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 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.     

The contribution of persistent soil seed banks and flooding to the restoration of alluvial meadows

Restoration of species-rich flood meadows impoverished by agricultural intensification is an important challenge. The relationships between flooding regime and soil seed bank were compared in three successive meadow communities (hygrophilic, mesohygrophilic and mesophilic) distinguished along a topographic and hydric gradient. Differences in flood duration and frequency between the three associations allowed the study of the contribution of floods to soil seed bank richness and density. No significant difference was found in species richness among the three soil seed banks, whereas the densities were significantly higher in the wettest community. The three seed bank compositions were clearly distinguished along the hydric gradient. In fact, the three seed bank types constituted a species poor version of the meadow communities to which they belong. Flood contributions appear to play a minor role in seed bank enrichment. Thus, seed dispersal by flood water would probably be insufficient to enable the restoration of alluvial meadows.     

Effects of topsoil removal,seed transfer with plant material and moderate grazing on restoration of riparian fen grasslands

Question: How do moderate grazing, topsoil removal and hay transfer affect species diversity and abundance on a eutrophic fen grassland site? Location: Northern Germany. Method: A three-factorial field experiment with the factors grazing, topsoil removal and hay transfer of diaspore-rich material was established in 2001. Soil nutrients and seed bank were analysed at the beginning of the experiment, species composition and vegetation development was monitored for four years (2002–2005). Results: Topsoil removal had a significant effect on the abundance of different plant species groups: resident vegetation of agricultural grasslands was suppressed, while clonal reed species were facilitated in recolonising the area. The establishment of regionally rare and endangered species of nutrient-poor fens and wet meadows introduced with hay was achieved mainly on plots with topsoil removal, with the exception of Rhinan-thus angustifolius, which also established on plots with intact topsoil. Effects of grazing after four years of experiments were of minor influence on species composition. Conclusion: The establishment of target plant species of nutrient-poor fens is most successful when both an adequate number of viable diaspores and suitable sites for germination and establishment are available. In our experiment this was achieved by the combination of topsoil removal and hay transfer. We recommend this combination, together with continuous management (grazing/cutting), for further restoration in fen grasslands.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Effect of flooding and draw‐down disturbance on germination from a seashore meadow seed bank

Abstract. The objective of this study was to investigate the effects of flooding and draw‐down on the germination from the coastal grassland seed banks and to determine whether the effect of flooding varies between the delta and the seashore. Seed bank samples were collected from three shore transects in SW Finland, two on the shore of the Baltic Sea and one on the delta of River Kokemäenjoki. Samples were germinated in non‐flooded and flooded conditions for over a month, after which both treatments were maintained in non‐flooded conditions. A total of 9267 seedlings of 47 species germinated and mean density of seeds in the soil was ca. 84 000/m². Most of the seedlings were monocots (98%) and perennials (98%). Ca. 30–40% of the species found in the above‐ground vegetation had a seed bank including the majority of the most abundant species. The number of seeds and species richness increased as the organic layer became thicker. The organic layer was thicker in the seashore samples and the seed bank was significantly larger than in the delta. The flooding and draw‐down treatment significantly increased the number of germinating seedlings in the seashore and also increased species richness in two transects, one in the delta and the other in the seashore. Two species, Schoenoplectus tabernaemontani and Typha latifolia, had significantly higher germination in the flooded treatment than in the non‐flooded. Apparently, many species in these coastal grasslands have adapted to flood disturbance and for seeds of some species flooding may work as a positive signal, possibly breaking dormancy.     

