Seasonal variation in the seed bank of a Patagonian grassland in relation to grazing and topography

Abstract. The germinable seed bank of Festuca spp., Carex patagonica, and other annuals and perennials in a semiarid Patagonian grassland was analyzed every three months. The effects of grazing, topography and microsites with respect to established grass tussocks on the germinable-seed bank were also analyzed. The total germinable-seed bank was larger in summer after seed rain. At this time of the year most of the seeds were from annuals and perennial dicots. Seeds of the dominant Festuca spp. were the main components of the graminoid seed bank, which was homogeneously distributed in patches of bare soil. In spring, i.e. some months after the seed rain, the germinable-seed bank of most of the species was greatly reduced, while the seed bank of Carex patagonica did not change significantly. At this time of the year, the graminoid seed bank was heterogeneously distributed in space, with the seeds accumulating in wind-protected microsites. In the grazing treatments, the seed bank of the perennial grasses was reduced proportionally to the decrease of the plant cover. On the other hand, the germinable-seed bank of C. patagonica increased with the grazing treatments, in relation to the increase of vegetation cover. Topographical position had two types of effect on the seed bank of perennial grasses. One was the increase of the germinable-seed bank, after the seed rain, on the relatively warmer northeast facing slopes, which was related to a higher seed rain in these places. The other was an increased seed loss on slopes. The different seed-bank strategies observed in this grassland are compared and discussed in relation to strategies described for other grasslands.     

Size traits and site conditions determine changes in seed bank structure caused by grazing exclusion in semiarid annual plant communities

1. Contrasting patterns of change in the seed bank of natural grasslands are frequently found in response to grazing by domestic herbivores. Here, we studied the hypotheses that a) patterns of change in seed bank density and composition in response to grazing depend on spatial variation in resource availability and productivity, and b) that variation among species in patterns of seed bank response to grazing is linked to differences in species size traits (i.e. size of plant, dispersal unit and seed). 2. Effects of sheep grazing exclusion on the seed bank were followed during five years in a semiarid Mediterranean annual plant community in Israel. Seed bank density and composition were measured in autumn, before the rainy season, inside and outside fenced exclosures in four neighboring topographic sites differing in vegetation characteristics, soil resources and primary productivity: Wadi (dry stream terraces, high productive site), Hilltop, South‐ and North‐facing slopes (less productive sites). 3. Topographic sites differed in seed density (range ca 2500–18000 seed m^?2) and in seed bank response to grazing exclusion. Fencing increased seed density by 78, 51 and 18% in the Wadi, South‐ and North‐facing slopes, respectively, but had no effect in the Hilltop. At the species level, grazing exclusion interacted with site conditions in determining species seed bank density, with larger or opposite changes in the high productive Wadi compared to the other less productive sites. 4. Changes in seed bank structure after grazing exclusion were strongly related to species size traits. Grazing exclusion favored species with large size traits in all sites, while seed density of tiny species decreased strongly in the high productive Wadi. Species with medium and small size traits showed lesser or no responses. 5. The size of plants, dispersal units and seeds were strongly correlated to each other, thus confounding the evaluation of the relative importance of each trait in the response of species to grazing and site conditions. We propose that the relative importance of plant size vs seed size in the response to grazing changes with productivity level.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

SPATIAL PATTERNS OF SEED DISPERSAL AND POSTDISPERSAL SEED PREDATION OF CRYPTANTHA FLAVA (BORAGINACEAE)

I examined the spatial patterns of seed dispersal and postdispersal seed predation of the semidesert perennial Cryptantha flava (A. Nels.) Payson (Boraginaceae) at two sites in north-eastern Utah. Most flowers mature only one seed (nutlet) which is permanently retained within a pubescent calyx. The calyx and enclosed seed abscise from the plant as a unit. These dispersal units are effectively dispersed by wind as evidenced by the highly directional seed shadows and the long distances some of them travel (up to 31.3 m). Potential seed predators at the sites include five species of rodents, of which Peromyscus maniculatus is the most common, and two species of ants, Pogonomyrmex occidentalis and an undescribed species of Conomyrma. There were no strong spatial patterns of postdispersal seed predation. More seeds were removed from dishes placed at the bases of fruiting adults than from dishes ≥ 1.0 m away in one of three experiments. More seeds were removed from under shrubs or clumps of grass than in the open in one of four experiments. After 3–4 days, there was a consistent tendency for more seed removal from high density (75 seeds per .25 m²) quadrats than from low density (75 seeds per 6.25 m²) quadrats, but the difference was not always significant. There was a similar nonsignificant difference between high- and low-density quadrats exposed for 21 days. The pubescent calyx greatly discourages seed predation by ants, and probably also reduces predation by rodents. In addition, by increasing the surface area of the dispersal unit, the calyx may facilitate dispersal by wind.     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.     

