Recovery of pristine boreal forest floor community after selective removal of understorey, ground and humus layers

In boreal spruce forests that rarely experience extensive disturbances, fine-scale vegetation gaps are important for succession dynamics and species diversity. We examined the community implications of fine-scale gap disturbances by selective removal of vegetation layers in a pristine boreal spruce forest in Northern Finland. The aim was to investigate how the speed of recovery depends on the type of disturbance and the species growth form. We also wanted to know if there appeared changes in species composition after disturbance. Five different treatments were applied in the study: Control, removal of the ground layer (bryophytes and lichens), removal of the understorey layer (dwarf shrubs, herbs and graminoids), removal of both the ground and understorey layers, and complete removal of the vegetation and humus layers above the mineral soil. The vegetation recovery was monitored in terms of cover and species numbers over a 5-year period. Understorey layer cover, composed mainly of clonal dwarf shrubs, recovered completely in 4 years in treatments where the humus layer remained intact, whereas ground layer cover did not reach the control level in plots from where bryophytes and lichens were removed. Recovery was faster in terms of species number than species cover. Bryophytes, graminoids and dominant dwarf shrubs appeared in all disturbed plots quickly after disturbance. Seedlings of trees appeared exclusively in disturbed plots. Graminoids dominated after the removal of humus layer. The results indicate that the regeneration of forest floor after small gap disturbance occurs mainly by re-establishment of the dominant species. Although destruction of the humus layer leaves a long-lasting scar to the forest floor, exposing of mineral soil may enhance the sexual reproduction of dominant species and the colonization of weaker competitors.     

Experimental Restoration of Native Vegetation in Indiana Dunes National Lakeshore

We investigated the effects of prescribed fire, herbicide treatment, and sod removal on the eradication of exotic grasses and the establishment of native plant species in 24 experimental restoration plots in three razed residential sites within the boundary of Indiana Dunes National Lakeshore. During 1992–1995, herbicide treatment and sod removal decreased the combined cover of Poa pratensis (Kentucky blue grass) and Agropyron repens (quackgrass) significantly (from 82% to 13%, and 85% to 8%, respectively), whereas fire did not suppress such exotic lawn grasses. In 1993, several opportunistic species, represented by Cyperus spp. (umbrella sedges), Digitaria sanguinalis (crab grass), and Ambrosia artemisiifolia (common ragweed), filled the gaps left by the removal of lawn grasses. For the same period, Detrended Correspondence Analysis revealed a clear vegetation divergence between the control-fire plots and the herbicide-sod removal plots. While Poa pratensis and Agropyron repens continued to dominate the control and fire plots, the planted native species, represented by Schizachyrium scoparium (little blue-stem), Sorghastrum nutans (Indian grass), Rudbeckia hirta (black-eyed Susan), and Monarda punctata (horsemint), began to dominate in the herbicide and sod removal plots from 1994. In both herbicide and sod removal plots, the ground cover of grasses (68%) was much higher than the forbs (10%). The herbicide plots, where exotic species were removed but nitrogen-rich top soils were not removed, showed a higher diversity of planted native species than the sod removal plots (where both exotic species and top soils were removed) and the control-fire plots (where neither was removed). This finding suggests that an optimum but not excessive concentration of soil nitrogen is needed to support a maximum species diversity in such infertile substrate as sandy soil. In addition, the decrease in potassium in all plots, regardless of treatment, suggests that potassium may become a limiting factor for our restored native vegetation.     

Effects of reclamation measures on plant colonization on lignite waste in the eastern Pyrenees,Spain

