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.     

Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau

Background and Aims

Information on soil seed bank processes is crucial for understanding vegetation dynamics. Despite the documented importance of soil seed banks in many ecosystems, their role is not fully understood in some sensitive habitats, such as the alpine meadows of the Tibetan Plateau.

Methods

We studied the seasonal dynamics of the germinable soil seed bank under four disturbance intensities in an alpine meadow on the Tibetan Plateau as well as seed size distribution relative to disturbance intensity. Composition of the seed bank was compared with that of the standing vegetation.

Results

Density of buried seeds increased with disturbance intensity, but species richness and species diversity decreased. Seed density and species richness of the seed bank varied seasonally in all layers (0–2, 2–7, 7–12 cm) and the whole (0–12 cm). The species composition of seed bank was not significantly influenced by season. There was no trend in seed size distribution as disturbance increased. Seasonal seed bank turnover rates increased with increase in disturbance. The result of the NMDS showed that species composition of seed bank and vegetation exhibited a fairly uniform pattern in each season.

Conclusions

Although as a whole the species composition of the vegetation and seed bank showed a relatively low degree of similarity in each season, similarity was highest in the most disturbed habitat. There was no alteration in species composition of seed bank regardless of disturbance intensity, but seed density decreased as disturbance increased. Disturbances in alpine plant communities might increase persistence of regeneration niches. Regeneration from the seed bank together with vegetative reproduction contributed to aboveground vegetation in highly disturbed habitats. Clonal species played an important role in regeneration of vegetation in slightly disturbed areas, where there was little contribution of ruderals from soil seed banks.     

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.     

Comparison of aboveground vegetation and soil seed bank composition at sites of different grazing intensity around a savanna-woodland watering point in West Africa

Grazing removes a plant’s aboveground vegetative and reproductive tissues and can modify the soil seed bank, potentially impacting the restoration of preferred species. Knowledge about aboveground vegetation and species composition of soil seed bank and the processes that contribute to vegetation recovery on and surrounding watering points subjected to grazing is lacking. Successful restoration strategies hinge on addressing these knowledge gaps. We assessed the effects of livestock grazing on aboveground vegetation and soil seed bank characteristics along a river bank and surrounding areas subject to different grazing intensities and draw implications for restoration. Plots (50?×?50 m) were established along five transects representing differing levels of grazing intensity. Soil samples were taken from three layers within each plot to determine soil properties and species composition of soil seed bank using the seedling emergence method. Heavy grazing resulted in the disappearance of perennial grasses, a reduction in species diversity and a decrease in soil nutrients with increased soil depth. Overall, the similarity between the extant aboveground vegetation and flora within the soil seed bank was low. The soil seed bank was dominated by herbaceous species and two woody species, suggesting that many woody species are not accumulating in the soil. With increasing soil depth, the seed density and richness declined. Canonical correspondence analyses (CCAs) showed that emerged seedlings from the soil seed bank were significantly influenced by soil carbon, organic matter, total nitrogen, total potassium and soil cation exchange capacity. This finding suggests that current grazing practices have a negative impact on the vegetation surrounding watering points; hence there is a need for improved grazing management strategies and vegetation restoration in these areas. The soil seed bank alone cannot restore degraded river banks; active transfer of propagules from adjacent undisturbed forest areas is essential.     

The dynamics of three shrub species in a fire-prone temperate savanna: the interplay between the seed bank, seed rain and fire regime

A model was developed to assess how the seed rain and fire regime affect seed bank dynamics and seedling establishment of three native shrub species (Acanthostyles buniifolius, Baccharis pingraea and Baccharis dracunculifolia) with different regeneration strategies, in temperate South American savanna. Seed bank and seed rain were quantified for each species under different fire regimes, and their relative roles in regeneration were evaluated. All species had short-term persistent seed banks and high annual variability in seed production. A high proportion of seeds deposited in the seed rain produced seedlings after fire; few entered the soil seed bank. Fire killed a high proportion of the seeds in the soil seed bank. Seedlings derived from the seed rain had a higher probability of surviving for 2 years than seedlings emerging from the soil seed bank. In the absence of fire, establishment depended on germination both from the seed rain and the soil seed bank, whereas with annual fire, establishment was primarily dependent on germination of seeds arriving in the annual seed rain, regardless of species’ regeneration strategies. These results help to explain changes in the vegetation of South American temperate savannas as a result of changes in fire regime and grazing management during the last 50 years. By revealing the crucial roles of the soil seed bank and seed rain in regeneration, this study provides vital information for the development of appropriate management practices to control populations of shrub species with different regeneration strategies in South American temperate savannas.     

