The Influence of Time on the Soil Seed Bank and Vegetation across a Landscape‐Scale Wetland Restoration Project

Wicken Fen National Nature Reserve (NNR) in Cambridgeshire, U.K. is a wetland of international importance isolated in a landscape dominated by arable farming. The prospect of species extinctions within the NNR led to the creation of the Wicken Fen Vision, an ambitious project that will eventually expand the reserve boundary by the purchase and restoration of c.50 km² of arable land. We sampled three fields from each of three distinct age‐categories of restoration land (5, 15, and 60 years post‐arable), and three fields within the adjacent, undrained NNR, to determine (1) differences in seed bank composition across age‐categories, (2) relationships between restoration age, the seed bank and standing vegetation, and (3) changes in species traits across age‐categories. Historic arable management contributed to an apparent “vertical mixing” effect in the seed bank of the youngest two age‐categories, with associated and significant differences in species functional traits across the study area. Almost all plants associated with NNR vegetation were absent from restoration area seed banks and standing vegetation. Seed bank species common to all ages‐categories exhibited a bias toward moderate to high Ellenberg F (moisture) values, persistent seed banks, and lateral vegetative spread. Relatively short (c. 6 years) periods of drainage and plowing impact heavily upon seed bank diversity and soils, resulting in a lack of predrainage vegetation, even after decades of subsequent restoration adjacent to intact, species‐rich habitat. However, the seed banks of highly degraded fields can contribute toward the creation of novel wetland vegetation assemblages over time and under suitable environmental conditions.     

Ecological correlates of seed persistence in soil in the north-west European flora

1 Using the data in a recently published seed bank database for north-west Europe, we describe how a species' seed bank behaviour can be characterized by a single 'longevity index', and investigate how representative the information in the database is of the north-west European flora. We also test the hypotheses that seeds of short-lived species are more persistent than those of long-lived species, and long-lived seeds are characteristic of species living in disturbed habitats.
2 The data are not representative of the north-west European flora as a whole; they are a fair reflection of a research effort that has been largely directed towards grassland and arable weeds. Data for rare species, non-agricultural habitats in general and wetlands, rocky habitats and native woodland in particular, are scarce or absent.
3 Annuals and biennials almost always have more persistent seeds than related perennials, and this difference is most striking when, as in Anagallis arvensis and Aphanes arvensis , the short-lived species have moved well away from the 'core' habitat of the family. Confamilial monocarps and polycarps do not differ consistently in seed mass.
4 Gradients of habitat disturbance are accompanied by predictable changes in seed persistence, which are themselves often (but not always) accompanied by parallel shifts in seed size. These results suggest that increasing habitat disturbance (i.e. increasing density-independent mortality) always selects for increased seed persistence, confirming both theory and previous analyses. However, increased seed persistence is not always associated with reduced seed size. This is because persistence depends not only on seed size, but on other traits, many of them physiological. In many habitats the probability of seed burial is strongly linked to seed size and shape, but in arable habitats cultural practices have broken this link.     

Assessing soil seed bank persistence in flood‐meadows: The search for reliable traits

Abstract. To assess seed bank persistence of target species in endangered flood‐meadows (alliances Cnidion and Molinion), we investigated established vegetation and soil seed bank of 46 plots for 3 yr and 2 yr, respectively. As traits of seed persistence we calculated various continuous indices that refer to the frequency and abundance of species in above‐ground vegetation and at different soil depths. Furthermore, we tested the significance and soundness of easily observed traits such as maximum seed density per plot and seed attributes (mass, size and shape) as predictors of soil seed bank features. In linear regression, SAI, the seed accumulation index, showed the best agreement (R²= 0.64) with the seed longevity index that was derived from the database by Thompson et al. (1997) for a set of 115 species. The second best predictor (R²= 0.39) of the seed longevity index was maximum seed density per plot in the lower soil layer (5–10 cm). Depth distribution indices and seed attributes showed weaker but still significant relations. The dynamic character of flood‐meadows was reflected by a large proportion of species with a strong tendency to accumulate seeds in the soil relative to their importance in above‐ground vegetation. Most of these species have a ruderal strategy, exploiting gaps after flood disturbances, while the dominants of flood‐meadows tended to have short‐lived seed banks. Compared to other grassland types, a relatively large proportion of rare and endangered target species can be expected to form long‐term persistent seed banks. However, only under marginal conditions that facilitate seed survival in the soil (e.g. fallow) are these persistent seed banks likely to contribute to restoration. We conclude that easily observed traits of persistence such as seed weight, size and shape do not meet the accuracy needed in scientific and practical applications. Thus, there is a crucial demand for further seed bank studies in poorly investigated habitats and of rare species.     

