Seed bank spatial pattern in a temperate secondary forest

Abstract. Seed bank spatial pattern was studied in a secondary forest dominated by Fagus sylvatica and Betula celtiberica in the Urkiola Natural Park (N Spain). Soil samples were taken every 2 m in a regular grid (196 points) and divided into two fractions (0–3 cm and 3–10 cm deep). The viable seed bank was studied by monitoring seedling emergence for ten months. The effect of different factors on seed bank composition and patterning was analysed using constrained ordination as a hypothesis testing tool. Furthermore, the existence of spatial autocorrelation was evaluated by geostatistical analysis. Seed density was high, 7057 seed.m^?2, with a few species dominating. Species composition in the various layers were significantly correlated. The seed bank showed significant spatial structure, which was partially explainable by the spatial structure of the canopy and understorey vegetation. Spatial clumping from 0–8 m was observed in seed bank density and composition, mainly due to the pattern of two abundant taxa Juncus effusus and Ericaceae. The Ericaceae seed bank was related to the spatial distribution of dead stumps of Erica arborea. J. effusus was not present in the above‐ground vegetation, which indicates that its seed bank was formed in the past. As expected, the seed bank of this forest reflects its history, which is characterized by complex man‐induced perturbations. The seed bank appears to be structured as a consequence of contrasting driving forces such as canopy structure, understorey composition and structural and microhabitat features.     

Seed bank offers potential for active restoration of mountain meadows

The nitrogen‐fixing legume Lupinus polyphyllus invaded semi‐natural mountainous grasslands across Europe during the last decades. This invasion resulted in degraded habitats through changes in the structure and function of the mountain meadow vegetation. In our study, we analyzed (1) the effects of increasing cover of L. polyphyllus on the seed bank of mountain meadows, and (2) the potential of the seed bank of these stands for active restoration of mountain meadows in terms of species composition and number. We conducted a seed bank analysis on 84 plots with increasing cover of L. polyphyllus in three mountain‐meadow types of the Rhön Biosphere Reserve, Germany. Seedlings from 119 species germinated from the seed bank samples, including 17 Red List species but only a few seedlings of L. polyphyllus. The species composition of the seed bank matched distinct patterns of the three meadow types, but differed from the species composition of the current aboveground vegetation in a nonmetric multidimensional scaling ordination. While the influence of L. polyphyllus on the current vegetation was visible, no effects on the seed bank were apparent. L. polyphyllus had no influence on total seed density, seed density of typical mountain‐meadow species, or species numbers in the seed bank. Only the seeds of the Red List species were significantly related to the cover of L. polyphyllus. We conclude that the seed bank offers potential for active restoration of species‐rich mountain meadows, but species absent from the seed bank have to be added by other measures.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Impact of Formica exsecta Nyl. on seed bank and vegetation patterns in a subalpine grassland ecosystem

The mound building ant Formica exsecta Nyl. is widely distributed in grassland ecosystems of the Central European Alps. We studied the impact of these ants on seed bank and vegetation patterns in a 11 ha subalpine grassland, where we counted over 700 active ant mounds. The mounds showed a distinct spatial distribution with most of them being located in tall‐grass, which was rarely visited by ungulates (red deer; Cervus elaphus L.). Heavily grazed short‐grass, in contrast, seemed to be completely avoided by ants as only few mounds were found in this vegetation type. The species composition of the ant mound and grassland seed banks was quite similar, i.e. from 15 common plant species 12 were found in both seed bank types. We found the same proportions of myrmecochorous seeds in ant mound and grassland soil samples. In contrast, the number of seeds was 15 times higher in mound compared with the grassland soil samples. Also, the vegetation growing on ant mounds significantly differed from the vegetation outside the mounds: graminoids dominated on ant mounds, herbaceous and myrmecochorous species in the grassland vegetation. We found significant continuous changes in vegetation composition on gradients from the ant mound centre to 1 m away from the mound edge. Overall, F. exsecta was found to have a considerable impact on seed bank and vegetation patterns in the grassland ecosystem studied. These insects not only altered grassland characteristics in the close surrounding of their mounds, but also seem to affect the entire ecosystem including, for example, the spatial use of the grassland by red deer.     

