Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Biodiversity management of fens and fen meadows by grazing,cutting and burning

Question: Can the biodiversity of fens in Europe and North America be maintained through the use of grazing (especially cattle grazing), fire, and/or cutting? Location: European and North American fens. Methods: This paper is a review of the literature on the effects of grazing, fire and cutting on fens, to explore the relationship between management and biodiversity in fens. Results: A reduction of cattle grazing, mowing and burning in fens has led to a reduction in biodiversity in fens. The vegetation of abandoned fens shifts to trees and shrubs after 10–15 years, which shade the smaller and rarer species of these wetlands. While careful use of fire is used to manage fens in North America, it is not widely used in European fens, perhaps because the peat of drained fens may catch fire. Cattle grazing cannot be considered a natural disturbance in North America, since cattle did not evolve on that continent. In Europe, cattle do not generally graze in unaltered fens, but they do use slightly drained fen meadows. Conclusions: Three approaches have been used to control the dominance of tall woody and herbaceous species in abandoned fens, including the re‐introduction of cattle, mowing, and burning. Overgrazing results in a permanent reduction in biodiversity, therefore cattle re‐introduction must be approached cautiously. In Europe, but not in North America, mowing has been an important management tool, and mowing has been successful in maintaining species richness, particularly in fens that have been mowed annually for centuries. Fire has been the most common and successful management tool in North America although it is not effective in removing shrubs that have become large. Because the problems and solutions are similar, the literature of both European and North American fen management can be analyzed to better assess the management of fens on both continents. Many management questions require further study and these are listed in the paper.     

Effects of removal of organic matter on the productivity of heathlands

Abstract. Effects of milling, mowing and sod cutting on productivity in a Deschampsia grass heath and a Molinia-Deschampsia grass heath were studied from 1977 to 1986. The sum of above-ground biomass, litter and accumulated humified matter (TOM) in both types was ca. 70 ton/ha and the primary productivity 3–5 ton ha^-1yr^-1. Mulching did not affect the annual production. Sod cutting reduced the productivity to 1–2 ton ha^-1yr^-1; on the Molinia-Deschampsia site this reduction lasted for at least a decade, while the Deschampsia heath started to recover from sod cutting soon and increased in productivity again. Annual mowing both with and without removal of hay reduced the production as well, particularly on the Molinia-Deschampsia heath. Lower production was either the result of phosphorus depletion - Molinia-Deschampsia site - or nitrogen depletion - Deschampsia site. In the mowing treatments there was also a depletion of carbohydrate reserves.     

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.     

Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows

We examined effects of abandonment on species diversity and species composition by comparing 21 calcareous fen meadows in the pre-alpine zone of central and northeastern Switzerland. Meadows were divided into three classes of successional stages (mown: annually mown in late summer, young fallow: 4–15 years, and old fallow: >15 years of abandonment). In each fen, we measured litter mass in four 20 cm×20 cm plots, as well as (aboveground) biomass and species density (number of species per unit area) of bryophytes and vascular plants. Bryophyte biomass was reduced in abandoned fens, whereas litter mass and aboveground biomass of vascular plants increased. Species density of both taxonomic groups was lower in abandoned than in mown fens. Young and old successional stages were not different except for bryophytes, for which old successional stages had higher species density than young stages. We used litter mass and aboveground biomass of vascular plants as covariables in analyses of variance to reveal their effects on species density of both taxonomic groups. For bryophytes, litter mass was more important than vascular plant biomass in explaining variance of species density. This indicates severe effects of burying by litter on bryophyte species density. For species density of vascular plants, both vascular plant biomass and litter mass were of similar importance in explaining the decreased species density. Canonical correspondence analyses showed that abandonment also had an effect on species composition of both bryophytes and vascular plants. However, young and old successional stages were not different indicating fast initial changes after abandonment, but slow secondary succession afterwards. Furthermore, indicator species analysis showed that there was no establishment of new species after abandonment that might dramatically alter fen communities. Re-introduction of mowing as a nature conservation strategy may thus be very promising – even for old fallows.     

