On the water relations of four heath species

Four species of heath, occurring in the heathlands of Brittany, are compared regarding their water relations: Calluna vulgaris, Erica ciliaris, E. cinerea and E. tetralix. E. cinerea is unable to establish itself in wet heathland because of its intolerance of prolonged waterlogging. It is the Erica species best adapted to dry habitat conditions in Brittany. E. ciliaris canot establish itself in dry heathland and is less tolerant of waterlogging than E. tetralix. E. tetralix is the species best adapted to wet heathland, being tolerant of waterlogging, but can also establish itself in dry heathland. Of the three E. species it has the widest ecological range. Calluna is tolerant of both wet and dry conditions and has a wide ecological range. E. cinerea is typical of dry- and E. tetralix of wet heathland. Although both species did best in moist aerated soil in experimental cultures, neither is abundant in mesophilic heaths where E. ciliaris is dominant. One explanation may be competition for aerial space. E. cinerea and E. tetralix both have an upright growth, whereas E. ciliaris rapidly adopts a straggling bushy habit, with long rooting branches. E. ciliaris thus establishes large interpenetrating clumps. With increasing dryness E. ciliaris disappears and may be replaced by E. cinerea and, with increasing wetness and especially waterlogging, E. tetralix will take over.     

The occurrence of former heathland vegetation in the coastal areas of the south-east Baltic sea,in particular Lithuania: a review

Previously published pollen diagrams from the coastal area of the south-east Baltic Sea including parts of the Polish and Lithuanian peninsulas, Mierzeja Wilana (Frische Nehrung) and Kuri Nerija (Courland Spit) respectively, document the former presence of heathland vegetation and an associated burning practice. This type of human influence can be dated back to ca. 1200 B.P. at the latest, and was maintained until the 18th century. It is suggested that this former Calluna-dominated vegetation belonged to the known west to north-west European heathland vegetation. Several dwarf shrub taxa, such as Genista spp. and Erica tetralix, normally connected to heathland vegetation, may have had a former wider distribution.     

Seed-shedding in a Scottish heath community

Abstract. For a 28-week period in late 1987 and early 1988, a study of seed-shedding by several heath species was carried out at the Muir of Dinnet in northeastern Scotland. The dominant species in the heath is Calluna vulgaris. Seed-shedding in Calluna began in early September 1987 and was completed in April 1988, with the period of maximum shedding falling between early November and late December 1987. The total numbers of seeds/m² deposited in stands of Calluna in its four growth-phases were: pioneer, 18 910; building, 169010; mature, 198580; degenerate, 33900. Substantial loss of potential seeds results from the shedding of immature flowers. A control area close to, but outside the area of Calluna dominance had a deposition rate of 770 Calluna seeds/m², indicating sufficient seed to colonise nearby available habitats. Seed rain was also recorded for several other heathland species: Erica cinerea, E. tetralix, Carex spp. and Betula pendula. Seeds of Erica cinerea were deposited in all the Calluna stands, 32670/m² in the pioneer stand, 17600/m² in building, 3720/m² in mature, 210/m² in degenerate (120/m² in the control area). Numbers were greatest at the start of the sampling period, declining thereafter. This applied also to Betula seeds. Erica tetralix occurred in the degenerate Calluna stand and yielded 640 seeds/m² (400/m² in the control area). Seeds of Carex spp. were obtained in the control: 4110/m². The method of sampling has a significant effect on the figures obtained. A method using tube collectors, emptied frequently, is recommended.     