Abstract. This paper deals with vegetation establishment on waste dumps in the montane and submontane belts of the eastern Pyrenees, where submediterranean Quercus humilis forests, Pinus sylvestris forests and mesoxerophilous pastures make up most of the landscape; these are considered as target communities for reclamation. The waste consists of marl and lime regolith, very poor in nutrients and structure (Poch et al. 1989). The dumps were terraced in 1985–1986; some were left for spontaneous re‐colonization and others were reclaimed in various ways. In 1992 we took 36 vegetation samples on the dumps, based on the point quadrat procedure, to evaluate the colonization status and the differences between reclamation techniques. The samples were analysed on the basis of their species composition (ecological groups and multivariate analysis) and structural aspects (coverage, diversity, etc.). Reclamation treatment results varied widely, most of the dumps showing a low degree of naturalness. Colonizing vegetation ranged from a mixture of opportunist and stress‐resistant taxa, forming poorly covered surfaces on the dumps where colonization has been poor, to dense grasslands dominated by one (or a few) introduced competitive grass species, where reclamation procedures had been intensive. Sowing treatments, where pasture species were sown onto dumps, produced intermediate results, as resulting vegetation cover was similar to control plots and naturalness was low, but in a few cases they yielded more interesting swards, fairly dense and diverse, and including high numbers of spontaneous species. Choice of sown species and proximity to undisturbed vegetation accelerate succession.     

Plant diversity,species turnover and shifts in functional traits in coastal dune vegetation: Results from permanent plots over a 52‐year period

Question: What is the relationship between plant diversity and species turnover in coastal dune vegetation plots? How is the long‐term change in species composition of vegetation plots related to shifts in functional traits, and what does it tell us about the dominant processes? Location: Coastal dunes, the Netherlands. Methods: Our data set comprised 52 years of vegetation data from 35 permanent plots in grassland/scrub/woodland vegetation. Vegetation dynamics were described in terms of changes in species composition and abundance, and shifts in 13 functional traits related to resources capture and forage quality, regeneration and dispersal. Results: Species turnover in the plots was high, because of local extinction and colonization. Species‐rich plots were more stable in terms of species abundance and composition compared with species‐poor plots. Over time, the plots converged with respect to their abiotic conditions, as reflected by Ellenberg indicator values – indicating that the prevailing process was succession. The high species turnover reflected high invasibility: accordingly, the relative importance of annuals increased. Most newcomer annuals, however, were competitive generalists of little conservation value. The functional trait analysis allowed us to unravel the complexity of effects of disturbances and succession, and yielded information on the processes driving the observed vegetation dynamics. Conclusions: In this study, small‐scale species turnover was negatively related to species diversity, indicating more stability in species‐rich communities. Regarding shifts in trait diversity, unifying filters appeared to be more dominant than diversifying filters. Counteracting this homogenization process poses a challenge for nature management.     

Plant colonization windows in a mesic old field succession

Abstract. Closed canopy vegetation often prevents the colonization of plant species. Therefore the majority of plant species are expected to appear at the initial phase of post‐agricultural succession in mesic forest environment with moderate levels of resources. This hypothesis was tested with data from the Buell‐Small Successional Study, NJ, USA, one of the longest continuous fine‐scale studies of old‐field succession. The study started in 1958, including old fields with different agricultural histories, landscape contexts, and times of abandonment. In each year of the study, the cover values of plant species were recorded in 48 permanent plots of 1 m² in each field. We analysed the temporal patterns of colonization at plot scale and related these to precipitation data and other community characteristics. The number of colonizing species decreased significantly after ca. 5 yr, coinciding with the development of a continuous canopy of perennial species. However, species turnover remained high throughout the whole successional sequence. The most remarkable phenomenon is the high inter‐annual variation of all studied characteristics. We found considerable temporal collapses of vegetation cover that were synchronized among fields despite their different developmental stages and distinctive species compositions. Declines of total cover were correlated with drought events. These events were associated with peaks of local species extinctions and were followed by increased colonization rates. The transitions of major successional stages were often connected to these events. We suggest that plant colonization windows opened by extreme weather events during succession offer optimum periods for intervention in restoration practice.     