Effect of three mound-building ant species on the formation of soil seed bank in mountain grassland

The effect of three ant species (Lasius flavus, Formica spp., Tetramorium caespitum) on soil seed bank formation was studied in temperate mountain grassland. Seed removal experiments, analysis of soil seed content and seed survival experiments were carried out to evaluate the influence of ground ants on the seed fate. In the seed removal experiment seeds of 16 species, including 5 species with elaiosome-bearing seeds (myrmecochores), were exposed and their removal followed for 39 h. On average, ants removed 63.8% of myrmecochorous seeds and 10.9% of seeds without adaptation to ant dispersal. Analysis of soil seed content revealed that myrmecochores, in spite of expectations that they would accumulate in nests of seed dispersing ants, were most abundant in the soil of control plots. Evidence on seed relocation to the ant nests was obtained from a comparison of mounds of seed dispersing and seed non-dispersing ant species, as more seeds were found in the mounds of Formica spp. and Tetramorium caespitum (seed dispersers) in comparison with the mounds of Lasius favus (non-disperser).The soil seed bank of the compared microhabitats (control plots and mounds of 3 ant species) differed in their species composition, seed abundance and vertical distribution. The most distinct qualitative differences were between seed flora of control plots and mounds of Tetramorium caespitum. Control plots had approximately 30,000 propagules per m², which was double the number of seeds found in the ant mounds. In control plots, abundance and diversity of seeds steeply declined with depth; this trend was not observed in the mounds probably due to bioturbation. In the seed survival experiment, more seeds (2 out of 3 species) survived in control plots, which may also contribute to the higher seed abundance in this microhabitat.This study showed that seed relocation by ants does not contribute significantly to seed bank build-up at this study site. Ants may, however, increase the regeneration success of myrmecochores, mainly by dispersal for distance and placement in a larger spectrum of microsites, in contrast to species not adapted for myrmecochory.     

围封沙质草甸土壤种子库与幼苗库的特征

采用野外定点取样和萌发实验相结合的方法,研究了科尔沁沙地围封沙质草甸土壤种子库和幼苗库的特征.结果表明:土壤种子库总密度为13854±1651有效种子数/m2,而幼苗库的总密度为808±164株/m2,仅为土壤种子库总密度的6%,土壤种子库与幼苗库总密度之间存在显著的差异 F=36.0,n=219,P<0.001 .土壤种子库与幼苗库的植物种数分别为37和44,其中一、二年生草本植物分别占种子库与幼苗库植物种数的62%和50%,而多年生草本植物所占的比例分别为24%和41%,灌木所占的比例很小,分别为14%和9%.土壤种子库与幼苗库种类组成上的相似性系数为0.667,表现出较小的异质性.土壤种子库密度与幼苗库密度之间存在显著的相关性,其间关系可用三次曲线来描述.     

The persistence of calcareous grassland species in the soil seed bank under developing and established scrub

The relationships between the composition of the soil seed bank, the field layer vegetation, and the scrub canopy were investigated along a 69 m transect, grading from incipient woodland, through scrub, into intensively rabbit-grazed calcareous grassland. The results are used to assess the persistence of species associated with open calcareous grassland in the seed bank under developing scrub. Scrub age, composition and density, changed along the transect from the woodland to open grassland. A total of 35 forb and grass species were found in the field layer. The pattern evident in the scrub layer was also reflected in the herbaceous vegetation. The field layer in the most closed portion of the transect, where the scrub was oldest, was dominated by shade-tolerant species normally associated with woodland habitats. The abundance of these species decreased along the transect as the scrub age declined, and the field layer became increasingly dominated by species typical of open grassland. A total of 47 species germinated from the seed bank. Few species were recorded in the seed bank along the entire length of the transect. Overall, the seed bank was dominated by Hypericum perforatum and Centaurium erythraea, which accounted for 38.2% and 28.6% of emerging seedlings respectively. As with a number of similar studies, the composition of the seed bank had a low correspondence with the composition of the field layer vegetation. The results also emphasise that the composition of the seed bank can be viewed as an ecological palimpsest, with germinable seed of species from each stage of the old-field succession occurring in the soil. The seed bank is an important component in the re-vegetation of an area after disturbance such as scrub removal. This study supports the findings of previous research in showing that relatively few characteristic calcareous grassland species form persistent seed banks. The soil seed bank would therefore appear to be of limited value in the restoration of such grassland following scrub removal.     

