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.     

Establishment and persistence of target species in newly created calcareous grasslands on former arable fields

The effects of different restoration measures and management variants on the vegetation development of newly created calcareous grasslands were studied in southern Germany from 1993 to 2002. In 1993, fresh seed-containing hay from a nature reserve with ancient calcareous grasslands was transferred onto ex-arable fields with and without topsoil removal. Nine years after start of the restoration, the standing crop was lower and the cover of bare soil was higher on topsoil-removal sites than on sites without soil removal. Topsoil removal had a positive effect on the proportion of target species (class Festuco-Brometea), because the number and cover of productive meadow species (class Molinio-Arrhenatheretea) were reduced. Persistence of hay-transfer species and the number of newly colonizing target species were highest on topsoil-removal sites. On plots with and without soil removal, species richness and the number of target species increased quickly after hay transfer and were always higher on hay-transfer plots than on plots that had not received hay in 1993. In 2002, differences induced by hay transfer were still much more pronounced than differences between management regimes. Management by mowing, however, led to higher species richness, a greater number of target species and a lower number of ruderals in comparison to no management on restoration fields without soil removal. A detrended correspondence analysis (DCA) indicated that vegetation composition of the hay-transfer plots of the restoration fields still differed from the vegetation of ancient grasslands in the nature reserve. Vegetation of an ex-arable field in the nature reserve (last ploughed in 1959) showed an intermediate successional stage. In general our results indicate that the transfer of autochthonous hay is an efficient method for the restoration of species-rich vegetation, which allows not only quick establishment but also long-term persistence of target species.     

Heat increases germination of water-permeable seeds of obligate-seeding <Emphasis Type="Italic">Darwinia</Emphasis> species (Myrtaceae)

We examined the response of seeds to heat in four geographically restricted and one widespread species of shrubby Darwinia from the fire-prone region of southeastern Australia. These shrubs are killed by fire and rely on seed germination after a fire to maintain populations. We replicated the germination trials across several sites and several fruiting seasons for most species. Seeds had a high level of viability and were largely dispersed in a dormant state, except in D. glaucophylla, where seed dormancy varied significantly across fruiting seasons. The indehiscent fruit of all species readily imbibes moisture when wet and seeds are not considered to be ‘hard-seeded’. All species had increased seed germination in response to a limited range of heating temperatures (generally 80–100°C). Higher temperatures killed increasing proportions of seeds. This pattern was broadly consistent across species, population and seasons, although the proportion of seeds whose germination was promoted by heat varied from high (D. diminuta, D. fascicularis, D. glaucophylla) to moderate (D. biflora, D. procera). Our work highlights the importance of heat as a mechanism for influencing germination in species that are not hard-seeded. Consequently, soil temperatures during a fire should strongly influence post-fire germination levels in Darwinia. The roles of other cues that promote germination, i.e. smoke, seasonal temperatures and their interactions with heat, remain to be investigated.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Restoration of high species density in calcareous grassland: the role of seed rain and soil seed bank

Abstract. A small-scale restoration experiment concerning a species-rich calcareous grassland was conducted over a 7-yr period. 20 yr of abandonment (1971-1990) resulted in both a strong reduction of the number of species and an absolute dominance of the shrub Crataegus monogyna and the grass Brachypodium pinnatum - which contributed 98% of the above-ground herb phytomass. The plot (3 m x 6 m) was cleared in September 1990 and thereafter yearly mown in autumn. Small soil seed bank samples were taken immediately after clearing and again in the autumn of 1996. The inner part of the plot showed a reduced soil seed bank compared to the plot underneath the vegetation outside the plot in 1990. Six years later an increased number of seeds in the soil in this inside area could be recorded mainly in the upper soil layer, although the amount was only 40% compared to the outside or control area. Ca. 20% of the species present in the restored vegetation were recorded in seed traps during the main seed dispersal period in 1997. Restoration of high species richness in the plot was a fast process during the first few years: from seven species in 1990 towards 39 three years later. This was significantly faster than the restoration process of the calcareous grassland at the same slope 20 yr earlier. This was probably due to the increased size of the community species pool at the site as a result of appropriate management during the last decades. Relative species number increase was highest at the largest scale during this experiment, viz. at the total plot area. The success of restoration of species-rich, dry grassland vegetation largely depends on the interaction between several conditions, e.g. seed availability, either originating from the seed rain or soil seed bank, favourable germination conditions and a successful development to a mature, seed producing individual. Germination and establishment strongly depend on suitable vegetation structure as a result of sustainable management.     

