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.     

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.     

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.     

Long-term recovery of bracken (Pteridium aquilinum (L.) Kuhn) after asulam spraying

This paper investigates the effects of a single asulam application, sprayed from the air, on the rhizome biomass, bud density, fronds and carbohydrate reserves of bracken (Pteridium aquilinum (L.) Kuhn) using a time sequence approach. Regression models were used to investigate how these characteristics varied with time after spraying, and were used, where appropriate, to calculate the time taken for full recovery after treatment. Frond density and biomass recovered in approximately eight years, bud numbers in seven, but rhizome biomass and total carbohydrate reserves required 10 to 12 years to recover. The consequences of these results are compared with predictions from a computer model and discussed in relation to the best timing of re-treatment and the management needed for long term control.     

The effects of cutting and herbicide treatment on Pteridium aquilinum encroachment

Abstract. Pteridium aquilinum (bracken) encroachment is an important factor in the loss of certain habitats in the United Kingdom. However, no information exists as to whether prevention of encroachment is a cost‐effective strategy for Pteridium management. Conventional methods for the control of Pteridium (cutting, asulam application) were tested at one site (Levisham) to quantify their ability to prevent or delay encroachment and to affect the vigour of the Pteridium at the edge of the stand. The effects of encroachment and asulam application on the vegetation present were monitored at a second site (Ramsley), where techniques commonly used for moorland restoration were employed in combination with asulam application. Cutting once per year or a single application of asulam delayed the advance of the Pteridium front. At Levisham, the untreated front advanced 2.7 m in 5 yr, while in the same period the cut front advanced 0.88 m and the sprayed front was 1.5 m behind its initial position. At Ramsley, the untreated front invaded 1.8 m in 5 yr, and the sprayed front was again 1.5 m behind its starting position. Both spraying and cutting reduced frond biomass, frond cover and rhizome biomass. Herbicide spraying prevented the loss of Calluna vulgaris, though the restoration treatments had little effect. The merits of a balanced targeting of control on encroaching fronts or Pteridium at the stand level are discussed for different situations.     

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 mode of action of asulam [methy] (4-aminobenzenesulphonyl) carbamate] in bracken

Foliage-application of 4.4 kg/ha asulam [methyl (4-aminobenzenesulphonyl) carbamate] to field bracken reduced frond density by 97.1, 87.6 and 77.8%, one, two and three years respectively, after treatment. Examination of the rhizome system from these plots revealed that a high proportion of the rhizome apices and frond buds were killed, the remainder being damaged or remaining dormant. Autoradiography of bracken plants grown from sporelings or fragments and treated with 14 C-ring-labelled asulam revealed that uptake was rapid and progressive with time, whilst translocation was generally basipetal showing a typical source-sink pattern, with intense accumulation in the apices and buds. Optimum penetration and basipetal translocation was achieved when treatments were applied to almost fully-expanded fronds; application to immature fronds resulted in predominantly acropetal movement. Basipetal translocation was extensive in field bracken, accumulation occurring in the buds distant from the base of the treated fronds. There was evidence that asulam was relatively persistent in the rhizome system, 59 and 8% of the applied activity remaining in the rhizome, one and ten months respectively, after treatment. Hydrolysis of asulam produced small amounts of sulphanilamide (<10%) and traces of sulphanilic acid, 4-aminophenol and benzene sulphonamide. Studies on the effect of asulam on the incorporation of 14 C-labelled precursors into RNA and protein indicate that inhibition of RNA and protein syntheses may be major sites of action of asulam.     

Effect of herbicides on bracken rhizome survival

Glyphosate sprayed at 1.1, 2.2, 4.5 or 6.7 kg/ha on to bracken decreased the total carbohydrate content of the storage rhizome 20 months after spraying, the decrease being greater when spraying was carried out at the beginning rather than at the end of August. Asulam, benazolin and metribuzin, although reducing frond numbers, showed no significant effect on the rhizomes over the same period.     

Seasonal variations in the carbohydrate content of bracken

The amounts of dry matter, mobile carbohydrate and reserve carbohydrate have been determined in bracken fronds and buds, frond-bearing rhizomes and storage rhizomes over a period of 13 months. Dry matter and reserve carbohydrate levels are closely linked, especially in the rhizomes, falling from May to the beginning of July and rising until complete frond death at the end of September; but variations occur between frond-bearing and storage rhizomes. The carbohydrate reserve of frond-bearing rhizome is rapidly exhausted by the developing fronds and further carbohydrate is drawn from the storage rhizomes, but a concurrent fall in mobile carbohydrate suggests that the rate of conversion of reserve carbohydrate is slower than the rate of translocation to the fronds. By August, there is an accumulation of carbohydrate in the frond-bearing rhizome, which is followed by a fall in September and this again suggests low enzyme activity in the storage rhizome, but also that sink strengths in bracken are in the order (1) developing buds, (2) storage rhizome, (3) frond-bearing rhizome.
The responses of bracken to cutting and herbicide-application are discussed in relation to sink strengths and a low level of enzyme activity in the storage rhizome.     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

Control of bracken

Mechanical methods of bracken control are now being replaced by the use of chemicals. Trials indicate the value of asulam and glyphosate in reducing frond numbers of bracken when sprayed in late July or early August in the west of Scotland. There is still a need to find a bracken eradicant chemical rather than a control chemical. Research is also required to investigate the results of removing the bracken cover on the ecology of the treated areas. The effects of sudden exposure of the hitherto protected grass to extremes of climate are not known.     

