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.     

Vegetation and seed bank in a calcareous grassland restored from a Pinus forest

Question: What is the importance of the seed bank in the maintenance of the restoration potential of a 60‐year‐old abandoned calcareous grassland overgrown by Pinus trees? Location: ‘Les Pairées’, province of Luxembourg, Belgium. Methods: The seed bank and the above‐ground vegetation were surveyed in three adjacent stands, previously forming a unique calcareous grassland: a 60‐year‐old Pinus forest, a four‐year‐old clear‐cutting and a typical calcareous grassland. Floristic diversity was compared among stands and between vegetation and seed bank. Results: The species richness of the vegetation and the seed bank was significantly lower in the Pinus forest. More floristic similarities were found between the clear‐cutting and the calcareous grassland. Seed bank was essentially transient, dominated by annual species. Its correspondence with the above‐ground vegetation was weak. Conclusion: Very few calcareous grassland species have persisted in the Pinus stand. Four years after clear‐cutting, the stand was nearing restoration towards a calcareous grassland. Seed longevity in the soil was not the most explicative factor. Dispersal of propagules from adjacent sources was also important.     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

Temporal Responses of Propagule Banks during Ecological Restoration in the United Kingdom

Successful ecological restoration is expected to be accompanied by change in the propagule bank. We tested for temporal change in the soil propagule bank at two Bracken (Pteridium aquilinum)‐infested sites in the United Kingdom (acid grassland and heathland), each with replicate experiments. A combination of bracken control (cutting, spraying, and combinations) and vegetation restoration treatments (seeding, fertilizer, harrowing) were applied. Soil propagule banks were sampled in 1998, 4–5 years after the start, and in 2003 after a further 5 years. We used univariate and multivariate statistical methods to investigate the response of the propagule bank to experimental treatment in space and time. Few effects were found in 1998, but after a further 5 years, the propagule bank size and composition changed significantly, implying that propagule bank development lags behind vegetation development. The effect of treatment on the propagule bank differed within sites. Thus, ecosystem development is occurring at different speeds and directions even in closely adjacent areas. Coupling between the propagule bank and vegetation changed between sampling times with either some new coupling or decoupling. At both sites, the propagule bank contained propagules of the target community and thus propagule bank development during restoration provides increased potential for vegetation recovery. However, it can take a considerable time for management effects to be detected in the propagule bank. Moreover, the effect or speed of effect is spatially variable. Continuing application of restoration treatments is recommended at heathland where there is a deep bracken litter layer.     

Germinable soil seed banks in abandoned grasslands in central and western Norway and their significance for restoration

Questions: Could the seed bank increase biodiversity during restoration of abandoned, species‐poor, formerly cultivated vegetation? Is it possible to identify how climate, soil and former and present management and vegetation affected the seed bank? Location: The study sites were eight abandoned grasslands, four in Orkdal, central Norway and four in Gaular, western Norway. Methods: 144 seed bank samples were collected from three depths. Each sample was sown and placed in a greenhouse. After three months, the trays were dried and stored at 4°C in a dry place for two months. This was repeated twice. Results: There was a separation of the two regions along the first DCA axis in both the seed bank and in the vegetation analysis and also a clear separation of the seed bank from the vegetation along the second axis. These results are caused by differences in former management as well as temperature, precipitation and soil type between Gaular and Orkdal. We found more annuals, short‐lived species and species demanding light open conditions in the seed bank than in the vegetation probably because these species have the capacity for producing persistent seeds. Most of the species found only in the seed bank were found in very few samples and with few individuals. Conclusion: These results suggest that it may be difficult to increase vegetation biodiversity through restoration of grasslands such as those investigated if the natural soil seed bank is the main seed source.     

Can soil seed banks contribute to the restoration of dune slacks under conservation management?

Questions: Does the soil seed bank resemble the former early successional stages of a dune slack system more than the established later successional vegetation? Does it have the potential to contribute to the conservation of a highly endangered habitat? Location: Dune slacks at Newborough Warren, UK. Methods: The composition of the soil seed bank in two depth layers was determined using the seedling emergence method between March 2004 and April 2005. Long-term monitoring data on the floristic composition of the established vegetation were obtained from the national conservation agency, and additional monitoring was undertaken in 2003. Floristic composition, seed weights, seed longevity of component species and Ellenberg indicator values were used to compare the seed bank and established vegetation. Results: The soil seed bank was diverse and contained typical dune slack species, species of early successional stages and species of conservation interest. A comparison between the composition of the seed bank and historical data on the composition of the established vegetation showed that the seed bank reflects earlier successional stages more closely than the current aboveground vegetation. This study increases the scarce information currently available on the seed bank ecology of several species, including two orchid species. Conclusions: The soil seed bank can be expected to contribute to vegetation change after disturbance. Stimulation of germination from the seed bank through management may contribute to the conservation of both characteristic and threatened species typical of dune slacks.     