The invasion of Pteridium aquilinum and the impoverishment of the seed bank in fire prone areas of Brazilian Atlantic Forest

Several studies have documented that fires are widespread in the tropics. Because fire in the Atlantic Rain Forest is rare, fire events and their consequences at the community level have not been evaluated. This study describes the composition of different seed banks in areas of Atlantic Forest. The study was carried out in the National Park of Tijuca in the state of Rio de Janeiro. Four different areas were chosen considering their differences on fire history. Fifteen soil surface samples (23 cm (centimeter) diameter ×7 cm height at 5 cm deep) were obtained from random locations within each site. The dominant family at all study sites except in the most preserved one were Dennstaetiaceae, followed by Melastomataceae and Poaceae. The family Dennstaetiaceae was exclusively represented by Pteridium aquilinum (Klf.) Herter. There were no significant differences among areas when comparing the number of seedlings excluding P. aquilinum. However, the number of P. aquilinum was higher in the most disturbed area, while Melastomataceae and Rubiaceae were more frequent in a less impacted area. Our results suggest that the seed bank in disturbed areas of the Atlantic Forest possibly will not contribute for forest restoration after disturbance.     

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.     

The effect of cold stratification and light on the seed germination of temperate sedges (Carex) from various habitats and implications for regenerative strategies

The germination responses of 32 temperate Carex species were tested in light and darkness at five constant temperatures and under one fluctuating temperature regime, before and after cold-wet stratification. Using a linear logistic regression model, the probability of germination tested across all species was found to be significantly higher after stratification, in light and at the fluctuating temperature. In addition, the probability increased with temperature. Stratification increased germination in 28 species and had very little or no effect on four species. There was almost no germination in darkness prior to stratification, and the germination in light was considerably higher in all but two species compared with that in darkness. Thus, it can be concluded that the Carex species tested have broadly similar germination response patterns. The fact that Carices can be released from high levels of primary dormancy by low-temperature stratification implies that they are spring germinators. A light requirement after stratification in the major fraction of seeds and the capability of almost all investigated sedges to respond to fluctuating temperatures make it likely that persistent seed banks are formed. Additionally, sedges generally seem to have a high temperature requirement for germination which prevents them from emerging at the very beginning of the growing season. Regeneration by seed is probably largely restricted to gaps resulting from late spring disturbances where buried seeds have an opportunity to germinate and grow. Differences in germination were apparent between species occupying different habitats. Overall germination was significantly higher in wetland species than in dry-site species, probably owing to the greater capability of wetland species to respond to fluctuating temperatures. Differences in germination between forest and open-site species can be attributed to the higher capability of forest sedges to respond to low temperatures and temperature fluctuations. The influence of seed weight on germination was not significant in the 18 species adapted to wet, open habitats. There was, however, a tendency for the germination percentages to be low for large-seeded Carices. The interpretation of habitat differences is difficult due to a positive correlation between seed weight and dry habitats.     

Predicting future community composition from random soil seed bank sampling – evidence from a drained lake bottom

Questions: What is the accuracy and reliability of the commonly used random soil sampling methodology for predicting seedling density, species richness and composition of the emerging seedling community? Location: Lake Kraenepoel, western Belgium. Methods: We compared density, species composition and observed and rarefactioned species richness of the seedling community emerging on a soft water lake bed exposed after drainage with the seedling community germinating in the laboratory from random soil samples in the same plots. Results: Seedling density did not differ between the two methods and there was a significant correlation between seedling density on the exposed lake bed and in the soil samples. This indicates that future seedling density can be reliably predicted based on soil sampling, in particular for the most abundant species. The most frequently occurring and abundant species among the seedlings in the soil samples were also the most frequent and abundant species germinating on the exposed lake bed. In contrast, species richness was much higher on the exposed lake bed than in the soil samples, and this difference was still significant for annual species after correction for differences in sampling intensity by rarefaction. We found no correlation between the number of species retrieved by the two methods. Although seedlings of rare and target species emerged on the lake bed, random soil sampling clearly failed to detect seeds of most of these species. Conclusions: Random soil sampling at a commonly used intensity and using the standard germination conditions can accurately predict future total seedling density and the density of the most abundant species. However, the method is not reliable for predicting the probability of establishment of populations of uncommon species. When executing a seed bank study, sampling intensity and germination conditions need to be adapted to the nature and the level of detail of the research question to be answered.     

