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 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 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.     

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.     

What affects the probability of biological invasions in Antarctica? Using an expanded conceptual framework to anticipate the risk of alien species expansion

Successful alien species invasion depends on many factors studied mostly in post invasion habitats, and subsequently summarized in frameworks tailored to describe the studied invasion. We used an existing expanded framework with three groups of contributing factors: habitat invisibility, system context and species invasiveness, to analyze the probability of alien species invasions in terrestrial communities of Maritime Antarctic in the future. We focused on the first two factor groups. We tested if the expanded framework could be used under a different scenario. We chose Point Thomas Oasis on King George Island to perform our analysis. Strong geographical barrier, low potential bioclimatic suitability and resource availability associated with habitat invasibility significantly reduce the likelihood of biological invasion in Antarctica. An almost full enemy release (low pressure of consumers), the high patchiness of the habitat, and the prevalence of open gaps also associated with habitat invasibility increase the possibility of invasion. The dynamics of functional connectivity, propagule pressure and spatio-temporal patterns of propagule arrival associated with human activity and climate change belonging to the system context contribute to an increase in the threat of invasions. Due to the still low land transport activity migration pathways are limited and will reduce the spread of alien terrestrial organisms by land. An effective way of preventing invasions in Antarctica seems to lie in reducing propagule pressure and eliminating alien populations as early as possible. The expanded conceptual framework opens up wider possibilities in analyzing invasions taking place in different systems and with multiple taxa.     

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 “Trojan horse” strategy: Seed fungal endophyte symbiosis helps to explain the invasion success of the grass,Poa annua,in Maritime Antarctica

Aim

Poa annua L. (annual bluegrass) is presently the sole invasive vascular plant species to have successfully established in Maritime Antarctica, where it poses a significant conservation threat to native plant species. However, the reasons for its success in the region have yet to be established. Here, we determined whether the invasiveness of P. annua, and its competitiveness with the native Antarctic hairgrass Deschampsia antarctica, is influenced by symbioses formed with seed fungal endophytes, and whether plants derived from seeds from four global regions differ in their performance.

Locations

Four regions (Maritime Antarctica, sub-Antarctica, South America and Europe).

Methods

Endophyte frequency was measured in P. annua seeds collected from the four regions. The germination, survival, biomass accumulation, flowering and competitiveness with D. antarctica of P. annua plants grown from endophyte-uncolonised and uncolonised seeds was determined in the laboratory. The effects of endophytes on P. annua seed germination and survival and seedling osmoprotection were also assessed in the Maritime Antarctic natural environment using locally-sourced seeds.

Results

Endophytes were at least twice as frequent in seeds from Maritime Antarctica than in those from other regions. A higher proportion of endophyte-colonized seeds germinated and survived than did uncolonised seeds, but only when they originated from Maritime Antarctica. Seed endophytes increased the competitiveness of P. annua with D. antarctica, but only for plants grown from Maritime Antarctic seeds. In the field, endophyte-colonized seeds from Maritime Antarctica germinated and survived more frequently than uncolonised seeds, and osmoprotection was higher in seedlings grown from colonized seed.

Main Conclusions

The findings indicate beneficial effects of seed endophytes on invasion-related traits of P. annua, such as survival, germination success and flowering. Together with vegetative and reproductive traits facilitating the colonization process, the seed-fungal endophyte symbiosis can be invoked as an important factor explaining the invasiveness of P. annua in Maritime Antarctica.     

Alpine species seed longevity in the soil in relation to seed size and shape – A 5-year burial experiment in the Central Alps

Long-term persistent seeds are generally expected to be small and spherically shaped. In contrast, transient seeds are often large, flattened and elongated. We hypothesized that alpine species follow these trends, and that most species of unstable alpine habitats have the potential to form long-term persistent seed banks. The latter would enable them to delay germination until favourable conditions are present in their environment thereby increasing the likelihood for establishment of seedlings. We selected nine alpine species of more or less unstable habitats (i.e. Achillea moschata, Artemisia genipi, Anthyllis vulneraria ssp. alpicola, Geum reptans, Linaria alpina, Oxyria digyna, Saxifraga aizoides, Saxifraga oppositifolia, and Trifolium pallescens) for this study. Seed longevity was estimated by a 5-year burial experiment in the field with seed excavations after 1 winter, 2 years and 5 years. Germination experiments in the growth chamber and viability tests were performed after each excavation. We detected longevity patterns ranging from transient to long-term persistent. Two species were classified as transient. All other species have the potential to form persistent seed banks, with almost half of the investigated species showing a trend for long-term persistence. Despite contradictory reports, long-term persistent seed banks are an important life history trait of species of unstable alpine habitats. We found that seed shape and hard-seededness are good predictors of seed persistence of alpine species. Seed size seems to be less important. The data from this study support the general hypotheses describing the relationship among seed size, seed shape and seed persistence in the soil.     

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 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.     

Does governance play a role in the distribution of invasive alien species?