Resilience,persistence and relationship to standing vegetation in soil seed banks of semi‐arid Australian old fields

Questions: Do soil seed banks of semi‐arid grasslands reassemble after abandonment from cultivation? Do seeds of native and exotic species persist in the soil? Does time since abandonment affect compositional similarity between the vegetation and seed bank? Does the seed bank contribute to resilience in the vegetation? Location: Native grasslands in northern Victoria, Australia. Methods: Seed bank sampling was conducted in spring and autumn over 3 yrs, across a 100‐yr chronosequence. Species richness, composition and germinant density were determined using the seedling emergence method. Seed persistence was assessed by comparing seed densities in spring and autumn. Seed bank composition was compared with the vegetation. Results: The spring seed bank was dominated at all stages by sedges and rushes; hence, native species richness and seed density were largely unaffected by abandonment. In autumn, grassland species contributed more to the seed bank, but richness was reduced after abandonment and showed little recovery, although seed density partially recovered. Seed bank composition showed some recovery in both seasons. Most species had low persistence in the soil. Compositional similarity between the vegetation and seed bank was greater in old fields than uncultivated grasslands in spring, but not autumn. Conclusions: Resilience varied among seed bank parameters and seed banks had low functional importance. Patterns in the seed bank followed, rather than caused, those in the vegetation. Thus, vegetation recovery cannot rely on the seed bank and persistent seeds were not the key mechanism of resilience in the vegetation.     

Seed bank diversity in mesic grasslands in relation to vegetation type,management and site conditions

Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow‐pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above‐ground vegetation, edaphic factors and land‐use history? 3. Are there differences in C‐S‐R plant strategy types and seed longevity under different management regimes? Location: Lahn‐Dill Highlands in central‐western Germany. Methods: Above‐ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above‐ground vegetation, site conditions and land‐use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above‐ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R‐strategy compared to hay meadows. In contrast, in hay meadows and meadow‐pastures proportions of S‐strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re‐establish from the seed bank after disappearance from above‐ground vegetation.     

Fire severity affects vegetation and seed bank in a wetland

Questions: How does the severity of prescribed fires affect vegetation and seed bank in a wetland? Location: A fire‐prone reed swamp in northern Japan (250 ha, 40°49′N, 141°22′E, <10 m a.s.l.). Methods: Vegetation, biomass and seed bank were monitored for the 2 yr after annual prescribed fires were discontinued. Plant communities were placed into three categories based on fire severity: high (H) – fire consumed litter completely; moderate (M) – fire removed standing litter but left wet fallen litter; and low (L) – fire incompletely removed standing litter and did not remove fallen litter. Soil samples were collected in autumn 2007 and early summer 2008, and germinable seed bank was investigated by greenhouse trials. Results: High fire severity increased diversity in the next growing season by the establishment of short herbs in the standing vegetation. The biomass of forbs and grasses was greater in H where Phragmites australis biomass was reduced. The density of seed bank was >30 000 seeds m^?2 throughout all the treatments. Perennial plants were dominant in the vegetation, while annuals, biennials and rushes were dominant in the seed bank. Small seeds were more abundant in the soil than in the litter. Qualitative and quantitative similarities between seed bank and the vegetation were low, and tended to be higher in H. Conclusions: Fire contributed to the development of diverse standing vegetation via the positive effects on seed bank dynamics, and can be considered a tool to maintain species‐rich marshes.     

