The impact of site conditions and seed dispersal on restoration success in alluvial meadows

Abstract. We studied the restoration success of flood plain meadows in the northern Upper Rhine valley, where between 1988 and 1992, 35 ha of arable land was converted into grassland and subsequently managed for nature conservation. Remnant populations of typical alluvial meadow species were found in old meadows and along drainage ditches that dissect the whole area. We analysed the site conditions and phytosociological relevés in old and new meadows. Small differences in site parameters between old and new meadows contrasted with a clear floristic differentiation between the two meadow types. The vegetation of old meadows was much more differentiated along prevailing environmental gradients than the vegetation of new meadows. Despite the favourable site conditions for the re‐establishment of species‐rich meadows on the former arable land, restoration success was limited to the vicinity of remnant stands. In contrast to old meadows, indicator species of new grassland were still typical species of regularly disturbed ruderal and arable habitats, often capable of building up a persistent seed bank. The precise mapping of 23 target species revealed that even wind dispersal predominantly leads to re‐establishment in the close circumference of parent plants. We found no indication that regular flooding, hay‐making and autumnal grazing had an impact on recolonization of newly created grassland. Even under favourable conditions for the re‐establishment of target species, restoration success in alluvial meadows proved to be strongly dispersal limited. We discuss the implications of our findings for future restoration management in grasslands.     

Seed preferences by rodents in the agri‐environment and implications for biological weed control

Post‐dispersal seed predation and endozoochorous seed dispersal are two antagonistic processes in relation to plant recruitment, but rely on similar preconditions such as feeding behavior of seed consumers and seed traits. In agricultural landscapes, rodents are considered important seed predators, thereby potentially providing regulating ecosystem services in terms of biological weed control. However, their potential to disperse seeds endozoochorously is largely unknown. We exposed seeds of arable plant species with different seed traits (seed weight, nutrient content) and different Red List status in an experimental rye field and assessed seed removal by rodents. In a complementary laboratory experiment, consumption rates, feeding preferences, and potential endozoochory by two vole species (Microtus arvalis and Myodes glareolus) were tested. Seed consumption by rodents after 24 h was 35% in the field and 90% in the laboratory. Both vole species preferred nutrient‐rich over nutrient‐poor seeds and M. glareolus further preferred light over heavy seeds and seeds of common over those of endangered plants. Endozoochory by voles could be neglected for all tested plant species as no seeds germinated, and only few intact seeds could be retrieved from feces. Synthesis and applications. Our results suggest that voles can provide regulating services in agricultural landscapes by depleting the seed shadow of weeds, rather than facilitating plant recruitment by endozoochory. In the laboratory, endangered arable plants were less preferred by voles than noxious weeds, and thus, our results provide implications for seed choice in restoration approaches. However, other factors such as seed and predator densities need to be taken into account to reliably predict the impact of rodents on the seed fate of arable plants.     

Evaluation of Initial Restoration Measures during the Restoration of Calcareous Grasslands on Former Arable Fields

In 1993, experiments on the restoration of calcareous grasslands on ex‐arable fields were started in order to provide new habitats for species of a small nature reserve with ancient grasslands north of Munich (Germany). The effects of diaspore transfer by the application of seed‐containing hay on vegetation establishment were studied on restoration fields with and without topsoil removal for 5 years. The aim of the study was to assess plant diversity for the evaluation of restoration success by different methods including determination of species with viable seeds in the hay by germination tests, phenological investigations on hay‐transfer source sites at the time of harvest, and vegetation analyses on the restoration sites. Total seed content of the hay and the number and composition of plant species with viable seeds were affected by the time of harvesting and differed between a site which had been used as arable field until 1959 and ancient grassland sites. Nevertheless, the number of established hay‐transfer species showed only few differences between restoration fields. The proportion of species transferred to restoration fields in relation to the number of species with viable seeds in the hay was between 69 and 89%. Five years after the hay transfer, the proportion of the established species was still between 58 and 76%. Up to now, topsoil removal had no significant effect on the number of established hay‐transfer species. After triple hay application the absolute number of transferred grassland species was higher than on sites with single hay application, but restoration efficiency was lower because many of the species with viable seeds in the hay did not establish. In general, our results showed that the transfer of autochthonous hay is a successful method to overcome dispersal limitation in restoration projects.     

