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.     

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.     

Seed banks of Erica australis and Calluna vulgaris in a heathland subjected to experimental fire

Abstract. Soil samples were collected before and after an experimental fire on a heathland in the province of León (Spain). The seed banks were assessed by counting the numbers of seedlings appearing of Erica australis and Calluna vulgaris. A total of 2285 germinable seeds/m² before burning and 1177 germinable seeds/m² after the fire were estimated for Erica australis, and 90 and 690 germinable seeds/m², respectively, for Calluna vulgaris. After the fire the number of germinable seeds/m² of Erica australis had decreased, whilst there was a considerable increase in the number of germinable seeds/m² of Calluna vulgaris. Neither seedlings nor sprouts of Calluna vulgaris had appeared in the field plots 10 months after the fire. Erica australis did recover mainly by sprouting after fire.     

Early stages of Calluna vulgaris regeneration after burning of coastal heath in central Norway

Questions: How does Calluna vulgaris regenerate after burning wet and dry heaths of different age? Location: Central Norway. Methods: Patches of Calluna dominated old and young wet heath and old dry heath were burned. The mean temperatures reached in the categories of heath were measured. Cover, frequency, height and number of seedlings of Calluna were studied in plots. Results: The mean burning temperatures varied from 680 to 740°C, and were highest in wet heath that had not been burned for at least 50 years. Regeneration of Calluna was evident in the year of burning, but only from seeds. Even heath with a short burning interval lacked vegetative regeneration. Calluna cover increased yearly and exceeded 50 % three years after burning. Regeneration was evident in the year of burning, and was highest in heath with a short burning interval. The height increased regularly at all sites, but was most rapid in dry heath, probably because of better microclimatic conditions. There were more seedlings in the heath with a short burning interval. Conclusions: Calluna only regenerates from seeds afterburning in central Norway. The density of Calluna increased rapidly, which is positive for the conservation and management of coastal heaths in central Norway.     

Calluna and its associated species: Some aspects of co-existence in communities

Summary Using heath communities (dominated byCalluna vulgaris) as an example, an attempt is made to identify those characteristics of other species wiich enable them to co-exist with the dominant. In the course of its life-span,Calluna passes through a series of growth phases (pioneer, building, mature, degenerate) during which there are marked changes in the condition affecting associated species. These display a variety of ‘complementary strategies’ which enable them to co-exist withCalluna during a part, or the whole, of the sequence. Changes with time in the flora of even-aged stands ofCalluna, produced as a result of management by burning, are accounted for in similar terms. It is also shown that invasion of heath communities by tree species (e.g.Betula spp.,Pinus sylvestris), which may eventually eliminateCalluna, is related to the sequence of growth-phases. Contribution to the Symposium on plant species and plant communities, held at Nijmegen, 11–12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff-to whom the author conveys the good wishes of Scottish botanists and conservationists, and particularly those of the University of Aberdeen. Nomenclature follows Clapham, Tutin & Warburg (1962) for vascular plants, Watson (1968) for bryophytes, and Duncan (1970) for lichens. Following common practiceCalluna vulgaris is referred to simply asCalluna. Grateful acknowledgement is made to several present and former research students who have kindly permitted the use of their data: particularly P. Barclay-Estrup, Edith M. French, Gong Wooi Khoon, C.J. Legg, Evelyn W. Paterson, S.D. Ward. I am especially grateful to C.J. Legg for valuable discussions on the subject of this paper.     

The size and composition of soil seed-banks in remnant patches of three structural rainforest types in North Queensland