Dispersal,diversity and distribution patterns in pioneer vegetation: The role of river‐floodplain connectivity

Question: Are there changes in dispersal patterns in floodplain pioneer vegetation with effects on seedling number, species richness and species composition along a gradient of declining river‐floodplain connectivity? Location: Middle Elbe river floodplain, Germany. Methods: An experiment with five treatments was set up along a gradient of declining river‐floodplain connectivity, partitioning seedlings into three groups: (1) emerging solely from water dispersed seeds, (2) from wind/animal dispersed seeds and (3) from the soil diaspore bank. Two controls were established: without any manipulation and exclusion of all seeds. The results were compared with those of vegetation and soil sampling to evaluate the representativeness of the experimental sites in terms of species composition, diversity, seedling number and soil parameters. Results: Water dispersal and the soil diaspore bank were the major dispersal strategies shaping floodplain pioneer vegetation at the Middle Elbe river. The number of seedlings, species richness and the variation in species composition in these habitats depend on the degree of connectivity. The seedling number and species richness is highest in sites of permanent or almost permanent exchange with the main channel, where water dispersal additionally contributes to the number of seedlings grown from the soil seed bank. Conclusion: The results underline the importance of river‐floodplain ecotones as sink habitats for water‐dispersed seeds. Considering the strongly reduced river‐floodplain interactions due to dykes and other engineering structures, management strategies are necessary to improve connectivity and the renewal of fluvial land forms.     

Restoration success of low‐production plant communities on former agricultural soils after top‐soil removal

Abstract. The success inrestoring seven low‐production vegetation types on former agricultural soil after top‐soil removal was investigated. The colonization and establishment of target species in permanent plots was recorded during the first nine years after restoration measures were taken. For each permanent plot abiotic site conditions were used to determine which of the vegetation types could persist there. A comparison of the actual vegetation in the permanent plots with reference releves of the selected vegetation type revealed a gradually increasing similarity during consecutive years for five vegetation types. This was due to the occurrence of an increasing number of target species and the number of permanent plots they occurred in. However, nine years after top‐soil removal a large number of the target species were still lacking from the vegetation in the permanent plots, although most were present in the local species pool. Seed dispersal therefore seems to be a major limiting factor for restoration of these low‐production vegetation types on formerly agricultural soils.     

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.     

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.     

Frugivorous birds mediate sex‐biased facilitation in a dioecious nurse plant

Abstract. Facilitation by dispersal occurs if the nurse plant acts as a focus which is actively selected by seed dispersers and enhances the fitness of the facilitated plant. Sex‐biased facilitation may be produced if seed dispersers tend to concentrate the seeds under female, fruit‐bearing plants of dioecious species more often than under conspecific males. Juniperus sabina is a dioecious shrub with a prostrate growth form from Mediterranean high mountains that modifies many microhabitat characteristics related to seedling establishment and survival. Soil water availability, maximum soil temperature in summer, organic matter and total nitrogen content, were different on open ground as compared with beneath J. sabina shrubs, irrespective of its sex. Other studied characteristics such as soil bulk density and soil compaction after rain did not differ between the microhabitats considered. Some species, such as Juniperus communis, Pinus nigra, Helleborus foetidus and Euphorbia nicaeensis, are spatially associated to J. sabina shrubs, strongly suggesting a facilitative role. The anemochorous P. nigra and myrme‐chorous H. foetidus and E. nicaeensis did not associate preferentially to any sex of J. sabina. Only J. communis, an endozoochorous species sharing the same bird dispersers as J. sabina, presented a female‐biased spatial association with the nurse plant. Seed dispersal mediated by birds attracted by the fruit‐rewarding females of J. sabina explains the sex‐biased spatial pattern of Juniperus communis.     

Seedling recruitment in flood‐meadow species: The effects of gaps,litter and vegetation matrix

Question: Is the failure of establishment of rare flood‐meadow species in habitat restoration primarily due to seed or microsite limitation? How do sown species respond to artificially created gaps and added litter at two neighbouring sites with similar physical conditions but contrasting vegetation matrix (young arable fallow field vs species‐poor meadow sward)? Location: Upper Rhine valley, southwestern Germany, 85 m a.s.l . Methods: Seeds of six typical flood‐meadow species were sown in four treatment combinations of the factors gap creation and litter addition. Seedling recruitment was monitored for three years. Results: Five of the six species established successfully at both sites largely irrespective of treatments, indicating seed limitation. Only in the small‐seeded Arabis nemorensis, which was revealed to be strictly gap‐dependent at the meadow site, could an obvious microsite limitation be shown. The non‐significance of gap treatments in all other species at the relatively high productive meadow site is probably due to biomass removal by mowing in early summer. Only at the extremes of the seed size spectrum did the results meet predictions of plant ecological theory, such as the strict gap dependence of small‐seeded species in closed swards or the positive to neutral response of large‐seeded species to litter layers. Conclusions: Species identity was revealed to be the major factor influencing differences in recruitment. Due to the lack of a general trend in the response towards treatments the results support conceptual models that describe the interplay of facilitation and interference as a highly dynamic equilibrium, driven by variable abiotic and biotic marginal conditions.     