Transfer seeds,hay, or soil blocks? The importance of the completeness of biological inputs to address dispersal and establishment limitations during the restoration of plant assemblages in floodplain grasslands

This study investigated different techniques of grassland restoration to overcome dispersal or establishment limitation, which are key processes influencing early-successional plant community assembly. A fully randomized in situ experiment was set up in a former arable land in a floodplain along the Garonne river (south-western France) to test for the effect of (1) the type and completeness of the biological input (any biological material—seed, hay, and soil—transferred to the site under restoration), (2) soil disturbance by deep tillage, and (3) their interaction on plant community dynamics for 5 years. All inputs influenced the plant community structure and composition. The effects depended on the type of the input but not necessarily in link with its completeness. High density and diversity seed mixture led to high levels of richness and relative abundance of target species like soil blocks, the input considered as the most complete. During the first year, hay transfer mostly influenced community assembly through negative litter effects. Delayed germination of several species contributed to buffer these early effects. This study supports the importance of dispersal limitation during early succession in degraded grassland ecosystems whereas soil disturbance had only subtle effects on the seed bank and standing plant community, indicating that competition and establishment limitation were of secondary importance in our study system. Our results suggest that even low-frequency immigration events (e.g. when a few seeds are transferred within hay) can be of great importance providing that a sufficiently long time period is considered for grassland restoration.     

Does the seed bank contribute to the restoration of species‐rich vegetation in wet dune slacks?

Questions: What is the contribution of the seed bank to restoration of species‐rich vegetation in oligotrophic wet dune slacks? Does the restoration management affect the seed bank? Location: Calcareous coastal dune slacks at the west coast of The Netherlands. Methods: Species composition of the seed bank and the above‐ground vegetation was sampled in dune slacks that had a variable extent of groundwater level rise in combination with either topsoil removal or mowing. Results: The seed bank had a high potential for restoration of species‐rich vegetation: 60 species were found in the seed bank of which 14 were characteristic of oligotrophic, wet dune vegetation. While topsoil removal almost completely removed the seed bank, groundwater level rise did not permanently submerge the seed bank of species of oligotrophic, wet conditions. Changes in abundance in the established vegetation were unrelated to species abundance in the seed bank. Of all new species establishments in the vegetation relevés, 76% occurred where the species was not found in the seed bank. The chance that presence of a species in the seed bank led to establishment the following year was only 11%. Conclusion: The seed bank was not the dominant source for newly establishing species following the large disturbance that was induced by restoration management. Changes in species abundance after the restoration impact were not related to species abundance in the seed bank, but to ongoing succession and current year dispersal. To attain a high number of new establishments, restoration projects should preferably be planned in the proximity of refuge populations, rather than relying on the seed bank alone.     

Effect of Hay Transfer on Long-Term Establishment of Vegetation and Grasshoppers on Former Arable Fields

The transfer of seed‐containing hay is a restoration measure for the introduction of plant species of local provenance. We investigated the effect of hay transfer on species richness and on long‐term establishment of target plant and grasshopper species on former arable fields with and without topsoil removal in comparison to reference sites in a nature reserve. Plant species richness, the number of target plant species, and Red List plant species were significantly positively affected by hay transfer, both on the scale of whole restoration fields and on permanent plots of 4 m². Eight years after the start of the restoration, only few of the transferred plant species had disappeared and some target species were newly found. Grasshoppers were affected not by hay transfer but by topsoil removal. The proportion of target grasshopper and plant species and Red List grasshopper species was higher on topsoil removal sites with low standing crop and high cover of bare soil than on sites without soil removal. On topsoil removal sites without hay, however, plant species richness was very low because of the slow natural dispersal of the target species. Vegetation and grasshopper communities still differed between restoration fields and the nature reserve. Nevertheless, our results indicate that the transfer of autochthonous seed‐containing hay is a successful method to establish species‐rich grasslands with a high proportion of target species.     