Nutrient use efficiency in evergreen and deciduous species from heathlands

Summary The nutrient (N, P) use efficiency (NUE: g g^–1 nutrient), measured for the entire plant, of field populations of the evergreen shrubs Erica tetralix (in a wet heathland) and Calluna vulgaris (in a dry heathland) and the deciduous grass Molinia caerulea (both in a wet and a dry heathland) was compared. Erica and Calluna are crowded out by Molinia when nutrient availability increases. NUE was measured as the product of the mean residence time of a unit of nutrient in the population (MRT: yr) and nutrient productivity (A: annual productivity per unit of nutrient in the population, g g^–1 nutrient yr^–1. It was hypothesized that 1) in low-nutrient habitats selection is on features leading to a high MRT, whereas in high-nutrient habitats selection is on features leading to a high A; and that 2) due to evolutionary trade-offs plants cannot combine genotypically determined features which maximize both components of NUE.Both total productivity and litter production of the Molinia populations exceeded that of both evergreens about three-fold. Nitrogen and phosphorus resorption from senescing shoots was much lower in the evergreens compared with Molinia. In a split-root experiment no nutrient resorption from senescing roots was observed. Nutrient concentrations in the litter were equal for all species, except for litter P-concentration of Molinia at the wet site. Both Erica and Calluna had a long mean residence time of both nitrogen and phosphorus and a low nitrogen and phosphorus productivity. The Molinia populations showed a shorter mean residence time of N and P and a higher N- and P-productivity. These patterns resulted in an equal nitrogen use efficiency and an almost equal phosphorus use efficiency for the species under study. However, when only aboveground NUE was considered the Molinia populations had a much higher NUE than the evergreens.The results are consistent with the hypotheses. Thus, the low potential growth rate of species from low-nutrient habitats is probably the consequence of their nutrient conserving strategy rather than a feature on which direct selection takes place in these habitats.     

Hierarchical and spatially aggregated plant cover data

Most plant species are spatially aggregated and here the importance of taking the spatial variation into account when analyzing plant cover data is demonstrated in a general stochastic model where both the within-site and the among-site spatial variation of species cover data are parameterized. Using a generalised binomial distribution (or Pólya–Eggenberger distribution), where the among-site variation in mean cover is modeled by a zero-inflated beta distribution, it is possible to adequately analyze hierarchical plant cover data and link the estimates to the underlying ecological processes. The model is demonstrated in a case-study of pin-point cover data of Erica tetralix from 1148 wet heathland plots at 84 Danish sites, and it is shown that both parameter estimates and the conclusions of hypotheses testing critically depend on the correct modeling of the observed spatial variation. Finally, statistical power simulations of plant cover measurements are presented, which will be useful for planning ecological experiments and monitoring programs.     

Leaching of nitrate and ammonium in heathland and forest ecosystems in Northwest Germany under the influence of enhanced nitrogen deposition

To study the impact of high atmospheric nitrogen deposition on the leaching of NO₃^– and NH₄⁺ beneath forest and heathland vegetation, investigations were carried out in adjacent forest and heathland ecosystems in Northwest Germany. The study area is subjected to high deposition of nitrogen ranging from 15.9 kg ha^–1 yr^–1 in bulk precipitation to 65.3 kg ha^–1 yr^–1 beneath a stand of Pinus sylvestris L. with NH₄–N accounting for 70–80% of the nitrogen deposited. Considerable leaching of nitrogen compounds from the upper horizons of the soil, mostly as nitrate, occurred at most of the forest sites and below a mixed stand of Calluna vulgaris (L.) Hull. and Erica tetralix, but was low in a Betula pubescens Ehrh. swamp forest as well as beneath Erica tetralix L. wet heath and heath dominated by Molinia caerulea(L.) Moench. Ground water concentrations of both NO₃–N and NH₄–N did not exceed 1 mg L^–1 at most of the sites investigated.     

The relation between above- and belowground biomass allocation patterns and competitive ability

Summary In a 2-year experiment, the evergreen shrubsErica tetralix andCalluna vulgaris (dominant on nutrient-poor heathland soils) and the perennial deciduous grassMolinia caerulea (dominant on nutrient-rich heathland soils) were grown in replacement series in a factorial combination of four competition types (no competition, only aboveground competition, only belowground competition, full competition) and two levels of nutrient supply (no nutrients and 10 g N+2 g P+10 g K m⁻² yr⁻¹). Both in the unfertilized and in the fertilized treatmentsMolinia allocated about twice as much biomass to its root system than didErica andCalluna. In all three species the relative amount of biomass allocated to the roots was lower at high than at low nutrient supply. The relative decrease was larger forMolinia than forErica andCalluna. In the fertilized monocultures biomass of all three species exceeded that in the unfertilized series.Molinia showed the greatest biomass increase. In the unfertilized series no effects of interspecific competition on the biomass of each species were observed in either of the competition treatments. In the fertilized mixtures where only belowground competition was possibleMolinia increased its biomass at the expense of bothErica andCalluna. When only aboveground competition was possible no effects of interspecific competition on the biomass of the competing species were observed. However, in contrast with the evergreens,Molinia responded by positioning its leaf layers relatively higher in the canopy. The effects of full competition were similar to those of only belowground competition, so in the fertilized series belowground competition determined the outcome of competition. The high competitive ability ofMolinia at high nutrient supply can be attributed to the combination of (1) a high potential productivity, (2) a high percentage biomass allocation to the roots, (3) an extensive root system exploiting a large soil volume, and (4) plasticity in the spatial arrangement of leaf layers over its tall canopy. In the species under study the allocation patterns entailed no apparent trade-off between the abilities to compete for above- and belowground resources. This study suggests that this trade-off can be overcome by: (1) plasticity in the spatial arrangement of leaf layers and roots, and (2) compensatory phenotypic and species-specific differences in specific leaf area and specific root length.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

