Conservation of the Dartford warbler on english lowland heaths: A review

The Dartford warbler Sylvia undata is a rare bird of lowland heaths in southern England. Based on the results from a recent study, the present essay aims to describe the practical steps conservationists might take to promote its future in the British avifauna. There is a need for greater safeguarding of sites by acquisition or protection from undesirable developments, while on surviving heaths encroachment of birch, bracken and pines will have to be resisted or even reversed. The maintenance of an ideal age range of vegetation is considered and selective propagation of gorse is suggested as a way to improve some sites for Dartford warblers. Consideration is given to translocating birds and to providing food artificially in winter, though both these possibilities are not considered to be ideal. Recommendations are made for the maintenance and development of sufficient knowledge on the Dartford warbler to aid its conservation.     

Losses of lowland heath through succession at four sites in Breckland,East Anglia,England

Although it has been generally accepted that plagio-climax communities, such as lowland heaths, may be lost through successional processes, there have been few attempts to document the scale of these losses. This paper, therefore, attempts to assess the loss of open heathland habitat at four sites in the Brecklands of East Anglia, and discusses successional change in relation to site management. Colonisation by scrub (birch and Scots pine) rather than bracken was found to be the major threat.     

Restoration of bracken-invaded Calluna vulgaris heathlands: Effects on vegetation dynamics and non-target species

The coastal heathlands of north-western Europe are endangered habitats of great conservation value. Invasion by bracken Pteridium aquilinum is a major challenge for conservation and restoration of these heathlands, including the under-studied northern regions. Today, the herbicide asulam is the most widely applied bracken control measure, but increasing focus on organic farming and nature conservation calls for alternative, preferably mechanical, approaches. In a 7-year replicated field experiment in western Norway, we investigated efficiencies of the four bracken control measures asulam, Gratil, annual cutting and biannual cutting, in restoring the characteristic heathland vegetation structure and species composition. We specifically tested herbicide effects on diversity and composition of non-target species. Effects of treatments over time were evaluated by repeated measures ANOVA, and for multivariate data, Principal Response Curves. Our results show that UK based control methods are largely applicable to bracken at its northern limit in the European heathland habitat. Asulam resulted in the fastest reduction in cover but cutting proved equally efficient long-term. Community compositions progressed towards desired heathland vegetation, but successional trajectories differed. Asulam had unintended effects on a number of heathland species not predictable by species characteristics or functional groups. Gratil failed to have any long-term effects. In summary, cutting is as efficient as herbicide application in reducing bracken, and more so in restoring northern heathland vegetation over time.     

Re-creation of heathland on improved pasture using top soil removal and sulphur amendments: Edaphic drivers and impacts on ericoid mycorrhizas

Understanding the factors that drive successful re-creation and restoration of lowland heaths is crucially important for achieving the long-term conservation of this threatened habitat type. In this study we investigated the changes in soil chemistry, plant community and interactions between Calluna vulgaris and symbiotic ericoid mycorrhizas (ERM) that occurred when improved pasture was subjected to one of three treatments (i) acidification with elemental sulphur (ii) acidification with ferrous sulphur (iii) removal of the topsoil. We found that the soil stripping treatment produced the greatest reduction in available phosphate but did not decrease soil pH. Conversely, acidification with elemental sulphur decreased pH but increased availability of phosphate and potentially toxic cations. The elemental sulphur treatment produced plant communities that most closely resembled those on surrounding heaths and acid grasslands. The most important driver was low pH and concomitant increased availability of potentially toxic cations. Plant community development was found to be little related to levels of available soil phosphate, particularly at low pH. The elemental sulphur treatment also produced the best germination and growth of C. vulgaris over 4-5 years. However, this treatment was found to inhibit the development of symbiotic relationships between C. vulgaris and ERM. This may affect the long-term persistence of re-created vegetation and its interactions with other components of heathland communities.     

Long-term heathland restoration on former grassland: The results of a 17-year experiment