Restoration of wooded meadows ‐ a comparative analysis along a chronosequence on Öland (Sweden)

Abstract. Wooded meadows on the Baltic Island of Öland result from traditional agricultural management over centuries which has led to a species‐rich vegetation with high species diversity. Today, nearly all of these meadows have been abandoned and became rapidly overgrown by deciduous shrub and tree species forming a closed canopy which resulted in a rapid and strong decrease in species numbers of the herb layer. Recent efforts aim to restore overgrown wooded meadows by cutting single shrubs and trees to open the canopy. However, the effects of abandonment as well as of any restoration management in wooded meadows have rarely been documented until now. Mechanisms driving succession after restoration such as the dispersal potential of the respective species over time and space have not been analysed yet. Therefore, a chronosequence was studied which included a traditionally managed wooded meadow, an overgrown meadow which has been abandoned for more than 100 yr and a meadow which was restored 36 yr ago by cutting and is now grazed. We analysed the soil seed bank of the 3 meadows in comparison with the established vegetation and endozoochorous seed dispersal by cattle and sheep. After abandonment 87% of the typical grassland species vanished from the established vegetation and were replaced by species characteristic of woodland and disturbed grassland communities. The mean number of species decreased from 52 species per plot (4 m²) to 18 species. Mean species number and number of seeds in the seed bank declined significantly from the traditionally managed to the overgrown meadow. Most of the grassland species were assigned to a transient seed bank type while only 1/3 could be classified as having a short‐term persistent seed bank. Thus, restoration of wooded meadows cannot rely on the soil seed bank. Endozoochorous seed dispersal by cattle and sheep was shown for 15% of the species with seed densities per 100 g air dried dung of 737 and 767, respectively. Movement of animals between ancient and restored wooded meadows is recommended since many of the species only occurred in low densities and therefore, will probably not be found in the dung samples.     

Land use history and site location are more important for grassland species richness than local soil properties

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. Grazing ex‐arable fields contain a fraction of the plant species richness confined to semi‐natural grasslands. Still, they have been suggested as potential target sites for re‐creation of semi‐natural grasslands. We asked to what extent does fine‐scale variation in soil conditions, management history and site location effect local plant diversity in grazed ex‐arable fields. We examined local soil conditions such as texture, pH, organic carbon, nitrogen (N) and extractable phosphate (P) and effects on plant richness in ten pairs of grazed ex‐fields and neighbouring semi‐natural grasslands in different rural landscapes. Each grassland pair where in the same paddock. A multivariate test showed that site location and land use history explained more of differences in species richness than local soil property variables. Plant species richness was positively associated to grazed ex‐fields with low pH, low N and P levels. Sites with high plant richness in semi‐natural grasslands also had more species in the adjacent grazed ex‐fields, compared to sites neighbouring less species‐rich semi‐natural grasslands. Although both soil properties and species richness were different in grazed ex‐fields compared to semi‐natural grassland, the site location within a landscape, and vicinity to species‐rich grasslands, can override effects of soil properties. In conclusion, if properly located, ex‐arable fields may be an important habitat to maintain plant diversity at larger spatio‐temporal scales and should considered as potential sites for grassland restoration.     

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.     

The Influence of Time on the Soil Seed Bank and Vegetation across a Landscape‐Scale Wetland Restoration Project