Population dynamics of Gentiana pneumonanthe and Rhynchospora fusca during wet heathland restoration

Abstract. The population dynamics and reproductive strategies of two rare wet heathland species, Gentiana pneumonanthe and Rhynchospora, were studied in experimental permanent plots in a wet heathland near Bremen, NW Germany, to assess how effective sod cutting and mowing are in promoting these species. In one experiment, small plots (0.25 m² - 4 m²) were sod-cut or mown; in the second, one large plot 30 mx 50m) was sod-cut. The development of the vegetation and the number of shoots of the two target species were recorded annually. Sod cutting lead to the highest shoot numbers of Gentiana in the long run, whilst mowing was more effective at the beginning of the experiment. Seedlings and adult shoots slowly became more and more abundant after sod cutting. In contrast, Rhynchospora soon formed closed stands after sod-cutting, first through seed germination and then through fast clonal growth of the established individuals. Moist soil or short inundations promoted the germination and seedling establishment of Gentiana, whereas drought negatively affected seedling recruitment. Long periods of inundation severely reduced the population at first, but high numbers of seedlings were found in the following growing season. For the disturbance-dependent Gentiana and Rhynchospora, the availability of gaps within the vegetation is of crucial importance. To promote existing populations, we suggest small-scale sod cuttings which create gaps without disturbing existing flowering individuals too much. For degenerated stands of wet heathland we recommend large-scale sod cutting to activate the seed bank. Additionally, seed introduction may be helpful to encourage the development of a wet heathland with characteristic floristic composition.     

Experimental assessment of initial revegetation on abandoned paths in temperate deciduous forest

Question: Do soil treatments and addition of seed facilitate rapid vegetation restoration on forest paths excluded from trampling? Location: Six mesophilic mixed deciduous Querco‐Fagetea forests in Flanders, northern Belgium. Methods: Enclosures on paths were excluded from trampling by fencing. In a full factorial design, plots were subjected to seeding, soil scarification, addition of organic material and inoculum. The following two years, seedling establishment and growth were sampled during spring and summer. Soil treatments and seeding effect was tested and seedling occurrence was analysed in relation to species' origin. Results: Spontaneous re vegetation was significant in all plots since fencing. Throughout the observation period seedling cover and height continued to increase. Seeding had an overall effect on seedling density, cover and height. Some soil treatment interactions significantly enhance re vegetation, although each individual soil treatment had no significant effect. Regardless of the seeded individuals, invading species mostly originated from the surrounding area and the seed bank. Conclusion: The preliminary results of this experiment imply that seeding is the only treatment which has a positive effect on re vegetation success in all circumstances, provided that the exclusion from trampling is effective.     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

Long-Term Effects of Reclamation Treatments on Plant Succession in Iceland

The long‐term effects (20–45 years) of reclamation treatments on plant succession are examined at two localities in Iceland that were fertilized and seeded from 1954 to 1979 with perennial grasses or annual grasses, or left untreated. The areas that underwent reclamation treatments had significantly higher total plant cover (7–100%) than the untreated control plots (<5%), and floristic composition was usually significantly different between treated and untreated plots. Dwarf‐shrubs (Calluna vulgaris and Empetrum nigrum), bryophytes, biological soil crust, grasses, and shrubs characterized the vegetation in the treated plots, but low‐growing herbs that have negligible effects on the environment, such as Cardaminopsis petraea and Minuartia rubella, and grasses characterized the control plots. The seeded grass species had declined (<10%, the perennials) or disappeared (the annuals) but acted as nurse species that facilitated the colonization of native plants. It seems that by seeding, some factors that limit plant colonization were overcome. Soil nutrients, vegetation cover, litter, and biological soil crust were greater in the treated areas than the control plots. This may have enhanced colonization through an increase in soil stability and fertility, increased availability of safe microsites, increased moisture, and the capture of wind‐blown seeds. This study demonstrates the importance of looking at the long‐term effects of reclamation treatments to understand their impact on vegetation succession.     

Dynamics of submersed aquatic vegetation on shallow soft bottoms in the Baltic Sea