The effect of increased nutrient availability on leaf turnover and aboveground productivity of two evergreen ericaceous shrubs

Summary Leaf turnover and aboveground productivity in relation to nutrient availability were studied in the evergreen shrubs Erica tetralix and Calluna vulgaris. In monospecific stands of these species four levels of nutrient (NPK) availability were created during three growing seasons. Percentage survival and life expectancy of Erica leaves decreased with increasing nutrient availability. For Calluna there was no effect. Winter mortality of Erica leaves was smaller than growing season mortality. These was no difference for Calluna. The timing of leaf mortality of both species was not affected by nutrient treatment. At the end of the experimental period current year leaf biomass, total biomass and current year second year and third year biomass of both species showed a significant increase with increasing nutrient availability. The relative increase was greater for Calluna, except for second and third year biomass. Stem production and stem mortality of both species increased with increasing nutrient availability. The increased stem mortality resulted also for Calluna in an increased leaf turnover (per unit ground area) with increasing nutrient availability. Nutrient cycling in ecosystems dominated by these species will increase with increasing nutrient availability, because of increased leaf and stem turnover and productivity. This phenotypic effect is similar to the effect of the shift in dominance between different species which occurs along natural gradients of nutrient availability.     

Effects of fertilization and cutting on the chemical composition of vegetation and soils of mountain heathlands in Spain

Abstract. In the mountains of northern Spain, patches dominated by Calluna vulgaris are scarce and they may disappear or change as a result of continued lack of management and possibly increasing nutrient availability through atmospheric deposition. The effects in the soil properties and in the composition of Calluna vulgaris and Erica tetralix shoots on heathlands dominated by Calluna and Erica subjected to fertilization and experimental cutting were studied in three mountain passes in northern Spain. A total of 90 1‐m² plots received different combinations of cutting and twice the estimated atmospheric deposition of nitrogen (5.6 g.m^?2.yr^?1) as ammonium nitrate. One of the dominant ericaceous species (Calluna and Erica) was selectively cut by hand at ground level and their nitrogen shoot content were compared in the presence or absence of the other. Treatments were carried out in April 1998. In each plot one soil sample was taken in the original situation and 12, 24 and 36 months after the treatments. Soil properties such as organic matter, total nitrogen, available phosphorus and pH were determined. In every plot five shoots of Calluna and Erica were also taken to analyse total nitrogen content in the original situation and 12, 24 and 36 mo after the treatments. Nitrogen addition does not necessarily lead to increased levels in the soil, and a clear pattern was not found in the three areas. A gradual decrease in available phosphorus content was detected in the three areas until two years after treatment, although values tend to recover in two of the areas in the third study year. An increase in organic matter content was observed in all areas. It is concluded that increased nutrients alone, at twice the rate of the estimated current atmospheric deposition for the area, which is relatively low, will not alter significantly the soil characteristics of the mountain heathland stands. A clear increase in plant N‐content is observed in the fertilized plots in comparison with the non‐fertilized ones and Calluna always has higher nitrogen content than Erica. This increase is most pronounced one year after the treatments started in one of the areas and after two years in the other two areas. In some cases the elimination of one species is seen to favour nitrogen increase in the other.     