The size and species composition of the soil seed-bank in a remnant patch of each of three structurally and floristically distinct rainforests (Complex Mesophyll Vine Forest, Complex Notophyll Vine Forest and Semi-Evergreen Vine Thicket) were assessed. Seeds of 94 species germinated from 12 surface soil samples collected from each site. All three seed-banks were composed mostly of herbs characteristic of roadsides and agricultural land, and pioneer rainforest trees and shrubs. Agglomerative classifications indicated that the seed-bank samples from each rainforest remnant had a characteristic species composition and could be distinguished reliably from seed-bank samples drawn from other sites. Seeds of species present in the standing forest were poorly represented in the seed-banks except for one long-lived pioneer tree, Dendrocnide photinophylla, at one site. The seed-bank from the seasonally dry vine thicket was significantly larger (4000 seeds m^-2) than those from the two moister sites (400–600 seeds m^-2, contained more seeds of roadside and agricultural herbs, and fewer seeds of rainforest pioneer and secondary shrubs and trees. We suggest three explanations for the different seed-bank structure observed in the seasonally dry forest site. First, with increased deciduousness in rainforests, seed-banks are increasingly subject to invasion and domination by seeds of rapidly maturing herbs. Second, long-lived seeds that germinate in canopy gaps would be less likely to accumulate under deciduous forests because they would he exposed annually to conditions suitable for germination. Third, chronic disturbance by cattle and pigs produces sites suitable for the establishment of rapidly maturing herbs, and possibly disperses their seeds into the forest.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Nitrogen mineralization in heathland ecosystems dominated by different plant species

Net N mineralization rates were measured in heathlands still dominated by ericaceous dwarf shrubs (Calluna vulgaris or Erica tetralix) and in heathlands that have become dominated by grasses (Molinia caerulea or Deschampsia flexuosa). Net N mineralization was measuredin situ by sequential soil incubations during the year. In the wet area (gravimetric soil moisture content 74–130%), the net N mineralization rates were 4.4 g N m^–2 yr^–1 in the Erica soil and 7.8 g N m^–2 yr^–1 in the Molinia soil. The net nitrification rate was negligibly slow in either soil. In the dry area (gravimetric soil moisture content 7–38%), net N mineralization rates were 6.2 g N M-2 yr^–1 in the Calluna soil, 10.9 g N m^–2 yr^–1 in the Molinia soil and 12.6 g N m^–2 yr^–1 in the Deschampsia soil. The Calluna soil was consistently drier throughout the year, which may partly explain its slower mineralization rate. Net nitrification was 0.3 g N m^–2 yr^–1 in the Calluna soil, 3.6 g N m^–2 yr^–1 in the Molinia soil and 5.4 g N m^–2 yr^–1 in the Deschampsia soil. The net nitrification rate increased proportionally with the net N mineralization rate suggesting ammonium availability may control nitrification rates in these soils. In the dry area, the faster net N mineralization rates in sites dominated by grasses than in the site dominated by Calluna may be explained by the greater amounts of organic N in the soil of sites dominated by grasses. In both areas, however, the net amount of N mineralized per gram total soil N was greater in sites dominated by Molinia or Deschampsia than in sites dominated by Calluna or Erica. This suggests that in heathlands invaded by grasses the quality of the soil organic matter may be increased resulting in more rapid rates of soil N cycling.     

Soil seed banks of woodland, heathland, grassland, mire and montane communities, Cairngorm Mountains, Scotland

The size and species composition of soil seed banks were assessed at 111 altitudinally diverse sites in the Cairngorm Mountains. Mean densities of germinable seeds varied from 83 000 m^–2 in Scots pine (Pinus sylvestris L.) woodland at 230–490 m to 200 m^–2 in moss (Racomitrium lanuginosum (Hedw.) Brid.) heath at 1000–1120 m. Seed banks were dominated by Calluna vulgaris (L.) Hull, not only wherever it was prominent in the vegetation, but also at some sites with less than 5% cover of parent plants in the ground vegetation. Many species conspicuous in the vegetation were under-represented in or absent from the seed bank and surface vegetation generally was more species rich than was the underlying seed bank, especially in high montane communities. Multiple regression was used to examine the relationship between the density of buried Calluna seeds and the abundance of parent plants in the vegetation, site altitude and the organic matter content of the soil. The model fitted to woodland communities accounted for 95% of the variation in seed density. The heathland model was less predictive but still explained 52% of the variation in seed bank size. In mire communities there was no relationship, collective or individual, between buried seed density and the measured environmental variables, possibly due to variations in the duration and frequency of waterlogging at these sites. The potential role of seed banks for initiating the recolonisation of disturbed ground is discussed. Densities of buried seeds at most Calluna-dominant sites were probably sufficient to generate successful recolonisation but the prospects for recovery were poor at other sites, particularly in graminaceous communities at 800 m or higher.     

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 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.     