Invasive alien species (IAS) constitute a major threat to global biological diversity. In order to control their spread, a detailed understanding of the factors influencing their distribution is essential. Although international trade is regarded as a major force structuring spatial patterns of IAS, the role of other social factors remains unclear. Despite studies highlighting the importance of strong governance in slowing drivers of biodiversity loss such as logging, deforestation, and agricultural intensification, no study has yet analyzed its contribution to the issue of IAS. Using estimates of governance quality and comprehensive spatiotemporal IAS data, we performed multiple linear regressions to investigate the effect of governance quality upon the distribution of species listed under “100 of the worst” IAS in 38 Eurasian countries as defined by DASIE. Our model suggested that for countries with higher GDP, stronger governance was associated with a greater number of the worst IAS; in contrast, for the lowest GDP countries under analysis, stronger governance was associated with fewer of these IAS. We elucidate how the quality of governance within a country has implications for trade, tourism, transport, legislation, and economic development, all of which influence the spread of IAS. While our findings support the common assumption that strengthening governance benefits conservation interventions in countries of smaller economy, we find that this effect is not universal. Stronger governance alone cannot adequately address the problem of IAS, and targeted action is required in relatively high‐GDP countries in order to stem the influx of IAS associated with high volumes of trade.     

Açaí palm management disturbs seed rain and soil seed bank of an Amazonian estuarine forest

The exploitation of non-timber forest products has been proposed as a sustainable way to exploit tropical forests, but such an opportunity remains to be proved. Here, we examine the impact of intensive açaí palm (Euterpe oleracea) management on the seed rain and soil seed bank in an estuarine forest landscape with a long history of forest management by locals in the Amazon region. Seed rain (100 80 cm² collectors) and soil seed bank (100 30 cm² samples) were monitored through a year across 20 forest stands, covering a gradient of açaí stem density (50–3575 açaí stems per ha). Seed rain and bank were dominated by açaí seeds (85.5%–85.8%) and by excluding them, seed rain and bank were low density and species poor, capturing a tiny subset from the local (17.91%–19.40%) and landscape woody flora (11.82%–14.55%). Moreover, autochthonous and vertebrated-dispersed predominated as well as those from tree species considered useful by locals. Overall, açaí stem density positively affected açaí seed abundance in the seed rain and negatively affected seed pools in the bank in relation to abundance of seeds and vertebrated dispersed, while adult tree species richness and density and accessibility to forest stands were associated with more diversified seed pools. Thereby, forest stands are exposed to different levels of açaí management (high vs. low intensity) supported taxonomically distinct seed pools. Our results suggest that açaí intensification disturbs seed rain and soil seed bank with potential impacts on forest regeneration and the forest integrity standards required to consider açaí fruit production as sustainable according to current legislation.     

Trend in aliens species in the Mediterranean. An answer to Galil, 2009 «Taking stock: inventory of alien species in the Mediterranean Sea»

The number of marine alien species reported in Galil (Biol Invasions, 2009) is incomplete. Recent literature suggests that the number of aliens is nearly 1,000 and that the introduction rate is way off the 10 species per year suggested by Galil (Biol Invasions, 2009).     

Spatial heterogeneity in soil particle size: does it affect the yield of plant communities with different species richness?

Aims Soil heterogeneity is ubiquitous in many ecosystems. We hypothesized that plant communities with higher species richness might be better adapted to soil heterogeneity and produce more biomass than those with lower richness. This is because there is niche differentiation among species and different species can complement each other and occupy a broader range of niches when plant species richness is high. However, no study has tested how soil particle heterogeneity affects the yield of plant communities, and whether such effects depend on the spatial scale of the heterogeneity and the species richness within the communities.Methods In a greenhouse experiment, we sowed seeds of four-species or eight-species mixtures in three heterogeneous treatments consisting of 32, 8 or 2 patches of both small (1.5mm) and large quartz (3.0mm) particles arranged in a chessboard manner and one homogeneous treatment with an even mixture of small and large quartz particles.Important findings Biomass production was significantly greater in the communities with high species richness than those with low species richness. However, soil particle heterogeneity or its interactions with patch scale or species richness did not significantly affect biomass production of the experimental communities. This work indicates that plant species richness may have a bigger impact on plant productivity than soil particle heterogeneity. Further studies should consider multiple sets of plant species during longer time periods to unravel the potential mechanisms of soil heterogeneity and its interactions with the impacts of species richness on community yield and species coexistence.     

How much of the invader's genetic variability can slip between our fingers? A case study of secondary dispersal of Poa annua on King George Island (Antarctica)

We studied an invasion of Poa annua on King George Island (Maritime Antarctic). The remoteness of this location, its geographic isolation, and its limited human traffic provided an opportunity to trace the history of an invasion of the species. Poa annua was recorded for the first time at H. Arctowski Polish Antarctic Station in the austral summer of 1985/6. In 2008/9, the species was observed in a new locality at the Ecology Glacier Forefield (1.5 km from “Arctowski”). We used AFLP to analyze the genetic differences among three populations of P. annua: the two mentioned above (Station and Forefield) and the putative origin of the introduction, Warsaw (Poland). There was 38% genetic variance among the populations. Pairwise Ф_PT was 0.498 between the Forefield and Warsaw populations and 0.283 between Warsaw and Station. There were 15 unique bands in the Warsaw population (frequency from 6% to 100%) and one in the Station/Forefield populations (which appears in all analyzed individuals from both populations). The Δ(K) parameter indicated two groups of samples: Warsaw/Station and Forefield. As indicated by Fu's Fs statistics and an analysis of mismatch distribution, the Forefield population underwent a bottleneck and/or founder effect. The Forefield population was likely introduced by secondary dispersal from the Station population.