Restoration of species‐rich flood‐plain meadows from abandoned arable fields in NE France

Abstract. Attempts to restore species‐rich flood‐plain meadows from abandoned arable fields in the valley of the river Meuse, NE France, were studied. The study area was sown with a commercial seed mixture, composed of Phleumpratense, Festuca pratensis, Lolium perenne and Trifolium repens. The above‐ground vegetation in the study area 1, 2 and 3 yr after restoration was compared to (1) the vegetation present during the previous 5‐yr fallow stage and (2) target flood‐plain meadows. Before restoration, the above‐ground fallow vegetation was dominated by ruderal and annual species, while only very few meadow species were present. Sowing led to tall, dense vegetation, mainly dominated by the sown species. Ruderal and annual species had decreased 3 yr after restoration, but target species were still poorly represented. Species richness was significantly lower in the sown site than in the semi‐natural target meadows and the vegetation had a different composition. Analysis of the soil seed bank of the restored meadow showed that only a few meadow species were present and that it was dominated by a few ruderal species. Three years after sowing, the vegetation of our experimental site is moving slowly towards the target communities but impoverished seed sources seem to limit the success of this restoration operation and will lead to under‐saturated communities.     

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.     

The role of the soil seed bank in vegetation recovery on an oceanic island severely damaged by introduced goats

Question: Are the seed banks of an isolated subtropical oceanic island capable of naturally regenerating vegetation either with species of the historical forest community or with the existing grassland community after severe damage to the vegetation by goats? Location: Nakoudojima Island, Bonin Archipelago (Ogasawara Shoto), Japan. Methods: Soil samples were collected at 0–5 cm and 5–10 cm depths from seven plots in forests, grasslands, artificially matted areas and bare land. Soil seed banks were assessed using the seedling emergence method followed by the hand‐sorting of ungerminated seeds. We determined the size and composition of the seed banks in upper soil layers of plots and compared the seed banks to the standing vegetation. Results: A total of 12 220 seedlings belonging to 42 species from 20 families germinated. Total mean seed density (0–5 cm depth) was low in all plots within forest, grassland, and heavily degraded vegetation types (34.7 ± 8.6 to 693.5 ± 123.6, 58.6 ± 7.8 to 107.1 ± 10.0, and 1.1 ± 0.5 to 7.2 ± 2.3 seeds/m², respectively). Forbs and graminoids dominated the seed banks of grassland and forest plots including Cyperus brevifolius, Gnaphalium pensylvanicum, Oxalis corniculata and Solanum nigrum, and these alien species comprised 90% of the density of the seed bank. There was little correlation between seed banks and standing vegetation of the island (Sørensen similarity coefficient values 0.26 to 0.45). Conclusions: If natural regeneration occurs from the seed bank of the island, future vegetation will not move toward the original forest community, because the seed bank is dominated by non‐native herbaceous grassland species. Though isolated, a few forest remnants with low species richness could be an important source for the natural re‐establishment of forest on the island; however, seed availability may be limited by either poor dispersal or pollination so that woody species will probably recover very slowly on this goat‐impacted island.     

Characteristics of the soil seed bank in Mediterranean temporary ponds and its role in ecosystem dynamics

Species in temporary ponds overcome periods of unfavorable weather conditions by building up a large seed bank. With this strategy, the species diversity of ponds is preserved and information on their dynamics and structure is retained. Little is known about the characteristics, spatial patterns and role in the vegetation dynamics of the soil seed banks of Mediterranean temporary ponds, which are regarded as priority habitats under protection. We studied two sites of western Crete: Omalos, a mountain plateau at 1,060 m a.s.l. and Elafonisos, located near the coast at 60 m a.s.l. The seed bank was surveyed along transects using the germination method. Aboveground vegetation was measured on quadrats along the same transects. Canonical Correspondence Analysis (CCA) was run to define the zonation patterns. High density and species richness were recorded in both sites, with an average of 75,662 seeds/m² found in Omalos and 22,941 seeds/m² in Elafonisos. The community composition of both sites was remarkably different but in both locations perennial species were inconspicuous while annuals, prevailed in the seed banks. An important array of protected or rare species as well as several others which were absent from the vegetation were hosted in the soil seed banks, thereby rendering a low similarity between their composition. Soil seed banks in these ecosystems indicated a spatial heterogeneity that mirrored the aboveground vegetation distribution, sorted along the moisture gradient by their tolerance to flooding. Soil seed banks play a key role in the vegetation recovery after summer drought. The acts of preserving the soil seed bank and ensuring a transient flooding regime are essential to protect the unique vegetation communities of Mediterranean temporary ponds.     