SPATIAL AND TEMPORAL VARIATION IN THE SEED BANK OF A SEMIARID GRASSLAND

The spatial and temporal variability in the seed bank of a semiarid grassland in Colorado was evaluated using soil cores. Spatial variability in the soil storage of germinable seeds was assessed by sampling two shortgrass plant communities on sites with the same climatic conditions but differing in soil texture. Differences between communities were largely the result of annual plant seeds. Eight sampling dates over two years were used to assess temporal variability, which was more important to the storage of germinable seeds than spatial variability. Differences in the numbers of seeds stored were found between the two sampling years, and seasonally within years. The number of seedlings that emerged from the samples ranged from 122–2,748/m². A poor correspondence was found between the species composition of the plant communities and the storage of germinable seeds; however, the species composition of the seeds produced on the sites tended to have a high similarity with the seedlings that emerged. Most of the species had a transient rather than a persistent presence in the seed bank.     

Flooding, soil seed bank dynamics and vegetation resilience of a hydrologically variable desert floodplain

1. This paper explores soil seed bank composition and its contribution to the vegetation dynamics of a hydrologically variable desert floodplain in central Australia: the Cooper Creek floodplain. We investigated patterns in soil seed bank composition both temporally, in response to flooding (and drying), and spatially, with relation to flood frequency. Correlations between extant vegetation and soil seed bank composition are explored with respect to flooding. 2. A large and diverse germinable soil seed bank was detected comprising predominantly annual monocot and annual forb species. Soil seed bank composition did not change significantly in response to a major flood event but some spatial patterns were detected along a broad flood frequency gradient. Soil seed bank samples from frequently flooded sites had higher total germinable seed abundance and a greater abundance of annual monocots than less frequently flooded sites. In contrast, germinable seeds of perennial species belonging to the Poaceae family were most abundant in soil seed bank samples from rarely flooded sites. 3. Similarity between the composition of the soil seed bank and extant vegetation increased following flooding and was greatest in more frequently flooded areas of the floodplain, reflecting the establishment of annual species. The results indicate that persistent soil seed banks enable vegetation in this arid floodplain to respond to unpredictable patterns of flooding and drying.     

Phenology of Festuca pallescens in relation to topography in north-western Patagonia

Abstract. The phenological changes in populations of Festuca pallescens (St. Yves) Parodi at different topographic positions and exposure along an altitudinal gradient (600 - 1100 m) were investigated during two growing seasons in northwestern Patagonia. Stepwise multiple regression analysis was used to describe the relationship between phenology and environment during the entire growing season. Analysis of variance was also performed at each sample date to detect significant environmental factors influencing phenology at different sites. The sum of maximum air temperatures was identified as the environmental variable best correlated with the seasonal variation of phenological events of Festuca pallescens over the period of two growing seasons, explaining 93.2 % of the total variance. Significant differences between sites were observed at each sample date. Main effects of altitude and topographic position and two-way interactions between altitude and topographic position, and topographic position and exposure were also detected as significant. Phenology was delayed at increased altitude. Differences in phenology between topographic sites at the same altitude were not detected during the entire growing season and were only observed in the reproductive phase. At this time, the phenology was significantly delayed at high topographic positions on the slopes as compared with low and mid positions. At high altitudes in the valley (950 m a. s. 1.), where steep slopes and humid conditions prevail, phenology was delayed on western exposures and low positions. The results adequately summarize and quantify the effect of spatial and temporal environmental variation on the phenological development of Festuca pallescens in northwestern Patagonia.     

