Temporal effects of agri-environment schemes on ditch bank plant species

Many of the agri-environment schemes (AES) implemented in the Western Peat District of the Netherlands have as their objective the conservation of the diversity of ditch bank plants. We investigated the effects of AES on ditch bank species in this area, using a dataset collected by 377 farmers who managed and monitored ditch banks during a 10-year period. We found that species richness has increased minimally over the last 10 years in ditch banks. Yet, we found no differences in increases in time between ditch banks with and without AES. In both ditch bank types plant species composition changed to species with higher nitrogen tolerance. Furthermore, species that disperse over long distances by water increased, whereas species with no capacity to disperse over long distances declined in both ditch bank types. This indicates that changes in vegetation composition in ditch banks are affected by other factors than AES.     

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

Temporary floating island formation maintains wetland plant species richness: The role of the seed bank

In wetlands, fluctuating water levels create opportunities for recruitment of new individuals from seed banks, and drawdown periods often favor the establishment of species adapted for life in shallow water. In this study, floating island formation functioned similarly to drawdowns in water level by creating patches of sediment that were less inundated relative to the surrounding deep water marsh. The disturbance of floating island formation (i.e., mats of sediment and vegetation rising vertically in the water column) also resulted in reduced cover of the dominant rooted, floating-leaved macrophyte, thereby creating temporary gaps for the establishment of other species. To assess how floating island formation influences species richness and composition of wetland plant assemblages relative to surrounding deep water marsh, field surveys of plant percent cover on and off of islands were conducted over 2 years, along with a controlled greenhouse seed bank experiment in which levels of inundation were manipulated. Five plant species were present in deep water marsh compared to 22 in surrounding on floating islands. Plant assemblages on floating islands consisted primarily of emergent species, while floating-leaved perennials dominated the deep water marsh. Species richness and assemblage composition in the greenhouse seed bank experiment differed among water level treatments in a manner consistent with differences observed in field surveys. Assemblages germinating under minimal inundation treatments were more species rich (3.5–4.3 species per sample) and contained more emergent species (>450 individuals m⁻²) than did those germinating under flooded conditions (2.8–2.9 species per sample and <405 individuals m⁻²). This study, in addition to being the only reported seed bank study of temporary (i.e., seasonal) floating islands, demonstrates that islands altered levels of inundation favoring the germination of more species-rich, emergent wetland plant assemblages. Because these islands persisted long enough for several species to set seed, their formation may be one mechanism by which the seed bank is replenished and populations of otherwise uncommon species are maintained.     

Spatial variation in ditch bank plant species composition at the regional level: the role of environment and dispersal

Questions: Can patterns of species similarity on ditch banks be explained by environmental and dispersal factors and, if so, to what extent? Does the pattern of distance decay differ among different species groups (all species versus target species of conservation interest; species of different dispersal type)? Location: Krimpenerwaard, the Netherlands. Methods: In 2006‐2007, ditch bank vegetation data on 130 terrestrial herbaceous species were collected on 72 plots. Species similarity was measured and related to environmental distance (soil type and nutrient level) and dispersal distance (geographic distance and limitation of dispersal by water, wind and agricultural activities) as explanatory factors using multiple regression on distance matrices (MRM). Differences in rates of distance decay in species similarity among different subsets of data (species groups) were investigated using randomization tests. Results: In all species, patterns of similarity of composition are influenced mainly by variations in dispersal, while for target species these are due to combined effects of environmental and dispersal variation. Compared with species using other dispersal mechanisms, water‐dispersed species had half the rate of distance decay. Conclusions: For all species considered here, dispersal limitation seems more responsible for the spatial variation in species composition than environmental determinism. Conservation management focused on plant species diversity would be more successful in areas adjacent to those where a similar management regime is already in force. For target species of conservation interest, besides dispersal limitation, environmental determinants like nutrient level are also important. As a means of conserving such target species, therefore, focusing on reducing nutrient levels and facilitating species dispersal will be more effective than current management practices, which mainly focus on reducing fertilizer inputs.     

Ecological correlations between the persistence of the soil seed bank and several plant traits,including seed dormancy

It was empirically showed that seed size and life history correlate with the formation of a soil seed bank. Although no empirical data are available that indicate a close relationship between seed dormancy and the soil seed bank, dormancy has been considered essential to the formation of a soil seed bank. I have considered the formation of the soil seed bank and survival of seeds for more than a year in the soil, and the persistence and survival of the seed bank for more than 5 years. These periods were derived from the definition of a persistent seed bank and the criterion for seed banks of long-term persistence. Plant traits that are closely related to the formation or persistence of a seed bank and their relationships to dormancy were analysed using two pre-existing databases of seed longevity in soil and comparative ecology. The integrated database comprised 18 plant traits and seed bank formation or persistence data. This approach was used to identify more reliable general empirical rules. The results of a regression tree analysis and common statistical tests of plant traits indicated that only life history and seed size were closely related to seed bank formation, and dormancy was not essential for the formation and persistence of a seed bank. However, the contribution of dormancy differed slightly between dormancy types. Scarification or dry storage requirements to break dormancy slightly enhanced the formation and persistence of a seed bank, whereas a chilling requirement decreased the formation and persistence of a seed bank. In contrast, fluctuating temperature requirements clearly contributed to the formation and persistence of a seed bank.     