Abstract. Persistence and colonization of submersed aquatic plant species were studied in permanent plots (20 cm × 20 cm) at three shallow sites adjacent to Askö island, in the northern Baltic Sea. The study started in 1991 at two of the sites, in 1992 at a third site and continued until 1997. Two major weather‐induced disturbances occurred during the study: a long, stormy period during a mild winter and a cold winter with extreme low water levels. The stormy period caused a large loss of species from plots (95 ± 5% SE) at the most wave‐exposed site, resulting in a low species number per plot (0.8 ± 0.2) the following summer. During the three following years the mean species number increased to 3.6 ± 0.2. The cold winter caused high species turnover and increased species number per plot at the two most wave‐exposed sites. The species turnover at the sheltered site was highest in the two years with low water level in May, possibly due to increased waterfowl herbivory. Annuals, loosely anchored and highly reproductive species had significantly higher plot colonization rates and lower persistence than perennials, well‐anchored species and species with none or low reproduction. The extent of lateral growth had no significant effect on colonization or persistence. Although these disturbances have a large effect on the dynamics, species mobility was also high in other years. Relative to other, similar, studies in terrestrial vegetation mean persistence in plots was remarkably low and mean mobility and species turnover rates were very high.     

Experimental analysis of patch dynamics and community heterogeneity in tallgrass prairie

Effects of fire and small-scale soil disturbances on species richness, community heterogeneity, and microsuccession were investigated in a central Oklahoma tallgrass prairie. In the fall of 1985, 0.2 m² soil disturbances were created on burned and unburned tallgrass prairie. Vegetation on and off disturbances was sampled at monthly intervals over two growing seasons. During the first growing season, the cover of forbs and annuals, and species richness were significantly greater on versus off disturbances, but these differences did not persist through the second year. The variation in species composition among disturbed plots (heterogeneity) was significantly greater compared to undisturbed areas throughout the study. Fire had no consistent effect on richness and heterogeneity of vegetation on soil disturbances but fire reduced heterogeneity on undisturbed vegetation. Rate of succession, based on an increase in cumulative cover of perennial grasses over time, did not differ among treatments during the first growing season. During the second year, rate of succession was significantly greater on burned soil disturbances compared to unburned soil disturbances. These results suggest that while small-scale soil disturbances have primarily short-lived effects on grassland community structure, disturbances do help to maintain spatial and temporal variation in tallgrass prairie communities. Unlike in undisturbed vegetation, however, species richness and heterogeneity on soil disturbances were little effected by fire, but the rate of colonization onto disturbances appeared to be enhanced by fire.     

Low impact of climate change on subalpine grasslands in the Swiss Northern Alps

While phenological shifts and migration of isolated species under climate change have already been observed on alpine summits, very few studies have focused on community composition changes in subalpine grasslands. Here we use permanent plots monitored since 1954 and precisely located phytosociological censuses from 1970 to study compositional changes of subalpine grasslands in two distinct regions of the Swiss Northern Alps. In both areas, warming trends during the monitoring period were associated with changes in land management (abandonment of goat and sheep pasturing or grazing replaced by mowing). Old and recent inventories were compared with correspondence analyses (CA). Ecological indicator values, community‐affinities and biological traits of the species were used to infer the factors responsible for triggering the observed changes. In both regions, subalpine grasslands were stable with smaller changes than have previously been observed in alpine environments. Only a few species appeared or disappeared and changes were generally limited to increasing or decreasing frequency and cover of certain taxa. At one site, grazing abandonment favored fallow species. Some of these species were located at their upper altitudinal distribution limits and may have spread because of rising temperatures. In both areas, declining species were predominantly alpine and low‐growing species; their decline was probably due to increased competition (e.g., shadow) with more vigorous subalpine taxa no longer limited by grazing. We conclude that vegetation communities can respond rapidly to warming as long as colonization is facilitated by available space or structural change. In the subalpine grasslands studies, changes were mainly driven by land management. These communities have a dense vegetation cover and newly arriving herbaceous species preferring warmer conditions may take some time to establish themselves. However, climate disturbances, such as exceptional drought, may accelerate community changes by opening gaps for new species.     