Effect of Gibberellic Acid on Dark and Light Germination at Different Temperatures of Calluna, Ledum andRhododendron Seeds

The aim of the experiments was to study the effects of gibberellic acid (GAs) on the germination of Calluna vulgaris L., Ledum palustre L. and Rhododendron lapponicum (L.) Wahlenb. seeds under different environmental conditions. Under continuous light from white fluorescense tubes (3000 lux), untreated seeds of Calluna were partly dormant at all temperatures studied (9, 15, 21, 27, 27/9, 8/16 hours). Percentage of dormant seeds increased, however, with decreasing temperature, and it varied also from seed lot to seed lot. Untreated seeds of Ledum were dormant in light at 9° and 15°C but not at higher temperatures. Untreated seeds of Rhododendron were completely dormant in light at temperatures from 13° to 24°C. Seeds of all species were completely dormant in darkness both at 15° and 27°C. GA₃ stimulated greatly the germination of all species under all studied environmental conditions. The used concentrations (0.2–3.2 mM) gave nearly 100% germination in most cases. At 9°C the dormancy in some seed lots of Calluna and Ledum was only partly broken by the used concentrations of GA₃.     

Root turnover as determinant of the cycling of C,N, and P in a dry heathland ecosystem

Root production and turnover were studied using sequential core sampling and observations in permanent minirhizotrons in the field in three dry heathland stands dominated by the evergreen dwarfshrub Calluna vulgaris and the grasses Deschampsia flexuosa and Molinia caerulea, respectively. Root biomass production, estimated by core sampling, amounted to 160 (Calluna), 180 (Deschampsia) and 1380 (Molinia) g m^-2 yr^-1, respectively. Root biomass turnover rate in Calluna (0.64 yr^-1) was lower compared with the grasses (Deschampsia: 0.96 yr^-1; Molinia 1.68yr^-1)). Root length turnover rate was 0.75–0.77 yr^-1 (Deschampsia) and 1.17–1.49 yr^-1 (Molinia), respectively. No resorption of N and P from senescing roots was observed in either species. Input of organic N into the soil due to root turnover, estimated using the core sampling data, amounted to 1.8 g N m^-2 yr^-1(^Calluna), 1.7 g N m^-2 yr^-1 (Deschampsia) and 19.7 g N m^-2 yr^-1 (Molinia), respectively. The organic P input was 0.05, 0.07 and 0.55 g P M^-2 yr^-1, respectively. Using the minirhizotron turnover estimates these values were20–22% (Deschampsia) and 11–30% (Molinia) lower.When the biomass turnover data were used, it appeared that in the Molinia stand root turnover contributed 67% to total litter production, 87% to total litter nitrogen loss and 84% to total litter phosphorus loss. For Calluna and Deschampsia these percentages were about three and two times lower, respectively.This study shows that (1) Root turnover is a key factor in ecosystem C, N, and P cycling; and that (2) The relative importance of root turnover differs between species.     

An integrated assessment of ecosystem carbon pools and fluxes across an oceanic alpine toposequence

Alpine ecosystems are predicted to be severely affected by climate change. Cold, wet oceanic-alpine environments may also accumulate large total ecosystem carbon (C) pools, but have rarely been investigated. We assessed C pools and fluxes on a toposequence of oceanic-alpine habitats from blanket mire and boreal Calluna heath to Racomitrium heath, Nardus snowbed and alpine Calluna heath. We quantified C pools in vegetation and soils for each habitat and compared these with C inputs from net primary production (NPP) and outputs via decomposition, measured in a 3-year litter bag experiment. We also investigated principle drivers (temperature, moisture, community composition) of C pool and flux differences between habitats. Total ecosystem C pools were large; 11–26 kg C m^?2 in alpine habitats and 50 kg C m^?2 in blanket mire. Within the alpine zone C storage was greatest in the snowbed. Litter decomposition was slow in all habitats (k?=?0.09–0.29 y^?1) while NPP was within the range reported for continental alpine systems. C pool sizes and C fluxes did not vary consistently with altitude but reflected topographic gradients of temperature and moisture within the alpine zone. Oceanic-alpine ecosystems contain large stores of C which may be vulnerable to the effects of climate change.     

