Nurse effect of Bolax gummifera cushion plants in the alpine vegetation of the Chilean Patagonian Andes

Abstract. It has been proposed that in the harsh arctic and alpine climate zones, small microtopographic variations that can generate more benign conditions than in the surrounding environment could be perceived as safe sites for seedling recruitment. Cushion plants can modify wind pattern, temperature and water availability. Such modifications imply that cushion plants could act as ‘nurse plants’ facilitating the recruitment of other species in the community. This effect should be more evident under stressful conditions. We tested these hypotheses comparing the number of species that grow inside and outside Bolax gummifera cushions at two elevations (700 and 900 m a.s.l.) in the Patagonian Andes of Chile (50°S). At both elevations, and in equivalent areas, the number of species was registered within and outside cushions. A total of 36 and 27 plant species were recorded either within or outside B. gummifera cushions at 700 and 900 m a.s.l., respectively. At 700 m a.s.l., 33 species were recorded growing within cushions and 29 outside them, while at 900 m a.s.l. these numbers were 24 and 13 respectively. At both elevations there were significantly more species growing within than outside cushions, and the proportion of species growing within cushions increased with elevation. Thus there is a nurse effect of cushion plants and it is more evident at higher elevations. Shelter from wind and increased soil water availability seem to be the factors that increase plant recruitment within cushions.     

Variation in vegetation and seed bank in a Chilean semi‐arid community affected by ENSO 1997

Abstract. We followed plant cover and soil seed density of shrubs and herbs in two markedly contrasting years with regard to annual rainfall in two opposite‐facing slopes (mesic vs xeric) and an intervening, relatively wet, ravine of a typical creek in semi‐arid Chile. During the ENSO year 1997 cover of ephemerals increased in all three sites; 43% vs 8% on the xeric slope 75% vs 26% on the mesic slope and 57% vs 32% in the ravine. The number of species was almost three times higher on the xeric slope (27 vs 10 species), increased by 47% on the mesic slope (28 vs 19) and by 14% in the ravine (24 vs 21). Cover of shrubs plus perennial grasses increased from 52% in 1996 to 59% in 1997 on the xeric slope, but no significant cover changes were found on the mesic slope or the ravine. On the xeric slope peak density of seeds was 4500/m² in 1996, while during the ENSO year it was 24000/m². On the mesic slope equivalent values were 3000 and 17000 seeds/m² while in the ravine figures were 8500 and 27000 seeds/m². The strong responses of ephemerals and seed bank to the ENSO‐driven wet year of 1997 demonstrate the importance of this phenomenon in replenishing, recovering and probably maintaining ephemeral vegetation in this region.     

Seed size,establishment sites and species coexistence in a Chilean rain forest

Abstract. Seedling densities on the forest floor and on elevated microsites (logs and stumps) were compared for eight woody species in a temperate rain forest in southern Chile. Degree of association with elevated microsites varied significantly between species, showed no systematic relationship with reported shade tolerance, but was significantly negatively correlated with seed mass. Large-seeded Podocarpus nubigena established preferentially on undisturbed forest floor sites, whereas seedlings of small-seeded species such as Nothofagus nitida and Laurelia philippiana were found mainly on fallen logs and stumps. The abundance of large seedlings and saplings of N. nitida on logs/stumps, and the growth forms of canopy trees, confirm that recruitment of this species occurs mainly on decaying wood. The relationship between seed size and microsite preferences may be caused by effects of seed size on (1) ability to establish in forest floor litter and (2) retention of seeds on logs. Seedling occupancy of logs and stumps varied with state of decay. Few seedlings of any species were present on logs in the early stages of decay. N. nitida established earlier than the other species, attaining maximum abundance on wood in the middle decay classes. Species richness and overall seedling abundance were highest on wood in advanced stages of decay. Seed size differences are suggested as a determinant of differential utilization of forest floor heterogeneity, and hence of plant species coexistence.     