Long‐term influence of sod cutting depth on the restoration of degraded wet heaths

Temperate heaths have an unfavorable conservation status in most European biogeographical regions. Increasing nitrogen levels promote competitive grass species such as Molinia caerulea, which is a main threat to heathland conservation in Europe. This article investigates the long‐term influence of sod cutting and the resulting changes in soil properties on the heath composition, integrity, and structure. In 15 nature reserves across the northern half of Belgium, we used (1) a large number of plots (203); (2) a broad range of sod cut depths (2–40 cm), and (3) a temporal dimension that describes how long the effects of sod cutting persist (census up to 19 years after sod cutting). Multivariate analyses were used in order to explore the influence of sod cut depth and time after sod cutting on the soil and vegetation properties. There was a positive relationship between sod cut depth and soil pH and water level, and a negative relationship with Al³⁺, NH₄⁺, and total organic matter (TOM). However, only a limited number of typical (target) species appeared after sod cutting, and then only weakly. Most of the time they remained a minor component of the restored vegetation. Moreover, M. caerulea reappeared and its cover significantly increased during the years following sod cutting. Although we were able to show that sod cut depth has a differential effect on soil properties and vegetation recovery, it also appeared that sod cutting does not restore wet heaths in the long term when applied in regions with high nitrogen deposition.     

Using Precision Prairie Reconstruction to Drive the Native Seeded Species Colonization Process

Restoring forb richness to the northern tallgrass prairie (U.S.A.) is often problematic. A potential solution is the establishment of native forb‐seeded patches that can serve as colonization sites. This study was designed to determine the following: (1) the success at which native forbs sown in small patches can colonize the surrounding vegetation matrix and (2) whether soil amendments (C additions, P fertilization, and seed bank reduction) applied to the seeded sites can facilitate such colonization (patch quality). Colonization was investigated at (1) the immediate surrounding (1 m belt transect, BT) of the seeded sites and (2) in disturbed but not seeded patches located 3 m from the seeded site (out‐plots). Soil amendments did not affect colonization, but native forb density in the BT and out‐plots was correlated to the density of forbs in the seeded sites (r² = 0.3, p < 0.01). Seeded native forb density in the out‐plots was higher than in the adjacent BT (19 vs. 5 plants/m²), suggesting that competition from the matrix vegetation may be more important in forb colonization than distance from the source. Taller forbs with larger seeds showed advantages in colonization success. This study showed that the colonization of matrix vegetation with native forbs from seeded patches is a viable method for prairie reconstruction but requires the availability of disturbed sites. In most prairies, animal‐generated soil disturbances are common. In their absence, disturbances of the vegetation matrix will need to be added to the management plan.     

Restoration success of low‐production plant communities on former agricultural soils after top‐soil removal

Abstract. The success inrestoring seven low‐production vegetation types on former agricultural soil after top‐soil removal was investigated. The colonization and establishment of target species in permanent plots was recorded during the first nine years after restoration measures were taken. For each permanent plot abiotic site conditions were used to determine which of the vegetation types could persist there. A comparison of the actual vegetation in the permanent plots with reference releves of the selected vegetation type revealed a gradually increasing similarity during consecutive years for five vegetation types. This was due to the occurrence of an increasing number of target species and the number of permanent plots they occurred in. However, nine years after top‐soil removal a large number of the target species were still lacking from the vegetation in the permanent plots, although most were present in the local species pool. Seed dispersal therefore seems to be a major limiting factor for restoration of these low‐production vegetation types on formerly agricultural soils.     

Recovery of small denuded patches of the dominant NSW coastal saltmarsh species (Sporobolus virginicus and Sarcocornia quinqueflora) and implications for restoration using donor sites

Summary Efforts to restore coastal saltmarsh habitats through transplantation of saltmarsh species from donor sites are becoming more frequent. As sections removed are generally small there is little information on the ability of these areas to recover naturally after removal. The present study provides an understanding of the potential of dominant saltmarsh species in NSW (Saltcouch, Sporobolus virginicus and Samphire, Sarcocornia quinqueflora) to regenerate naturally after such small‐scale removal (25 cm × 25 cm plots) at separate sites for each species. The increase in percentage cover of vegetation within the denuded plots was measured over 21 months along a gradient within the marsh (between the mangrove and terrestrial boundaries) and then compared to non‐denuded plots. Recovery of Saltcouch was slow with an estimate of 5–6 years to reach the surrounding cover levels. Similarly, recovery of Samphire on upper levels of the shore is estimated at 4–5.5 years. On the lower level on the shore, however, Samphire had reached a cover that was within the range of the surrounding vegetation by the end of the 21‐month study period.     

