Vegetation re‐establishment on land previously subject to control of Pteridium aquilinum by herbicide

Abstract. Vegetation development after conventional bracken control with herbicide in the uplands is often slow and rarely results in the development of the target community. Two experiments were established on contrasting sites to investigate cost‐effective methods for the re‐establishment of vegetation dominated by Calluna vulgaris (heather), which is the usual target community seen as the desired replacement for Pteridium aquilinum in the uplands by many land managers. At one site, where grazing intensity was low and shelter was sufficient to prevent rapid Pteridium litter loss, the necessary treatments to establish Calluna were to disturb the litter and add a source of Calluna seeds. A difference between continued Pteridium control or a relaxation of control was not observed over the time‐scale of the experiment. At the second site, where grazing intensity was high and shelter minimal, vegetation dominated by Rumex acetosella had developed on a thin but dense litter layer. To enhance Calluna regeneration it was necessary to prevent sheep grazing, disturb the remaining litter layer and add a source of Calluna propagules. A nurse crop hindered Calluna establishment due to its vigorous growth. At neither site was an initial application of fertilizer an aid to Calluna establishment. It is possible to devise methods for the re‐establishment of Calluna at sites where previous attempts at Pteridium control alone have not had the desired effect on vegetation development. However, the precise choice of methods used is dependent on the grazing management at the site. High levels of livestock grazing prevented Calluna establishment, but some trampling may aid litter breakdown. The ability to adjust stocking rates and access to the site by machinery are critical factors to take into account in producing an effective strategy for long‐term Pteridium control and its replacement by target vegetation.     

Control of Pteridium aquilinum: meta-analysis of a multi-site study in the UK

Background and Aims

A great deal of money is spent controlling invasive weeds as part of international and national policies. It is essential that the funded treatments work across the region in which the policies operate. We argue that experiments across multiple sites are required to validate these programs as results from single sites may be misleading. Here, the control of Pteridium aquilinum (bracken) is used as a test example to address the following four questions. (1) Does the effectiveness of P. aquilinum-control treatments vary across sites? (2) Is the best treatment identified in previous research (cutting twice per year) consistent at all sites, and if not why not? (3) Is treatment performance related to P. aquilinum rhizome mass, litter cover or litter depth at the various sites? (4) Does successful P. aquilinum control influence species richness?

Methods

Pteridium aquilinum‐control treatments were monitored for 10 years using six replicated experiments and analysed using meta-analysis. Meta-regressions were used to explore heterogeneity between sites.

Key Results

The effectiveness of treatments varied between sites depending on the measure used to assess P. aquilinum performance. In general, cutting twice per year was the most successful treatment but on some sites other, less expensive treatments were as good. The effectiveness of treatments at different sites was not related to rhizome mass, but the effectiveness of most applied treatments were inversely related to post-control litter. Effective treatment was also associated with high species richness.

Conclusions

It is concluded that successful development of national weed control programs requires multi-site experimental approaches. Here, meta-analyses demonstrate that variation in effectiveness between sites could be explained in part by pre-specified variables. Reliance on data from a single site for policy formulation is therefore clearly dangerous.Key words: Bracken, Pteridium aquilinum, weed, control, herbicide, litter dynamics, species diversity, meta-analysis     

A multi-site assessment of the effectiveness of Pteridium aquilinum control in Great Britain

Questions: The objectives of this study were to assess the effect of Pteridium aquilinum (bracken) control treatments, at the national scale, and the impact of restoration practices, at the local scale, on P. aquilinum performance. Hypotheses: 1. Geographical location (locally between and within sites) affects the control of P. aquilinum through time. 2. Are the P. aquilinum control treatments successful at all sites, and if so which ones? 3. The treatments applied at the individual site level to restore vegetation influences the performance of P. aquilinum through time. Location: Four geographically distinct acid grassland and heathland sites infested with P. aquilinum across Great Britain. Methods: Six main-plot, bracken control treatments were applied to all sites with site-specific vegetation restoration treatments. Response variables (P. aquilinum cover, frond length and density) were monitored twice yearly, in June and August between 1993 and 2003. Results: Between- and within-site spatial variation was found, although impact is perhaps less than suggested from shorter-term data. Despite local variation all sites responded similarly to bracken control treatments; asulam treatment resulted in a rapid reduction in frond performance followed by a continued recovery taking approximately ten years to return to untreated values. Cutting treatments tended to have a slower impact at the start but an increasing one over time, especially cutting twice per year. Restoration treatments had a limited impact; the only significant effect in August was grass seeding on frond length at Sourhope. In June only, the plots where sheep were fenced out showed a significant reduction in P. aquilinum cover at Peak. Conclusions: Long-term control of Pteridium aquilinum at all sites and on all measures was best achieved using a continuous cutting treatment, preferably twice per year.     