The use of diachronic spatial approaches and predictive modelling to study the vegetation dynamics of a managed heathland

According to the EU Habitats Directive, heathlands are a semi-natural habitat type of community interest. This status aims at conserving these habitats, especially where and when they are threatened by various changes, including natural vegetation succession. We present results of a study of the dynamics of a typical dry heathland plot located in the Fontainebleau massif (France). An exhaustive observation of vegetation changes were made on this area of four hectares between 2000 and 2008, employing a spatial approach. We recorded the expansion of Molinia caerulea (L.) Moench at the expense of Ericaceae. The potential future vegetation of the site was modelled using Markov chains coupled to a GIS programme. This model predicted a gradual change in the floristic composition of heathland in favour of M. caerulea at the expense of Calluna vulgaris (L.) Hull and Erica tetralix L., and the expansion of Pinus sylvestris L. The study demonstrates how spatial methods can contribute to the design of reliable management methods of habitats such as the heathlands.     

Limiting climatic factors and habitats of Erica tetralix at the eastern edge of its distribution range

We determined the limiting climatic factors, as well as the preferred habitats, of Erica tetralix L. at the eastern limit of its distribution range in the eastern Baltic region. It was found that E. tetralix in this region is a typical bog woodland plant preferring the wettest Sphagnum‐rich sites. Northern Atlantic wet heath fragments, base‐rich fens and species‐rich Nardus grasslands were other habitats for the species. The species composition in E. tetralix habitats resembled that found in appropriate habitats within its main distribution range, although the habitats in the eastern Baltic region lack many Atlantic floristic elements characteristic of wet heath. A generalised linear model describing the climatic niche of E. tetralix in Latvia suggests that mild winters, which exhibit a combination of a shallow maximum depth of soil freezing and a large number of days when the air temperature exceeds 0°C, as well as abundant annual and winter precipitation, are the main factors allowing the presence of E. tetralix in Latvia. The climate parameters at the eastern limit of the species’ distribution range are consistent with those recorded as being suitable for E. tetralix elsewhere. Our results reveal a strong relationship between oceanity and the distribution of E. tetralix. The climatic niche model suggests more climatically suitable areas in the Coastal Lowland geobotanical region where the species could potentially be found, most likely in bog woodlands. The future prospects to restore open areas of northern Atlantic wet heath with E. tetralix in Latvia are poor due to the decline in traditional land‐use; these areas have turned into bog woodlands or been converted into agricultural land.     

Plant nutrient mobilization in temperate heathland responds to elevated CO2, temperature and drought

Temperate terrestrial ecosystems are currently exposed to increased atmospheric CO₂ and progressive climatic changes with increased temperature and periodical drought. We here present results from a field experiment, where the effects of these three main climate change related factors are investigated solely and in all combinations at a temperate heathland. Significant responses were found in the top soils below the two dominant species (Calluna vulgaris and Deschampsia flexuosa). During winter incubation, microbial immobilization of N and ammonification rate decreased in response to warming in Deschampsia soil, and microbial immobilization of N and P decreased in warmed Calluna soil. Warming tended to increase microbial N and P in Calluna but not in Deschampsia soil in fall, and more microbial C was accumulated under drought in Calluna soil. The effects of warming were often counteracted or erased when combined with CO₂ and drought. Below Deschampsia, the net nitrification rate decreased in response to drought and, while phosphorus availability and microbial P immobilization decreased, but nitrification increased in response to elevated CO₂. Furthermore, leaf litter decomposition of both species decreased in response to drought. These complex changes in availability and release of nutrients from soil organic matter turnover and mineralization in response to elevated CO₂ and climate change may influence the future plant carbon sequestration and species composition at temperate heathlands.     

Recovery after Experimental Cutting and Burning in Three Shrub Communities with Different Dominant Species

The aim of this study is to compare the recovery dynamics in three shrub communities subjected to experimental burning and cutting, and situated on an altitudinal gradient. Climatic features are different in each area, but all had the common characteristic of very homogeneous vegetation cover before the disturbances, with only one shrub species clearly dominant, a different taxon in each area, and with different regeneration strategies. The first area was a heathland dominated by Calluna vulgaris, situated at an altitude of 1600 m, with a continental climate (mean annual precipitation 1320 mm). The second area was a heathland dominated by Erica australis, located at an altitude of 1000 m (mean annual precipitation 840 mm). The third area was a Cistus ladanifer shrubland, located at 900 m altitude, with a Mediterranean climate similar to that of the previous area, but with lower mean annual precipitation (470 mm). Erica australis recovers by vegetative resprouting, but Cistus ladanifer is an obligate seeder, as is Calluna vulgaris in these areas. Each experimental disturbance was carried out over 100 m² in each area. Post-fire recovery is faster in Cistus ladanifer: 2 years after burning there was 40% cover vs. less than 20% in the other two species. However, recovery after cutting was similar for Cistus ladanifer and Erica australis. Calluna vulgaris recovers very slowly, with cover values below 20% even 10 years after both disturbances. Cover of dominant shrub species is negatively correlated with cover of herbaceous species. So different recovery of dominant species lead a different community dynamic in each area.     