Annuario Della Società Botanica Italiana 1937

Abstract

The European heathers Erica manipuliflora, Erica multiflora and Erica vagans (sect. Gypsocallis) are related species characterized by a glabrous ovary, exerted, muticous anthers and a pale pink, urceolate corolla. The existence of three species is the most accepted treatment for this group, but there are uncertainties about this taxonomic interpretation. In this article, we used seed morphological characters to assess the systematics of the group. Seeds were studied by means of scanning electron microscopy techniques. E. multiflora has a distinct seed type, 1–1.5 mm long, flattened, yellowish with a striate-reticulate primary sculpture. E. vagans and E. manipuliflora have smaller seeds, brown with a reticulate pattern. Seeds of E. vagans are globose, 0.5 mm wide, while E. manipuliflora has mostly ellipsoid seeds 0.7 mm long. However, seeds of E. manipuliflora from the Adriatic coasts are closer to those of E. vagans, and this may support the existence of E. manipuliflora subsp. anthura.     

Plant recruitment in the High Arctic: Seed bank and seedling emergence on Svalbard

Abstract. Composition and density of the soil seed banks, together with seedling emergence in the field, were examined on Svalbard. 1213 soil samples were collected from six drymesic habitats in three regions representing various stages of colonization from bare moraines to full vegetation cover and spanning a range of typical nutrient and thermal regimes. Of the 165 vascular plant species native to Svalbard, 72 were present as mature plants at the study sites and of these 70% germinated seed. Proglacial soil had 12 seedlings per m², disturbed Dryas heath 131, intact Dryas heath 91, polar heath 715, thermophilic heath 3113, and a bird cliff 10437 seedlings. Highest seed bank species richness was at the thermophilic heath (26 species). Seedlings of 27 species emerged in the field, with fewer seedlings in disturbed habitats (60 seedlings per m²) than in intact Dryas heath (142), suggesting that an absence of ‘safe sites’ limited seedling establishment in disturbed habitats. Measurement of seedling emergence in the field increased awareness of which species are able to germinate naturally. This may be underestimated by up to 31% if greenhouse trials alone are used, owing partly to unsuitability of greenhouse conditions for germination of some species and also to practical limitations of amount of soil sampled. Most thermophilic species failed to germinate and some species present at several sites only germinated from the thermophilic heath seed bank, suggesting that climate constrains recruitment from seeds in the High Arctic.     

Community dynamics in relation to management of heathland vegetation in Scotland

The paper describes studies of post-fire succession in heathland vegetation in N.E. Scotland, dominated by Calluna vulgaris. A preliminary model (Legg, 1978) suggested good agreement between simulation of succession on the basis of a Markov chain and observations of stands at different stages of development after burning, at least in the earlier stages. Vegetation transitions are currently being recorded in permanent plots on burnt areas. First results confirm the view that (a) the post-fire succession has the properties of a Markov process, (b) this type of model remains valid when constructed from records of actual transitions, rather than data obtained by inference from evidence of transition. Comparing successional events in stands where, at the time of burning, the Calluna population was in pioneer-, building-, mature-and degenerate phases, shows that transition matrices generally agree with the Markov hypothesis, but not in the case of stands where Calluna was degenerate when burnt. The composition of establishing vegetation 1 year after fire is not confined to species normally associated with the early stages of succession, but reflects the composition of the stand before burning. Redevelopment after fire is described in terms of an initial floristic composition of species with strategies permitting early re-establishment, selected by the recurrence of the fire factor. Subsequent transitions represent changes in their relative abundance due to differing growth properties and competitive interactions. This interpretation applies only under conditions of recurrent incidence of fire (normally once in 10–15 yr). If fire does not recur, Calluna stands pass into the degenerate phase, where changes in the nature of relay floristics may come into play (e.g. with tree colonization).Nomenelature follows Clapham, Tutin & Warburg (1962) for vascular plants; Smith (1978) for bryophytes.     