科尔沁沙地围封沙质草甸土壤种子库特征的研究

科尔沁沙地沙质草甸围封 5年后土壤种子库中植物种类有 38种 ,其中一、二年生草本植物 2 4种 ,多年生草本植物 9种 ,灌木 5种。土壤种子库密度为 (1385 5± 16 5 1)粒·m-2 ,其中 ,一、二年生草本植物所占的比例最大为 96 5 8% ,前 5种一、二年生草本植物黄蒿、灰绿藜、猪毛菜、狗尾草和画眉草占总数的 87 2 0 % ,而多年生草本植物和灌木类种子库密度分别占0 94 %和 2 4 8%。若以单子叶和双子叶植物统计 ,单子叶植物种子数占 13 13% ,双子叶植物的种子数占 86 87% ,其中豆科植物仅占 2 83%。土壤种子库中所有植物种的密度与其相对应的地面植物种的密度呈显著非完全相关性 ,其关系可用三次曲线来表示。从种子库组成分析 ,已围封 5年的沙质草甸植被仍处在黄蒿 -杂类草混合植被阶段。     

A laboratory assessment of the relative susceptibility of Molinia caerulea (L.) Moench and Calluna vulgaris (L.) Hull to a range of herbicides

There is a concern that Molinia caerulea (L.) Moench may be increasing in upland moorland communities at the expense of Calluna vulgaris (L.) Hull and other ericaceous species. In order to develop a strategy for the control of Molinia, laboratory studies were carried out to determine the dose-response relationships of seven graminicides and glyphosate on both Molinia caerulea and Calluna vulgaris. Plants were grown under laboratory conditions and sprayed with increasing doses of herbicide using a precision sprayer. Results were highly variable, indicating the great morphological plasticity of these two species, making determination of the ED₅₀ difficult. Glyphosate, quizalofop-ethyl and sethoxydim successfully reduced various measures of Molinia growth to 50% of control levels (ED₅₀ estimates ranged from 0.41–0.67 kg a.i. ha^-1, 0.18–1.20 kg a.i. ha^-1and 0.37–0.49 kg a.i. ha^-1respectively). In contrast, only glyphosate reduced Calluna growth significantly (ED₅₀s ranged from 0.025–0.45 kg a.i. ha^-1), whilst selective herbicides left it undamaged. A comparison of the results for Molinia and Calluna suggests that there is no application rate of glyphosate which will reduce Molinia effectively whilst leaving Calluna undamaged. Recommendations for field testing of selective herbicides are discussed.     

Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow

Theeffects of light competition and litter on seedling recruitment and theperformance of established individuals were examined in Violapalustris. This polycarpic perennial plant was a common component ofspecies-rich fen meadows in northwestern Germany until the middle of thiscentury, but today is considered to be regionally endangered. From summer 1996until summer 1998 a bi-factorial field experiment combining three standing croptreatments (mowing, thinning, control) with two litter layer treatments (litterremoval, no litter removal) was carried out in an abandoned fen meadow toinvestigate the effects of these factors both on V.palustris and on aboveground species composition and diversity.MANOVAs (multivariate analysis of variance) revealed significant treatmenteffects for the performance of V. palustris related toexperimental manipulation. Mowing increased vital attributes including thenumber of rosette leaves, the percentage of individuals with chasmogamous andcleistogamous flowers and the number of rhizomes. In contrast, the length of theleafstalk, mean leaf area and the length of rhizomes were negatively associatedwith mowing. Litter removal significantly increased the number of V.palustris seedlings. The removal of the litter layer already resultedin an increase in aboveground species richness in the first year of theexperiment. After three years of experimental manipulation, both abovegroundspecies richness and diversity were positively associated with mowing.Redundancy Analysis showed that typical wet meadow species (Cardaminepratensis, Lychnis flos-cuculi, Lotuspedunculatus) and species of mesotrophic fens (Violapalustris, Agrostis canina, Potentillapalustris) were positively influenced by mowing. It can be concludedthat V. palustris recruitment and growth is promoted by aregular disturbance regime on fen meadows. The conservation of many typicalspecies of wet meadows and fens in northwestern Germany depends on moderategrazing or cutting. These management types prevent successional changes whichlead to an increase in standing crop and the development of a litter layer, bothof which can cause local extinction.     

Site‐specific responses of native and exotic species to disturbances in a mesic grassland community

Abstract. Grassland communities are increasingly recognized as disturbance‐dependent ecosystems, yet there are few replicated, multi‐site studies documenting vegetation responses to varying frequencies and types of grassland disturbance. Even so, land managers frequently manipulate disturbance regimes in an attempt to favour native grassland plants over exotic species. We conducted a factorial experiment testing three frequencies of clipping combined with litter accumulation, litter removal, and soil disturbance within the highly threatened California coastal prairie plant community. We monitored the response of native/exotic, grass/forb plant guilds once a year for four years. More frequent clipping reduced cover of exotic grasses and favoured exotic forbs, whereas native species were largely unaffected by clipping frequency. Litter accumulation, litter removal, and soil disturbance did not affect vegetation composition. Effects of litter accumulation may take longer than our experiment allowed, and soil disturbance due to our treatments was not sufficiently strong to show consistent effects relative to mammalian soil disturbance. Treatment response of some plant guilds differed among sites, highlighting the importance of replicating experiments at several sites before recommending conservation management practices.     