The effects of cutting pre-treatments on the performance of asulam on bracken (Pteridium aquilinum)

The performance of asulam was assessed from a field experiment in which bracken (Pteridium aquilinum) was subjected to a variety of cutting pre-treatments prior to spraying with the herbicide. Both cutting bracken twice in the year prior to asulam application, and spraying frond regrowth 10 wk after a single mid-June cut, improved the performance of asulam. Summer cutting removes nutrients and dry matter from the system and reduces frond height, but increases frond density and the number of active buds on the rhizome system. The enhanced efficiency of asulam on pre-cut plots may be due to any one of these factors, or a complex interaction between some (or all) of them. In areas of tall bracken a cutting pre-treatment has the important practical advantage of reducing the height of the canopy, making the subsequent use of ground spraying equipment easier.     

The effects of control on the biomass, carbohydrate content and bud reserves of bracken (Pteridium aquilinum (L.) Kuhn), and an evaluation of a bracken growth model

This paper examines the initial effects of bracken control on frond numbers and biomass, and the biomass, carbohydrate reserves and bud densities of bracken stands cut once per year, twice per year, subject to a single application of asulam or left untreated. The seasonal dynamics of these parameters are displayed; carbohydrate and biomass are both removed from the rhizome system to produce frond tissue, and are replenished at the end of the growing season. Asulam application reduced densities of both active and dormant buds, and both frond biomass and density. It did not significantly reduce rhizome biomass or carbohydrate reserves in the two years after treatment. Cutting, either once or twice per year reduced both rhizome biomass and rhizome carbohydrate reserves, as well as bud densities, though the latter were reduced in proportion to biomass. Cutting twice a year reduced the production of fronds, both in numbers and biomass. The collected data were used to evaluate a model of bracken growth, and subsequently to improve estimates of some of the model parameters. The model simulations of control treatments were compared to field data. The effects of cutting once per year and spraying with asulam were predicted accurately, but the bracken stand was more resilient to cutting twice per year than would be expected from model predictions. The combination of cutting and spraying is discussed as a potential tool in land management and the deficiencies of the model are discussed in relation to the need for future research into the biology of bracken.     

Effects of the litter layer of Pteridium aquilinum on seed banks under experimental restoration

Questions: Does the litter layer of Pteridium aquilinum (bracken) act as a barrier to certain species in the seed bank? Does bracken control/restoration treatment affect seed transfer through the litter layer? Location: Five experiments at three sites across the UK covering two major vegetation types; acid‐grassland and heath‐land. Methods: At each experiment a range of bracken control and vegetation restoration treatments were applied for about ten years. The seed bank was sampled in both the bracken litter and the soil. The cover (%) of each species in the vegetation and the bracken litter abundance (cover and depth) was also estimated. Results: The bracken litter layer acts as an inert barrier as it contained a large proportion of seeds available in the litter‐soil profile (38%– 67% of the total). Bracken litter depth and cover also influenced significantly the seed bank composition in both the bracken litter and the soil. These effects were site‐specific, and species‐specific. The application of treatments changed significantly the balance between seed inputs and outputs in the bracken litter layer for some species. This was either a positive or negative response relative to the untreated control plots. Conclusion: For heathland and acid‐grassland restoration, the bracken litter layer may be an important seed source, but it must be disturbed particularly before seed addition.     

Effects of increased temperature and nitrogen availability on the displacement of Calluna vulgaris by Pteridium aquilinum

Abstract. Atmospheric greenhouse gas concentrations will continue to rise for the foreseeable future, and, due to radiative forcing, surface temperatures will also increase. If soils warm sufficiently, this will result in increased litter decomposition rates, while nitrogen availability will also rise due to increased deposition. A field-based experiment manipulating air temperatures using polythene tents and nitrogen availability by adding 50 kg N ha^-1 yr^-1 has shown that this will result in further encroachment of Pteridium into Calluna moorland. Although this experiment, treating plots on and around the boundary between Pteridium and Calluna, did not show any statistically significant effect on the movement of the boundary, there were clear indications of Calluna dieback at the boundary. In addition, the frequency of Pteridium fronds increased in the Calluna-dominated areas, while Calluna became less abundant in the pure Pteridium plots. Calluna dieback was not due to litter accumulation as is frequently assumed, but the dieback is likely to be caused by shading of Calluna by the increased growth of Pteridium fronds. Botanical diversity was higher in the Pteridium plots, but considering the importance of grouse shooting for the maintenance of Calluna moorland, displacement of Calluna by Pteridium may result in a lower conservation and economic value of moorlands.     