Wicken Fen National Nature Reserve (NNR) in Cambridgeshire, U.K. is a wetland of international importance isolated in a landscape dominated by arable farming. The prospect of species extinctions within the NNR led to the creation of the Wicken Fen Vision, an ambitious project that will eventually expand the reserve boundary by the purchase and restoration of c.50 km² of arable land. We sampled three fields from each of three distinct age‐categories of restoration land (5, 15, and 60 years post‐arable), and three fields within the adjacent, undrained NNR, to determine (1) differences in seed bank composition across age‐categories, (2) relationships between restoration age, the seed bank and standing vegetation, and (3) changes in species traits across age‐categories. Historic arable management contributed to an apparent “vertical mixing” effect in the seed bank of the youngest two age‐categories, with associated and significant differences in species functional traits across the study area. Almost all plants associated with NNR vegetation were absent from restoration area seed banks and standing vegetation. Seed bank species common to all ages‐categories exhibited a bias toward moderate to high Ellenberg F (moisture) values, persistent seed banks, and lateral vegetative spread. Relatively short (c. 6 years) periods of drainage and plowing impact heavily upon seed bank diversity and soils, resulting in a lack of predrainage vegetation, even after decades of subsequent restoration adjacent to intact, species‐rich habitat. However, the seed banks of highly degraded fields can contribute toward the creation of novel wetland vegetation assemblages over time and under suitable environmental conditions.     

Limited Effects of Large‐Scale Riparian Restoration on Seed Banks in Agriculture

I examined the effect of riparian forest restoration on plant abundance and diversity, including weed species, on agricultural lands along the Sacramento River in California (United States). Riparian forest restoration on the Sacramento River is occurring on a large‐scale, with a goal of restoring approximately 80,000 ha over 160 km of the river. In multiuse habitats, such as the Sacramento River, effects of adjoining habitat types and movement of species across these habitats can have important management implications in terms of landscape‐scale patterns of species distributions. Increased numbers of pest animals and weeds on agricultural lands associated with restored habitats could have negative economic impacts, and in turn affect support for restoration of natural areas. In order to determine the distribution and abundance of weeds associated with large‐scale restoration, I collected seed bank soil samples on orchards between 0 and 5.6 km from adjacent restored riparian, remnant riparian, and agricultural habitats. I determined the abundance, species richness, and dispersal mode of plant species in the seed bank and analyzed these variables in terms of adjacent habitat type and age of restored habitat. I found that agricultural weed species had higher densities at the edge of restored riparian habitat and that native plants had higher densities adjacent to remnant riparian habitat. Weed seed abundance increased significantly on walnut farms adjacent to restored habitat with time since restored. I supply strong empirical evidence that large areas of natural and restored habitats do not lead to a greater penetration of weed species into agricultural areas, but rather that weed penetration is both temporally and spatially limited.     

Seed Removal and Predation as Factors Affecting Seed Availability of Tree Species in Degraded Habitats and Restoration Plantings in Rainforest Areas of Queensland,Australia

The post‐dispersal removal or predation of seeds of native tree species was investigated in Queensland, Australia, at degraded habitats and rainforest restoration sites where direct seeding might be used to facilitate tree regeneration (old fields or open habitats, lantana thicket, rainforest edge, and 5‐ and 10‐year‐old restoration plantings). Seed removal/predation was assessed in relation to tree seed weight and canopy density of the habitats during the wet season period. Results indicated that seed removal/predation imposes limitations on seed availability, particularly for small seeded species. In most situations, larger seeds were less removed/predated, most likely due to the more limited range of large seed consumers. The use of large, hard‐coated seeds may potentially reduce seed loss in open situations (from both seed removal and desiccation), unless large seed consumers frequent the site. Canopy cover exerted an influence on seed removal/predation, though trends varied in relation to site and the time of season. Broadcast sowing of seed under planted tree canopies at the more advanced stages of closure may in some areas result in higher seed removal/predation. Likewise, seeding in areas dominated by woody weeds may result in high seed losses to consumers such as rodents. Results suggested that undertaking direct seeding to coincide with the maximal period of fruit production may in some situations be beneficial to minimize seed loss. Overall, site context, canopy cover, and species selection appear to be important considerations when aiming to reduce loss of seeds to animal seed consumers in restoration work.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Successful restoration of abandoned terraced vineyards and grasslands in Southern Switzerland

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

Large herbivores as mobile links between isolated nature reserves through adhesive seed dispersal