European lowland heaths have declined by up to 80% due to land use change and lack of management. There has been considerable research into the restoration of this threatened habitat. However, long-term outcomes of restoration are poorly understood, especially in situations where past agricultural land use imposes severe constraints on community re-assembly. In 1989 a large-scale experiment was established to examine the effectiveness of five treatments to restore heathland on formerly productive grassland: (i) natural regeneration; (ii) herbicide application to facilitate regeneration; (iii) cultivation and application of seed-rich heathland vegetation; (iv) soil removal and incorporation of heathland topsoil; and (v) heathland translocation. After 17 years the pH of the unamended agricultural soil remained significantly higher than that of the adjacent heathland. All treatments showed different trajectories of vegetation change in the long-term. Natural colonisation by heathland species was slow due to seed limitation, resulting in formation of an acid grassland community. Heathland community assembly was not facilitated by destruction of the initial grassland with herbicide. Incorporation of topsoil had an intermediate effect on pH reduction. This may explain the subsequent failure of the plant community to assemble in the anticipated proportions, and the dominance of leguminous scrub species (Ulex spp.). Turf translocation was effective in reducing pH to the required range and restoring the heathland community in the long-term. However, this technique should only be considered as a means of ‘rescue’ when habitat destruction is otherwise unavoidable. The only practical and sustainable means of increasing heathland extent on former farmland is the application of seed-bearing vegetation cut as part of routine management. However, this technique needs refining in order to establish the full range of characteristic heathland species.     

Long term effects of enhanced nitrogen deposition on a lowland dry heath in southern Britain

Experimental additions of ammonium sulphate to a nitrogen-poor dry heathland have been carried out since 1989. There are four nitrogen treatments: a control (receiving artificial rain only), a low treatment which receives an additional 7.7 kg N ha^?1 yr^?1, a high treatment receiving 15.4 kg N ha^?1 yr^?1 and an alternating treatment which receives either the control or the high nitrogen additions, in alternate years. The estimated background deposition at the study site is 13–18 kg N ha^?1 yr^?1, a value similar to the critical load that has been suggested for the conversion of lowland heath to grassland. Over the past 5 years there have been significant stimulations in shoot growth, flowering, canopy density and litter production. Flowering, in particular, strongly reflects nitrogen additions in the alternating treatment. Current models of the response of dry Calluna heathlaud to enhanced nitrogen deposition suggest that higher tissue nitrogen levels will occur and will be accompanied by heightened sensitivity to secondary stresses. This may in turn lead to canopy breakdown and replacement by grassland. The application of nitrogen at deposition rates only slightly in excess of the critical load over five years has produced small, non-significant increases in shoot nitrogen content. However, there is clear evidence of a large positive effect on shoot growth, flowering, litter production and canopy density of Calluna. The observation of these responses at the application rates used in this study supports the current proposals for critical loads of nitrogen for lowland heaths.     

Impact of litter quality on mineralization processes in managed and abandoned pasture soils in Southern Ecuador

Tropical regions are currently undergoing remarkable rates of land use change accompanied by altered litter inputs to soil. In vast areas of Southern Ecuador forests are clear cut and converted for use as cattle pastures. Frequently these pasture sites are invaded by bracken fern, when bracken becomes dominant pasture productivity decreases and the sites are abandoned. In the present study implications of invasive bracken on soil biogeochemical properties were investigated. Soil samples (0-5 cm) were taken from an active pasture with Setaria sphacelata as predominant grass and from an abandoned pasture overgrown by bracken. Grass (C₄ plant) and bracken (C₃ plant) litter, differing in C:N ratio (33 and 77, respectively) and lignin content (Klason-lignin: 18% and 45%, respectively), were incubated in soils of their corresponding sites and vice versa for 28 days at 22 °C. Unamended microcosms containing only the respective soil or litter were taken as controls. During incubation the amount of CO₂ and its δ¹³C-signature were determined at different time intervals. Additionally, the soil microbial community structure (PLFA-analysis) as well as the concentrations of KCl-extractable C and N were monitored. The comparison between the control soils of active and abandoned pasture sites showed that the massive displacement of Setaria-grass by bracken after pasture abandonment was characterized by decreased pH values accompanied by decreased amounts of readily available organic carbon and nitrogen, a lower microbial biomass and decreased activity as well as a higher relative abundance of actinomycetes. The δ¹³C-signature of CO₂ indicated a preferential mineralization of grass-derived organic carbon in pasture control soils. In soils amended with grass litter the mineralization of soil organic matter was retarded (negative priming effect) and also a preferential utilization of easily available organic substances derived from the grass litter was evident. Compared to the other treatments, the pasture soil amended with grass litter showed an opposite shift in the microbial community structure towards a lower relative abundance of fungi. After addition of bracken litter to the abandoned pasture soil a positive priming effect seemed to be supported by an N limitation at the end of incubation. This was accompanied by an increase in the ratio of Gram-positive to Gram-negative bacterial PLFA marker. The differences in litter quality between grass and bracken are important triggers of changes in soil biogeochemical and soil microbial properties after land use conversion.     