The invasion of Pteridium aquilinum and the impoverishment of the seed bank in fire prone areas of Brazilian Atlantic Forest

Several studies have documented that fires are widespread in the tropics. Because fire in the Atlantic Rain Forest is rare, fire events and their consequences at the community level have not been evaluated. This study describes the composition of different seed banks in areas of Atlantic Forest. The study was carried out in the National Park of Tijuca in the state of Rio de Janeiro. Four different areas were chosen considering their differences on fire history. Fifteen soil surface samples (23 cm (centimeter) diameter ×7 cm height at 5 cm deep) were obtained from random locations within each site. The dominant family at all study sites except in the most preserved one were Dennstaetiaceae, followed by Melastomataceae and Poaceae. The family Dennstaetiaceae was exclusively represented by Pteridium aquilinum (Klf.) Herter. There were no significant differences among areas when comparing the number of seedlings excluding P. aquilinum. However, the number of P. aquilinum was higher in the most disturbed area, while Melastomataceae and Rubiaceae were more frequent in a less impacted area. Our results suggest that the seed bank in disturbed areas of the Atlantic Forest possibly will not contribute for forest restoration after disturbance.     

Evaluation of a bracken (Pteridium aquilinum (L.) Kuhn) growth model in predicting the effects of control strategies across a range of climatic zones in Great Britain

A nationwide bracken control experiment provided data over a three-year period for testing the accuracy of a bracken growth model (BRACON). Objective assessments of model validity identified the model as a reasonably accurate predictor of bracken stand dynamics given the range of environmental conditions currently prevailing in Great Britain, in relation to (1) cutting regimes and (2) spraying asulam. Ranking of treatment efficacy at individual sites was closely reflected in model predictions. Predicted response of the rhizome system to cutting treatments underestimated bracken resilience. Failure to consider the consequences of frond regrowth in the latter portion of the growing season as a means to offsetting energy loss was identified as a potential explanation for this discrepancy. The role of the model as a practical management tool is discussed with particular reference to relating model predictions, based upon generalised environmental data (40 km plusmn; 40 km grid cells), to bracken management at individual sites.     

The effects of cutting and asulam on numbers of frond buds and biomass of fronds and rhizomes of bracken Pteridium aquilinum

The effects of cutting and the herbicide asulam for the control of a dense uniform stand of bracken Preridium aquilinum were investigated experimentally over a 6 —yr period in Norfolk, England. Bracken is a particularly difficult species to control by either mechanical or herbicidal means because of its extensive and complex rhizome network, containing large reserves of dry matter and a large number of frond buds. Cutting bracken in the summer removed dry matter from the system and reduced frond vigour but did not reduce the number of frond buds to a critical level. Whereas the first few cuts substantially reduced the standing crop of fronds, relatively stable standing crops of fronds were observed in future years despite continued cutting. Asulam caused a 99% reduction in the standing crop of fronds in the year after spraying, but in the absence of further treatment, fronds recovered to pre-spray levels after 6 yr. Asulam caused severe localised damage to frond buds and rhizome apices, but otherwise the rhizome network remained intact. New sections of frond bearing rhizome (short shoots) developed on storage rhizome (long shoots) deep in the soil of both the asulam plots and those cut.     

Effects of restoration treatments on the diaspore bank under dense Pteridium stands in the UK

Abstract. The diaspore bank (seeds of higher plants and spores of ferns and bryophytes) was assessed between 3 and 5 yr after experiments to control Pteridium aquilinum (bracken) and restore appropriate vegetation were initiated at two contrasting locations in the UK. We tested the response of the diaspore bank using univariate and multivariate analysis of variance. The two approaches were complementary and together improved the interpretation of these results. There were considerable differences in the diaspore banks of the two sites and among the experimental locations within sites. Within each experiment there were differences in species composition, with species that were (1) common to both diaspore bank and vegetation, (2) restricted to the diaspore bank and (3) restricted to the vegetation. There is a possibility of increasing the biodiversity of the developing vegetation if some of the species present in the diaspore bank can be germinated. This was especially true for ferns where four species were found in the spore bank which were not present in the vegetation. There were few significant effects of management treatment on the diaspore bank as the experiments had been in progress for only 3 to 5 yr, but a few species had different densities in the different treatments (Betula pubescens, Juncus effusus and some bryophytes). The greatest correlation between vegetation and diaspore bank was found at the top hierarchical level (entire dataset) and this progressively reduced with scale. We interpret this as a landscape/species pool effect: as the scale of the study reduces the correlation between diaspore bank and vegetation also reduces, at least over the time scale of our study. The relevance of these results for restoration ecology is discussed briefly.     