Stand dynamics of the shade-tolerant conifers Podocarpus nubigena and Saxegothaea conspicua in Chilean temperate rain forest

Abstract. Population structures were used to infer regeneration patterns of the conifers Podocarpus nubigena and Saxegothaea conspicua and associated angiosperm tree species in six stands in the temperate rainforests of the coastal ranges of southern Chile. On poorly-drained, nutrient-poor and high altitude sites, where most of the associated species were fairly shade-intolerant and light-crowned, dense all-aged populations and the presence of numerous saplings beneath the canopy suggested continuous regeneration of both P. nubigena and S. conspicua. On more favourable sites, where several of the associated angiosperms were highly shade-tolerant and densecrowned, both conifers, particularly P. nubigena, were less abundant, and their regeneration from seed appeared to be sporadic. S. conspicua's great longevity (≥ 750 yr) and frequent vegetative reproduction are probably crucial to its persistence in competition with shade-tolerant broadleaved species in undisturbed stands on favourable sites. Results of this study, and a review of related literature on southern temperate forests are not consistent with the hypothesis that heavily-shaded, infrequently-disturbed habitats are an evolutionary refuge for conifers. Sites likely to have high leaf area indices and infrequent disturbance are better exploited by shade-tolerant angiosperms.     

Persistent soil seed bank and standing vegetation at a high alpine site in the central Chilean Andes

The persistent soil seed bank (viable seeds >1 year) and standing vegetation were investigated in the upper alpine belt (3250 m) in the Andes of central Chile, 33° S. Nine species (eight in standing vegetation) were found in a total persistent seed bank of 899 seeds m⁻². Seven additional species were represented by physically intact, non-viable seeds. Over 90% of the persistent seed bank was concentrated in Montiopsis sericea (Portulacaceae), Pozoa coriacea (Umbelliferae), Phacelia secunda (Hydrophyllaceae) and Oxalis compacta (Oxalidaceae). Examination of the seed/cover ratio revealed different propensities for persistent seed bank formation among species, and annuals formed persistent seed banks more frequently than perennial species. Abundance in the standing vegetation had predictive value for abundance in the persistent seed bank only when non-persistent seed bank species in the standing vegetation were discarded from the analysis. At the local scale, species diversity in the persistent seed bank and standing vegetation were correlated, but compositional similarity was low. Secondary down-slope dispersal promoted by frost heaving in combination with runoff, and life-form correlates are discussed as possible factors accounting for poor correspondence between the persistent seed bank and the standing vegetation at a local scale. The high Andean seed bank is similar to or larger than that reported for two Arctic tundra sites, but smaller than for a northern hemisphere subalpine site. If seed bank size is considered in relation to plant cover, the Andean seed bank greatly exceeds that of one Arctic site. Our study constitutes the first demonstration of a sizable persistent seed bank at an alpine site in the South American Andes and in southern hemisphere temperate mountains in general. Received: 26 May 1998 / Accepted: 21 November 1998     

The role of the permanent soil seed bank in early stages of a post-agricultural succession in the Inland Pampa,Argentina

We studied the soil seed bank composition in four old fields of different ages, after abandonment from agriculture. Complete seed bank composition was assessed by direct seed separation from soil samples and identification to species. Most species found in the seed bank were not important in the present seral communities. Seed of the species that dominated the early succession were generally not found. Additionally, there were very few propagules rather than on the germination of in situ propagules. We suggest that pampean grasses evolved under that the course of post-agricultural succession will depend strongly on the pattern of arrival of exogenous propagules rather than the germination of in situ propagules. We suggest that pampean grasses evolved under disturbances of low intensity and/or a disturbance regime dominated by small gaps, in which open areas could be rapidly colonized from the edges and/or by remnant vegetative propagules. The changes produced by the introduction of agriculture triggered the invasion by exotic species adapted to the new disturbance regime.     

