El Niño effects on soil seed bank dynamics in north-central Chile

The soil seed bank was monitored in four 75×75 m plots over 6 years (1990–1995) in an arid thorn scrub community in north-central Chile. Sixty-six species were identified. Total seed densities ranged from 2,000 to 42,000/m². Average mass of shrub seeds was significantly greater than that of other growth-forms. Between 70 and 90% of the seeds were less than 1 mg, with those in the 0.51–1.00 mg size class being most numerous. Seed densities were highly variable between years as well as within years, but were also closely associated with plant cover patterns and rainfall regime. Higher seed densities were found in wet years, and in samples taken in early summer and early autumn (i.e., after seed set); the lowest seed densities were in late winter (i.e., after annual plant germination). The annual plant species with the highest cover were also the most abundant in the soil seed bank and exhibited the largest seed density fluctuations. In general, seed densities were 5- to 10-fold higher during the 1991–1992 El Niño/southern oscillation (ENSO) years than non-ENSO years, showing the importance of this phenomenon for seed bank replenishment in the arid region of Chile.     

Does invasion by an alien plant species affect the soil seed bank?

Questions: How does invasion affect old‐field seed bank species richness, composition and density? How consistent are these effects across sites? Does the soil seed bank match vegetation structure in old‐fields? Location: Menorca, Balearic Islands, Spain, western Mediterranean basin. Methods: We monitored seed germination in soils from old‐fields that were both uninvaded and invaded (legacy effect) by the annual geophyte Oxalis pes‐caprae. We also added O. pes‐caprae bulbs to uninvaded soils to test O. pes‐caprae interference with seedling emergence (competitive effect). We compared species composition in the seed bank with that of the vegetation. Results: Species richness in the seed bank and in the vegetation was not significantly different between invaded and uninvaded areas. Uninvaded areas did not have larger seed banks than invaded areas. More seedlings, especially of geophytes, emerged when O. pes‐caprae bulbs were added to the soil. Species similarity between invaded and uninvaded areas was higher in the seed bank (74%) than in the vegetation (49%). Differences in species composition were as important as differences among sites. The degree of species similarity between the seed bank and the vegetation was very low (17%). Conclusions: Despite invasion by O. pes‐caprae not affecting species richness, the variation in the seed bank species composition in invaded and uninvaded areas, and the differences between the seed bank and the mature vegetation, highlights that even if the invader could be eradicated the vegetation could not be restored back to the exact composition as found in uninvaded areas.     

Can a seed bank maintain the genetic variation in the above ground plant population?

There are indications that a persistent seed bank can protect small and isolated plant populations from local extinction. Genetic mechanisms contributing to this phenomenon are the increase of local effective population size – and hence the decrease of genetic drift – through a reservoir of persistent seeds, and the accumulation of intergenerational genetic diversity in the seed bank. To find evidence for these mechanisms, we conducted two formal meta-analyses. First, we analyzed 42 published habitat fragmentation studies and investigated whether the degree of genetic differentiation between fragmented plant populations was mediated by seed longevity. Second, we reviewed 13 published studies reporting the genetic diversity of both the seed bank and the above ground plants, aiming at comparing genetic diversity contained in the seed bank with the above ground vegetation. We conclude that a persistent seed bank may indeed mitigate the consequences of habitat fragmentation and protect a species from genetic drift and population genetic differentiation. We found no evidence, however, of high levels of genetic diversity accumulating in the soil seed bank. If genetic differences are present between the standing crop and the seed bank, they are very likely the result of local selection acting either directly or indirectly as a filter on the alleles present in the seed bank. We finally suggest that 1) the role of the seed bank should not be neglected in habitat fragmentation studies and 2) it is not very fruitful to continue comparing seed bank genetic diversity with above ground plant genetic diversity, unless this is performed under different selection regimes.     

Fire,aridity and seed banks. What does seed bank composition reveal about community processes in fire‐prone desert?

Questions: The relationship between fire, aridity and seed banks is poorly understood in plant community ecology. We tested whether there was a close correspondence between the seed bank and standing vegetation composition with time‐since‐fire in a desert. We also examined whether longer‐lived species showed seed limitation relative to more ephemeral species, as this could influence grass‐woody ratios in a major biome. Location: Dune hummock grasslands/shrublands of central Australia. Methods: The effects of time‐since‐fire on floristic and functional group composition were examined by comparing plots unburned since 1984 against plots that had been burned in 2002. Three methods were used to quantify seed abundances: a germination trial using heat and smoke application, a flotation method, and a sieving method. Results: Seed bank densities were very low (<3000 m^?2). Species similarity between the seed bank and standing vegetation was high at sites recently burned (0.86) and low in sites long‐since burned (0.52). The relative abundance of ephemeral species in the seed bank peaked in recently burned plots, but the relative abundance of seeds of woody species did not match the pattern of abundance in the standing vegetation. Remarkably, the dominant perennial grasses and woody species were either absent from the seed bank or present at extremely low abundances. Discussion: Differences in the relative abundance of ephemeral species between standing vegetation and seed bank relate to the post‐fire succession process. The small soil pool of seed from woody species may be explained by allocation to belowground carbohydrate storage over seed production. Field observations suggest, however, that production of strongly dormant seed can be prolific and that high levels of seed predation make this system strongly seed‐limited. The discovery of this seed bank syndrome indicates that shifts in grass‐woody ratios can be driven by the juxtaposition of unpredictable seed rain and fire events in these desert dunes. However, estimates of grass‐woody ratios due to changing fire regimes will be difficult to predict.     