Long‐term farming systems and last crop sown shape the species and functional composition of the arable weed seed bank

Questions: The assembly of arable weed communities is the result of local filtering by agricultural management and crop competition. Therefore, soil seed banks can reflect the effects of long‐term cumulative field management and crop sequences on weed communities. Moreover, soil seed banks provide strong estimates of future weed problems but also of potential arable plant diversity and associated ecological functions. For this, we evaluated the effects of different long‐term farming systems under the same crop rotation sequence on the abundance, diversity and community assembly of weed seed bank, as well as on the functional diversity and composition. Location: DOK (biodynamic [D], bioorganic [O], conventional [K]) long‐term trial, Therwil, Switzerland. Methods: The effects of long‐term contrasted farming systems (i.e., biodynamic, organic, conventional, mineral and unfertilised systems) and last crop sown (i.e., wheat and maize) were evaluated on different indicators of species and functional diversity and composition of the weed soil seed bank. Results: The results showed significant influences of 40 years of contrasted farming systems on the diversity and composition of the seed bank, with higher diversities being found in unfertilised and organic farming systems, but also higher abundances than those found under conventional systems. Organic farming also allowed higher functional richness, dispersion and redundancy. Different farming systems triggered shifts in species and functional assemblies. Conclusions: The results highlight the importance of organic management for the maintenance of a diverse arable plant community and its functions. However, such results emphasise the need for appropriate yearly management to reduce the abundance of settled weediness and prevent affecting crop production. The farm management filtered community composition based on functional traits. Although the soil seed bank buffers the long‐term farming and crop sequence, the last crop sown and, thus, the yearly management were important determinants of seed bank composition.     

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.     

Seed bank classification and its importance for the restoration of species-rich flood-meadows

Abstract. This paper focuses on the persistence of seeds in the seed bank of a formerly fertilized flood-meadow into which seeds from an original Alopecurus pratensis-Sanguisorba officinalis community have been introduced. The longevity of seeds in the seed bank of the original community is also estimated. The established vegetation was compared with the seed bank, as divided into two layers (0 - 5 cm and 5–10 cm); this allowed a classification of species into three groups, with (1) transient, (2) short-term persistent and (3) long-term persistent seed bank. The majority of the species of the unfertilized flood-meadow community have a transient or short-term persistent seed bank with seeds showing a large variance in shape i.e. flattened or elongate. Because of this, the characteristic flood-meadow species will disappear soon after the beginning of fertilizer application and will not re-establish from the seed bank, once the fertilizer application is ceased. The formerly fertilized flood-meadow contains many ruderal and arable weed species, the seeds of which tend to be compact or round. Significantly more seeds were found in the seed bank of the formerly fertilized flood-meadow under cattle-grazing than under sheep-grazing and no grazing.     

Cover crops and compost prevent weed seed bank buildup in herbicide‐free wheat–potato rotations under conservation tillage

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Variation in Capsella (shepherd's purse): an example of intraspecific functional diversity

The extent of functional trait diversity is quantified for 157 different Capsella bursa-pastoris (L.) medic (shepherd's purse) accessions. These individuals encompass replicate progeny generated from seed of 53 different Capsella 'maternal lines' that were isolated at random as they emerged from soil cores (used to estimate baseline seed bank numbers and weed diversity) at 34 different arable sites across the United Kingdom. The replicate progeny were subject to ex situ characterisation for traits determining life history and architecture. Seven leaf-type classes were identified and representative parents of each leaf type were distinguishable using four different simple sequence repeat markers. Life-history traits were only loosely associated with leaf shape, and cluster analysis grouped the accessions into three broad types: small plants that flowered early with intermediate reproductive output; large plants, with intermediate time to flowering and a high reproductive output; late-flowering plants, of intermediate size and low reproductive output. The most common leaf-type variants (83% of accessions) demonstrated a short time to flowering (ca. 70 days), while rarer variants included those that flowered after 140 days, accumulated more nitrogen and produced less seed: possibly representing advantageous and disadvantageous traits (respectively), in modern arable rotations. A wide trait variation was therefore found in Capsella bursa-pastoris despite decades of agricultural intensification, the range of time-to-flowering for C. bursa-pastoris being as broad the mean flowering times of the commoner annual and winter annual arable species. We propose the use of traits, rather than species, as the accounting unit to quantify functional biodiversity in arable systems.     