Efficacy of bracken (Pteridium aquilinum (L.) Kuhn) control treatments across a range of climatic zones in Great Britain

Bracken control field experiments were conducted at six locations across Great Britain. The effects of various cutting and herbicide management regimes upon the seasonal dynamics of bracken fronds and rhizomes were examined over a three year period. This enabled a national overview towards bracken control to be constructed. Initially, spraying with asulam was the most effective treatment in reducing frond biomass and density but was least effective in reducing rhizome biomass. Differential reductions in rhizome biomass were observed in relation to cutting frequency, with cutting twice yearly giving superior control. The national trends confirmed a number of previous observations from independent single-site studies; however, others were contradicted. Comparable management options were ranked consistently between the current and former investigation, however, the initial importance of differences in cutting frequencies did not agree between studies. This multiple-site study improves understanding of the consequences of a national bracken control programme by reducing the influence of confounding site-specific factors, and recommendations for the most appropriate bracken control techniques are made. The extent to which individual sites reflect a national trend in response to bracken control is considered and sites are compared. The hierarchy of treatments identified at the national scale was found to apply generally within individual sites. However, several responses which proved significant at the countrywide level were not so clearly defined at the site scale. Cutting once yearly was the only management regime which appeared to give different bracken control between sites. All other treatments gave similar responses between sites. This result was found in the Scottish Borders, during the second year of control, when frond biomass and density (relative to untreated plots) were greater than that recorded at other sites. This contrast was not found in subsequent monitoring. In terms of rhizome biomass depletion, poorer control was achieved following cutting once yearly at the northern sites (Mull, Scottish Borders, Lake District) compared with the southern sites (Clwyd, Breckland, Devon). The implications of experimental results are discussed in relation to increased cost effectiveness of national bracken control programmes.     

The effects of Fungi on the Gametophytes of Pteridium aquilinum (L.) Kuhn

I. The prothalli have been shown to be convenient subjects forthe study of some fungal relationships with green plants. Purecultures have been established from all the populations examinedby germination of young spores, and it has been shown that duringdormancy the number of spores which can germinate in pure culturedecreases more rapidly than the number which can germinate incultures containing contaminant fungi. Previous reports of thestimulatory effect of fungi on prothallial growth and developmenthave been confirmed and extended. 2. Prothalli growing on mineral agar have been shown to be potentiallysusceptible to the attack of a wide variety of pathogens ofhigher plants, but only two naturally occurring species of fungipathogenic to prothalli have been found during an examinationof growth on a variety of soils from bracken-infested areasin the west of Scotland. 3. The aetiology of the disease incited by Botrytis cmerca inprothalli has been shown to be similar to that of this diseasein higher plants. The pathogen is some-times restricted to lesionsof limited size around the sites of successful infections andthis is associated with changes in the cell walls of the host.A protopectinase solution prepared from a culture of germinatingspores by Brown's method induced symptoms in the prothallialcells similar to those induced by the attack of the fungus.The cell-attacking properties of the solution were greatly reducedwhen the protopectinase content was inactivated by low heatfor a short time, but they were not entirely destroyed. Thisconfirms previous indications that more than one complex maybe involved in the attack, but since the second moiety demonstratedis at least partially thermolabile the evidence does not supportjudgement of its relative significance in the attacking process.     

蕨菜乙醇提取物的Ames试验研究

目的：研究蕨菜乙醇提取物的致突变作用,为蕨菜的开发利用提供一定依据;方法：制备蕨菜乙醇提取物,采用Ames试验方法,以50μg/皿敌克松作为诱变剂,分别添加提取物50、100、200、400、800μg/皿,统计回变菌落数;结果：受试物各剂量组回变菌落数均未超过自发回变菌落数的2倍,且无剂量-反应关系;结论：在本实验条件下,蕨菜乙醇提取物无致突变作用.     

The effects of sod cutting and additional liming on potential net nitrification in heathland soils

The effects of sod cutting, a common restoration measure to remove excess nutrients from grass-dominated heathlands, on nitrification were studied in dry and wet Dutch heathlands and in incubation experiments. In the field, soil ammonium and nitrate concentrations were measured after treatment by sod cutting, with or without additional liming. Potential net nitrification was measured by incubating soil samples of all treatments with extra ammonium in a climate chamber at pH 6. Potential net nitrification of heaths dominated by Molinia caerulea was significantly higher than that of dwarf-shrub dominated heaths. Sod cutting of the former areas significantly decreased potential net nitrification, whereas in the latter areas no differences were found. Liming of sod-cut soils greatly increased potential net nitrification and the accumulation of ammonium in the soil up to toxic concentrations could be prevented. Our results show that the combination of sod cutting and liming would create suitable soil conditions for the germination and establishment of endangered plant species of dry and wet heathlands. The success of restoration projects of these areas can thus be increased.     