Long-term monitoring of the restoration and development of limestone grasslands in north western Germany: Vegetation screening and soil seed bank analysis

We analysed the long-term development of limestone grasslands in West- and Central Europe (Mesobromion: Gentiano-Koelerietum) under conditions of restoration comparing vegetation changes with the composition of its corresponding soil seed bank over a time period from 1992 to 2005. Because of optimal climatic, edaphic and geomorphologic conditions restoration of vegetation developed gradually into the optimal phase of the typical limestone grassland vegetation type. However, the development of the soil seed bank compared to that of the actual vegetation cover is much slower, and even after 13 years all three study plots did not reach the optimal phase compared to soil seed bank composition from well-developed control vegetation nearby. This finding agrees with the assumption that after changes of management limestone grasslands need decades or even centuries to develop into self-sustaining species-rich habitats. We also monitored the orchid Ophrys insectifera as target species and characteristic species of this type of plant association over the last 27 years. Here, population density increased continuously over time indicating that vegetation restoration efforts of the last decades have had a positive effect.We further documented that the data obtained for the soil seed bank largely depend on the applied method used for the characterization of the soil seed bank. The seedling emergence and the rinsing method provided different results, and in case of our study area the rinsing method was mostly superior. The rinsing method detected up to 20fold higher total seed numbers.     

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.     

Spatial relationships between the standing vegetation and the soil seed bank in a fire-prone encroached dehesa in Central Spain

Postfire vegetation regeneration in many fire-prone ecosystems is soil seed bank dependent. Although vegetation and seed bank may be spatially structured, the role of prefire vegetation patterns and fire in determining postfire vegetation patterns is poorly known. Here, we investigated the spatial patterning of species abundance and richness in the vegetation and seed bank of a Mediterranean encroached dehesa in Central Spain. The seed bank was studied with and without a heat shock simulating a spatially homogeneous fire. Semivariograms and cross-semivariograms showed that species richness in the vegetation was aggregated in patches, mainly of herbs, with highest values corresponding to high herb cover and low tree cover. Species richness in the seed bank was also structured in patches, but the spatial pattern was weak. Seedling density of germinates in the seed bank also showed weak spatial pattern. Heating increased overall germination and species richness, and the intensity of the spatial pattern of species richness, particularly of herbaceous species. However, seed bank density patterns disappeared after heat shock because of increased germination of shrubs without spatial pattern. Our results document that the spatial structure of plant richness in the vegetation may persist after fire due to the spatial patterns of herbaceous species in the seed bank, and that postfire species richness patterns can arise independently of fire intensity patterns. However, the spatial structure of the vegetation after fire can be altered by the feedback between shrub encroachment and an eventual fire because of the ubiquitous germination of shrubs.     

Seed bank assembly follows vegetation succession in dune slacks

Question: Is the seed bank in dune slacks during the whole successional range mainly composed of early successional species or does it vary according to the changing vegetation? Location: Belgium, western part of the coast. Methods: We investigated the soil seed bank composition using a seedling germination method in a chronosequence of 20 dune slacks, ranging in age from five to 55 yr. Results: Both seed density and species richness in the seed bank were very low in the first successional stages and increased with age, mainly as a result of increasing seed production. The similarity between seed bank and vegetation was higher in older slacks. A comparison of characteristics between seed bank and vegetation showed that the seed bank was, to a large extent, composed of later successional species. Occurrence patterns of individual species also showed that seeds become incorporated in the soil after the species has established in the vegetation. Conclusion: The seed bank is not likely to be the driving force for successional changes in the vegetation, and successional changes rely on dispersal. Some early successional species persist in the seed bank, but generally only in low numbers. The results also confirm that most typical dune slack species do not form persistent seeds, so that re‐establishment from the seed bank is not to be expected when the species has disappeared from the vegetation.     