Restoration of wooded meadows ‐ a comparative analysis along a chronosequence on Öland (Sweden)

Abstract. Wooded meadows on the Baltic Island of Öland result from traditional agricultural management over centuries which has led to a species‐rich vegetation with high species diversity. Today, nearly all of these meadows have been abandoned and became rapidly overgrown by deciduous shrub and tree species forming a closed canopy which resulted in a rapid and strong decrease in species numbers of the herb layer. Recent efforts aim to restore overgrown wooded meadows by cutting single shrubs and trees to open the canopy. However, the effects of abandonment as well as of any restoration management in wooded meadows have rarely been documented until now. Mechanisms driving succession after restoration such as the dispersal potential of the respective species over time and space have not been analysed yet. Therefore, a chronosequence was studied which included a traditionally managed wooded meadow, an overgrown meadow which has been abandoned for more than 100 yr and a meadow which was restored 36 yr ago by cutting and is now grazed. We analysed the soil seed bank of the 3 meadows in comparison with the established vegetation and endozoochorous seed dispersal by cattle and sheep. After abandonment 87% of the typical grassland species vanished from the established vegetation and were replaced by species characteristic of woodland and disturbed grassland communities. The mean number of species decreased from 52 species per plot (4 m²) to 18 species. Mean species number and number of seeds in the seed bank declined significantly from the traditionally managed to the overgrown meadow. Most of the grassland species were assigned to a transient seed bank type while only 1/3 could be classified as having a short‐term persistent seed bank. Thus, restoration of wooded meadows cannot rely on the soil seed bank. Endozoochorous seed dispersal by cattle and sheep was shown for 15% of the species with seed densities per 100 g air dried dung of 737 and 767, respectively. Movement of animals between ancient and restored wooded meadows is recommended since many of the species only occurred in low densities and therefore, will probably not be found in the dung samples.     

Distribution of Herbaceous Species in the Soil Seed Bank of a Flood Seasonality Area,Northern Pantanal,Brazil

To identify factors affecting the spatial distribution of soil seed banks of herbaceous species in the Pantanal floodplain, Brazil, the aims of this study were: to characterize the seed bank in terms of the abundance, richness and composition of germinated seeds; to relate these characteristics to flood duration, elevation, chemical and physical properties of the soil and to examine the seed bank’s spatial pattern. Soil samples were collected at 14 points and were then placed in a greenhouse to allow germination to occur. Each sample point had the flood level monitored, the elevation measured and the soil properties identified. A total of 1710 seedlings from 26 species were recorded, of which Echinodorus tenelus was the most abundant (24.9%). Cyperaceae and Poaceae were the richest floristic families (5 species each), with Alismataceae the most represented in terms of number of individuals (36% of the total). Duration of flooding plays an important role in determining Pantanal soil seed banks. The flood pulse influences the abundance (r = 0.79; P = 0.006; partial), richness (r = 0.61; P = 0.02; partial) and composition of the soil seed bank (Pillai trace = 0.552; P = 0.027), carrying the seeds to areas where the duration of flood is longer. Except for aluminium, the soil characteristics expressed by the first principal component of PCA exert indirect positive effects on the seed bank. This is because this component was correlated with the duration of inundation (r = 0.76). Elevation and the toxic effect of aluminium do not vary sufficiently to be able to influence seed bank characteristics. The correlograms show that soil seed banks have no discernible spatial pattern, even though most species are dispersed hydrochorically. This suggests that, at the scale of the study, the tendency for flooding to cause homogeneous dispersion has no influence on seed‐bank spatial structure, because of the complexity of flood‐plain geomorphology. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Association between cacti and nurse perennials in a heterogeneous tropical dry forest in northwestern Mexico