Question: Do large herbivores contribute to the dispersal of plant seeds between isolated habitats by epizoochory? Location: Nature reserves in Flanders, Belgium. Methods: Epizoochory was studied by brushing plant seeds from the fur of 201 domesticated large herbivores (Galloway cattle, donkeys and horses), grazing in 27 Flemish nature reserves. Several herbivores were examined after transport between different nature reserves as part of the seasonal grazing system in Flanders, allowing detection of seed dispersal both within and between reserves. The seedling emergence method was used to identify the dispersed plant species. Results: In total, 6385 epizoochorous seeds from 75 species germinated, yet the real seed quantity was underestimated by the seedling emergence method. A wide variety of seed morphology, seed weights and plant heights was represented among the dispersed species, many of which had a transient seed bank. There was a gradual turnover in epizoochorous species composition in the course of the vegetation season, and seed dispersal occurred both within and between different nature reserves. Conclusions: Domesticated large herbivores, as models for wild mammals in the present and the past, are important dispersers of many plant species. Through seasonal grazing, the herbivores function as ‘mobile link organisms’, connecting isolated nature reserves through seed dispersal, possibly influencing vegetation development and long‐term survival of plant populations. As such, large herbivores are important instruments in ecological restoration, especially in fragmented ecosystems.     

The effects of management on vegetation trajectories during the early‐stage restoration of previously arable land after hay transfer

The restoration of floodplain grasslands has benefited from many studies of the underlying mechanisms. Among the operational tools that resulted, hay transfer is now used increasingly to alleviate the effects of limited seed dispersal and recruitment. To improve this method, we still need to understand how it can affect restoration trajectories, and particularly their direction and magnitude during the early stages of restoration. Based on concepts from the field of community ecology theory, we investigated the effects of early‐stage management through grazing or mowing on restoration trajectories after soil harrowing and hay transfer. We established a randomized block design experiment and quantified several community‐related metrics to formalize restoration trajectories for 3 years after hay transfer on a previously arable alluvial island in southwestern France. Whatever the management treatment, the species richness and evenness were significantly higher in hay‐inoculated than in control plots. This effect was linked to the recruitment of species originating not only from the reference grassland through hay transfer, but also from the seed bank, a well‐known effect of soil harrowing. Although generally oriented toward the reference grassland, the origin, direction, and magnitude of the trajectory of hay‐inoculated plots all depended on the management applied. Sheep grazing applied at the same time as hay transfer enhanced the recruitment of reference species as from the first experimental year, because it controlled aboveground competition and maintained the window of opportunity open for a sufficiently longer period of time. Our findings show that the type of management applied simultaneously to hay transfer influences the origin of a grassland trajectory, while its direction and magnitude are dependent on the management applied in subsequent years. Grazing immediately after hay transfer may be appropriate to accelerate the recruitment of species from the reference grassland.     

Recovery of native grass biodiversity by sowing on former croplands: Is weed suppression a feasible goal for grassland restoration?

Grassland restoration on former croplands offers good opportunity to mitigate the loss of grassland biodiversity. Weed suppression can be another benefit, which becomes increasingly important because of the high recent rate of abandonment of arable lands in Central and Eastern Europe. Our aim was to evaluate the usefulness of sowing two low-diversity seed mixtures followed by annual mowing, a frequently used restoration technique, in weed suppression. We found that rapidly forming cover of sown grasses effectively suppressed short-lived weeds and their germination except in the first year. The detected dense seed bank of short-lived weeds points out the possibility and threat of later weed infestation. In the short run perennial weeds cannot be suppressed easily by sowing and annual mowing. We found that the effectiveness of seed sowing followed by mowing in weed suppression can be different on sites with different history or seed mixture. Rapidly establishing perennial weeds, such as Agropyron species were only detected in former alfalfa fields; Cirsium arvense was found in former cereal and sunflower fields but not in former alfalfa fields. We found that the rate of weed suppression and success was influenced by the seed mixtures used. In several alkali restorations the high proportion of perennial weeds was detected in year 3. In loess restorations, much lower scores were typical. This was likely caused by the different seed mixture used. The loess seed mixture contained seeds of a clonally spreading tall-grass, Bromus inermis, which could compete more effectively with clonally spreading weeds, than could short grass species with or without tussock forming. Our findings indicate that post-restoration management require carefully designed actions that are fine-tuned addressing specific threats at the site level.     