Disturbance of suburban Fagus forests by recreational activities: Effects on soil characteristics,above‐ground vegetation and seed bank

Questions: How does recreational disturbance (human trampling) affect soil characteristics, the performance of the understorey vegetation, and the density and species composition of the soil seed bank in Fagus sylvatica forests? Location: Suburban forests near Basel, northwestern Switzerland. Methods: We compared various soil characteristics and the performance of the understorey vegetation in six beech forest areas frequently disturbed by recreational activities with those in six undisturbed control areas, in spring 2003. In the same forest areas, the soil seed bank was investigated using the seedling emergence method. Samples were obtained from soil cores in January 2003. Results: We found substantial changes in soil compaction, above‐ground vegetation and in the soil seed bank due to recreational activities. In frequently visited areas, soil compaction was enhanced which caused a decrease in cover, height and species richness of both herb and shrub layers. Compared with control areas, the number of trampling‐tolerant species of the seed bank was significantly higher in disturbed areas, and total species richness tended to be higher in disturbed than in control areas. Furthermore, the similarity in species composition between the above‐ground vegetation and seed bank was significant lower in disturbed than in control areas. Conclusions: The intensive use of suburban forests for recreational activities, mainly picnicking, affects the vegetation of natural beech forests. Our study indicates that a restoration of degraded forest areas from the soil seed bank would result in a substantial change of the vegetation composition.     

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.     

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.     

Assessing soil seed bank persistence in flood‐meadows: The search for reliable traits

Abstract. To assess seed bank persistence of target species in endangered flood‐meadows (alliances Cnidion and Molinion), we investigated established vegetation and soil seed bank of 46 plots for 3 yr and 2 yr, respectively. As traits of seed persistence we calculated various continuous indices that refer to the frequency and abundance of species in above‐ground vegetation and at different soil depths. Furthermore, we tested the significance and soundness of easily observed traits such as maximum seed density per plot and seed attributes (mass, size and shape) as predictors of soil seed bank features. In linear regression, SAI, the seed accumulation index, showed the best agreement (R²= 0.64) with the seed longevity index that was derived from the database by Thompson et al. (1997) for a set of 115 species. The second best predictor (R²= 0.39) of the seed longevity index was maximum seed density per plot in the lower soil layer (5–10 cm). Depth distribution indices and seed attributes showed weaker but still significant relations. The dynamic character of flood‐meadows was reflected by a large proportion of species with a strong tendency to accumulate seeds in the soil relative to their importance in above‐ground vegetation. Most of these species have a ruderal strategy, exploiting gaps after flood disturbances, while the dominants of flood‐meadows tended to have short‐lived seed banks. Compared to other grassland types, a relatively large proportion of rare and endangered target species can be expected to form long‐term persistent seed banks. However, only under marginal conditions that facilitate seed survival in the soil (e.g. fallow) are these persistent seed banks likely to contribute to restoration. We conclude that easily observed traits of persistence such as seed weight, size and shape do not meet the accuracy needed in scientific and practical applications. Thus, there is a crucial demand for further seed bank studies in poorly investigated habitats and of rare species.     

The persistence of calcareous grassland species in the soil seed bank under developing and established scrub

The relationships between the composition of the soil seed bank, the field layer vegetation, and the scrub canopy were investigated along a 69 m transect, grading from incipient woodland, through scrub, into intensively rabbit-grazed calcareous grassland. The results are used to assess the persistence of species associated with open calcareous grassland in the seed bank under developing scrub. Scrub age, composition and density, changed along the transect from the woodland to open grassland. A total of 35 forb and grass species were found in the field layer. The pattern evident in the scrub layer was also reflected in the herbaceous vegetation. The field layer in the most closed portion of the transect, where the scrub was oldest, was dominated by shade-tolerant species normally associated with woodland habitats. The abundance of these species decreased along the transect as the scrub age declined, and the field layer became increasingly dominated by species typical of open grassland. A total of 47 species germinated from the seed bank. Few species were recorded in the seed bank along the entire length of the transect. Overall, the seed bank was dominated by Hypericum perforatum and Centaurium erythraea, which accounted for 38.2% and 28.6% of emerging seedlings respectively. As with a number of similar studies, the composition of the seed bank had a low correspondence with the composition of the field layer vegetation. The results also emphasise that the composition of the seed bank can be viewed as an ecological palimpsest, with germinable seed of species from each stage of the old-field succession occurring in the soil. The seed bank is an important component in the re-vegetation of an area after disturbance such as scrub removal. This study supports the findings of previous research in showing that relatively few characteristic calcareous grassland species form persistent seed banks. The soil seed bank would therefore appear to be of limited value in the restoration of such grassland following scrub removal.     