Abstract. The Sierra de La Laguna, Baja California Sur harbors the only tropical dry forest of the Peninsula. An inventory of the perennial plants was carried out at the eastern part of this mountain range. Plant spatial distribution was analyzed considering the species composition and vertical stratification of plant clusters in a study plot of 2 500 m². A plant cluster was defined as a group of at least three individuals with a maximum distance between plants of 1 m. Soil physical-chemical characteristics and soil surface temperatures were compared between shade and sun conditions. The tropical dry forest of Baja California Sur presents a highly aggregated pattern of species distribution. Most of the perennial plants grow clumped, yet not all of the six cactus species recorded were significantly associated with them. Pattern analysis revealed that only Stenocereus thurberii and Ferocactus peninsulae were aggregated and associated to perennial plants. There is no replacement pattern between cacti and perennial plants. Nutrient content, including soil fertility, between shade and sun conditions does not seem to explain the patchy distribution of perennial plants, while the differences in solar radiation between these conditions appear to be restrictive and determinant of this spatial pattern of distribution.     

Spatial patterns of the germinable soil seed bank of coexisting perennial-grass species in grazed shrublands of the Patagonian Monte

We compared the size and spatial pattern of the germinable soil seed bank (GSB) of the three dominant perennial-grass species in the arid Patagonian Monte of Argentina. These species differ in plant functional traits, seed morphology, and in their preference by grazers. During three consecutive years, we sampled the soil seed bank at 1 m intervals and the intercepted diameter of perennial-grass species and shrub patches along 50-m linear transects at three sites with different grazing pressure. Soil samples were incubated at field capacity during 12 weeks and emerged seedlings counted. The largest-seeded and most xerophytic Stipa speciosa did not form a GSB. The other two grass species with more mesophytic traits and smaller seeds than S. speciosa formed different GSB. Poa ligularis formed a smaller and more heterogeneous GSB than Stipa tenuis. Grazing had not effect on GSB size of P. ligularis but increased the proportion of seeds under dense shrub canopies. In contrast, grazing reduced the size and also increased the proportion of the GSB of S. tenuis under shrub canopies. We conclude that the size and the spatial pattern of the GSB of the dominant perennial-grass species in the arid Patagonian Monte were related not only to seed and plant traits but also to the spatial distribution of grass plants and their exposure to grazers.     

Seed bank diversity in mesic grasslands in relation to vegetation type,management and site conditions

Questions: 1. Do different management types (i.e. hay meadow, silage meadow, meadow‐pasture, pasture) have different impact on the size and composition of the seed bank of mesic grassland (Arrhenatheretalia)? 2. How strong is the effect of management on the seed bank in relation to above‐ground vegetation, edaphic factors and land‐use history? 3. Are there differences in C‐S‐R plant strategy types and seed longevity under different management regimes? Location: Lahn‐Dill Highlands in central‐western Germany. Methods: Above‐ground vegetation and the soil seed bank of 63 plots (at 21 sites) in mesic grasslands were studied. Differences between management types in quantitative seed bank traits and functional characteristics were tested by ANOVA. The impact of management, above‐ground vegetation, site conditions and land‐use history on seed bank composition were analysed by partial CCA. Results: Management had no significant impact on species richness and density of the seed bank but significantly influenced their floristic composition and functional characteristics. CCA revealed that even after adjustment for soil chemical parameters and above‐ground vegetation management still had significant impact on seed bank composition. ANOVA revealed that silage meadows contained higher proportions of R‐strategy compared to hay meadows. In contrast, in hay meadows and meadow‐pastures proportions of S‐strategy were higher than in silage meadows. Conclusions: The type of grassland management has little impact on quantitative seed bank traits. Management types with a high degree of disturbance lead to an increase of species following a ruderal strategy in the seed bank. Irrespective of management type only a limited proportion of characteristic grassland species is likely to re‐establish from the seed bank after disappearance from above‐ground vegetation.     