Does aboveground vegetation composition resemble soil seed bank during succession in specialized vegetation on gypsum soil?

This paper evaluates the aboveground vegetation in relation to the soil seed bank throughout a 60-year succession process following agricultural abandonment in a semi-arid Mediterranean gypsum habitat. There is little information regarding the relationship between these two community components in the context of succession on semi-arid gypsum soils. Aboveground vegetation and the corresponding seed bank of gypsum plant communities were sampled through a chronosequence of 24 abandoned fields. Generalized linear models were used to model seed species richness and density, redundancy analyses to model the effect of time since abandonment and the effect of soil physicochemical parameters on seed bank species composition, and Mantel tests to analyze resemblance between above- and belowground species composition. In this last case, the effect of time since abandonment was controlled using a partial Mantel test. Mantel correlograms using time intervals instead of distances were used to describe the resemblance of above- to belowground species occurrence in different aged fields. No significant variability in seed species richness, seed density, or species composition due to time since abandonment was found. Differences in seed species composition were mainly due to small spatial scale predictors such as slope and soil calcium content. High correlations between species composition in the soil seed bank and the aboveground vegetation were detected during succession. The lack of a significant trend in aboveground species replacement over time was also reflected in seed bank composition. We concluded that the rapid establishment of strict gypsophyte species relied mainly on the long-term persistence of these species in the seed bank.     

Spatial structure of the soil seed bank of Poa annua L.—alien species in the Antarctica

Poa annua L. (annual bluegrass) is the only non–native flowering plant species that has successfully established a breeding population in the maritime Antarctic and has been shown to maintain a soil seed bank. The characteristic of the spatial structure of the Antarctic population of this species is the formation of distinct dense clumps—tussocks. In the temperate zone the species is only loosely tufted. We focused on the characteristics of seed deposition associated with the tussocks and some aspects of the spatial heterogeneity of the soil seed bank of P. annua in the Antarctic. We wanted to assess the microspatial structure of the soil seed bank of annual bluegrass at Arctowski Station. Therefore we compared the number of seeds deposited underneath and in the vicinity of P. annua clumps. Our results indicate that P. annua in the Antarctic maintains a soil seed bank comparable to species typical for the polar tundra. The microspatial structure of P. annua soil seed bank in the Antarctic is highly associated with the presence of tussocks. Seeds are deposited underneath the tussock rather than in the vicinity of the clump. Our results also indicate that seeds are able to survive the Antarctic winter and readily germinate under optimal conditions.     

How much genetic variation is stored in the seed bank? A study of Atriplex tatarica (Chenopodiaceae)

We investigated to what extent the soil seed bank differed genetically and spatially in comparison to three consecutive life history stages (seedlings, mature plants, and fruiting plants) in a natural population of Atriplex tatarica. Representatives of particular life history stages from twenty subunits within a large population were randomly collected and subjected to allozyme analysis. Comparison of population polymorphism among various life history stages showed significant differences in observed heterozygosity (H(O)) and F statistics (F(IS) and F(ST)), but nonsignificant ones in the cases of number of alleles per polymorphic locus (A) and gene diversity (H(S)). These results indicate an increasing number of heterozygotes, a decreasing level of inbreeding and an increase of the partitioning genetic diversity among populations with increasing population age. Spatial autocorrelation was used to calculate f, the average co-ancestry coefficient between individuals within distance intervals of two meters along a 39 m long transect. Significant positive fine scale genetic structure was detected in mature and fruiting plants but not in soil seeds and seedlings stages. The results of the presented study on A. tatarica indicated that significant differences exist in genetic differentiation, differentiation in allele frequencies and spatial autocorrelation among early (soil seeds and seedlings) and late (mature and fruiting plants) life history stages but not within early and late ones. This pattern suggests that, rather than storing genetic variability in the soil or germination and establishment success, self-thinning might be the major microselective force in populations of A. tatarica.     

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

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

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.     

