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.     

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.     

The ecological implications of bracken control to plant/animal systems

The importance of considering bracken as part of a dynamic soil/plant/animal system is emphasized and the extent, distribution and successional relationships of the fern in the uplands are discussed. Pteridietum is classified into six types according to its pattern of local distribution. Bracken communities as animal habitat are considered in relation to distribution types, frond density and canopy cover. As a direct source of food for the vertebrates bracken is considered to be of little or no value. As an indirect source of food bracken communities may be important in providing habitats for invertebrates, though in terms of numbers of species the insect fauna of vigorous Pteridietum is not great compared with that of other communities that may replace it. Pteridietum provides shelter and cover for a number of small mammals but in these respects it is of very limited value to the larger grazing animals. It is estimated that in Scotland, under the economic conditions of recent years, bracken herbicide is unlikely to be used on more than about 40,470 ha (100,000 acres) in the combined land uses of agriculture, forestry and sport; i.e. less than 25% of the Scottish bracken coverage is at risk. At this level of control, provided some precautions are taken, it is considered that the wild vertebrates are unlikely to be seriously affected; indeed in certain situations some benefits may be derived.     

Decomposition of bracken litter

Investigations on the decomposition of bracken petioles, over a five-year period on six adjacent soil types, including moder-type humus, mull and peat, are reviewed. Changes in gross physical features, chemical composition, pH and dry weight are outlined. The succession of colonizing fungi is described and related to fungal activities.
Until the petioles were buried in the litter layer, decomposition occurred at different rates on the various sites, the rate on moder > mull > peat, but the sequence of events was similar. Large proportions of readily leached components were removed in the first few months, but 95% loss of dry matter was estimated to occur only after 11–23 years. The majority of fungi were species cosmopolitan on litter, the population becoming less specialized as decay advanced. The succession resembled those on some other woody tissues, lignin and cellulose decomposers predominating before sugar fungi. From field observations and laboratory experiments, the Basidiomycete Mycena galopus (Pers. ex Fr.) Kummer appeared to be the most active of the fungal decomposers.
Some ecological and economic implications of the decomposition of bracken litter are briefly discussed, including its effect on soil type, and advantages of bracken compared with straw as bedding for farm animals.     

The hemicelluloses of bracken Part II. A galactoglucomannan

A galactoglucomannan has been isolated from the stem tissues of a fern, bracken (Pteridium aquilinum). It had [] −35° and, on hydrolysis, yielded mannose, glucose, and galactose in the molar ratios of 60:15:1. Mild hydrolysis with acid released galactose only. The d.p., determined by periodate-oxidation and Smith-degradation studies, was 43–45 for a doubly branched molecule. The methylated galactoglucomannan had a d.p. of 25–30 by ebulliometry. Methylation analysis, in combination with other evidence, indicated that the hemicellulose had β-(1→4)-linked -glucopyranose and -mannopyranose residues in the ratio of 1 to 4, with residues of the latter contiguous. -Mannopyranosidic and -galactopyranosidic residues were present as non-reducing end-groups in the ratio of 7 to 1, but there was no evidence of non-reducing -glucopyranosidic residues. The branching was to 6-positions on the main glucomannan chain and was probably -(1→6). The galactoglucomannan is similar to hemicelluloses isolated from softwoods, but the former has a lower d.p.     

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.     

The effects of fungi on bracken

Laboratory experiments on fungal pathogens of the gametophyte generation are described and the possible causative agents associated with disease of bracken colonies in the field discussed.     

Chemotaxonomy and phytochemical ecology of bracken

Variations in flavonoids extracted from fronds suggest that Pteridium aquilinum (L.) Kuhn is a monospecific genus. A wide range of secondary plant substances also occur in bracken including sesquiterpenoids, echydones, cyanogenic glycosides, tannins and phenolic acids.     

The history and ethnobotany of bracken

The fossil record shows that Pteridium has been present in all the temperate stages of the Quaternary, but that for most of this period it formed a component of the herb layer of deciduous woodlands. Bracken did not begin to reach its present abundance and importance until widespread forest clearance began with the arrival of Neolithic man about 5000 years ago. Some of the factors that have played a role in the history of bracken are illustrated by a discussion of the spread of bracken in Scotland that occurred contemporaneously with the change from cattle- to sheep-farming in the 18th century. It is shown that man has played a dominant role in influencing the spread of bracken in Scotland, but that bracken was considered of great positive economic value. The ethnobotany of bracken is discussed, and it is suggested that it was once an important source of potash for the glass, soap and bleaching industries. The use of the plant as fuel, thatch, litter, compost and food, and for medicinal purposes is considered, and mention is made of various minor uses of the plant. Unfortunately it is difficult to estimate the amount of bracken consumed by these various employments, but it is suggested that the increased abundance of the plant may have been exaggerated because what was once a useful resource has now become a pest.     