Tree invasion of a montane meadow complex: temporal trends,spatial patterns,and biotic interactions

Questions: Do spatial and temporal patterns of encroachment of Pinus contorta and Abies grandis in a montane meadow suggest strong biotic controls on the invasion process? Location: Forest–meadow mosaic, 1350 m a.s.l., Cascade Range, Oregon, US. Methods: We combined spatial point pattern analysis, population age structures, and a time‐series of stem maps to quantify spatial and temporal patterns of conifer invasion over a 200‐yr period in three plots totaling 4 ha. Results: Trees established during two broad, but distinct periods (late 1800s, then at much greater density in the mid‐1900s). Recent invasion was not correlated with climatic variation. Abies grandis dominated both periods; P. contorta established at lower density, peaking before A. grandis. Spatially, older (≥90 yr) P. contorta were randomly distributed, but older A. grandis were strongly clumped (0.2‐20 m). Younger (<90 yr) stems were positively associated at small distances (both within and between species), but were spatially displaced from older A. grandis, suggesting a temporal shift from facilitation to competition. Establishment during the 1800s resulted in widely scattered P. contorta and clumps of A. grandis that placed most areas of meadow close to seed sources permitting more rapid invasion during the mid‐1900s. Rapid conversion to forest occurred via colonization of larger meadow openings – first by shade‐intolerant P. contorta, then by shade‐tolerant A. grandis– and by direct infilling of smaller openings by A. grandis. Conclusions: In combination, spatial and temporal patterns of establishment suggest an invasion process shaped by biotic interactions, with facilitation promoting expansion of trees into meadows and competition influencing subsequent forest development. Once invasion is initiated, tree species with different life histories and functional traits can interact synergistically to promote rapid conversion of meadow to forest under a broad range of climatic conditions.     

Harvester ants (Messor barbarus) as disturbance agents in Mediterranean grasslands

Questions: 1. Do disturbances by harvester ants (Messor barbarus L.) affect soil properties? 2. Do they alter seed distribution? 3. Do they show a different species composition? 4 Are these changes related to seed size (length and weight)? Location: Mediterranean grasslands in central Spain, near Madrid. Methods: We recorded autumn seed banks and spring vegetation in middens, trunk trails and controls. Soil properties were also measured in middens and controls. The effect of seed weight and length was analysed after transforming data into phylogenetically independent contrasts. Results: Soil under middens is more silty and higher in potassium, organic matter and pH. Seed density and seed bank species richness increased greatly in middens, while vegetation species richness was significantly lower in comparison to control samples. Abundance changes in this disturbance type were positively correlated with seed weight, both in seed bank and vegetation. In contrast, we only detected a slight increase in bare ground on foraging trunk trails, with no clear effects on seed bank and vegetation composition. Conclusions: Midden development is a mechanism that generates heterogeneity which favours the survival of certain large‐seeded species mainly predated by ants in Mediterranean grasslands. This effect may partially neutralize the dominance of small seeded species expected from the seed predation process.     

Plant colonization of a bare peat surface: population changes and spatial patterns

Abstract. Changes in size and spatial arrangement of plant populations established on an initially bare peat surface were described over a period of 5 yr by following plant individuals on a 1-cm grid in an area of 10 m x 25 m. The spatial pattern of populations and association between species was analyzed statistically. The study site was very slowly colonized by 14 perennial plant species. The early successional stage was dominated by Carex rostrata, with a clumped spatial distribution, and the homogeneously distributed Eriophorum vaginatum and Pinus sylvestris. Both the growth in size of populations and changes in their spatial distribution were interpreted as a result of species dispersal ability, tolerance to severity of the substrate and pattern of reproduction.     