Soil seed banks in Pinus ponderosa forests in Arizona: Clues to site history and restoration potential

Question: How does the relationship between the viable soil seed bank species composition and the above‐ground vegetation in northern Arizona Pinus ponderosa forests differ under varying historical land use disturbances (low, intermediate, high)? Is above‐ground vegetation correlated to the viable soil seed bank immediately following soil disturbance from restoration thinning treatments? Location: Northern Arizona, USA. Methods: Soil seed bank samples were taken along replicated transects and collected with a 5‐cm diameter bulk density hammer. Samples included a 5 ‐cm diameter O‐horizon sample (at varying depths) plus the underlying mineral soil to a depth of 5 cm. The seedling emergent method was used to quantify seed bank species composition and density. The herbaceous and shrub plant community was quantified along the same transects using the point intercept method. Results: Early‐successional or ruderal species were common in the soil seed bank at all three disturbance sites. Non‐native species, notably Verbascum thapsus, were more numerous (up to 940 seeds/m²) under high disturbance with overgrazing and logging, and less common or absent under low disturbance. Most viable seeds were found in the O‐horizon and the upper 5 cm of mineral soil; there was little correlation between species in the soil seed bank and the above‐ground vegetation. Conclusions: We recommend that restoration plans be geared toward minimizing activities, such as severe soil disturbance, that may promote the spread of non‐native invasive species, and that manual seeding be explored as an option to restore plant species diversity and abundance.     

The contribution of rewetting to vegetation restoration of degraded peat meadows

Question: What is the contribution of a rise in groundwater level to vegetation restoration of degraded peat meadows compared to abandonment only? Location: Abandoned peat meadows in the central part of The Netherlands. Methods: Comparison of species composition and species abundance of vegetation and seed banks of reference and rewetted peat meadows, using plant trait and seed bank analysis. Results: Vegetation of rewetted meadows shared on average only 27% of their species with the reference meadow, while this was 50% on average for species in the seed bank. Rewetted meadows had a lower total number of species and a lower number of wet grassland and fen species present in the vegetation, but had higher species richness per m², although evenness was not affected. Rewetting increased the dominance of species of fertile and near neutral habitats, but did not result in an increase of species of wet or waterlogged habitats. Re-wetted meadows were dominated by species relying mainly on vegetative reproduction and species with a low average seed longevity compared to the reference meadow. Conclusion: Rewetting was not effective as a restoration measure to increase plant species diversity or the number of wet grassland and fen species in the vegetation. If no additional restoration management is applied, the seed bank will be depleted of seeds of species of wet grassland or fen habitats, further reducing the chances of successful vegetation restoration.     

Can atmospheric input of nitrogen affect seed bank dynamics in habitats of conservation interest? The case of dune slacks

Questions: Does the increased atmospheric deposition of nitrogen, which can have major effects on the established vegetation of nutrient‐poor habitats, also impact germination from the soil seed bank? Location: Coastal dune slacks at Newborough Warren, Wales, UK. Methods: The effects of nitrogen addition (15 kg.ha^‐1.a^‐1) on seed germination from the soil seed bank were investigated using the seedling emergence method between September 2004 and February 2005. Results: More seedlings emerged from fertilised samples than unfertilised controls. Most species showed enhanced germination after fertilisation with nitrogen, with seedling numbers statistically significantly greater in nitrogen addition samples in a quarter of species abundant enough for analysis. Species that responded positively to fertilisation were species with low Ellenberg indicator values indicative of infertile sites. Conclusions: Most species showed increased germination after fertilisation with nitrogen, including early successional species normally growing in nutrient‐poor conditions. This suggests that the increased atmospheric deposition of nitrogen probably not only impacts on established vegetation, but also has the potential to alter seed bank dynamics.     

Shrubland Restoration Following Woody Alien Invasion and Mining: Effects of Topsoil Depth, Seed Source, and Fertilizer Addition

Invasion by woody alien plants, construction, and mining operations are among the major disturbances degrading vegetation in the Cape Floristic Kingdom, South Africa. The aim of this study was to assess whether native fynbos shrubland vegetation could be restored following dense alien invasion and disturbance by mining. An area supporting dense alien trees was cleared and topsoil was stripped and stockpiled to simulate mining disturbance. A field trial investigated the effects of topsoil depth, seed mix application, and fertilizer on native species recruitment and vegetation development over a three‐year period. Soil‐stored seed banks contributed 60% of the species recruited, indicating that areas invaded for three decades have good restoration potential. The addition of a fynbos seed mix, which included serotinous overstory species, improved both the richness and structural composition of the vegetation. Most species sown in untopsoiled plots established, but survival and growth was low compared to topsoil plots. Poor growth in combination with a lack of soil seed bank species, indicate that restoring a diverse and functional cover of indigenous vegetation on subsoil is not possible in the short‐term. Soil amelioration is required to improve rooting conditions and initiate ecosystem processes. Shallow and deep topsoil treatments yielded high plant density, richness, and projected canopy cover, but canopy cover was higher in deep topsoil plots throughout the trial. Fertilizer addition increased canopy cover in untopsoiled and shallow topsoil plots via an increase in alien annual species. Fertilizer addition ultimately may lead to increased native vegetation cover in untopsoiled areas, but as it increased proteoid mortality on deep topsoil plots, it is not recommended for sites where topsoil is available. A species‐rich and structurally representative fynbos community may be restored on topsoiled areas provided that the native disturbance regime is simulated and seeds of major structural guilds not present in the soil seed bank are included in the seed mix.     