Productivity relations in a Carex-dominated ecosystem

Summary Seasonal and total primary productivity was measured for a Carex meadow in southern Quebec, Canada. Forty-five one-meter² plots were sampled for dry weight biomass, species composition, structure (species density, diversity, height) and soil parameters including macronutrient concentrations (Ca, K, Mg, Na, N, P), pH, organic matter, and water depth. Shoot net productivity and litter decomposition rates were computed for 20-day intervals May–September, inclusive. Relationships between all parameters were examined by principal components analysis.Dominant species included Carex lacustris, C. aquatilis, Calamagrostis canadensis, and Typha angustifolia. For a 130-day growth period, mean shoot net productivity was 6.3 g·m^-2· da^-1 and terminal standing crop 807 g·m^-2. Terminal standing crop was very close to above ground biomass predicted by the Gorham equation based on thermal relations for Carex ecosystems and to total accumulated litter mass (779 g·m^-2). Seasonal production showed a strong bimodal pattern with peak productivities in mid-June (15.3 g·m^-2·da^-1) and mid-July (4.3 g·m^-2·da^-1). Decomposition of the previous year's litter was 81% complete by late September.Soil fertility, fire incidence, and topographic position were the three most important gradients resolved by principal components analysis. The first component distinguished sediment-rich Typha angustifolia communities near open water from oligotrophic stands of Carex spp. on central areas of the meadow. Production levels correlated closely with extractable soil calcium (r=0.40^**) and phosphorus levels (r=0.39^**). Species diversity and stem density related inversely to productivity on this component. Fire incidence (component II) had a marked effect on species diversity due to surface scarification and removal of litter mass. Component III was a topographic gradient separating composition, and community structure.Magnesium and sodium levels decreased from upland to open water. Soil phosphorus increased markedly at water's edge related to mineral input by sedimentation. Pattern of N, P, K, and Ca coincided closely with total shoot production and litter mass levels suggesting closed biotic cycles of these elements.A model accounting for species diversity levels in Carex meadow was formulated based on the assumption that high productivity results in competitive species elimination.     

Community dynamics of evergreen broadleaf forests in southwestern Japan. II. Species composition and density of seeds buried in the soil of a climax evergreen oak forest

The species composition and density of seeds in the soil of a climax evergeen oak forest in the Kasugayama Forest Reserve, Nara, Japan were studied by directly extracting seeds from soil samples and using soil samples in planting boxes as the basis of germination tests. A total of 33 species were identified from all seeds collected in 6 stands: 11 evergreen broadleaf, 15 deciduous broadleaf, 2 coniferous, 2 liana, 1 herb, and 2 grass species. Species producing sap fruits and dry fruits accounted for 60% and 40% of the total number of species, respectively. The species composition of all the seeds was independent of the species composition of the forest vegetation. The mean density of the seeds was 22,134 seeds/m²·5 cm.Eurya japonica, of which seeds were found in all soil samples, was the most abundant species, followed byCryptomeria japonica, Ilex micrococca, andBoehmeria longispica. Pioneer species such asMallotus japonicus, Zanthoxylum ailanthoides, andAralia elata were found in all soil samples in spite of the paucity of adult trees in the forest. Seeds of evergreen oaks were relatively aboundant but no viable seeds were found. ViableE. japonica, I. micrococca, Symplocos prunifolia, andB. longispica seeds were abundant.     

The influence of ammonium nitrate on the root growth and ericoid mycorrhizal colonization of Calluna vulgaris (L.) Hull from a Danish heathland

Conversion of European heathlands to grassland has been reported as a response to increased nutrient availability, especially of nitrogen; a direct effect upon mycorrhizal colonization has been proposed as an likely explanation.This hypothesis was tested in a random block experiment with four blocks and four replicates on a Danish inland heath, Hjelm Hede. Ammonium nitrate was applied (0, 35, 50 and 70 kg N ha^–1 year^–1) to a stand of Calluna vulgaris (L.) Hull four times annually for 2 years. Calluna roots were sampled on four occasions in the 2nd year of the nitrogen treatment. The extent of ericoid mycorrhizal colonization was determined by direct observation of the roots using a line-intersection method. The nitrogen content of the current-year shoots of Calluna increased when they were treated with nitrogen. Nitrogen fertilization had no significant effects on ericoid mycorrhizal colonization of Calluna nor on root biomass. The seasonal variation in mycorrhizal colonization of the Calluna roots was highly significant. The spatial variability of mycorrhizal colonization, both in replicated plots and in the two contrasted soil horizons – the mor layer and the bleached sand – within the plots, were considerable. I conclude that heather decline under enhanced nitrogen input is unlikely to be caused by a direct impact on the ericoid mycorrhizae of Calluna. Received: 1 December 1998 / Accepted: 7 December 1999