Abandonment alters community composition and canopy structure of Swiss calcareous fens

Abstract. We compared the plant species composition, productivity and canopy structure of seven mown sites to a chronosequence of 20 abandoned calcareous fens in northeastern Switzerland. Cessation of mowing led to an 18% decline in overall plant species richness and the diversity of most functional groups. Abandonment did not lead to marked increases of above‐ground productivity, but rather selectively favoured certain functional groups. On abandoned fens biomass of grasses increased nearly threefold, at the expense of biomass of Cyperaceae and Juncaceae, which declined by 30% compared to mown fens, while forb biomass remained unaffected. Litter mass increased more than 15‐fold in fallows, while canopy height increased by 50%. The foliage in abandoned fens was oriented more horizontally and had a lower overall cover. However, these successional changes were never dependent upon the age of the fallow. Furthermore, nearly all traits differed significantly on regional and local spatial scales, suggesting that floristic and (meso‐)climatic differences obscure or override successional trajectories in these species‐rich wetlands.     

Germination characteristics of some plant species from calcareous fens in southern Germany and their implications for the seed bank

Seeds of 25 plant species from calcareous fen hay meadows were exposed to different experimental conditions and their germination was characterised. Constant temperature inhibited germination especially in Cyperaceae . Both gibberellic acid and potassium nitrate failed to terminate dormancy. Increased germination rates were found in dicot species after treatment with gibberellic acid. Temperature fluctuations increased germination of Cyperaceae as well as dicotyledons. Treatment with gibberellic acid removed the chilling requirement in some of the species. Dormancy of small seeds with thin seed coats was broken by the application of gibberellic acid or fluctuating temperature; large thick-coated seeds were unaffected by gibberellic acid. No obligatory darkness requirement was found in any species; three species germinated irrespective of light treatments. All other species achieved higher percentage germination in daylight or in red (670 nm) light. Permanent darkness and far-red light (730 nm) reduced germination drastically. The results indicate that germination characteristics of the species investigated can be related to their seed bank types.     

Long‐term influence of sod cutting depth on the restoration of degraded wet heaths

Temperate heaths have an unfavorable conservation status in most European biogeographical regions. Increasing nitrogen levels promote competitive grass species such as Molinia caerulea, which is a main threat to heathland conservation in Europe. This article investigates the long‐term influence of sod cutting and the resulting changes in soil properties on the heath composition, integrity, and structure. In 15 nature reserves across the northern half of Belgium, we used (1) a large number of plots (203); (2) a broad range of sod cut depths (2–40 cm), and (3) a temporal dimension that describes how long the effects of sod cutting persist (census up to 19 years after sod cutting). Multivariate analyses were used in order to explore the influence of sod cut depth and time after sod cutting on the soil and vegetation properties. There was a positive relationship between sod cut depth and soil pH and water level, and a negative relationship with Al³⁺, NH₄⁺, and total organic matter (TOM). However, only a limited number of typical (target) species appeared after sod cutting, and then only weakly. Most of the time they remained a minor component of the restored vegetation. Moreover, M. caerulea reappeared and its cover significantly increased during the years following sod cutting. Although we were able to show that sod cut depth has a differential effect on soil properties and vegetation recovery, it also appeared that sod cutting does not restore wet heaths in the long term when applied in regions with high nitrogen deposition.     

Heat shock increases the reliability of a temperate calcareous grassland seed bank study

Reliable assessment of the density and species richness of the viable seeds in the soil is essential to estimate the probability of successful restoration of a particular plant community. Since temperate calcareous grasslands contain several thermophilous species typical of fire prone habitats, heat shock can be expected to break dormancy of target species for calcareous grassland restoration. We tested the effect of heat shock on the estimated density and species richness of seeds in soil samples. Heat shock treatments between 70 and 110°C resulted in both a higher estimated seed density and estimated species richness, in particular of specialist calcareous grassland species. Moreover, 25 species germinated exclusively from the heat-treated samples. This indicates that several species would have been missed by handling soil seed bank samples following the standard germination protocol, resulting in an underestimation of the restoration potential.