Germination and establishment of seedlings in different phases of theCalluna life cycle in a Scottish heathland

Previous authors have suggested that cyclical succession takes place inCalluna-dominated heathland, and is determined by the different growth phases of the dominant. Characteristic species are said to invade at different stages of the life cycle ofCalluna, particularly in the initial and pioneer phases and in the degenerate phase. In the last phase, gaps are formed due to separation and dying of the oldest frame branches.This paper aims to analyze in which microhabitat and in which phase of theCalluna life cycle seedling emergence and establishment is possible.More seedlings appear in pioneerCalluna than in matureCalluna stands in experimentally manipulated micro-sites. In pioneerCalluna establishment is affected only by application of litter, in matureCalluna by increased illumination. The differences between treatments disappeared after three years, but not the difference in establishment between stands.Seedling emergence from species sown experimentally was different in the successive phases ofCalluna. Highest emergence was in the pioneer stand.In a seedling survey also most seedlings appeared and established in the pioneer phase. The number of recently germinated seeds was high in all phases. However, seedling survival was very low, except in the pioneer stands. This study does not of itself show evidence for cyclical succession inCalluna-dominated heath vegetation. Colonization of gaps by seedlings ofCalluna in mature or degenerate phases is possible, but must be an infrequent occurrence.Nomenclauture follows Clapham, Tutin & Warburg (1962).We wish to thank Dr G. R. Miller for stimulating discussions during the start of the experiments. E. de Hullu wishes to thank the Botany Department in Aberdeen for their hospitality during the research period.     

The Holocene vegetation history of northern West Jutland, Denmark

Holocene sediments of three closed Danish lake basins (Solsø, Skånsø, Kragsø) were used for the inference of post-glacial vegetational dynamics in former heathland areas in northern West Jutland, Denmark. The sites were selected to represent the major geomorphological units of West Jutland. The Holocene history of each lake basin was investigated by mapping of sediment distribution, analysis of loss-on-ignition, coarse inorganic matter, humus content, mineral magnetics, δ¹³C, pollen and selected other microfossils. These techniques were supplemented by plant macrofossil analysis at one site. Holocene terrestrial vegetational development was inferred at each site from analyses of pollen and microscopical charred particles. Chronologies were provided by numerous ¹⁴C dates. Stratigraphies of wet ground and terrestrial pollen and spore types were zooned by stratigraphically constrained cluster analysis. Based on the resultant site pollen asemblage zones (site PAZ), regional PAZ were proposed. Using modern analogues, Holocene floristic richness was estimated from pollen richness in the microfossil assemblages. The results support the hypothesis that disturbance is one of the most important mechanisms behind the maintenance of floristic richness. In particular, the response of estimated floristic richness to the intensity of vegetational fires followed the predictions of the Intermediate Disturbance Hypothesis. A period of elevated palynological richness and inferred vegetational disturbance was identified at all sites between 6000 and 5200 BC (calendar years). Using correspondence analysis (CA), the major gradient in the terrestrial pollen sequences was identified as a light-shade gradient, and CA first axis sample scores were used as a supplement to standard AP/NAP pollen ratios as an indicator of the shade-tolerance/light-demand of Holocene terrestrial plant communities. In spite of different vegetational developments since 4000 BC, the timing of major changes towards more light-demanding vegetation types were broadly synchronous at the three sites. Using chord distance as a dissimilarity index, rates of palynological change suggest that the interval between 8000 and 7500 BC (calendar years) was the period of most rapid vegetational change during the Holocene, both in terrestrial as well as lacustrine ecosystems. While climatic forcing of the rapid events around 8000 BC is hypothesised, the synchronous timing of relatively rapid inferred change in lake and terrestrial vegetation around AD 600 may reflect changes in climate as well as in land-use. Reducdancy analysis was used to develop a model between fire intensity (inferred from microscopical charred particles) and vegetational response, as reflected by pollen assemblages. Formulated at one site and tested at the two other sites, the model explains regional Calluna-heathland expansions as a result of vegetational burning. Similarly, declines in heathland cover are explained by lack of maintenance by fire. Regional vegetational development in northern West Jutland is reconstructed and special consideration is given to heathland history. The Holocene heathland development is interpreted as resulting from its importance for grazing. It is hypothesized that on poor soils, Calluna-dominated heathland was a better grazing resource than grass-dominated pasture, due to the winter-grazing offered by Calluna and the low palatibility of dominant grasses on poor soils. This hypothesis is relevant for the explanation of the variation in timing of heathland expansions on the different soil types represented by the study sites.     