Density of insect‐pollinated grassland plants decreases with increasing surrounding land‐use intensity

Pollinator declines have raised concerns about the persistence of plant species that depend on insect pollination, in particular by bees, for their reproduction. The impact of pollinator declines remains unknown for species‐rich plant communities found in temperate seminatural grasslands. We investigated effects of land‐use intensity in the surrounding landscape on the distribution of plant traits related to insect pollination in 239 European seminatural grasslands. Increasing arable land use in the surrounding landscape consistently reduced the density of plants depending on bee and insect pollination. Similarly, the relative abundance of bee‐pollination‐dependent plants increased with higher proportions of non‐arable agricultural land (e.g. permanent grassland). This was paralleled by an overall increase in bee abundance and diversity. By isolating the impact of the surrounding landscape from effects of local habitat quality, we show for the first time that grassland plants dependent on insect pollination are particularly susceptible to increasing land‐use intensity in the landscape.     

The ecophysiology of seed persistence: a mechanistic view of the journey to germination or demise

Seed persistence is the survival of seeds in the environment once they have reached maturity. Seed persistence allows a species, population or genotype to survive long after the death of parent plants, thus distributing genetic diversity through time. The ability to predict seed persistence accurately is critical to inform long‐term weed management and flora rehabilitation programs, as well as to allow a greater understanding of plant community dynamics. Indeed, each of the 420000 seed‐bearing plant species has a unique set of seed characteristics that determine its propensity to develop a persistent soil seed bank. The duration of seed persistence varies among species and populations, and depends on the physical and physiological characteristics of seeds and how they are affected by the biotic and abiotic environment. An integrated understanding of the ecophysiological mechanisms of seed persistence is essential if we are to improve our ability to predict how long seeds can survive in soils, both now and under future climatic conditions. In this review we present an holistic overview of the seed, species, climate, soil, and other site factors that contribute mechanistically to seed persistence, incorporating physiological, biochemical and ecological perspectives. We focus on current knowledge of the seed and species traits that influence seed longevity under ex situ controlled storage conditions, and explore how this inherent longevity is moderated by changeable biotic and abiotic conditions in situ, both before and after seeds are dispersed. We argue that the persistence of a given seed population in any environment depends on its resistance to exiting the seed bank via germination or death, and on its exposure to environmental conditions that are conducive to those fates. By synthesising knowledge of how the environment affects seeds to determine when and how they leave the soil seed bank into a resistance–exposure model, we provide a new framework for developing experimental and modelling approaches to predict how long seeds will persist in a range of environments.     

The seed bank longevity index revisited: limited reliability evident from a burial experiment and database analyses

Background and Aims

Seed survival in the soil contributes to population persistence and community diversity, creating a need for reliable measures of soil seed bank persistence. Several methods estimate soil seed bank persistence, most of which count seedlings emerging from soil samples. Seasonality, depth distribution and presence (or absence) in vegetation are then used to classify a species'' soil seed bank into persistent or transient, often synthesized into a longevity index. This study aims to determine if counts of seedlings from soil samples yield reliable seed bank persistence estimates and if this is correlated to seed production.

Methods

Seeds of 38 annual weeds taken from arable fields were buried in the field and their viability tested by germination and tetrazolium tests at 6 month intervals for 2·5 years. This direct measure of soil seed survival was compared with indirect estimates from the literature, which use seedling emergence from soil samples to determine seed bank persistence. Published databases were used to explore the generality of the influence of reproductive capacity on seed bank persistence estimates from seedling emergence data.