Can spatial variation and inter-annual variation in precipitation explain the seed density and species richness of the germinable soil seed bank in a tropical dry forest in north-eastern Brazil?

The interaction between microhabitat and inter-annual variation in precipitation has an important role on the dynamics of the seed bank and can play a crucial role in survival and maintenance of plant populations in semi-arid environments. We hypothesized that the type of microhabitat and the inter-annual variability in precipitation can explain the richness and density of the seed bank in a semi-arid region in Brazil. The study was conducted in an area of tropical dry forest with shrub-tree physiognomy, locally called caatinga. We collected 35 soil samples in three distinct microhabitats, at the end of rainy and dry seasons, respectively, over three years, totalling 630 samples. The seed bank (richness and seed density) were determined by seedling emergence method. Over the three years, 79 species emerged from the seed bank, 64, 45 and 42 in riparian, non-riparian and rocky microhabitats, respectively. We recorded differences in species richness and average density between microhabitats and between years, with significant statistical interaction between them. Inter-annual precipitation explained 48% and 5% of the variation in richness and seed density, respectively. Spatial variation explained 7% of the species richness and 31% of the density. Our results show that the interaction between spatial variation and precipitation has an important role on the spatial and temporal heterogeneity of the richness and density of seed banks in dry environments.     

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.     

Seasonal wetlands on the Lake Wales Ridge, Florida: does a relict seed bank persist despite long term disturbance?

Wetlands maintain biodiversity and provide numerous ecosystem services, so the pressure to perform successful restoration consequently is high. However, restoration projects rarely include an in-depth assessment of wetland potential for recovery, and restoration techniques may not be tailored to site-specific concerns. This study examined the seed bank of disturbed wetlands slotted for hydrologic, but not vegetation, restoration to determine if a seed bank comparable to that of nearby undisturbed wetlands persisted despite long-term anthropogenic disturbance. We compared the aboveground vegetation and seed bank compositions under drained, drawdown, and flooded conditions between undisturbed and historically ditched (“disturbed”) wetlands. Disturbed and undisturbed wetlands shared fewer than 30 % of total aboveground species. While undisturbed wetlands were dominated by graminoids, disturbed wetlands had greater cover of forbs. The seed banks of disturbed wetlands had high species diversity, but their composition was dissimilar to that of nearby undisturbed wetlands. In total, the seed banks of both disturbance histories germinated 56 species; drained conditions had the fewest germinants while flooded conditions had the most. Germinant richness was significantly affected by disturbance, moisture, and their interaction; evenness was significantly affected by moisture, and Shannon diversity by disturbance. Because the seed bank of disturbed wetlands included many fast-growing wetland plants, passive vegetation restoration and active hydrologic restoration may result in wetlands overgrown with weedy species and with fewer conservative wetland plants. An understanding of the capacity for seed banks to re-vegetate wetlands post-restoration and approximate undisturbed wetlands is crucial to the overall success of restoration projects.     

Do surface plant and soil seed bank populations differ genetically? a multipopulation study of the desert mustard Lesquerella fendleri (Brassicaceae)

Seed banks are an important component of many plant populations, but few empirical studies have investigated the genetic relationship between soil seeds and surface plants. We compared the genetic structure of soil seeds and surface plants of the desert mustard Lesquerella fendleri within and among five ecologically diverse populations at the Sevilleta National Wildlife Refuge in Central New Mexico. At each site, 40 Lesquerella surface plants and 40 samples of soil seeds were mapped and genetically analyzed using starch gel electrophoresis. Overall allele frequencies of soil seeds and surface plants showed significant differences across the five populations and within three of the five individual populations. Surface plants had significantly greater amounts of single and multilocus heterozygosity, and mean surface plant heterozygosity was also greater at the total population level and in four of the five individual populations. Overall soil seed (bot not surface plant) homozygosity was significantly greater than predicted by Hardy-Weinberg expectations at the total and individual population levels. Although F-alpha estimates revealed similarly small but significant genetic divergence within each life-history stage, estimates of coancestry showed that fine-scale (0.5-2 m) genetic correlations among the surface plant genotypes were roughly twice those of soil seed genotypes. An unweighted pair group method with arithrnetic mean cluster analysis indicated that in the two geographically closest sites, the surface plants were slightly more genetically similar to each other than to their own respective seed banks. We also found weak and/or negative demographic associations between Lesquerella soil seed and surface plant densities within each of the five sites. We discuss the difficulties involved with sampling and genetically comparing these two life-history stages.     

What potential is there for regeneration of native species from the soil seed bank in Coast Tea Tree‐dominated scrub?