Restoration of ditch bank plant species richness: The potential of the soil seed bank

Abstract. Small‐scale landscape elements, such as ditch banks, play an important role in preserving plant species richness in agricultural landscapes. In this study, we investigate whether the seed bank might be useful for restoring the above‐ground plant species richness. We studied the vegetation and seed bank composition at six species‐rich and six species‐poor ditch banks, where agri‐environment schemes are running to maintain and enhance ditch bank plant diversity. We show that the number of species in the seed bank was low, regardless of the number of species in the established vegetation. Moreover, the seed bank was always dissimilar to the established vegetation. Target species for nature conservation were occasionally present in the seed bank at both species‐poor and species‐rich sites, but rarely so if the species was absent from the established vegetation. We conclude that the potential use of the seed bank for restoration of ditch banks is minimal. At present, plant species richness seems to be largely controlled by germination opportunities; high biomass and competition appear to hamper germination at species‐poor sites. We recommend continued nutrient reduction at such sites. Soil disturbance measures and deliberate sowing should also be considered.     

The response of ecosystem CO<Subscript>2</Subscript> exchange to small precipitation pulses over a temperate steppe

We evaluated the process of salt-marsh colonization in early successional stages of salt-marsh restoration and investigated how the sequence of species establishment related to different success factors. Vegetation data were collected by permanent plots from the restoration site and adjacent, reference salt marshes during three consecutive periods. Seed length, width and mass were used as dispersal traits, and Ellenberg moisture, salinity and nutrient indices as indicators of site suitability. Seed production in the reference site and seed bank in the restoration site were also investigated. The establishment of salt-marsh species within the restoration site was rapid (less than 5 years). The cover of plant species was not correlated between the restored and the reference sites at the first year of restoration, but this correlation was significant during the following years. Seed availability was more important in explaining the sequence of species establishment than salt and nutrient-limitation tolerance. The first colonizers are known as massive seed producers, with shorter seed length and lower seed mass, which probably increased buoyancy. Among dispersal and site traits, seed length and mass, and in a less extent salinity and nutrients, indicated a relationship with new colonizers. Despite few species have not (yet) appeared in vegetation and seed bank in the restoration site, the existence of an existing salt marsh adjacent to the restoration site is shown to be vital for fast colonization of newly created intertidal areas.     

Soil seed-bank composition reveals the land-use history of calcareous grasslands

We compared soil seed banks and vegetation of recent (established on abandoned arable fields) and ancient (continuously managed as pastures at least since 1830) calcareous grasslands if there is any impact of former arable field use. The study was carried out in two regions of Southern Germany with well-preserved dry grassland vegetation: the western Jurassic mountains (Kaltes Feld) and the climatically drier eastern part of Southern Germany (Kallmünz).Total number of species in the seed bank was similar in both regions, but species composition partly differed, reflecting phytogeographical differences between the regions. The total number of emerged seedlings showed a large disparity (5457 compared to 2523 seedlings/m² in Kaltes Feld and Kallmünz, respectively).Though there were differences in seed bank composition and size, we found a uniform pattern of plant traits (affiliation to phytosociological groups, Raunkiaer plant life-forms and seed longevity), which depended on the age of the grassland.The main conclusion is that seed banks in contemporary calcareous grasslands still reflect the history of former land use – in this case arable cultivation, even though it occurred a long time ago (up to 150 years). Indicators of former arable fields are germinable seeds of weeds which have persisted in the soil to the present. By contrast, weedy species are completely absent from the seed banks of ancient grasslands. Soil seed banks of recent grasslands may be of substantial conservation importance because they may store seeds of rare and endangered weed species such as Kickxia spuria, Silene noctiflora and Stachys annua, the majority of which have already gone extinct from the current vegetation of the study sites.