Hidden biodiversity in the Arctic – a study of soil seed banks at Disko Island,Qeqertarsuaq, West Greenland

Seed dispersal is a key process in plant community dynamics, and soil seed banks represent seed dispersal in time rather than in space. Despite their potential importance, seed bank dynamics in the Arctic are poorly understood. We investigated soil seed banks and corresponding plant community composition in three contrasting vegetation types in West Greenland, viz. dwarf shrub heaths, herb slopes and fell‐fields. Through germination testing, 31 species were documented in soil seed banks. All of these were herbaceous, while no dwarf‐shrub species, which represents the dominating growth form in the above‐ground vegetation, were emerging from the seed bank. Consequently, across vegetation types, the lowest similarity between seed bank and above‐ground vegetation was found in dwarf shrub heath. Nine plant species were exclusively found in seed bank, all of which were non‐clonal forbs. Seed bank size (total number of seeds) and species richness seemed to increase with the level of natural disturbance. Additionally, we examined the effect of different experimental light and temperature conditions on the quantity and diversity of germinating seeds. The difference in diversity in vegetation and seed bank at the species level will impact population dynamics, regeneration of vegetation after disturbances and its potential to respond to climate change.     

Importance of the soil seed bank of disturbed sites in Chilean matorral in early secondary succession

Abstract. The dynamics of the seed bank may provide clues to the process of recovery of the vegetation of disturbed sites. The role of the seed bank may be more important in areas with a seasonal climate than in areas where seedling recruitment is not limited to one season. We studied the seed bank and the seed rain in three sites of the Chilean mediterranean-climate region (33° 48'S) which differed in the degree of anthropic disturbance: a closed-canopy, second-growth forest; an open matorral; and an old-field. Additionally, we tested the germination of seeds from the soil and from the current-year seed crop. The seed bank varied considerably between the two years of study, although no change in the vegetation could be observed. Seed density and species richness were lower in 1989 than in 1988. The seed bank of the second-growth forest changed less between years, while the old-field showed the largest change. The highest seed rain occurred under shrub patches in the open matorral, while few seeds fell in the spaces between shrub clumps or in the old-field. In the forest, seed rain was low and correlated with species cover. Germination was low (0 - 15%) in tests using either soil samples or fresh seeds. These results indicate that matorral succession is a very slow process, limited mainly by low germination and low arrival of propagules to open areas. Most woody species have animal-disseminated fleshy propagules. The presence of established shrubs which may serve as perches or refuges for animals increases species richness in the seed rain and the seed bank.     

Microhabitat variations and seed bank-vegetation relationships in a desert wadi ecosystem

Due to extreme variability in patterns of rainfall, plant seed banks are an important component of desert habitats. Here I report on effects of standing vegetation and three different microhabitats (channel, bank and terrace) on the soil seed bank of a desert wadi ecosystem in the Eastern Desert of Egypt. A total of 450 soil samples at 45 stands were collected to represent the different wadi microhabitats. The germinable seed bank was estimated by controlled counts of seedling emergence. The floristic composition, functional properties and diversity of the soil seed bank, as well as its similarity with the standing vegetation varied among wadi microhabitats. Such variation could be attributed to differences in disturbance intensity among microhabitats (terrace < bank < channel) and variation of soil factors along the microtopographic gradient. Channel showed the highest species richness and size of soil seed bank, followed by bank and then terrace. Moreover the Shannon index of diversity of the seed bank and its similarity with standing vegetation were significantly greater in both channel and bank microhabitats than in terrace. At the level of plant functional groups, number of seeds of annuals was higher in both channel and bank than in terrace. Shrubs were more abundant in seed banks of channel compared to terrace. The size and species richness of seed bank were increased with the total plant cover, annual/perennial ratio and species richness of the standing vegetation.     

Modeling and Analysis of a Density-Dependent Stochastic Integral Projection Model for a Disturbance Specialist Plant and Its Seed Bank

In many plant species dormant seeds can persist in the soil for one to several years. The formation of these seed banks is especially important for disturbance specialist plants, as seeds of these species germinate only in disturbed soil. Seed movement caused by disturbances affects the survival and germination probability of seeds in the seed bank, which subsequently affect population dynamics. In this paper, we develop a stochastic integral projection model for a general disturbance specialist plant-seed bank population that takes into account both the frequency and intensity of random disturbances, as well as vertical seed movement and density-dependent seedling establishment. We show that the probability measures associated with the plant-seed bank population converge weakly to a unique measure, independent of initial population. We also show that the population either persists with probability one or goes extinct with probability one, and provides a sharp criteria for this dichotomy. We apply our results to an example motivated by wild sunflower (Helianthus annuus) populations, and explore how the presence or absence of a “storage effect” impacts how a population responds to different disturbance scenarios.     