Short rotation forestry – Earthworm interactions: A field based mesocosm experiment

Short rotation forestry (SRF) which consists of planting rapidly growing native and non-native tree species has been introduced to the UK to increase woody biomass production. A largely unknown aspect of SRF species is their interaction with soil fauna, of which the earthworm community is a major component. Earthworms have a pronounced impact on litter decomposition, nutrient cycling and tree growth. Conversely, tree litter and root chemistry can impact on the associated earthworm community development. The aim of this study was to determine direct interactions between SRF species and earthworms. A field-based mesocosm experiment was conducted using Betula pendula (birch) and Eucalyptus nitens (eucalyptus) with two earthworm species Lumbricus terrestris and Allolobophora chlorotica. The one year experiment revealed that native birch and non-native eucalyptus had a similar influence on L. terrestris population development. However, birch had a positive impact on A. chlorotica population establishment compared with eucalyptus. In the presence of earthworms, total tree biomass and leaf nitrogen concentration of eucalyptus were increased respectively by 25% and 27% compared with an earthworm-free control. In the presence of earthworms, surface litter incorporation was greater for both tree species (almost 5 times for birch and 3 times for eucalyptus) compared with controls. This work showed direct SRF-earthworm interactions which differed for tree species.     

Environmental change as a cause of natterjack toad (Bufo calamita) declines in Britain

An investigation was made into factors which have influenced the decline of natterjack toads Bufo calamita in Britain over the last 40 years. Natterjacks are limited in Britain to two types of habitats, coastal dunes and inland heaths. By comparing the relative declines of the species on these two habitats with other changes which have occurred on them, it became clear: (1) That Bufo calamita has declined much more extensively on heaths than on dunes: (2) that this selective decline cannot be attributed to climatic changes, public pressure or direct development; (3) that large-scale changes in the flora of heathland areas have occurred concurrently with natterjack declines. These changes, which in general have not been reflected on dunes, have occurred subsequent to altered patterns of land use on the heathlands. Cessation of grazing and increased forestry activity have permitted widespread encroachment by tall vegetation, which creates shade, both changes being directly undesirable for natterjacks, and more importantly, enabling the common toad Bufo bufo to enter as a successful competitor. This latter factor is thought to have been one of the most important causes of natterjack declines on heathland.     

Soil and vegetation changes in experimental birch plots on a Calluna podzol

A long-term experiment set up by G. W. Dimbleby to test the effect of birch as a soil improver on a heather moor podzol is described. After 30 yr, an acid woodland type of vegetation developed under the birch trees with Deschampsia flexuosa and Vaccinium myrtillus locally dominant. The mean annual dry matter production of stems and branches was approximately 2.9 t ha^?1. The raw humus layer removed as an experimental treatment had re-formed. Its pH, about 3.4, had remained unchanged for 15 yr. The mean extractable Ca content of the upper 3 cm horizon of the birch plots and of the control plots under heather had increased and on some of the birch plots the concentrations of exchangeable Ca were somewhat higher than those under heather. The N content of the Ol + Of, Oh and Ea horizons was higher under the birch than under heather but there was no evidence of movement of Fe out of the ironpan under the influence of birch. The profile under birch remained a fully differentiated podzol.     

Growth and Leaf Gas Exchange in Three Birch Species Exposed to Elevated Ozone and CO2 in Summer

We examined the effects of ozone and elevated CO₂ concentration in summer on the growth and photosynthetic traits of three representative birch species in Japan (mountain birch, Monarch birch, and white birch). Seedlings of the three birch species were grown in 16 open-top chambers and were exposed to two levels of ozone (6 and 60?nmol?mol^?1 for 7?h per day) in combination with two levels of CO₂ (370?C380 and 600???mol?mol^?1 for daytime) from July to October. No adverse effects of ozone were found in the Monarch birch or the white birch, but elevated ozone in summer reduced branch biomass and net photosynthesis, and accelerated leaf abscission, in the mountain birch. Elevated CO₂ promoted root development and thereby reduced the ratio of shoot dry mass (stem + branch) to root dry mass (S/R ratio) in the mountain birch and white birch. In contrast, there was no difference in dry mass between ambient and elevated CO₂ for the Monarch birch, due to downregulation of photosynthesis. Studies of the combined effect of CO₂ and ozone revealed that elevated CO₂ did not ameliorate the effect of ozone on mountain birch in late summer. In considering the ameliorating effect of CO₂ on ozone damage, it is necessary to take account of the species and the season.     

