extinction of locally endangered plant populations.     

Changes in chalk-grassland structure and species richness resulting from selective nutrient additions

Abstract. A series of fertilization experiments was carried out over a 5-yr period in a chalk grassland in Limburg (The Netherlands) as part of a study of the maintenance of species richness in species-rich grasslands. Phosphorus and nitrogen were shown to be the most limiting nutrients. Addition of both elements doubled above-ground production, and species richness dropped ca. 50 % in 0.01-m² subplots, relative to controls. However, neither the above-ground production nor plant growth-forms were sufficient to explain the observed changes in species richness. Small-scale structural heterogeneity of the vegetation is probably critical for maintaining high levels of richness. Historically, high nitrogen, low phosphorus conditions were rarely encountered in the Dutch landscape and few species appear adapted to these conditions. Among the chalk grassland species, Brachypodiumpinnatum seems well adapted to these conditions, where it dominates and excludes most other species. A detailed understanding of the small-scale processes responsible for maintenance of species richness is critically important in efforts to maintain the biodiversity of natural ecosystems.     

Vertical seed dispersal by earthworms: a quantitative approach

Earthworms are supposed to play an important role in the dynamics of the soil seed bank, however, experimental evidence and quantitative data are scarce To evaluate the impact of the earthworm species Lumbricus terrains L on the vertical transport of seeds in the soil, laboratory experiments have been earned out Moreover, the worm casts produced and their seed content have been quantified in two chalk grassland sites during one year
The experiment on seed transport has been earned out in cylinders with controlled numbers of worms and seeds at different depths in stenlized soil Seeds used in this experiment proved to be eaten by the earthworms in preceding palatability tests A significant translocation of the seeds in the cylinders with earthworms was recorded after an eight-week period In spring earthworm activity was mainly found m the upper soil layers in the cylinders, whereas in summer the activity was spread over a larger vertical distance The amount of worm casts collected on permanently marked plots in two different chalk grasslands was very high a minimal quantity of c 750 g nr² of worm casts were produced yearly The number of germinative seeds brought to the soil surface by worm casts was at least 60-100 nr² each year This means that earthworm activity has a substantial impact on the soil seed bank dynamics and hence on the possibility of plant recruitment in this species-nch ecosystem     

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.     

Snail faunas in the early stages of a chalk grassland succession

Snail faunas from 10 sites in chalk grassland on the South Downs are described. The sites represent a range of early stages in the succession of grazed chalk grassland to scrub. The faunas of the later stages sampled, with vegetation 12–45 cm high, are substantially different from those of the earlier stages (2.5–10 cm high), lacking most xerophil species, and having much higher proportions of species typical of woods and hedgerows.
The results are discussed in relation both to those from other studies of existing faunas, and also to the environmental interpretation of buried faunas from geological and archeologtcal studies.     

Food overlap in co-existing populations of the land snails Cepaea nemoralis (L.) and Cepaeahortensis (Müll)

Faecal analysis of adult Cepaea nemoralis and Cepaea hortensis from a mixed population on chalk grassland shows that the two snail species select the same plant material as food. Herbs are selected in preference to grasses and Urtica dioica is particularly favoured. C. hortensis has the more pronounced preference for senescent material. These results are discussed in relation to competition between the two species.     

Re-creating Semi-natural Communities: Effect of Sowing Rate on Establishment of Calcareous Grassland

Restoration ecologists are increasingly aware of the potential to re-create chalk grassland on abandoned farmland. Success is often hampered by lack of desirable species in the seed bank and by poor dispersal from nearby sites. In certain schemes, the input of seed may be essential. Locally collected seed is desirable but availability is limited. We examined whether lower sowing rates than currently recommended may be successfully utilized, facilitating more-efficient use of available seed. Experimental plots on former agricultural land were sown at different rates in a randomized complete block, and the vegetation was surveyed for two years. We compared species richness and cover for chalk grassland plants and weeds - species not associated with chalk grassland communities. Values for cover and abundance were matched with data for communities of the British National Vegetation Classification (NVC). Species richness for chalk grassland plants increased with sowing rate and with time, although after two years there was no significant difference between the treatments sown at 0.4, 1.0, and 4.0 grams of seed per square meter. Weed species decreased with increasing rate and time. After two seasons, the vegetation on all treatment plots was similar to that of recognized NVC chalk grassland communities, while the controls were dominated by weeds and showed signs of developing into species-poor grassland. Higher rates rapidly eliminated weeds, but even a small inoculum of seed seemed to significantly enhance establishment of desirable plants and to reduce weed cover. We conclude that lower sowing rates would enable the desired vegetation to become established successfully, under appropriate conditions and management regimes. Lower rates allow for the re-creation of sizable areas using local seed, and they minimize damage to donor sites.     