Botanical implications of bracken control

In addition to bracken, Pteridium aquilinum , and other ferns, only a restricted range of species among the natural flora suffer severe damage if sprayed with asulam in late summer. Of the plants associated with bracken in relatively well-drained situations those affected most by the herbicide include the three Ulex species, Rumex acetosella , young plants of Calluna vulgaris , some Compositae and grasses of the genera Poa, Holcus and Agrostis (though not A. setacea ). In damp and wet upland habitats, Cirsium palustre, Juncus effusus, Glyceria fluitans and a few other species are moderately susceptible to asulam but the majority of plants appear to be resistant or only slightly damaged.
Areas from which bracken has largely been removed tend to be invaded by such species as Deschampsia flexuosa, Holcus mollis, Chamaenerion augustifolium, Digitalis purpurea and Urtica dioica . If grazing pressure is low the absence of competition enables woody species to regenerate from seed. Colonization and subsequent spread of the perennial plants is influenced by what was present as ground flora before the bracken was killed and by the depth of bracken litter.
Asulam is considered to be a useful aid to the management of nature reserves where encroachment by bracken needs to be checked. Its most widespread use for bracken control is likely to be on upland farms to clear bracken from marginal land with the aim of increasing and improving pasture. Adjoining woodland and moorland habitats may suffer from resulting changes in land management. Moreover, indiscriminate aerial spraying of asulam over large areas may affect local populations of plants, notably ferns, in the more open areas not protected by a canopy of bracken.     

The ecological status of bracken

Information about the morphology of bracken ( Pteridium aquilinum (L.) Kuhn var. aquilinum ) and the chief nutrients in the frond at different times of the year introduce an account of litter production and its accumulation in relation to the behaviour of frond, root and rhizome systems.
Where litter gain exceeds loss there is a correlation between the thickness and/or kind of litter and the level of the root and rhizome systems in relation to the mineral soil surface: with increase of litter the bracken becomes progressively more dependent for its physical and chemical soil environment on its own débris and less on the underlying mineral soil. An example of the limit of complete dependence has not been examined, but degeneration of the community can take place before that stage is reached.
From a review of the chief factors affecting bracken the conclusion is reached that the woodland habitat is both favourable and restrictive: in it bracken is in equilibrium with its environment, at a high social status. The relationship with other plant communities depends largely on the degree of human interference to which each is subject. Dominant bracken when left alone, and where gain of litter exceeds loss, becomes the victim of its own success; local degeneration opens the way for entry by other species.     

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 morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

Factors Affecting the Restoration of Heathland and Acid Grassland on Pteridium aquilinum–Infested Land across the United Kingdom: A Multisite Study

The variability in the success of Pteridium aquilinum (bracken) control and vegetation restoration has been highlighted as a major issue in the United Kingdom. Experiments were set up at four different regional locations to assess bracken control at the national scale and the impact of restoration practices at the local scale. Bracken control treatments (cutting once or twice per year, a combination of cutting and asulam spraying, and asulam in year 1) were combined with site‐specific treatments designed to restore appropriate heathland or acid grassland vegetation. This article considers the effects on the developing understorey vegetation, testing the following hypotheses: (1) local differences between sites would affect community change; (2) treatments applied to control Pt. aquilinum (same at all sites) influences community change; and (3) treatments applied at the individual site level to restore vegetation influences community change toward the target vegetation. There were a considerable number of spatial effects. It is, therefore, difficult to develop a one‐size‐fits‐all policy for vegetation restoration within a national Pt. aquilinum control strategy. Few bracken control treatment effects were found, and, where they were detected, it was only at single sites. Thus, the development of target vegetation requires a combination of control and restoration treatments that take into consideration the aspects of that site. Only three species, Deschampsia flexuosa, Galium saxatile, and Campylopus introflexus, increased as a direct effect of the control treatments. Vegetation restoration was most successful in the cutting‐twice‐per‐year plots, the treatment with the greatest reduction in Pt. aquilinum cover.     