Safe sites for establishment of Nymphoides peltata seedlings for recovering the population from the soil seed bank

The population of the endangered plant species Nymphoides peltata in Lake Kasumigaura, Japan, has been declining drastically recently, and seedlings that emerge from the remaining soil seed bank have entirely failed to establish. The decline has followed artificial control of the water level in the lake. To examine the environmental requirements at the seedling establishment stage, we investigated the survival and growth of emerged seedlings at various microsites on the lakeshore by manipulating environmental conditions such as exposure to waves, inundation, and light availability so as to include the conditions of the presumed safe sites. Seedling survival at the early stage was affected most by environmental variables relating to the susceptibility to inundation. Survival until seedling establishment was profoundly affected by light availability as well as by susceptibility to inundation. In all, 136 genets were established in quadrats less prone to inundation and with sufficient light availability, which is consistent with the presumed safe-site characteristics of the original water regime: bare ground exposed during the spring water-level drawdown. Recovering the original water regime with seasonal drawdown will be indispensable for recovering a self-sustainable population. However, as first-aid measures, active restoration efforts through adaptive management using the remnant soil seed bank are urgently needed before the soil seed bank becomes exhausted.     

The role of the soil seed bank in the unpredictable habitat of Anthemis chrysantha J. Gay,a critically endangered species

The ability of Anthemis chrysantha to form a soil seed bank (SB) was investigated in order to understand better the adaptation of this rare annual species to its arid and unpredictable natural habitat. The natural seed bank population was analyzed in five consecutive plant cohorts (2006–2010) by sampling at two different times: in May, after the germination period, and in October, after the first dispersal episodes due to the beginning winter rainfall. In addition, to determine the persistence in the soil of the two achene morphs of this species’ (white and dark achenes), an artificial SB was created where achenes were buried and exhumed successively after each season, during two years. In all the cohorts studied, seedling emergence from May samples indicated the existence of a persistent seed bank (PSB). Moreover, the dark achenes were largely responsible for the permanent fraction because, after two years of burial in the artificial SB, 85.0% of them remained apparently healthy without having germinated, versus 9.9% of white achenes. Both types of buried achenes exhibited an annual conditional dormancy/non-dormancy cycle, induced by low winter temperatures. The PSB dynamics appeared to oscillate between the minimum values at the end of the germination period in spring (up to 2000 achenes per square meter) and the maximum values of the dispersal episodes in early autumn (up to 6000 achenes per square meter), with fluctuations of achene density due to the variability in annual rainfall. Hence, the SB showed a decline due to the failure of fruiting in the 2008 cohort, caused by drought, although the low value of ca. 560 achenes per square meter was able to establish the following population. Our study highlights the importance of the PSB, which, in “bad” years, may be critical for the persistence of this species.     

Comparison between the soil seed banks of a burnt and an unburnt Quercus pyrenaica Willd. forest

Soil samples from burnt and unburnt sites dominated by Quercus pyrenaica Willd. forest in León province (NW Spain) were collected, separated into an upper layer (0–2 cm depth) from a lower layer (2–5 cm), put in a greenhouse, and seeds allowed to germinate.A total of 670 identified seedlings comprising 56 species and belonging to 24 families were recorded. Most seedling were herbaceous perennials (hemicryptophytes: 59%), followed by annuals (therophytes: 23%). Many species that germinated from buried seeds were not observed as plants at any site, and came from exogenous communities. The principal means of seed dissemination were anemochory (45%) and autochory (23%).The number of species as well as seedlings was not significantly different between soils from the burnt and unburnt sites, but seedlings were more numerous in samples from the unburnt sites, as the number of seedlings was always highest in the upper soil layer. Germination behaviour of four particular species was characterized.Seedlings from the seed bank temporarily were found in the early stages of recovery of the burnt Quercus pyrenaica forests.     