Organisation of the soil seed bank in mixed chaparral

Seed densities, estimated from germinations under glasshouse conditions, were determined for fire-treated and control soils sampled from north and south facing stands of mixedAdenostoma fasciculatum-Ceanothus greggii chaparral that last burned 10, 17, 36, 62 and 86 years before 1987. A total of 53 species was recorded. Total germinable seed density was not influenced by aspect nor by an experimental fire treatment, however, densities varied significantly, but not predictably, over the time since fire sequence. Species diversity was significantly lower in the 86 year old stand, but neither diversity or richness was influenced by the fire treatment. Soils from north facing slopes supported a higher species diversity overall, and seed densities of six species were significantly higher on north than on south slopes.We identify two functional groups in the seed bank on the basis of the direct effects of fire on seed densities: a fire-dependent group, comprising 6 species and about 21% of the total seed bank, in which germinable densities increased after the fire treatment, and a fire-independent group, made up of 11 species and almost two-thirds of the seed bank. The fire-independent group is further separated into two: a shrub-centred sub-group (Crassula erecta, Filago californica, Pterostegia drymarioides, Streptanthus heterophyllus andCryptantha intermedia) maintained larger seed banks in older stands and was associated with shrub cover; and an opportunistic sub-group (Camissonia hirtella, Gnaphalium chilense, Mimulus pilosus andSenecio vulgaris) tended to have larger seed banks in younger stands and was associated with canopy gaps.Overall, the soil seed assemblage from this mixed chaparral does not appear to be tightly coupled to fire-generated opportunities for recruitment. The reason for this is that the fire-dependent group represents only 6 species. In addition, among the fire-independent species, the shrub-centred group of 5 annuals made up almost 40% of total soil seed density. This group shows ecological and taxonomic affinities with desert annuals.     

The impact of flooding regime on the soil seed bank of flood‐meadows

Abstract. We assessed the significance of flooding for the floristic composition of seed banks in flood‐meadows of the northern valley of the Upper Rhine. We compared three hydrological compartments of the alluvial plain, consisting of the regularly flooded land between the river and low summer dykes (functional flood‐plain), the occasionally flooded land between summer dykes and high winter dykes (hybrid floodplain) and the land behind the winter dykes, which is now only submerged by ascending groundwater (fossil flood‐plain). Due to their different flooding regime, the three compartments should differ with respect to the prevailing conditions of diaspore input. The seed density of soil samples increased with the duration of flooding in the three compartments, while species richness and the proportion of species not occurring in the vegetation was constant. The increase in seed density can be largely attributed to an increase of disturbance indicators, which are present in the above‐ground vegetation and capable of forming a long‐term persistent seed bank. No effects of flooding on the composition of seed banks in the three flood‐plain compartments were found. The differences in seed bank composition can be largely explained by corresponding differences in above‐ground vegetation and former and present‐day meadow management. Seeds of species absent from above‐ground vegetation can be attributed to the local species pool present in the immediate vicinity of the study plots. We discuss consequences of the results for the restoration of species‐rich flood‐plain meadows.     

Assessing soil seed bank persistence in flood‐meadows: The search for reliable traits

Abstract. To assess seed bank persistence of target species in endangered flood‐meadows (alliances Cnidion and Molinion), we investigated established vegetation and soil seed bank of 46 plots for 3 yr and 2 yr, respectively. As traits of seed persistence we calculated various continuous indices that refer to the frequency and abundance of species in above‐ground vegetation and at different soil depths. Furthermore, we tested the significance and soundness of easily observed traits such as maximum seed density per plot and seed attributes (mass, size and shape) as predictors of soil seed bank features. In linear regression, SAI, the seed accumulation index, showed the best agreement (R²= 0.64) with the seed longevity index that was derived from the database by Thompson et al. (1997) for a set of 115 species. The second best predictor (R²= 0.39) of the seed longevity index was maximum seed density per plot in the lower soil layer (5–10 cm). Depth distribution indices and seed attributes showed weaker but still significant relations. The dynamic character of flood‐meadows was reflected by a large proportion of species with a strong tendency to accumulate seeds in the soil relative to their importance in above‐ground vegetation. Most of these species have a ruderal strategy, exploiting gaps after flood disturbances, while the dominants of flood‐meadows tended to have short‐lived seed banks. Compared to other grassland types, a relatively large proportion of rare and endangered target species can be expected to form long‐term persistent seed banks. However, only under marginal conditions that facilitate seed survival in the soil (e.g. fallow) are these persistent seed banks likely to contribute to restoration. We conclude that easily observed traits of persistence such as seed weight, size and shape do not meet the accuracy needed in scientific and practical applications. Thus, there is a crucial demand for further seed bank studies in poorly investigated habitats and of rare species.     