An assessment of the bracken problem in relation to hill farming in Scotland

Abstract. The reasons for bracken encroachment in Scottish hill land are examined. The cost effectiveness of bracken control methods are discussed and it is shown how the real cost of aerial spraying, the most practical method, has increased in the last decade. The waiving of the follow-up requirements under the DAFS grant schemes may alter this cost situation in the future. The need for more accurate estimates of the extent of the bracken problem is highlighted by the possible link between bracken and cancer in humans. Recent surveys utilizing remote sensing techniques are described and the cost of a full bracken eradication programme is estimated to be c. £80 million which is contrasted with the current level of grant assistance of c. £135 000 a⁻¹. The use of bracken as biofuel is considered as an alternative to eradication.     

How do coniferous needle tannins influence C and N transformations in birch humus layer?

The aim of this study was to explore the response of C and N transformations in the humus layer under silver birch (Betula pendula Roth) to compounds, especially condensed tannins, of different molecular weight extracted and fractioned from Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) needles. Lighter fractions containing tannin monomers and dimers as well as many other compounds, and heavier fractions consisting predominantly of polymerized condensed tannins, were added to samples taken from the humus layer of birch stand. The effects of the spruce and pine fractions were mostly similar, but some differences in magnitude were observed; our results indicated that lighter fractions of pine were easier for microbes to degrade and use than lighter fractions of spruce. Lighter fractions of both tree species increased soil respiration and decreased net N mineralization, while heavier fractions inhibited respiration and increased net N mineralization. Microbial biomass C was not clearly affected by any of the treatments, but with some of the pine fractions the amount of N in the microbial biomass was increased. Comparison of the effects of fractions in birch and in spruce and pine soils, which were studied earlier, showed no major differences between the effects of the fractions in birch and in their own soils, but gave some indication of adaptation.     

Organic matter characteristics and C and N transformations in the humus layer under two tree species, Betula pendula and Picea abies

The aim of this study was to compare the effects of silver birch (Betula pendula Roth) and Norway spruce (Picea abies (L.) Karst.) on soil C and N transformations and on the characteristics of organic matter. Soil samples were taken from the humus layer of a replicated 35-year-old birch-spruce field experiment growing on Vaccinium myrtillus site type in middle-eastern Finland. The soil was a podzol and humus type mor. Soil pH was higher under birch (4.7) than under spruce (4.1). The C-to-N ratio was lower under birch (17) than under spruce (23). Per unit organic matter, microbial biomass C and N, net N mineralization and net nitrification were all higher in birch soil than in spruce soil. The rate of C mineralization (CO₂ production) was, however, the same regardless of tree species. Water-extracts were analyzed for the concentrations of dissolved organic C (DOC) and N (DON) and characterized according to molecular size distribution by ultrafiltration and according to chemical composition using a resin fractionation technique. The concentration of DON, in particular, was higher in birch soil than in spruce soil. The distribution of DOC and DON into different fractions based on molecular size or chemical composition was rather similar in both soils. The concentration of total phenolics, expressed as tannic acid equivalents, was higher in the humus layer under birch than in the humus layer under spruce, because the birch humus layer contained significantly more low-molecular weight (about <0.5 kD) phenolics than the spruce humus layer did. The concentration of proanthocyanidins (condensed tannins) was higher in spruce soil than in birch soil. The concentrations of the five most abundant phenolic acids showed that ferulic and p-coumaric acids were more abundant in spruce soil. Birch soil tended to contain slightly more nonvolatile sesquiterpenes than the spruce soil. The concentration of diterpenes was similar in both soils; but birch soil contained significantly more triterpenes, mainly sterols, than spruce soil did.     

Spatial Variability of Carbon Dioxide Emission by Soils in the Main Types of Forest Ecosystems at the Zvenigorod Biological Station of Moscow State University

Carbon dioxide (CO₂) emission from the soil surface in forest biogeocenoses of the Zvenigorod Biological Station of Moscow State University in summer varies on average from 120 to 350 mg C–CO₂/(m² h) and depends on the hydrothermal conditions (soil moisture and temperature) and the type of phytocenosis. The intensity of CO₂ emission in the biogeocenosis does not depend on its parcel structure and varies with respect to plant microgroups: it is maximum in oxalis pine–spruce and maple–lime forests and bracken spruce–birch forests and minimum in areas of forest fall without vegetation. The upper (from 0 to 20 cm thick) soil layer provides up to 50% of the total soil CO₂ emission. The role of microbial respiration in the total CO₂ emission from soils is determined by weather conditions and varies from 9–33% in a dry summer to 55–75% in a summer with favorable temperature and moisture.     