Key Results

There was no relationship between a species'' soil seed survival in the burial experiment and its seed bank persistence estimate from published data using seedling emergence from soil samples. The analysis of complementary data from published databases revealed that while seed bank persistence estimates based on seedling emergence from soil samples are generally correlated with seed production, estimates of seed banks from burial experiments are not.

Conclusions

The results can be explained in terms of the seed size–seed number trade-off, which suggests that the higher number of smaller seeds is compensated after germination. Soil seed bank persistence estimates correlated to seed production are therefore not useful for studies on population persistence or community diversity. Confusion of soil seed survival and seed production can be avoided by separate use of soil seed abundance and experimental soil seed survival.Key words: Arable weeds, Bifora testiculata, Carthamus lanatus, Centaurea solstitialis, longevity index, seed bank persistence, soil seed bank     

Plant functional trait responses to grassland succession over 25 years

Abstract. Many grasslands and arable fields have been abandoned in Europe in the last decades. So far, however, ecologists have not been able to establish one general, unifying theory for successionally induced changes in species composition following abandonment. In fact, the course of succession seems to be unique for each site and year. Here we focused on the analysis of plant functional traits in order to detect general trends in trait responses to succession which would prove independent of site characteristics. We studied permanent plot series of 14 grassland sites in SW Germany that had been abandoned for 25 yr. Species composition as well as the course of succession varied significantly among the grasslands. Response to succession was analysed for the following traits: ‘plant height’, ‘canopy structure’, ‘specific leaf area’ (SLA), ‘storage organ’, ‘vegetative spread’, ‘plant persistence’, ‘seed bank longevity’, ‘start of flowering’, ‘duration of flowering’ and ‘seed mass’. We compared a univariate with a multivariate approach. In the univariate approach, attributes of each trait were analysed separately employing GLM whereas in the multivariate approach all traits were handled together in NMS. Both analyses indicated similar trait responses to succession. There was a significant increase in the proportion of species characterized by the following attributes: plant height > 0.6 m, leaves distributed regularly along the stem, vegetative spread > 100 mm, start of flowering later than May, duration of flowering 1–2 months and seeds of either low or high mass.     

Soil seed-bank composition reveals the land-use history of calcareous grasslands

We compared soil seed banks and vegetation of recent (established on abandoned arable fields) and ancient (continuously managed as pastures at least since 1830) calcareous grasslands if there is any impact of former arable field use. The study was carried out in two regions of Southern Germany with well-preserved dry grassland vegetation: the western Jurassic mountains (Kaltes Feld) and the climatically drier eastern part of Southern Germany (Kallmünz).Total number of species in the seed bank was similar in both regions, but species composition partly differed, reflecting phytogeographical differences between the regions. The total number of emerged seedlings showed a large disparity (5457 compared to 2523 seedlings/m² in Kaltes Feld and Kallmünz, respectively).Though there were differences in seed bank composition and size, we found a uniform pattern of plant traits (affiliation to phytosociological groups, Raunkiaer plant life-forms and seed longevity), which depended on the age of the grassland.The main conclusion is that seed banks in contemporary calcareous grasslands still reflect the history of former land use – in this case arable cultivation, even though it occurred a long time ago (up to 150 years). Indicators of former arable fields are germinable seeds of weeds which have persisted in the soil to the present. By contrast, weedy species are completely absent from the seed banks of ancient grasslands. Soil seed banks of recent grasslands may be of substantial conservation importance because they may store seeds of rare and endangered weed species such as Kickxia spuria, Silene noctiflora and Stachys annua, the majority of which have already gone extinct from the current vegetation of the study sites.     

Assessing the risks of transgene escape through time and crop-wild hybrid persistence

Transgenes introduced into crops can escape in time, as well as space, via the seed bank. For annual plants, especially ruderals, seed bank behaviour may be the most important factor determining population persistence. Crop seeds may exhibit some dormancy and germination cueing in the soil but are expected to be less able to persist than their wild relatives, which often have considerable dormancy and longevity, as well as effective germination cueing responses. Crop-wild hybrids may have seed bank characteristics more suited to persistence, and maternal effects may favour persistence of hybrids having wild plants for their female parent. Escape of transgenes via crop-wild hybrids presents unique concerns not present for crops. Hybrids can undergo natural selection and may back-cross with wild plants. We suggest methods that can be used in conjunction with evaluation of the relative fitness of crop-wild hybrids that will determine the likelihood of back-crossing. Accurate assessment of escape in time and transgene persistence via crop-wild hybrids requires proper plant materials. We emphasize the use of null segregants as controls for transgenic crops and for generating crop-wild hybrid controls for transgenic hybrids. Since good empirical and theoretical understanding of how individual genes influence the fate of plants in different environments is lacking, evaluation of escape in time and the persistence of transgenes via crop-wild hybrids should be on a case-by-case basis.     