Nitrification in Dutch heathland soils

A survey was conducted over a range of 17 Dutch heathland locations, subdivided into 41 sites dominated by either dwarf-shrubs (Calluna vulgaris or Erica tetralix) or grass species (Deschampsia flexuosa or Molinia caerulea). Among the habitats of the dominant plant species relatively little differences in general soil properties were observed. The P status of Deschampsia sites was relatively high as well as the NO₃ ^–-N concentrations in the 0–10 cm layer (FH included) at the grass-dominated sites. At sites with a dead or degenerating dwarf-shrub vegetation, NH₄ ⁺-N concentrations reached very high levels.Net production of nitrate was observed during incubation of intact 0–10 cm soil cores (FH-layer included) in the laboratory for all sites, even though in some instances, particularly at Calluna and Erica sites, no nitrate was initially measured. Generally, a higher nitrification rate was found for the grass-dominated sites, and for Deschampsia in particular. The net production of nitrate was highly significantly correlated with net N mineralization, being a reasonable predictor of nitrification in a simple regression model (R²=0.47; P<0.001). Net nitrification was also significantly correlated with the NO₃ ^–-N initially present at the start of the growing season (R=0.65; P<0.001) and with the labile organic P content of the soil (R=0.65; P<0.001). By including initial NO₃ ^–-N and labile organic P, together with net N mineralization and pH, in a multiple regression model, net nitrate production could be predicted with a much higher precision (R²=0.75; P<0.001). Although apparent nitrification was not significantly correlated with pH, the latter contributed significantly to the multiple regression equation for the prediction of the former.The influence of the labile organic P pool may act via its positive correlation with microbial biomass, thus more or less reflecting the potential mineralization/nitrifying capacity of a particular site.     

The invertebrates of heather and heathland

WEBB, N. R., 1989. The invertebrates of heather and heathland. The southern heathlands of Great Britain have long been renowned amongst entomologists. The invertebrates fall into two groups: species which feed on heathland plants–about 40 in the case of Calluna–and a second group which depends on the particular physical conditions provided by heathland, such as hot dry sandy areas or the micro-topography of Sphagnum bogs. The ecology of representative examples will be described, and the effects of fragmentation and isolation of the heathlands on the distribution and abundance of invertebrates discussed.     

Regeneration of heathland plants following burning

Calluna-dominated heathlands are widespread in N. Britain, especially in the eastern and central part of Scotland. The plant communities of these heathland ecosystems are influenced by regular burning as a land management practice. In a well-controlled fire, regeneration takes place both vegetatively (from surviving stem bases or underground rhizomes) and from seed. But there is always a post-fire successon taking place even in this brief period of vegetation reoovery. In order to understand the biological mechanisms operating in the post-fire succession, the present work was carried out to examine regeneration behaviour of different species. Three ‘regenerative strategies’ were identified: I. Woody low-shrubs with slow bud development to begin with, followed by vigorous and profuse branching, becoming dominant after two years; II. Small herbaceous hemicryptophytes; sprouting not as prolific as in plants of group I, but more vigorous than in plants of group III; III. Geophytes with rapid development, important in the first 2–3 years.     

Diurnal variations in lipids of bracken fronds

The galactolipids, sulpholipid and phosphatidyl glycerol in the fronds of bracken show diurnal variations of 50–100%, with a maximum at about 1600 hr.     

Problems in heathland and grassland dynamics

A review of factors governing vegetation change in heathlands and grasslands is presented, with emphasis on soil factors. Climate, microclimate, fire, fauna, grazing by vertebrates, grazing by invertebrates, disease, dispersal, establishment, competition, allelopathy, stabilization, soil formation and podzolization are discussed with emphasis on inconsistencies and lacks in our present knowledge. Examples are mainly from NW European Calluna heath and related woodlands.Nomenclature follows Clapham, Tutin & Warburg (1962) for vascular plants; species not included in this work have authorities cited in the text.     

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.     

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.