The effects of the aggressive species Pteridium caudatum on the vegetation of Socorro Island: Restoration challenges and opportunities

Humans’ impacts on natural environments have become pervasive, and natural regeneration is often hampered by extreme soil degradation. This situation is exacerbated in insular habitats where introductions of non-native grazing mammals have left behind completely barren areas. Once mammals are removed, aggressive native or non-native plant species can grow vigorously. This leads to the question of whether biodiversity can return through native succession, or if other actions are needed to prevent invading pioneer species from causing successional arrest. This occurs on the remote Socorro Island, in the UNESCO World-Heritage Revillagigedo Archipelago, Mexico. To understand whether native plants are establishing at sites invaded by Pteridium caudatum, we evaluated natural regeneration in sites with different Pteridium cover in two contrasting zones of the island (dry and mesic). We also calculated plant stress indicators (midday leaf water potential, chlorophyl content and maximum quantum yield) for two native species—Dodonea viscosa and Psidium socorrense—growing in open areas and under Pteridium. Our results showed that Pteridium is a significant barrier to seedling establishment. Thus, restoration actions such as seeding and/or plantings should focus on areas without Pteridium cover. Certain local species can be used to restore degraded areas: P. socorrese and Waltheria americana could be used in dry zones, while D. viscosa could be sown in either dry or mesic zones. For adult plants of the two dominant shrubs, Pteridium cover neither imposes or ameliorates stress. Our results suggest that the restoration of plant diversity in the vast Pteridium-invaded areas of Socorro Island will require management actions, including progressive physical removal of Pteridium cover to activate natural regeneration. Pteridium removal can be accelerated by nucleation, planting adults of native shrubs to generate shading. This strategy could be viable if restoration programs involve conservation institutions and government agencies, who currently have critical human and technological resources on the Island.     

The effect of thiouracil on the viability of eggs and embryogeny in Pteridium aquilinum

Summary Gametophytes of Pteridium aquilinum fed for six days with 50 and 75 p.p.m. thiouracil ceased to produce archegonia, but archegonia already initiated completed their development. Eggs produced in the presence of 50 p.p.m. thiouracil were viable, but embryogenesis was retarded. Gametophytes fed with 75 p.p.m. thiouracil mostly remained barren after insemination, the eggs being inviable, but others produced outgrowths from the archegoniate region which were either gametophytic, or sporophytic with a tendency to revert morphologically and functionally to gametophytic tissue.Thiouracil administered after fertilization either prevented embryogenesis, or retarded it, the sporophytes then being deformed and often showing a tendency to produce gametophytic tissue. The embryo became less sensitive to short periods of thiouracil as it developed, possibly because of the appearance of relatively quiescent regions, serving as reservoirs of unaffected cells.The results are held to support the view that the genes responsible for sporophytic growth become activated during oogenesis, and that the cytoplasm of the mature egg already contains the information leading to this kind of growth.     

Legacy effects of afforestation on prairie plant and seed bank communities in a northern Canadian prairie

Afforestation resulting from fire suppression, modified grazing, plantation establishment and climate change poses a threat to northern prairie ecosystems. Trees alter the composition and function of plant and soil fauna communities and can compromise the restoration of afforested prairies. To evaluate the hypothesis that legacies of afforestation persist in restored prairie communities and decrease the potential for restoration, we examined the composition, structure, and diversity of plant and seed bank communities along a 20 year chronosequence of plantation tree removal from a northern fescue prairie in Riding Mountain National Park, Manitoba, Canada. Tree removal increased the abundance of weedy species in the plant and seed bank communities of restored prairies and plant diversity peaked and declined over the 20 year period of passive restoration. As a result, time since tree removal and the encroachment of invasive species were key in explaining the composition of restored prairie communities. Low correlation between the species composition of plant and seed bank communities, including the complete absence of Festuca hallii in restored treatments, demonstrated that legacies of afforestation compromised the potential of seed banks to facilitate prairie restoration. We conclude that tree removal alone is insufficient for the restoration of northern fescue prairies and that, in the absence of active management, the persistence of low-diversity plant and seed bank communities constitutes an important legacy of afforestation and an important barrier to future restoration.