Role of soil seed bank along a disturbance gradient in an alpine meadow on the Tibet plateau

We examined the role of the soil seed bank along a grazing disturbance gradient and its relationship with the vegetation of alpine meadows on the Tibet plateau, and discussed the implications for restoration. The seed bank had a high potential for restoration of species-rich vegetation; 62 species were identified in the vegetation and 87 in the seed bank, 39 species being common to both. Mean seed density was 3069–6105 viable seeds m⁻². The density of buried seeds increased significantly with increasing disturbance, indicating that restoration of disturbed areas is not seed limited. Seed density and species richness decreased with depth. The proportion of perennial species decreased with decrease in disturbance both in seed bank and in vegetation. A large portion of species with persistent seeds in the disturbed areas indicate that this seed type can be regarded a strategy of adaptation to current disturbances. Detrended correspondence analysis (DCA) showed significant differences of species composition between seed bank and vegetation, except for the seriously disturbed site. Our results suggest that the establishment of new species in severely disturbed areas is more dependent on the seed bank. By contrast, the restoration in less-disturbed and mature meadows does not rely on seed banks, and the establishment of the vegetation in these communities is more likely to rely on seed dispersal from the standing vegetation and on species with vegetative reproduction.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

The relationship between the soil seed bank and above-ground vegetation of a coastal barrier island

Abstract. The germinable soil seed bank is described from a coastal barrier island off the northwest coast of Florida, USA. Soil samples collected from seven vegetation types, recently deposited dredge spoil and unvegetated areas in autumn 1990 and spring 1991 were placed out in greenhouse trays. 110 taxa germinated from the samples with the largest number (41) being C₃ perennial dicots. The largest number of taxa germinated from dry (57) and wet (54) swales, the fewest (one species: Heterotheca subaxillaris) from strand. Similarity of seed bank densities to above-ground species cover was low (Jaccard's Index = 0.36), not different between vegetation types, but higher in the autumn than in the following spring. Compositional gradients in the seed bank and above-ground vegetation determined using DCA ordination were highly correlated and related to distance from mean high water, and plot elevation. At the landscape scale, the seed bank provided an equally clear delineation of vegetation types to that based upon the above-ground vegetation. The seed bank of low disturbance, late-succession vegetation types (wooded dunes, swales, marshes) was well developed (high species richness, emergent density, and percentage annual species) with the exception that the large-seeded woody species (i.e. Quercus spp.) were absent from the wooded dune seed bank. By contrast, a poorly developed and transient seed bank occurred in more frequently disturbed (extensive sand movement, salt spray), early successional dredge spoil, unvegetated areas and strand. These contrasts support a general pattern of increasing seed bank development and a persistent rather than transient seed bank with decreasing disturbance frequency, increasing time since disturbance and successional maturity.     

Seed bank versus seed rain in the regeneration of a tropical pioneer tree

Summary We used the tropical pioneer tree, Cecropia obtusifolia to evaluate the relative importance of different sources of seeds in the regeneration of species that depend on ephemeral sites. We studied seed production in a population established in a 5 ha plot, and dispersal, dormancy and seed predation in two recent treefall gaps (<1 year-old), two building or successional forest patches (10–15 since disturbed), and two mature forest patches (>35 years since disturbed) for a one year period at Los Tuxtlas (Mexico). Flowers and fruits were counted at monthly intervals. Annual fecundity per tree ranged from 1.4×10⁴ to 1.4×10⁷ seeds. Seeds were continuously available on the trees and on the ground. Average annual seed rain per m² (as measured by 0.5×0.5 m seed traps) varied from 184 to 1925 among the six sites. Distance to nearest seed source and patch type explained more than 60% of the seed rain variation among sites. Soil seed density, estimated by counting seeds from ten samples (78.5 cm²×10 cm deep) collected from each site in October and January, ranged among the six sites from 269 to 4485 seeds per m² in January and from 204 to 5073 in October. Soil seed viabilities were much lower (17.1% in October and 5.1% in January) than those of rain seeds (48.26%). Annual survivorships of 2.2% were estimated for seeds artificially sown on the soil surface of a gap and a mature patch, and 3.75% in a building patch. In two other experiments seed removal rates ranged from 27% to 98% in 4 days. Removal rates were significantly higher in gap and mature patches than in building patches. Ants (Paratrechina vividula) and grasshopper nymphs (Hygronemobius. sp.) were the main predators. We draw three main conclusions from our data: (1) Pathogens and predators determine low survivorship of C. obtusifolia's seeds in the soil and a rapid turnover rate (1.07 to 1.02 years) of its seed bank; (2) a continuous and copious seed production and an abundant and extensive seed rain replenish the soil seed pool in patches with different disturbance ages at least up to 86 m from nearest source; (3) more than 90% of the seeds contributing to C. obtusifolia seedling recruitment in gaps are less than one year-old. We discuss our results in the context of previous similar studies for tropical forests.     