Distribution and decline of endangered herbaceous heathland species in relation to the chemical composition of the soil

High atmospheric deposition of ammonium affects the physical and chemical status of the soil, increasing nitrogen availability, soil acidity and the mobilization of toxic metal ions. To investigate whether and how the decline of several herbaceous plant species in Dutch heathlands is associated with these processes, the chemical composition of the soil on which these species grow has been compared with the soil on which heathland species such asCalluna vulgaris (L.) Hull,Erica tetralix L. andMolinea caerulea (L.) Moench dominate. The discrimination between both soil types was primarily based on differences in pH (H₂O), pH (NaCl) and the aluminium/calcium ratio in the waterextracts. Within the group of endangered herbaceous heathland species these soil parameters also varied. This led to a division into 4 groups of species:

u

Dominating species growing on acid soils

Herbaceous species growing together with dominating species on acid soils

Herbaceous species growing together with dominating species on moderately acid soils

Herbaceous species growing together with dominating species on weakly acid soils.

This study indicated that, unlike the decline of heather species, the decline of herbaceous species is not likely to be due to increased competition from grass species as a result of eutrophication. Soil acidification and the changed mineral balance in the soil are most likely to be responsible for the decline of all three groups of herbaceous plant species. ei]R F Huettl     

Early regeneration of Calluna heathland under various fertilization treatments

Summary A sod-cutting and fertilization experiment was performed on a Calluna-dominated heathland in The Netherlands to determine appropriate management regimes for Calluna regeneration, and to further understand the nutrient responses of heathland species. Replicated permanent plots were analysed by multivariate techniques. Sod-cutting alone caused Calluna regeneration from its soil seed bank. A single fertilization at the start of the experiment caused initial vegetation differences which disappeared after a few years as the nutrients were lost from the system, except that one application of nitrogen enhanced the rate of Calluna regeneration. Repeated fertilization caused large differences in the vegetation: repeated nitrogen enhanced several bryophyte species while greatly inhibiting Calluna, repeated phosphate partly inhibited Calluna while greatly favouring several lichen species, and the most striking result of repeated calcium was also an increase in bryophytes, but the species were different from those favoured by nitrogen. Treatments which inhibited Calluna tended to increase species diversity because of the lessened Calluna dominance.     

Conservation of heathland by sheep grazing in Brittany (France): Importance of grazing period on dry and mesophilous heathlands

Heathlands are characteristic semi-natural ecosystems of western Europe. Nowadays they are threatened because of their lack of agricultural value. Management of the remaining heathland areas necessitates precise knowledge about different management tools that can be used, including grazing. The effects of sheep grazing on dry and mesophilous heathlands of Brittany were studied in relation to grazing season and duration. Management aims were to control shrubs and to maintain the dominance of heathers. The grazing pressures applied here were quite high, about 10–15 sheep ha⁻¹ yr⁻¹. Erica ciliaris appeared to be very sensitive to grazing at the end of summer. This implicates special care in the management of mesophilous heathland. In dry heathland, Erica cinerea did not show this sensitivity. Control of shrubs was achieved by most grazing regimes which led to a decrease of 11–22 cm in gorse height, except summer grazing which seemed to stimulate their growth.This study underlined the relevance of the use of sheep grazing as a heathland management tool but also the necessity of prior experiments to adapt these operations mainly in terms of grazing period to the type of heathland and the objectives put forward.