Shrub encroachment generally causes the loss of native species in herbaceous‐dominated communities. The ability of the original ecosystem to return to its pre‐encroachment state (i.e. its ecological resilience) will be partially contingent on the capacity of these species to regenerate from soil‐stored seed. Coast Tea Tree (Leptospermum laevigatum) has formed a dense scrub in many areas previously dominated by grassy woodland, and hence, managers need guidance about the effectiveness of strategies designed to recover the pre‐encroachment vegetation. In this context, we ask: what is the potential of species stored in the soil seed bank to return following Tea Tree removal? A germination experiment was undertaken using soil collected from dense stands of Tea Tree that had been long established. Heat/smoke was applied to soils to simulate the effects of a fire on the soil seed bank, while leaf litter treatments were used to mimic both undisturbed stands and stands where shrubs have been slashed where litter creates a physical barrier to emergence. We found the soil seed bank was dominated by exotic forbs (83% of all germinants) and contained few grasses. Heat and smoke decreased total species density but increased species diversity through the suppression of common exotics. Our data suggest that slashing would result in germination being dominated by exotic flora, but using fire would likely reduce that dominance. However, we conclude that recovery by much of the original flora after site occupation by Coast Tea Tree may be contingent on mechanisms other than soil‐stored seeds.     

The weed seed bank of soils in a landscape segment in southern Bavaria — I. Seed content, species composition and spatial variability

The purpose of the present investigation was to examine the composition and the spatial variability of the arable soil seed bank in one large, coherent landscape segment, where the cultivation practice of neighboring farms and the gradients in soil properties determine the distribution of the weed seeds in soil. To this end, a 50×50 m grid of 338 reference points was fixed throughout a 143 ha area in southern Bavaria. The seed bank contained a median density of 4950 seeds per m² able to germinate. These values showed remarkable variation throughout the investigated area. Thus, 27% of the sampling area contained less than 2500 seeds per m², another 23% were below 5000 seeds per m² and 19% ranged among 5000 and 10 000 germinating seeds per m². Among the remaining sites, 22% showed seed densities between 10 000 and 20 000 and 9% more than 20 000 seeds per m². To visualize the spatial distribution of the weed seed bank over the whole study area, the interpolation method kriging was used. For this purpose the spatial continuity must be described by means of a semivariogram analysis. The goodness of the interpolation results depends on the fit of the semivariogram model which shows the spatial dependency. Two important characteristic parameters of this model are the nugget and the range. The nugget expresses the sampling error and the local variability up to the 50 m sampling distance, which cannot be detected by the semivariogram. Therefore the interpolation cannot describe the seed bank variation on a scale below this threshold. In our investigations the nugget comprises about one third of the total variance. The estimation error increases towards the border of the investigated area. In general, this is due to missing sampling points at the field edges. The range observed in the present investigations was 340 m. Beyond this no more spatial variability exists and the measurements are independent of each other. The species occurring in more than five of the investigated samples were ordinated by Detrended Correspondence Analysis and classified by k-means cluster analysis of the ordination axes. Five weed communities were recognized from this analysis which distinctly differed in their relation to the soil type and to the preceding management.     

The weed seed bank of soils in a landscape segment in southern Bavaria – II. Relation to environmental variables and to the surface vegetation

In the first part of their investigations of the weed seed bank in a 110 ha landscape segment in southern Bavaria, Albrecht & Forster (1996) showed that there was a tremendous spatial variability in seed density as well as in species composition. The intention of the present paper was to elucidate the reasons for this variability. For this purpose, the influence of soil conditions, land use practices, and soil fauna, all of which are environmental factors potentially affecting the weed seed bank, were investigated.Highly significant correlations were observed between the soil seed bank, preceding management variables, and the available field capacity. In contrast, multiple regression analyses between seed banks and the chemical characteristics of the soil, soil fauna, and actual management showed either minor correlation or none at all. Thus, prior management and water supply must be supposed to have had the most decisive influence on the soil seed bank.Several environmental variables affect arable weed populations, not so much through the soil seed bank but through the germination, establishment, and reproduction phases in the field. For this reason the relations between the apparent weed flora and the potential vegetation in the soil also were investigated. The correlation of the soil seed content and the density of individuals on the soil surface was highly significant. The analysis for a single species revealed that the seed bank/field density-ratios of different taxa vary from below 10:1 up to 100:1 and more. This wide range may be related to the life-forms and ecological requirements characteristic for each species.Finally, comparisons of the environmental variables with the soil seed bank and surface vegetation allowed ranking of the variables as to their importance for these two stadia of plant population development. These investigations indicated that the preceding management by different farmers exerted a more decisive influence on the number of seeds in the soil. In contrast, the actual grain yield and N-content in a soil showed a closer relationship to the soil surface vegetation. With regard to the species composition, no distinct differences in the environmental relationships between the soil seed bank and the soil surface vegetation could be observed.