Secondary succession in disturbed Pinus-Quercus forests in the highlands of Chiapas,Mexico

We present floristic and structural data on serai plant communities (Old-Field, Grassland, Shrubland, and Early Successional, Mid-Successional, and Mature Forest) resulting from the current land use pattern in the Pinus-Quercus forests in the highlands of Chiapas, Mexico. The number of species ranged from 24 in the Old-Field stage up to 100 in the Mature Forest, and the shrub layer was richest in all the forested stands. An almost complete floristic replacement exists between the open and the forested communities. Quercus and Pinus dominate the canopy of old-growth stands, but their recruitment does not occur or is very low in the shade. Vigorous seedlings and saplings of these species are found only in the open serai stages or in forest gaps. Demographic and phytosociological data are used to indicate the successional role of dominant species in the forested stages. Some implications of the regional land use patterns for conservation and succession are discussed.     

Gene expression in the soybean seed axis during germination and early seedling growth

Copy-DNA clones have been obtained that distinguish eight messenger mRNAs, moderately abundant in the axes of the germinating soybean (Glycine max (L.) Merr.) seedling. These clones have been used to characterize the size of the mRNAs and to anlyze the accumulation of the mRNAs at different time points and in different parts of the axis during germination and early seedling growth. Three of the mRNAs accumulate to a substantial level by 9 h, a time point before either the beginning of growth or the accumulation of polyribosomes. Four other mRNAs reach a substantial level only at 24 h, a period when rapid seedling growth is occurring. Those mRNAs whose accumulation begins at 24 h were found only in the top (hypocotyl) half of the 24-h seedlings, while the remaining mRNAs were present also in the bottom half of the seedlings in different amounts. By 44 h, the bottom 0.5 cm of the seedlings, i.e., the region of meristematic growth, had little or none of the mRNAs, with the exception of one mRNA. These temporal and spatial observations indicate that many of the mRNAs are not involved simply in the general maintenance of ongoing cell proliferation, but that they may be related to differentiation during early seedling formation. Further, the early accumulating mRNAs may be functioning in regulating the onset of seedling growth.Abbreviations cDNA copy DNA - poly(A)⁺RNA polyadenylated RNA     

Species associations and nurse plant effects in patches of high-Andean vegetation

Abstract. In high-elevation communities of the southern Andes, plant cover is low due to severe environmental conditions and vegetation occurs mostly as isolated small (< 1 m²⁾ patches. Most patches are dominated by flat cushion plants. We evaluated patterns of plant species co-occurrence and species affinity for patches with and without cushion plants and different species richness. We mapped and recorded species composition of patches occurring within two 20 m × 20 m plots at the NE slope of Cerro Chall-Huaco, Nahuel Huapi National Park, Argentina. In these plots, we identified 32 and 24 plant species, and a maximum of 15 and 12 species per patch, respectively. The community was characterized by positive associations between species. Patches in which either of the common cushion plants Mulinum leptacanthum and Oreopolus glacialis occurred sustained richer communities than patches in which they were absent. Patches dominated by different cushion plants did not differ in species composition, but species differed in their affinities for patches with different numbers of species. Because richness increased with patch size and patch size with time, differential affinities of plant species suggest that successional changes take place in the patches. Some small herbaceous species appear to act as late colonizers, mostly restricted to species-rich patches. Flat cushion plants are considered ‘nurse plants’; they strongly modify micro-environmental conditions and allow establishment and survival of associated species.     