Cotyledon damage at the seedling stage affects growth and flowering potential in mature plants

Seedling herbivory is an important selective filter influencing patterns of plant community composition. Nevertheless, while many of the mechanisms governing seedling selection by herbivores are well established, the effects of tissue loss at the seedling stage on subsequent plant development are poorly understood. Here we examined how the removal of 50 or approximately 100% of cotyledon area from 7-d-old chalk grassland seedlings affected subsequent plant growth and flowering over a 100-d period. Cotyledon damage had a significant effect on growth during the establishment phase for six of the nine species. For two species, significant effects on plant growth were manifest in 100-d-old plants. Of the five species that flowered, three developed fewer inflorescences or flowered later as a consequence of cotyledon damage suffered as a seedling. Our results show that, in addition to the direct effect of herbivory on seedling mortality, more subtle sublethal effects may also influence plant establishment. Reduced growth as a result of cotyledon damage may have implications for plant competition during the establishment phase, and on subsequent reproductive success at maturity.     

Regional and local variation of spider assemblages (Araneae) from coastal grey dunes along the North Sea

Aim This study aims to determine the underlying causes of local and regional patterns of variation in community structure of spiders in coastal grey dunes, and especially whether ecological time constrains the species composition in young and isolated grey dune habitats. Location The study was conducted in coastal dunes from northern France (Boulonnais, Nord‐Pas‐de‐Calais), Belgium (Flemish coastal dunes) and the Netherlands [Amsterdam Water Supply (AWD) dunes and Dunes from the Provincial Water Company North‐Holland]. Methods Spiders were collected with pitfall traps in twenty‐eight grey dune patches in the four areas under investigation. Species composition and environmental parameters (vegetation structure, distance to the sea and the patch‐edge, eolic dynamics and lime richness) were determined. Assemblage composition was related to the regional and local environmental factors with the Primer software package to determine the assemblage‐determining parameters. Differences in species presence were analysed as a function of their habitat preference and distribution range. Results Differences in grey dune spider assemblage structure can mainly be attributed to differences in local sand dynamics and the region. Species from dynamic dunes are mainly present in grey dunes from Belgium and France, while species from non‐dunal xerothermic habitats (chalk grasslands and heathland) occur in both the Boulonnais and the north Holland dune region. These species are absent from geologically young or other xerotherm habitat isolated Flemish coastal dunes. Main conclusion The data show that regional variation in spider assemblage composition results from local landscape characteristics (dynamics in the dune area), the latitude and the connectivity to non‐dunal xerothermic habitats. The strong and moderate geological isolation of dune areas from the Flemish coast and the AWD dunes, respectively, results in the absence of (at least some) species that are primarily bound to heathland and/or chalk grassland. This indicates the importance of ecological time for the assemblage structure. The limited dispersal capacity of the heathland and/or chalk grassland species is probably the main reason for their absence.     

An 11400 year paleoecological history of a British chalk grassland

The first broad-based, paleoecological analysis of a sedimentary sequence on the British chalk, dating to the terminal Pleistocene, reveals a history of climatic, vegetational and faunal change. The past co-occurrence of currently allopatric species among molluscs, beetles and plants supports hypotheses of the impermanence of communities. Modern pollen rain data are utilized to refine the ecological interpretation of the fossil pollen data. The presence of the Windermere Interstadial (Allerod) and the Loch Lomond re-advance (Younger Dryas) are represented by decreased abundances of arboreal taxa and increased representation of cold grassland elements. Open grassland habitats appear to have been a continuous landscape element, at least locally, since the late-glacial period on the northern British chal Mands, although their species composition has changed greatly in the last 11400 yr.     

Permanent seed banks in chalk grassland under various management regimes: their role in the restoration of species-rich plant communities

A survey was carried out on four stands of a chalk grassland (open, intensively grazed, tall and scrub) in order to monitor the permanent soil seed bank under various management regimes responsible for the decline of the ecological interest of this ecosystem. There was no real similarity between the seed banks and the associated vegetation, except in the intensively grazed stand. When vegetation changes, seed banks appear to be quite stable and are mostly dominated by three species (Centaurium erythraea, Origanum vulgare, Hypericum perforatum). The seed bank seems to be quite useless in order to restore a species-rich community after both grazing intensification or abandonment. Because recolonization processes are the critical factor, the management of species-rich grassland has to be designed with a view to preserving a given plant community as well as to improving dispersion processes from a source to target areas.     