Inter-annual variation in the soil seed bank of flood-meadows over two years with different flooding patterns

Persistent soil seed banks are presumed to play a significant role in bridging and exploiting the effects of major flood disturbances in riparian systems. However, presently there is little empirical data on the correlation between flooding and seed bank dynamics. The objective of this study was to assess the impact of inter-annual flooding variation on soil seed bank dynamics in flood-meadows. We analysed seed bank composition at 46 plots along the northern Upper Rhine River, Germany, after two periods with contrasting flooding conditions (1996–1997: low and rare flooding; 1998–1999 high and very frequent flooding). Between both sampling occasions the total number of seedlings emerging from the seed bank decreased by 50% and average species-richness per sample by 30%, irrespective of the sampling depth. Similar decline occurred in mesophytes and wetland species as well as over different functional groups, with the exception of legumes. Considerable compositional changes in the seed bank were also indicated by shifts in DCA ordination and by the comparison of similarity matrices from both years with the Mantel test. The Mantel test also confirmed that compositional changes were more pronounced in the seed bank than in aboveground vegetation. There was poor correlation between the decline in total seed densities and species in the soil seed bank and environmental variables such as flooding duration and ecological groupings such as floodplain compartment and seed bank community types. Further distinct patterns of decline and persistence were evident only at the species level. While 21 species displayed a significant decline, only two species increased. Annual arable weeds and perennial ruderals with high temperature or nutrient requirements to break dark dormancy were amongst the most significantly declining species. In contrast, there was no decline in typical winter annuals and certain perennials that preferably germinate in autumn and fall into dormancy at the beginning of the warm period. Generally, differences in germination requirements and dormancy cycles of species gave the best explanation for the patterns of decline and persistence after intense flooding. Given these findings, the heavy depletion of the persistent soil seed bank we observed is most likely attributed to a post-flood germination flush due to very favourable recruitment conditions prevailing during the draw-down period after early summer flooding in 1999. The results of our study suggest that persistent soil seed banks are of outstanding importance to bridge and exploit the highly variable hydrological conditions that are typical of flood-meadows. Thus, they play a significant role in maintaining the floristic diversity of this habitat type over time and space. This revised version was published online in June 2006 with corrections to the Cover Date.     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

The interrelationship between productivity,plant species richness and livestock diet: a question of scale?

Question: What relationship exists between productivity, plant species richness and livestock diet? Are the results dependent on scale? Location: A sheep‐grazed Koelerio‐Corynephoretea sandy habitat of the northern upper Rhine (Germany) as a low productivity model system. Methods: The investigation was carried out for three years at a fine scale (2 m²) and for two years at a broad scale (79 m²). Productivity was measured by means of weighed above‐ground phytomass for fine scale and colour‐infrared (CIR) aerial photographs of the same system for fine and broad scales. For both scales, total numbers of vascular plant species and numbers of endangered vascular plant species were extracted from current vegetation relevés. Additionally, we obtained data on livestock diet (grazed phytomass, crude protein content). Results: Statistical analyses show an influence of the year on all variables; relationships between variables are not significant in every year. Species richness and number of endangered species are negatively related to productivity at fine scale while crude protein content and grazed phytomass are positively related to productivity. At the broad scale the diversity‐productivity relationship shows a ‘hump’ with highest species numbers in middle pioneer stages; numbers of endangered species are highest in all pioneer stages. Conclusions: We found a strong impact of scale and year on the diversity‐productivity relationship. It is inappropriate to analyse only small plots (2 m²), and it is necessary to study different years. This vegetation complex is dependent on grazing impact; thus there is an inversely proportional relationship between nature conservation value (high diversity) and livestock nutrition.     

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.     

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.