Regeneration of a Gentiana pneumonanthe population in an oligotrophic wet meadow

Abstract. The aim of this study was to evaluate the critical phases of the life cycle of Gentiana pneumonanthe, the marsh gentian, a threatened species of the Bohemian flora. The effect of various conditions on germination and seedling establishment and the possible effect of competition on the performance of the species were tested. Seeds were sown in plots which were subjected to four treatments in a randomized complete blocks experiment: unmanaged meadow, mown meadow, burned meadow, and meadow with cut sod. The significantly highest recruitment was found in plots with cut sod, the lowest in the unmanaged control plots. Seedling survival also differed among the treatments. In the following year surviving individuals were only found in plots with cut sod. The influence of neighbouring vegetation on target gentian individuals was evaluated by removing the surrounding vegetation and comparing the performance of these individuals with controls. The initial height of each individual was measured and used as a covariable. No significant effect of neighbouring vegetation on performance was detected. Thus, the establishment phase appears to be critical for population persistence and is also more influenced by the management regime than other stages of the life cycle.     

Shrubland Restoration Following Woody Alien Invasion and Mining: Effects of Topsoil Depth, Seed Source, and Fertilizer Addition

Invasion by woody alien plants, construction, and mining operations are among the major disturbances degrading vegetation in the Cape Floristic Kingdom, South Africa. The aim of this study was to assess whether native fynbos shrubland vegetation could be restored following dense alien invasion and disturbance by mining. An area supporting dense alien trees was cleared and topsoil was stripped and stockpiled to simulate mining disturbance. A field trial investigated the effects of topsoil depth, seed mix application, and fertilizer on native species recruitment and vegetation development over a three‐year period. Soil‐stored seed banks contributed 60% of the species recruited, indicating that areas invaded for three decades have good restoration potential. The addition of a fynbos seed mix, which included serotinous overstory species, improved both the richness and structural composition of the vegetation. Most species sown in untopsoiled plots established, but survival and growth was low compared to topsoil plots. Poor growth in combination with a lack of soil seed bank species, indicate that restoring a diverse and functional cover of indigenous vegetation on subsoil is not possible in the short‐term. Soil amelioration is required to improve rooting conditions and initiate ecosystem processes. Shallow and deep topsoil treatments yielded high plant density, richness, and projected canopy cover, but canopy cover was higher in deep topsoil plots throughout the trial. Fertilizer addition increased canopy cover in untopsoiled and shallow topsoil plots via an increase in alien annual species. Fertilizer addition ultimately may lead to increased native vegetation cover in untopsoiled areas, but as it increased proteoid mortality on deep topsoil plots, it is not recommended for sites where topsoil is available. A species‐rich and structurally representative fynbos community may be restored on topsoiled areas provided that the native disturbance regime is simulated and seeds of major structural guilds not present in the soil seed bank are included in the seed mix.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Relationships between seed banks and spatial heterogeneity of North American alvar vegetation

Abstract. This is the first quantitative study of seed bank characteristics in North American alvar habitats. We assessed seed bank density, species richness, and species composition in 75 plots distributed among five alvar sites in Bruce Peninsula National Park, Ontario, Canada, each of which displayed areas of high and low vegetation cover within the alvar and a fully forested perimeter area. Forested habitats immediately adjacent to alvar patches contained minimal seed banks for species restricted to the alvar patches. Open alvars contained less than 1% seeds from woody forest species. This suggests that forest is not invading adjacent alvar habitat via seeds and that adjacent forest does not contain a reservoir of alvar seeds. When compared to areas on the alvar with high vascular plant cover, areas with low cover contained a slightly smaller viable seed bank, but seed banks from high and low vegetation cover plots had similar species composition and species richness. High vegetation cover plots had slightly higher mean and maximum soil depths compared with low cover plots, but no differences in other physical and chemical parameters. Thus, spatial heterogeneity in plant cover is associated only weakly with heterogeneity in below‐ground factors. Despite the availability of seed and soil resources, vegetation dynamics are constrained in areas with low plant cover, and thus alvar community development seems to respond non‐linearly to resource availability.