A Bryophyte flora of Kent. II. Hepaticae

Abstract

The military training area on Salisbury Plain contains the largest area of chalk grassland in western Europe. The grassland swards, though relatively tall, are often rich in flowering plant species. The bryophyte flora of areas disturbed in five different ways (prehistoric earthworks, twentieth century rifle ranges, ant-hills, vehicle tracks and shell-holes) was compared to that of adjacent, relatively undisturbed grassland. There was no significant difference in bryophyte cover between quadrats on disturbed ground and the controls, but the disturbed sites supported more bryophyte species per quadrat. Of the 55 taxa recorded, 38 species were at least three times more frequent in the disturbed than the undisturbed sites, compared to four which were at least three times more frequent in the control sites. The species favouring disturbed conditions included several bryophytes characteristically associated with chalk soils in southern England, including some that fruit freely (e.g. Microbryum curvicollum, Tortula lanceola) and others that fruit very rarely (e.g. Abietinella abietina, Entodon concinnus). These results are discussed in relation to the conservation of bryophytes and other disturbance-tolerant and disturbance-dependent species on Salisbury Plain and in the wider context of the protection of the bryophytes of chalkland habitats.     

Increased solar UV-B radiation may reduce infection by arbuscular mycorrhizal fungi (AMF) in dune grassland plants: evidence from five years of field exposure

An area of coastal dune grassland, dominated by the gramineous species Calamagrostis epigeios and Carex arenaria, was exposed to enhanced levels of UV-B radiation during a five year period. These species showed reduced AM-fungal infection percentages in their roots. In C. epigeios AM infection was reduced by 18%, C. arenaria showed a reduction by 20%. The major effect of enhanced UV-B on AM associations was a reduction of the number of arbuscules. This indicates a reduction in the exchange of nutrients between the symbionts. Since the effect of UV-B on AM associations may result from altered flavonoid levels in the root exudates of the host plants, flavonoid levels in the roots were investigated. No detectable flavonoid concentrations were found in the roots of C. epigeios and C. arenaria. Less effective AM associations can have pronounced negative effects on biodiversity and nutrient dynamics of the dune grassland ecosystem. The possible mechanisms causing these indirect effects of elevated UV-B on below ground AM associations are discussed. We conclude that UV-B induced changes in plant hormone levels are more likely to be the mechanism reducing AMF infection than UV-B induced alterations in flavonoid concentrations in the root exudates of the host plant.     

Plant functional diversity enhances associations of soil fungal diversity with vegetation and soil in the restoration of semiarid sandy grassland

The trait‐based approach shows that plant functional diversity strongly affects ecosystem properties. However, few empirical studies show the relationship between soil fungal diversity and plant functional diversity in natural ecosystems. We investigated soil fungal diversity along a restoration gradient of sandy grassland (mobile dune, semifixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China, using the denaturing gradient gel electrophoresis of 18S rRNA and gene sequencing. We also examined associations of soil fungal diversity with plant functional diversity reflected by the dominant species' traits in community (community‐weighted mean, CWM) and the dispersion of functional trait values (FD_is). We further used the structure equation model (SEM) to evaluate how plant richness, biomass, functional diversity, and soil properties affect soil fungal diversity in sandy grassland restoration. Soil fungal richness in mobile dune and semifixed dune was markedly lower than those of fixed dune and grassland (P < 0.05). Soil fungal richness was positively associated with plant richness, biomass, CWM plant height, and soil gradient aggregated from the principal component analysis, but SEM results showed that plant richness and CWM plant height determined by soil properties were the main factors exerting direct effects. Soil gradient increased fungal richness through indirect effect on vegetation rather than direct effect. The negative indirect effect of FDis on soil fungal richness was through its effect on plant biomass. Our final SEM model based on plant functional diversity explained nearly 70% variances of soil fungal richness. Strong association of soil fungal richness with the dominant species in the community supported the mass ratio hypothesis. Our results clearly highlight the role of plant functional diversity in enhancing associations of soil fungal diversity with community structure and soil properties in sandy grassland ecosystems.     

Significant Achievements in Protection and Restoration of Alpine Grassland Ecosystem in Northern Tibet, China

Alpine grassland is a fragile ecosystem, and a large area of this grassland type has been severely degraded in Northern Tibet, to the extent that it has become the primary ecological problem in the region. Various levels of government, including the national central government, the Tibetan Autonomous Region government, and the Nagqu Prefecture government have worked together to achieve alpine grassland ecosystem protection and prevent grassland degradation. These efforts have resulted in significant ecological, social, and economic benefits in Northern Tibet.