The role of the soil seed bank in vegetation recovery on an oceanic island severely damaged by introduced goats

Question: Are the seed banks of an isolated subtropical oceanic island capable of naturally regenerating vegetation either with species of the historical forest community or with the existing grassland community after severe damage to the vegetation by goats? Location: Nakoudojima Island, Bonin Archipelago (Ogasawara Shoto), Japan. Methods: Soil samples were collected at 0–5 cm and 5–10 cm depths from seven plots in forests, grasslands, artificially matted areas and bare land. Soil seed banks were assessed using the seedling emergence method followed by the hand‐sorting of ungerminated seeds. We determined the size and composition of the seed banks in upper soil layers of plots and compared the seed banks to the standing vegetation. Results: A total of 12 220 seedlings belonging to 42 species from 20 families germinated. Total mean seed density (0–5 cm depth) was low in all plots within forest, grassland, and heavily degraded vegetation types (34.7 ± 8.6 to 693.5 ± 123.6, 58.6 ± 7.8 to 107.1 ± 10.0, and 1.1 ± 0.5 to 7.2 ± 2.3 seeds/m², respectively). Forbs and graminoids dominated the seed banks of grassland and forest plots including Cyperus brevifolius, Gnaphalium pensylvanicum, Oxalis corniculata and Solanum nigrum, and these alien species comprised 90% of the density of the seed bank. There was little correlation between seed banks and standing vegetation of the island (Sørensen similarity coefficient values 0.26 to 0.45). Conclusions: If natural regeneration occurs from the seed bank of the island, future vegetation will not move toward the original forest community, because the seed bank is dominated by non‐native herbaceous grassland species. Though isolated, a few forest remnants with low species richness could be an important source for the natural re‐establishment of forest on the island; however, seed availability may be limited by either poor dispersal or pollination so that woody species will probably recover very slowly on this goat‐impacted island.     

Distribution and effects of tree leaf litter on vegetation composition and biomass in a forest-grassland ecotone

Aims After abandonment of grasslands, secondary succession leads to the invasion by woody species. This process begins with the accumulation of tree litter in the forest–grassland ecotone. Our objectives were to determine the relationships between litter amounts and vegetation composition and cover along natural forest–grassland ecotones and to experimentally study the initial effects of tree litter accumulation on grassland vegetation and on microsite conditions.Methods We established 11 transects varying from 12 to 15 m in length in different forest–grassland ecotones in the Lahn-Dill highlands, Germany, and measured the mass and cover of tree litter and the cover and composition of vegetation at five sequential positions along each transect by using 1 m 2 plots with five replications. In a field experiment, we established plots subjected to different litter amounts (0, 200 and 600g m ?2) and evaluated changes in grassland vegetation, soil temperature and soil nutrient availability below the litter layer.Important findings Tree litter amounts decrease from 650 to 65g m ?2 across the forest–grassland ecotone. Vegetation changed from shrubs and annual species (adapted to more stressful conditions) in the forests edge to grasses, rosettes and hemirosette species (with higher competitive abilities) in the grassland. These anthropogenic forest–grassland ecotones showed abrupt edges, and the two adjacent ecosystems were characterized by different species pools and functional groups. In the field experiment, the presence of a litter layer reduced vegetation biomass and cover; the species richness was only reduced in the treatment with high litter (600g m ?2). Additionally, adding litter on top of vegetation also reduced thermal amplitude and the number of frost days, while increasing the availability of some nutrients, such as nitrogen and aluminium, the latter being an indicator of soil acidification. Adding a tree litter layer of 600g m ?2 in grassland areas had strong effects on the composition and diversity of grassland vegetation by reducing the cover of several key grassland species. In, or near, forest edges, litter accumulation rapidly changes established vegetation, microsite conditions and soil nutrients.