Relationships among the seed rain,seed bank and vegetation of a Hawaiian forest

Abstract. Hawaiian ecosystems are prone to invasion by alien plant species. I compared the seed rain, seed bank, and vegetation of a native Hawaiian forest to examine the potential role that seed ecology plays in allowing alien species to invade native forest. Absolute cover of seed plants in the forest was 126 %, annual seed rain was 5 713 seeds m^-2 yr^-1, and the mean density of seedlings emerging from the seed bank averaged across four seasons was 1 020/m². The endemic tree Metrosideros polymorpha was the most abundant species in the vegetation, seed rain and winter seed bank. Overall, native seed plants comprised 95 % of the relative cover in the vegetation and 99 % of the seeds in the seed rain, but alien species comprised 67 % of the seeds in the seed bank. Alien species tended to form persistent seed banks while native species formed transient or pseudo-persistent seed banks. Dominance of the seed bank by alien species with persistent seed banks suggests that aliens are favorably placed to increase in abundance in the vegetation if the forest is disturbed.     

Species attributes in early primary succession on volcanoes

Abstract. We predicted that plants that can establish on volcanic soils with similar disturbance histories will have similar growth characteristics. We tested this prediction by a multivariate analysis of 27 traits of 84 species found six years after an eruption on Mount St. Helens, Washington State, USA, and Mount Usu, Hokkaido, Japan. These traits include vegetative, life-history, phenological and seed-biology characteristics. Cluster analysis revealed five species groups: annual herbs, perennial forbs, graminoids, shrubs and trees. Each group has distinct vegetative, life-history, and seed-biology traits. Except for shrubs, which were lacking on Mount Usu, both floras were well represented in each group. On intensely disturbed sites on both volcanoes, perennial forbs, whose development is dependent primarily on well-developed below-ground organs and wind-dispersal, expanded their cover more rapidly than did graminoids. These graminoids generally produce gravity-dispersed seeds and have close-set rhizomes and/or shoots. These results suggest that species that can establish during the early stages of succession on each volcano have similar vegetative, life-history, and seed-biology traits.     

Restoration of high species density in calcareous grassland: the role of seed rain and soil seed bank

Abstract. A small-scale restoration experiment concerning a species-rich calcareous grassland was conducted over a 7-yr period. 20 yr of abandonment (1971-1990) resulted in both a strong reduction of the number of species and an absolute dominance of the shrub Crataegus monogyna and the grass Brachypodium pinnatum - which contributed 98% of the above-ground herb phytomass. The plot (3 m x 6 m) was cleared in September 1990 and thereafter yearly mown in autumn. Small soil seed bank samples were taken immediately after clearing and again in the autumn of 1996. The inner part of the plot showed a reduced soil seed bank compared to the plot underneath the vegetation outside the plot in 1990. Six years later an increased number of seeds in the soil in this inside area could be recorded mainly in the upper soil layer, although the amount was only 40% compared to the outside or control area. Ca. 20% of the species present in the restored vegetation were recorded in seed traps during the main seed dispersal period in 1997. Restoration of high species richness in the plot was a fast process during the first few years: from seven species in 1990 towards 39 three years later. This was significantly faster than the restoration process of the calcareous grassland at the same slope 20 yr earlier. This was probably due to the increased size of the community species pool at the site as a result of appropriate management during the last decades. Relative species number increase was highest at the largest scale during this experiment, viz. at the total plot area. The success of restoration of species-rich, dry grassland vegetation largely depends on the interaction between several conditions, e.g. seed availability, either originating from the seed rain or soil seed bank, favourable germination conditions and a successful development to a mature, seed producing individual. Germination and establishment strongly depend on suitable vegetation structure as a result of sustainable management.     

Altitudinal gradients in seed dynamics of Calluna vulgaris in eastern Scotland

Abstract. Relationships between seed deposition, size of soil seed banks and some of the environmental factors affecting them were assessed for Calluna vulgaris throughout its altitudinal range (150–960 m a.s.l.) in eastern Scotland. Seed rain was assessed using pitfall traps, collected every 5–10 wk for 3 yr; germinability was determined by laboratory incubation. Seed bank size was estimated, once only, by counting seedlings emerging from soil cores kept for 50 wk in a glasshouse. Seed deposition varied annually, was related to parent plant cover but always declined with altitude, falling sharply above 600 m a.s.l. Seed bank size was more closely correlated with the proportion of organic matter in the soil than with the amount of seed rain. Seed bank sizes declined gradually with altitude but did not differ significantly between four altitudinal zones. The mean density of buried seeds was less than half the mean annual seed rain at sites below 300 m a.s.l. but was over 200 × greater than annual seed rain above 800 m, suggesting that seeds buried at high altitudes remain viable for much longer than those at lower altitudes.