Effects of grazing on soil seed bank dynamics: An approach with functional groups

Abstract. The relationship between intensity and timing of cattle grazing on changes in the size and composition of the soil seed bank were investigated in a 3‐yr study in a Mediterranean grassland in northeastern Israel. Treatments included manipulations of stocking rates and of grazing regimes, in a factorial design. The retrieved soil seed bank community was rich in species, with 133 species accounting for 80% of the 166 species recorded at the site. Within the seed bank, 89% of the species were annuals. Seed bank dynamics was analysed in terms of plant functional groups and germination strategies. In terms of total seed bank density and including all functional groups, 42% of the seeds present in the soil did not germinate under watering conditions. The dormancy level differed greatly among functional groups. The seed bank of annual legumes, crucifers, annual thistles and annual forbs had a large fraction of non‐germinated seeds and characterized areas grazed early in the growing season under high and very high grazing intensity. These functional groups were considered to have a higher potential for persistent seed banks production. In contrast, short and tall annual grasses and tall perennial grasses, that were dominant in ungrazed or moderately grazed paddocks, generally had seed banks with a very small fraction of non‐germinated seeds. Seed bank densities varied widely between grazing treatments and years. Under continuous grazing, heavy grazing pressure reduced seed bank densities of grasses and crucifers in comparison to moderate grazing. The greatest reduction on the seed bank densities resulted from heavy grazing concentrated during the seed‐set stages.     

Spatial segregation of plant species caused by Retama sphaerocarpa influence in a Mediterranean pasture: a perspective from the soil seed bank

Retama sphaerocarpa is a leguminous shrub whose important role in the semi-arid regions of south-eastern Spain has already been assessed: shrub canopies reduce light intensity, and thus evapotranspiration; also their extensive radical system take water and nutrients from great volumes of soil, concentrating them in the understorey. Consequently, subcanopy vegetation benefits from these facilitation processes, increasing its productivity. However, these shrublands have been rarely studied at a broader scale, i.e. as a savannah-like system composed of a variable number of shrubs scattered in a herbaceous matrix. As the microenvironmental conditions associated to the understorey are rather different from those of the open spaces among shrubs, species composition of the herbaceous matrix is expected to change accordingly. Thus, R. sphaerocarpa would be an important and still unknown source of spatial heterogeneity to the system. Our main purpose was to evaluate, through the soil seed bank, the heterogeneity in the herbaceous community induced by this shrub species. Soil samples were collected around adult shrubs from three positions relative to the canopy: near the centre of the shrubs, at the edge of the understorey, and completely outside the canopy. Floristic composition was evaluated by germination under greenhouse conditions. The results show that each position has a different floristic composition, characterised by a group of different species. The herbaceous species associated with the external position have functional traits which enable them to resist water stress and herbivore pressure, such as hairs, CAM metabolism, early flowering, horizontal growth or tiny stature. The species associated with the central position lack those traits, and are more competitive in more mesic environments, rich in nitrogen. The lowest number of seedlings and species was found in the internal position, suggesting that in our study the facilitation process may have less importance for community dynamics due to less stressful environmental conditions.     

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.     

Similarities in the impact of three large invasive plant species on soil seed bank communities

Invasions by alien plant species may substantially alter soil seed bank communities. While decreases in seed bank species richness, diversity, and composition as a consequence of plant invasions have been reported, the characteristics of seed banks associated with different invasive species have not been compared in any detail. Here, we describe changes in the characteristics of soil seed banks invaded by three large herbaceous invasive plants, Fallopia japonica, Gunnera tinctoria, and Heracleum mantegazzianum. The study was carried out at the spatial scales of site and plot, to reduce variability in seed bank data. Information on seed bank persistence was inferred from seed depth (0–5, 5–10, and 10–15 cm) and from time of sampling (May and October). Despite differences in the reproductive strategy and geographic distribution of these invaders, as well as in the standing vegetation and habitat types examined, the seed banks of invaded areas were similar in composition and in the relative abundance of different species. Invaded seed banks were dominated by seeds of a few agricultural weed species and/or rushes, suggesting that common features of the invaders, including a large standing biomass, extensive litter production, and the formation of mono-species stands may result in comparable selection pressures that favors traits that are largely genera or species-specific. These findings have a direct relevance for the development of strategies aimed at restoring previously-invaded sites while also improving our understanding of the long-term implications of plant invasions.     

Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density

Immigration rates of species into communities are widely understood to influence community diversity, which in turn is widely expected to influence the susceptibility of ecosystems to species invasion. For a given community, however, immigration processes may impact diversity by means of two separable components: the number of species represented in seed inputs and the density of seed per species. The independent effects of these components on plant species diversity and consequent rates of invasion are poorly understood. We constructed experimental plant communities through repeated seed additions to independently measure the effects of seed richness and seed density on the trajectory of species diversity during the development of annual plant communities. Because we sowed species not found in the immediate study area, we were able to assess the invasibility of the resulting communities by recording the rate of establishment of species from adjacent vegetation. Early in community development when species only weakly interacted, seed richness had a strong effect on community diversity whereas seed density had little effect. After the plants became established, the effect of seed richness on measured diversity strongly depended on seed density, and disappeared at the highest level of seed density. The ability of surrounding vegetation to invade the experimental communities was decreased by seed density but not by seed richness, primarily because the individual effects of a few sown species could explain the observed invasion rates. These results suggest that seed density is just as important as seed richness in the control of species diversity, and perhaps a more important determinant of community invasibility than seed richness in dynamic plant assemblages.     

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.