The ecology of the seed in Trapa natans var. Japonica in a eutrophic lake

The production, density and environment of buried seeds and the temperature response of germinating seeds in Trapa natans var. japonica Nakai were ascertained. The mechanism of high reproductivity, tolerance of the plant for eutrophication and the stability of the stand in an eutrophic lake are discussed. The mean fresh weight of seeds was about 7.2 g, and the density of liver and dead seeds in April ranged 66.7 to 80.0 m^-2 and 26.7 to 97.8 m^-2, respectively. The threshold temperature that breaks dormancy and promotes germination in the lake was the low temperature, 8°C. The dormancy of buried seeds was almost broken under the field condition in January. The rate of germination obtained by the field experiment, and the estimation from the density of buried seed and the number of seeds germinated at the stand, were 87% and 60%, respectively. It was estimated that about 30% of the seedlings died before the floating leaf reached water surface. A rosette produced less than three seeds a year, which were fewer than for another Trapa, but the size of the former was larger. The number of seeds changed year by year, but the size-distribution pattern of seeds did not change with years, nor was it affected by the density of rosettes. The reason why T. natans var. japonica could succeed in eutrophic waters was attributed to reproductive strategy. The stand could recover by increasing the number of rosettes through the great branching ability of the main stem, even when the density of buried seeds was greatly decreased in the previous year.     

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.     

Spatial variability of the soil seed bank in a heterogeneous ephemeral wetland system in semi-arid Australia

Soil seed banks are an important component of plant community diversity in ephemeral wetlands, allowing many species to persist through unpredictable periods of flood and drought. Spatial variation of extant vegetation in such habitats commonly reflects patterns of flood history and often varies predictably between broadly differing hydro-geomorphic habitat types. Here we investigate whether spatial variation of soil seed banks is similarly controlled by fluvial processes at this scale. Results are presented from a seedling emergence trial using samples collected from a range of habitat types, and at different scales within these, in the ephemeral Narran Lakes system in semi-arid Australia. Composition and structure of soil seed banks varied significantly between habitat types reflecting broad differences in flood frequency. As predicted, germinable seed abundance was found to be highest in intermediately flooded habitats. Variability in soil seed bank composition at a local scale was also found to be influenced by hydrology with greater spatial heterogeneity evident in the river channel as well as amongst the least frequently inundated riparian and floodplain habitats.     

The role of ants and mammals in dispersal and post-dispersal seed predation of the shrubs Grevillea (Proteaceae)

The role seed predators play in influencing the dynamics of plant populations has been little studied in Australia. The interaction of ant dispersal and seed predation on the soil seedbank in six shrubby species of Grevillea from the Sydney region of southeastern Australia was examined in selective exclusion experiments, seed array trials and placement of single seeds on the ground.Two distinct seed types in Grevillea were examined and different seed dispersal and post-dispersal seed predation patterns were associated with each: (a) seeds lacking an elaiosome were not attractive to ants and annual seed losses of between 82 and 95% were found in vegetation unburnt for greater than 8 years. Native rodents, Rattus fuscipes, and macropods, Wallabia bicolor, were responsible for these seed losses; (b) seeds with an elaiosome were rapidly handled by ants. Two functional types of ants were recognised. Most encounters were by ants that were small (Local) relative to seed size and these ants simply removed the elaiosome in situ or moved seeds only small distances (<20 cm). Some 0–24% of ant/seed encounters were by large (Removalist) ant species that were capable of moving seeds back to nests. In addition, Rattus fuscipes and Wallabia bicolor consumed at least 32–68% of seeds of Grevillea species with an elaiosome.Ants may reduce the overall levels of seed predation where seeds moved by Removalist ant species escape predation and are deposited in safe sites, hence allowing more seeds to reach the persistent soil seedbank. Mammals do not consume all seeds when ants are excluded, allowing for the potential for some seed escape from predation after seeds are discarded by Local ant species.     

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.