Which traits promote persistence of feral GM crops? Part 1:implications of environmental stochasticity

Transgenes in plants affect life history traits including seed survival and germination. With stochastic matrix models we predict population-level consequences of transgene induced life history changes. We assess systematically which changes in life history traits, resulting from genetic modification, may increase the risk of invasion and persistence of feral crops or increase fitness in case of introgression from arable fields into conspecific, feral populations. We apply our method to feral populations of oilseed rape. Like many annual weeds, oilseed rape depends critically on disturbance; in undisturbed habitats it is generally outcompeted by perennials. The associated inherent variability and unpredictability render deterministic models inappropriate. With a stochastic matrix model we study population growth rate, elasticities and quasi-extinction times. Our results indicate that changes in survival in the seed bank impact population growth and persistence most. Less important are dormancy, fecundity and seedling survival. The predicted distribution of extinction times is highly skewed, with some patches persisting for decades.     

A model for small‐scale seed bank and standing vegetation connection along time

Since inter‐annual climatic variability influences composition and structure of seed bank and extant vegetation, it is expected that it also affects the relationship between both compartments at small scales along time. We hypothesize that seed bank and aboveground vegetation are closely and sequentially linked at small spatial and time scales. We conducted a field spatially explicit investigation to explore the spatial and temporal relationship between both compartments. Abundance, composition and spatial structure at different small‐scales of seed bank and aboveground vegetation were analysed through 100 permanent plots during two consecutive (dry and wet) periods of growth. Following a conceptual transition path model, we analysed changes in correlation values in composition and spatial aggregation between both compartments along time including seasonal variations of seed bank. Shape of spatial structures were evaluated using partial Mantel correlograms. Annuals and perennials guilds were studied separately. During the wet year, annuals increased their aerial spatial aggregation and cover, whereas the opposite happened for perennials. Density and spatial aggregation of the seed bank increased for both guilds in the two seasons following the rain period, especially in annuals. The clumped structure of the transient seed bank is a consequence of the addition of spatial structure of extant vegetation and the persistent seed bank. Expression of the persistent seed bank is much lower during the dry period for the annual guild. In spite of the strong yearly variability, the community maintained a highly structured spatiotemporal pattern. The mechanisms promoting this stability differed for annual and perennial guilds. Temporal persistence in perennials relied mainly in established plants longevity, whereas annual guild persistence depended solely on seed bank. This tight structure was coherent with the existence of successional dynamics in the community, although persistent seed bank could moderate the pace. Longer term studies of seed bank–standing vegetation dynamics are required to fully understand this process.     

Reproductive diversity and survival of the potential annual Diplotaxis harra (Forssk.) Boiss (Brassicaceae) in Egypt

Diplotaxis harra is a common annual species in the desert of Egypt, The investigated population was located in Wadi Hof near Helwan city. Additional water in relatively wet years cause these ephemeral plants to live longer and become perennials. The species has ephemeral, modular and coppiced life‐cycles and may shift from a r‐ to a K‐selected strategy. The germinable seed bank represented 15,6% of the total seed rain, out of which only 0,03% established as adult plants. About 72.7% of the adult plants were ephemeral. 27.0% modular and 0.3% coppiced plants. The highest contribution of the seed rain came from modular plants and the lowest contribution from coppiced plants. The survivorship curve of the coppiced plants exhibited a combination of Deevey type I and III curves. The ephemeral and modular plants showed steep curves. In contrast to ephemeral plants, the coppiced plants attained the highest reproductive value and the lowest reproductive effort, while the modular plants demonstrated intermediate values. The active phonological cycle started earlier in coppiced plants than in the modular and ephemeral plants. The period from the start of vegetative growth lo shoot‐die back extends from mid‐December to early June in coppiced plants, from February to May in modular plants, and from late February to early May in ephemeral plants. The living stumps of the coppiced plants remained dormant during summer and autumn months. Seeds collected from coppiced plants attained high germinability and viability, while those from ephemeral and modular plants attained low values. Seed longevity was higher in the modular plants (> 10 yr) than in seeds of ephemeral and coppiced plants (7–8 yr). The high energy content of the seeds produced from different cohorts over ten years of storage enabled the population to maximize the germinability. viability and longevity of seeds despite the unpredictable environment. The potential annual behaviour of D. harra is mainly attained by the persistence of viable seed bank and root‐shoot stock. The ability to shift from a r‐ to a K‐selected life‐cycle, phenological variations among the different cohorts of the population, variations in seed longevity and energy content strengthen intra‐population interactions.     