Impact of flooding on potential and realised grassland species richness

In order to reduce flood risk, river management policies advise floodplain restoration and the recreation of water retention areas. These measures may also offer opportunities for the restoration of species-rich floodplain habitats through rewetting and the restoration of flood dynamics. The potential to enhance biodiversity in such flood restoration areas is, however, still subject to debate. In this paper we investigate whether flooding along a small altered lowland river can contribute to the potential and realised species richness of semi-natural meadows. We compare the seed bank and vegetation composition of flooded and non-flooded semi-natural meadows and test the hypothesis that flooding contributes to an input of diaspores into the meadow seed banks, thereby promoting seed density and potential species richness. Furthermore we hypothesise that, where habitat conditions are suitable, flooding leads to a higher realised species richness. Results showed that seed densities in flooded meadows were significantly higher than in non-flooded meadows. The seed banks of flooded meadows also contained a higher proportion of exclusively hydrochorous species. However, the seed bank species richness, as well as the species richness realised in the vegetation did not differ significantly between flooded and non-flooded meadows. Finally, the seed bank and standing vegetation of flooded sites showed larger differences in species composition and Ellenberg nitrogen distribution than non-flooded sites. From these results we conclude that, although flooding does contribute to the density and composition of the seed bank, most imported seeds belong to only a few species. Therefore, it is unlikely that flooding substantially enhances the potential species richness. Furthermore, even if new species are imported as seeds into the seed bank, it seems unlikely that they would be able to establish in the standing vegetation. However, it is unclear which factors impede the establishment of imported species in the vegetation. The implications of our findings for flood meadow restoration are discussed.     

Spatial relationships between the standing vegetation and the soil seed bank in a fire-prone encroached dehesa in Central Spain

Postfire vegetation regeneration in many fire-prone ecosystems is soil seed bank dependent. Although vegetation and seed bank may be spatially structured, the role of prefire vegetation patterns and fire in determining postfire vegetation patterns is poorly known. Here, we investigated the spatial patterning of species abundance and richness in the vegetation and seed bank of a Mediterranean encroached dehesa in Central Spain. The seed bank was studied with and without a heat shock simulating a spatially homogeneous fire. Semivariograms and cross-semivariograms showed that species richness in the vegetation was aggregated in patches, mainly of herbs, with highest values corresponding to high herb cover and low tree cover. Species richness in the seed bank was also structured in patches, but the spatial pattern was weak. Seedling density of germinates in the seed bank also showed weak spatial pattern. Heating increased overall germination and species richness, and the intensity of the spatial pattern of species richness, particularly of herbaceous species. However, seed bank density patterns disappeared after heat shock because of increased germination of shrubs without spatial pattern. Our results document that the spatial structure of plant richness in the vegetation may persist after fire due to the spatial patterns of herbaceous species in the seed bank, and that postfire species richness patterns can arise independently of fire intensity patterns. However, the spatial structure of the vegetation after fire can be altered by the feedback between shrub encroachment and an eventual fire because of the ubiquitous germination of shrubs.     

Comparison of soil seed banks of habitats distributed along an altitudinal gradient in northern Iran

In this study we investigated the variations in soil seed banks along an altitudinal gradient in the Alborz mountains, Iran, covering three habitats from lower to upper altitudes: forest, forest-subalpine grassland ecotone and subalpine meadow. In each habitat from 1850 to 2400 m, 20 quadrats were established along four transects, and the above-ground vegetation and the germinable seed banks were determined. Results show that the similarity between seed bank and vegetation was lowest in the ecotone located at intermediate altitudes. Together with the contrasting highest density and species diversity of seeds at these altitudes, the ecotonal role of this habitat was confirmed.We found evidence that lower altitudes could act as storage for seeds of some species growing at higher altitudes; the role of the ecotone was more prominent as a reserve for the meadow plant seeds than the role of the forest as a reserve for seeds of the meadow and ecotone habitats. Soil seed banks, particularly from the ecotone, can be used for restoring vegetation in some degraded sites.     

Nested spatial patterns in seed bank and vegetation of Mediterranean old-fields

A nested sampling design was used to describe the spatial patterns for the species richness and composition in the seed bank and vegetation of three Mediterranean old-fields (1, 7, and 15 yr after the last ploughing). Three scales were examined hierarchically: sampling units within plots of 0.25 m² for the vegetation and of 0.05 m² for the seed bank, 100 m² plots within fields, and fields of 1000 m². In spite of the strong spatial variation among sampling units, species richness and composition of both seed bank and vegetation showed hierarchically structured patterns of heterogeneity, while each old-field was a homogeneous entity. These spatial patterns tended to be partially masked when the data were aggregated at the scale of the plot. Such results stress the use of a nested sampling design for studying variation in species richness and taxonomic composition in both vegetation and seed bank. This design, in combination with CCA, also showed that the vegetation showed a coarser grain than the seed bank, probably in relation to seed clumping.