Seed Bank Effects on Recovery after Disturbance in Reconstructed Tallgrass Prairies

Litter‐removing disturbances such as fire in grasslands temporarily increase available resources for plants, opening a window of opportunity for new establishment as communities recover. At this time, new individuals or species could be added to the community as a result of germination from the local seed bank. In reconstructed grasslands this may be problematic, as the seed bank may contain a suite of undesired species reflective of prior and surrounding land uses. In two, 25‐year‐old, low‐diversity reconstructed grasslands, we tested the effect of local seed bank establishment following litter‐removing disturbance using seedling removal plots (1 m²) and plots where natural seedling establishment was allowed. Following disturbance, the vegetation was either left intact or hayed to enhance seedling establishment (a common practice following inter‐seeding efforts). Although the seed bank and seedling community were dominated by resident grasses (Andropogon gerardii and Poa pratensis), recruitment from the seed bank increased species richness and reduced evenness through the addition of forb species (including Cirsium arvense) in one of the study sites. Haying temporarily altered the abundances of the dominant grasses, but did not consistently affect seedling recruitment. Disturbances that facilitate seed bank recruitment may promote establishment of undesired species within reconstructed grassland communities, and we need to take steps to better manage the contributions into and recruitment from the seed bank to reconstruct sustainable grasslands.     

Fire severity drives variation in post-fire recruitment and residual seed bank size of <Emphasis Type="Italic">Acacia</Emphasis> species

Very high-severity fires are a component of many fire-prone ecosystems, yet are often viewed as detrimental to vegetation. However, species in such systems are likely to have adapted to persist under a fire regime that includes high-severity fires. We examined how fire severity affects post-fire recruitment and residual seed banks of Acacia species and whether severity may affect plant responses to fire intervals. Nine sites of either high or low burn severity were identified after a large-scale mixed-severity fire in Warrumbungle National Park, south-eastern Australia. Transects were used to sample above-ground woody plant density. Seed bank size was surveyed by soil extraction from two depths and manual searching for seeds. Residual soil seed bank and recruitment were compared across the two burn severities. Acacia seedling density was higher in areas burnt at high severity, indicating that increased severity triggers increased germination from the seed bank. Size of residual seed bank was smaller after high-severity fire, but varied between species, with few Acacia cheelii seeds remaining despite high above-ground abundance. In contrast, A. penninervis retained a small residual seed bank. There was little evidence of negative effects on populations of Acacia species after high-severity burns. However, we found that high fire severity may impact on the ability of a species to persist in response to a subsequent short fire interval. Fire management for maintaining biodiversity needs to consider other key aspects of the fire regime, including severity and season, rather than focusing solely on fire frequency.     

Can the soil seed bank contribute to revegetation of the drawdown zone in the Three Gorges Reservoir Region?

The drawdown zone of the Three Gorges Reservoir Region was assumed to be completely formed in 2009 and the water level would range from ~145 m in flood season (summer) to ~175 m during non-flood season (winter). The soil seed bank is an important propagule source for vegetation restoration. In order to evaluate the potential of the soil seed bank to revegetate the drawdown zone of this region, we examined the quantitative relationships between the germinable soil seed bank and the established vertical and horizontal vegetation patterns. A total of 45 soil samples at four sites was collected to examine seed bank density, species richness, and composition using the seedling-emergence method. Forty-five species (from 20 families) germinated from the soil seed bank, and the average seed density was 4578 m⁻². The seed bank was dominated by annual plants, suggesting reestablishment of some above-ground species was plausible. However, most established woody plants and perennials were absent from the seed bank indicating a low probability of reestablishment for non-annuals through the seed bank. Thus, due to low species compositional similarity to extant vegetation and the dominance of annual plants, the soil seed bank had a low potential to restore pre-dam vegetation in the drawdown zone of the Three Gorges Reservoir Region, but its potential as a propagule source should be considered regarding the management of the drawdown zone for vegetation cover.     