Does seed mass drive interspecies variation in the effect of management practices on weed demography?

Optimizing the effect of management practices on weed population dynamics is challenging due to the difficulties in inferring demographic parameters in seed banks and their response to disturbance. Here, we used a long‐term plant survey between 2006 and 2012 in 46 French vineyards and quantified the effects of management practices (tillage, mowing, and herbicide) on colonization, germination, and seed survival of 30 weed species in relation to their seed mass. To do so, we used a recent statistical approach to reliably estimate demographic parameters for plant populations with a seed bank using time series of presence–absence data, which we extended to account for interspecies variation in the effects of management practices on demographic parameters. Our main finding was that when the level of disturbance increased (i.e., in plots with a higher number of herbicides, tillage, or mowing treatments), colonization success and survival in large‐seeded species increased faster than in small‐seeded species. High disturbance through tillage increased survival in the seed bank of species with high seed mass. The application of herbicides increased germination, survival, and colonization probabilities of species with high seed mass. Mowing, representing habitats more competitive for light, increased the survival of species with high seed mass. Overall, the strong relationships between the effects of management practices and seed mass provide an indicator for predicting the dynamics of weed communities under disturbance.     

Seed bank diversity in mesic grasslands in relation to vegetation type,management and site conditions

Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow‐pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above‐ground vegetation, edaphic factors and land‐use history? 3. Are there differences in C‐S‐R plant strategy types and seed longevity under different management regimes? Location: Lahn‐Dill Highlands in central‐western Germany. Methods: Above‐ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above‐ground vegetation, site conditions and land‐use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above‐ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R‐strategy compared to hay meadows. In contrast, in hay meadows and meadow‐pastures proportions of S‐strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re‐establish from the seed bank after disappearance from above‐ground vegetation.     

Life history traits as predictors of plant rarity, with particular reference to hemiparasiticOrobanchaceae

We made a comparison between life history and reproductive characteristics of a group of Dutch rare (30) and common (105) plant species, all dicotyledonous and insect-visited forbs. The traits life span, clonality, breeding system, seed production, seed dispersal, and soil seed bank longevity are considered. All trait values have been ranked according to their possible effect on the vulnerability of a species for extinction, where low values indicate a low risk and high values a high risk for a species. Rare and common plants differed significantly in four traits: seed production, breeding system, seed bank longevity and clonality. The discriminant analysis showed that 79% of the 135 species was correctly classified in the group they presently belong to. Especially species that are rare now but were much more common 50–100 years ago were classified as being common species, pointing at the detrimental effect of habitat loss these species encountered at which they have not yet been able to adapted to. The data set include eight hemiparasitic plant species (familyOrobanchaceae), of which six are endangered in The Netherlands and two more common. Both rare and common hemiparasites scored high traits values, indicating that they are vulnerable for extinction. The hemiparasites had several characteristics in common: they are all annuals or biennials that have no means of clonal reproduction. They all have non-clustered, zygomorphic flowers that produce nectar and have a precise pollen presentation and are mainly visited by (bumble) bees. Mean life span was significantly shorter in the eight hemiparasiticOrobanchaceae than in the non-hemiparasiticScrophulariaceae used for comparison (4 species). Overall, we concluded that hemiparasitic plants have a special combination of life history traits which makes them vulnerable for local and regional extinction.