The effect of glyphosate on frond regeneration, bud development and survival, and storage rhizome starch content in bracken

In addition to very effectively suppressing bracken frond regeneration for two years at levels of 2.24 kg a. i./ha and above, glyphosate had considerable effects on the rhizome system. Dry weight of frond-bearing rhizomes, number of living apices and developed buds, number and viability of dormant buds and starch content of storage rhizomes were all markedly reduced. The effects on dormant buds and starch content are particularly important when considering frond regeneration and as a consequence of these, glyphosate is likely to give long lasting control of bracken. Low levels of glyphosate (0.02 and 0.07 kg a. i./ha) did not stimulate bracken growth, while the addition of NH₄SCN apparently inhibited the action of the sub-lethal rate of 0.56 kg a. i./ha.     

What is a suitable management for Typha latifolia control in wet meadows?

Aim

Typha latifolia causes serious problems in wet meadows by overgrowing and suppressing other native plants. To determine suitable management for T. latifolia control, we addressed the following question: What are the effects of long-term cutting at different frequencies (once or twice per year and no management) and biomass removal on cover and other characteristics of T. latifolia, and on sward productivity and plant species composition?

Location

Malá Strana nature reserve, Jizerské hory Mountains, Czechia.

Methods

A long-term experiment arranged in a randomised block design with three blocks was established in 2005. Data were collected from five treatments: unmanaged control; cutting once a year in June without biomass removal and with biomass removal; cutting twice per year in June and August without biomass removal and with biomass removal. Percentage cover of T. latifolia and other vascular plant species was visually estimated and T. latifolia characteristics (tiller density, height, dry-matter biomass [DMB] yield and litter), sward height and DMB yield were measured during 2005–2018 at the end of June.

Results and Discussion

Regular cutting once or twice per year regardless of cut biomass removal led to reductions in tiller density, height, litter and DMB yield of T. latifolia. Biomass removal had only a slight tendency to affect T. latifolia characteristics. The higher frequency of cutting significantly decreased the mean T. latifolia cover, litter and DMB yield. Cutting once or twice per year regardless of biomass removal led to successive changes in plant species composition but had no effect on the species richness and evenness.

Conclusions

Cutting at least once per year without biomass removal seems to be sufficient to achieve a decrease in DMB yield and litter of T. latifolia plants, and thereby maintain the wet-meadow vegetation without loss of species richness and also preventing the overgrowth of shrubs and trees.     

Recovery of Moorland Vegetation after Aerial Spraying of Bracken (Pteridium aquilinum (L.) Kuhn) with Asulam

Bracken (Pteridium aquilinum) is a major weed of seminatural vegetation in Great Britain, as in many other countries. As a consequence, large areas have been subject to control with the intention of restoring the former vegetation. The use of aerial spraying with asulam, a narrow spectrum, carbamate herbicide, has become a common method of control. However, its long‐term efficacy has not been assessed in terms of either controlling the bracken or in producing suitable restored habitats. This study undertook such an assessment by comparing the results of a 2002 survey of the sites subject to bracken control by aerial spraying of asulam with previous (1990/1991 and 1994) surveys of the same sites. This showed that a single application of asulam was effective in eradicating bracken (<1% cover remaining) on a third of sites. However, on 10% of the sites, the bracken had regenerated completely (cover >80%) and on the remainder it was still present in patches (>20% of quadrats), often at high density. More than half the sprayed sites had seen good recovery of moorland vegetation, the target of the restoration, because they were now classified as having upland heathland vegetation within the National Vegetation Classification. Considerable amounts of bracken control are grant aided as part of agri‐environment schemes. These schemes should be adapted to encourage good practice, namely, intensive follow‐up treatment by spraying any emerging fronds, and to encourage treatment of previously sprayed areas rather than spraying of new areas in order to protect previous investment of grant aid.     

The impact of mechanical harvesting regimes on the aquatic and shore vegetation in water courses of agricultural areas of the Netherlands

It was demonstrated that a mechanical harvesting regime can influence the species composition of ditch vegetation at the community level. This effect, however, was very small compared with those of other factors such as the between-site and the within-site spatial variation, and several soil and water quality parameters. Cutting in November had the largest effect, in that it caused the greatest extremes in species cover. The vegetation was composed of 136 plant species. The semi-aquatic and aquatic species were less numerous than the terrestrial ones (52 versus 84). The total number per vegetation type and site ranged from 5 to 49; that of persistent plant species, from 4 to 22. Only 16% of the species was significantly influenced by the mechanical harvesting regime. The significant effects of the mechanical harvesting regime on plant species were related to plant-inherent factors. Mechanical harvesting repeated within a year, on one hand, opened up the vegetation, 1) freeing sites for colonization of new species, 2) improving the light climate for seedlings which had already colonized; on the other hand, it exhausted (carbohydrate) reserves of solitary species. Mechanical harvesting once a year in November had a contrasting effect in that it caused suffocation of the shore vegetation in spring by the not yet decomposed plant material harvested in autumn. The highest species richness was attained for the aquatic vegetation: on sand by cutting three times per year (in May, July and September), and on peat by cutting once a year (in November); and for the shore vegetation: on sand once a year (in May) and two times per year (in May and July), and on peat once a year (in May or November).