Relationships of marsh seed banks to vegetation patterns along environmental gradients

• 1 The relationship of the seed bank to the vegetation of a freshwater marsh was studied along gradients of water depth and soil organic matter content. Characters examined included standing crop, seedling density, and species composition, distribution and richness.
• 2 The seed bank differed from the vegetation in that only nine of twenty-seven species were present in both, abundant seed-bank species were uncommon as adults, and adults showed different distributions along a gradient of soil organic matter content whereas their seeds were most abundant in soils with high organic matter.
• 3 The seed bank resembled the vegetation in that separate multivariate analyses of the communities revealed that variation in the species composition of each was significantly correlated with water depth and soil organic matter content. Further, species richness in both communities decreased with water depth and increased with soil organic matter content. Lastly, the standing crop of the vegetation and the number of seedlings both decreased with water depth and increased with soil organic matter.
• 4 Consideration of spatial patterns and environmental gradients revealed more similarities between vegetation and seed banks than were obtained by comparing species lists. The results suggest that artificial stimulation of seed bank germination for management purposes will not produce vegetation changes as large as those suggested by differences in species lists.

     

A model for small‐scale seed bank and standing vegetation connection along time

Since inter‐annual climatic variability influences composition and structure of seed bank and extant vegetation, it is expected that it also affects the relationship between both compartments at small scales along time. We hypothesize that seed bank and aboveground vegetation are closely and sequentially linked at small spatial and time scales. We conducted a field spatially explicit investigation to explore the spatial and temporal relationship between both compartments. Abundance, composition and spatial structure at different small‐scales of seed bank and aboveground vegetation were analysed through 100 permanent plots during two consecutive (dry and wet) periods of growth. Following a conceptual transition path model, we analysed changes in correlation values in composition and spatial aggregation between both compartments along time including seasonal variations of seed bank. Shape of spatial structures were evaluated using partial Mantel correlograms. Annuals and perennials guilds were studied separately. During the wet year, annuals increased their aerial spatial aggregation and cover, whereas the opposite happened for perennials. Density and spatial aggregation of the seed bank increased for both guilds in the two seasons following the rain period, especially in annuals. The clumped structure of the transient seed bank is a consequence of the addition of spatial structure of extant vegetation and the persistent seed bank. Expression of the persistent seed bank is much lower during the dry period for the annual guild. In spite of the strong yearly variability, the community maintained a highly structured spatiotemporal pattern. The mechanisms promoting this stability differed for annual and perennial guilds. Temporal persistence in perennials relied mainly in established plants longevity, whereas annual guild persistence depended solely on seed bank. This tight structure was coherent with the existence of successional dynamics in the community, although persistent seed bank could moderate the pace. Longer term studies of seed bank–standing vegetation dynamics are required to fully understand this process.     

Ecosystem response to removal of exotic riparian shrubs and a transition to upland vegetation

Understanding plant community change over time is essential for managing important ecosystems such as riparian areas. This study analyzed historic vegetation using soil seed banks and the effects of riparian shrub removal treatments and channel incision on ecosystem and plant community dynamics in Canyon de Chelly National Monument, Arizona. We focused on how seeds, nutrients, and ground water influence the floristic composition of post-treatment vegetation and addressed three questions: (1) How does pre-treatment soil seed bank composition reflect post-treatment vegetation composition? (2) How does shrub removal affect post-treatment riparian vegetation composition, seed rain inputs, and ground water dynamics? and (3) Is available soil nitrogen increased near dead Russian olive plants following removal and does this influence post-treatment vegetation? We analyzed seed bank composition across the study area, analyzed differences in vegetation, ground water levels, and seed rain between control, cut-stump and whole-plant removal areas, and compared soil nitrogen and vegetation near removed Russian olive to areas lacking Russian olive. The soil seed bank contained more riparian plants, more native and fewer exotic plants than the extant vegetation. Both shrub removal methods decreased exotic plant cover, decreased tamarisk and Russian olive seed inputs, and increased native plant cover after 2 years. Neither method increased ground water levels. Soil near dead Russian olive trees indicated a short-term increase in soil nitrogen following plant removal but did not influence vegetation composition compared to areas without Russian olive. Following tamarisk and Russian olive removal, our study sites were colonized by upland plant species. Many western North American rivers have tamarisk and Russian olive on floodplains abandoned by channel incision, river regulation or both. Our results are widely applicable to sites where drying has occurred and vegetation establishment following shrub removal is likely to be by upland species.     

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

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