Soil microbial communities and activities in sand dunes of subtropical coastal forests

To understand the soil microbial activities and community structures in different forests in a sand-dune ecosystem, we conducted a study of 2 topographic conditions, upland and lowland, under a Casuarina forest. As well, in the lowland site, we compared forest soil microbial properties under 3 coastal forests (Casuarina, Hibiscus and mixed stand). The soil microbial biomass did not significantly differ between the upland and lowland Casuarina forest sites. At the lowland site, the soil microbial biomass was higher in the Hibiscus than Casuarina forest soil. Cellulase, xylanase, phosphatase and urease activities did not show a consistent trend by topography or vegetation. Analysis of phospholipid fatty acids (PLFAs) of bacteria and actinomycetes revealed a significant difference in microbial community structure by both topography and vegetation. PLFA content was higher at upland than lowland sites in the Casuarina forest. At the lowland site, the level of PLFAs was higher in Hibiscus than Casuarina forest soil. In addition, we examined the ratios 16:1ω7t/16:1ω7c and, cy17:0/16:1ω7c as indicators of physiological stress; the soil in the Casuarina forest had the highest values, which suggests that the microbial community in the Casuarina forest soil is under physiological stress or starvation conditions. Comparison of soil microbial properties suggest that planting Hibiscus may help to enrich soil fertility and increase microbial activities in coastal sand-dune Casuarina forest.     

Mobility and breakdown of asulam in the soil and the possible impact on the environment

Asulam reaching the soil either directly as spray drift or as rain washings from bracken fronds would, due to its rapid mobility in soil, be expected to leach down the soil profile into the drainage water and thereby contaminate streams and water catchments. The mobility of asulam in soil is influenced by pH, where undissociated asulam will leach less rapidly than the associated form (asulam pK_a 4·82). This is particularly relevant in the case of bracken as it can grow satisfactorily over a wide range of pH values (3·6 – 7·6). Adsorption coefficients expressed as Kd values [(μg asulam/g soil)/(μg asulam/ml solution)] were 2·54, 1·94 and 0·96 at pH readings of 4·0, 4·3 and 4·9 respectively. Another factor involved, the breakdown of asulam, was shown to be temperature dependent, 30·95, 7·65 and 2·50% remaining after 28 days at 5, 18 and 25°C respectively, thus underlining the influence of geographical regions and climatic conditions on asulam behaviour.     

Bracken effects on inorganic nitrogen leaching from an upland podzol

Leaching of inorganic N species to stream waters from upland areas of the UK is increasing, reflecting the increases in atmospheric deposition of nitrogen species due to increased levels of vehicular emissions. Bracken cover in UK uplands is also increasing overall ( Taylor, 1986 ), and the architectural nature of bracken has been shown to both increase deposition of atmospheric constituents to soils and change their chemical and physical nature. We have tested the nutrient status of upland podzols from the Lake District, Cumbria, UK, from under both moorland grass and bracken. The results show reduced levels of base cations and nitrate throughout the profile and decreases in the organic matter content of surface horizons as a consequence of bracken encroachment. We also report increased ammonium concentrations in lower soil horizons under bracken, possibly leading to increased leaching to stream waters. These results suggest either increased leaching of soil nutrients due to bracken encroachment or increased storage of nutrients within the bracken rhizome system. Results also suggest changes in either microbial activity or the microbial community of podzols, possibly due to the allelopathic nature of bracken. This may have caused changes in the soil nutrient dynamics, possibly accounting for some of the changes seen. We hypothesize that increased bracken cover within the British uplands has the potential to increase leaching of inorganic nitrogen to upland streams.     

Effects of increased salinity and an introduced predator on lowland amphibians in Southern China: Species identity matters

Approximately 30% of amphibian species are threatened due to a variety of factors affecting their habitats and physiology, yet contributions that interactions among factors make to population declines are not well-explored. Two factors, introduced mosquitofish (Gambusia affinis) and increased salinity, may coincide in lowland habitats used by breeding amphibians. Mosquitofish have been introduced worldwide and can be significant predators of amphibian larvae. Salinization of wetlands is an increasing problem globally due to (1) application of road deicing salts in temperate regions, (2) irrigation practices associated with intensive agriculture, and (3) saltwater intrusion due to sea-level rise. We investigated the effects of mosquitofish (G. affinis) and increasing salinity on five species of lowland wetland-breeding amphibians from southern China. We exposed anuran hatchlings to four salinity levels and two fish treatments and all combinations of the two factors in a series of experiments. Four of the species were susceptible to predation by mosquitofish, two were sensitive to increased salinity at concentrations >6% seawater, and one was tolerant of both increased salinity and mosquitofish. We found no interaction between the predator and increased salinity. Salinization and mosquitofish represent significant threats to lowland amphibians in this region and, coupled with the ongoing loss and degradation of lowland wetlands, portend a bleak future for lowland amphibian populations in the region.