Seed dispersal in both space and time is necessary for plant diversity maintenance in fragmented landscapes

Metacommunity theory emphasizes that seed dispersal not only limits but equally maintains plant diversity, though the latter receives little empirical attention. Discerning the temporal and spatial components of seed dispersal and understanding how their interaction shapes fragmented communities and maintains their diversity may be pivotal to further our ecological understanding of spatial and temporal seed dispersal and its implications for landscape‐scale conservation management. To investigate the relative importance of spatial and temporal seed dispersal and their roles in maintaining plant diversity, the herb layer and seed bank of grassland communities were inventoried in 77 sites across abandoned and intact rotational grazing networks in a 100 km² fragmented grassland landscape in the Stockholm archipelago (Baltic Sea, Sweden). Besides analysing alpha‐ and beta‐diversity patterns, nestedness analyses connect deterministic community changes and diversity losses with dispersal‐related life‐history traits and habitat specialization to identify the mechanism driving community changes and maintaining local diversity. The loss of rotational grazing networks caused community diversity declines via non‐random extinctions of spatially and temporally poor dispersers, particularly among grassland specialists. Temporal seed dispersal halted further community disassembly, maintaining diversity in the abandoned grazing networks. Spatial dispersal within the intact grazing networks was found to be an overriding, homogenizing agent conserving diversity in both the herb layer and seed bank. This empirical evidence establishes how spatial and temporal seed dispersal interact to maintain diversity in fragmented landscapes. Poorly connected grasslands appear limited by spatial dispersal, yet are maintained by temporal seed dispersal. In fragmented landscapes where grazing networks are rarely present, temporal rather than spatial seed dispersal may be more important in maintaining species diversity, since effective spatial dispersal may be significantly diminished. The grazing network's efficacy at boosting spatial dispersal and upholding community diversity presents a powerful management tool to conserve local and regional species diversity.     

Composition and seasonal flux of the soil seed bank of species-rich Themeda triandra grasslands in relation to burning history

Abstract. Most species-rich grasslands dominated by Themeda triandra in southeastern Australia have been ungrazed and frequently burnt for decades. The seedling emergence technique was used to determine the size and taxonomic composition of the soil seed bank of five grasslands that had different fire histories (i.e. burnt at 1 yr, 3 yr and > 10 yr intervals) and this was compared to the standing vegetation at each site. A nested sampling design (subplot, plot, site) was used to determine the effect of spatial scale on the patterns observed in both the vegetation and the seed bank. Temporal variation in the seed bank was assessed by repeated soil sampling over a two year period. 61 native and 30 exotic species were recorded in the vegetation. Richness varied more between sites than within sites. Sites were therefore internally homogeneous for species richness. However, no correlation between burning frequency and richness was found. DCA ordination separated the sites into distinct groups, but sites with similar fire history did not necessarily group closely. 60 taxa germinated from the soil seed bank, comprising 32 native and 28 exotic species; 11 species, mostly therophytes, were restricted to the seed bank. The richness of the seed bank was significantly lower than the vegetation at all spatial scales. No correlation between seed bank richness and fire history was found. The seed bank of species-rich grasslands is dominated by a limited number of widespread, highly clumped, annual, native and exotic monocots. Most native hemicryptophytes, and perennials in general, were represented in the soil by a transient seed bank. Only 12 % of study species, all therophytes, were considered to form large, persistent seed banks, the size of which was greater in unburnt grasslands at all times of the year. The distinct floristic patterns observed in the vegetation are less clearly represented in the seed bank. The seed bank represents a floristically distinct (and less variable) component of the vegetation when compared to the standing flora. The size of the long-term soil seed bank suggests that it has little functional importance for many native species and probably contributes little to seedling regeneration processes following disturbance. Altering established fire regimes is likely to only change the composition and small-scale richness of the existing site vegetation and will not re-integrate species previously lost from the vegetation due to past management. It is suggested that the maintenance of vegetation remnants and processes that encompass a range of long-term burning histories will be necessary if the flora is to be conserved in situ. Restoration of degraded grasslands cannot rely on the soil seed bank but rather, will be dependent on the reintroduction of propagules.     

Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

Spatial dynamics of <Emphasis Type="Italic">Ilex aquifolium</Emphasis> populations seed dispersal and seed bank: understanding the first steps of regeneration

The objective of this study was to analyse quantitatively the spatial distribution of holly (Ilex aquifolium L.) seed rain and seed bank, and to detect the relationships between these consecutive processes. We measured seed dispersal by birds and fallen fruits, and also density and viability of seed bank in two Ilex populations in central Spain. Analysis was made distinguishing the following microhabitats: holly woodland, edge of holly woodland, open grassland 10 m and 100 m from the woodland, fleshy fruit shrubs, dry fruit shrubs, and adjacent non-holly woodland. Spatial distribution of dispersed and in-soil seeds was measured by the clumping index. Seed rain and seed bank under holly woodlands were significantly higher than in the other microhabitats. Forest edges and fleshy fruit shrubs were the next microhabitats with the highest seed rain and seed bank density. Interannual and interlocality variations were not significant. The relative importance of the different dispersal methods varied between microhabitats, with a similar support of bird dispersed seeds and fallen fruits within the woodland and a greater influence of cattle dispersal in open areas. Seed spatial aggregation was significant in both dispersed seeds and soil seeds from holly woodlands and the edge of the forest. Aggregation under shrubs, grasslands and the adjacent forests evidenced a general random distribution of holly seeds (only in some cases clumping index was significant). Quantitative differences between seed rain and seed bank are important. Post-dispersal seed predation did not modify seed rain distribution, which was mirrored in the seed bank pattern. These two phases of holly regeneration had a heavy spatial influence, determined by the landscape structure and activity of the dispersal agents, that reflects a differential recruitment potential. Comparisons between both populations suggest that in the southern locality (Robregordo) holly has a weaker capacity to colonize open areas, and a stronger recruitment limitation due to propagule availability.     

Assessment of spatial discordance of primary and effective seed dispersal of European beech (Fagus sylvatica L.) by ecological and genetic methods

Spatial discordance between primary and effective dispersal in plant populations indicates that postdispersal processes erase the seed rain signal in recruitment patterns. Five different models were used to test the spatial concordance of the primary and effective dispersal patterns in a European beech (Fagus sylvatica) population from central Spain. An ecological method was based on classical inverse modelling (SSS), using the number of seed/seedlings as input data. Genetic models were based on direct kernel fitting of mother‐to‐offspring distances estimated by a parentage analysis or were spatially explicit models based on the genotype frequencies of offspring (competing sources model and Moran‐Clark's Model). A fully integrated mixed model was based on inverse modelling, but used the number of genotypes as input data (gene shadow model). The potential sources of error and limitations of each seed dispersal estimation method are discussed. The mean dispersal distances for seeds and saplings estimated with these five methods were higher than those obtained by previous estimations for European beech forests. All the methods show strong discordance between primary and effective dispersal kernel parameters, and for dispersal directionality. While seed rain was released mostly under the canopy, saplings were established far from mother trees. This discordant pattern may be the result of the action of secondary dispersal by animals or density‐dependent effects; that is, the Janzen‐Connell effect.     

Genetic basis and consequences of niche construction: plasticity-induced genetic constraints on the evolution of seed dispersal in Arabidopsis thaliana

Because seed dispersal influences the environment experienced by seeds, that environment can change as dispersal evolves. The evolutionary potential of dispersal can in turn change as dispersal evolves, if its expression of genetic variation depends on the postdispersal environment. We examined whether seed dispersion patterns have a detectable genetic basis (and therefore evolutionary potential) and determined whether that genetic basis changed depending on one postdispersal environmental factor: conspecific density. We grew replicates of 12 ecotypes of Arabidopsis thaliana at high and low density and measured seed dispersion patterns and maternal traits associated with dispersal under controlled conditions. We found density-dependent ecotypic variation for maternal traits that influence dispersal. Significant genetic variation for postdispersal sibling density was detected only when plants were grown at high density, suggesting that if dispersal evolves to result in lower postdispersal densities, the expression of genetic variation for dispersal would be reduced. This dynamic could lead to a plasticity-induced constraint on the evolution of dispersal. The ability of organisms to alter the environment they experience and the ability of that environment to evolve can alter evolutionary dynamics by augmenting or reducing evolutionary potential and thereby facilitating or constraining evolutionary responses to selection.     

The Effects of Habitat on Dispersal Patterns of an Invasive Thistle, Cynara Cardunculus

Dispersal of offspring is a critical step in the spread of invasive species, yet dispersal patterns are seldom well studied, inhibiting effective management and ecological understanding of invasions. Dispersal patterns can be affected by characteristics of the parent plant and by climatic or site characteristics, including community vegetation structure, but these factors have not been studied in the context of plant invasion processes. Cynara cardunculus (L.) is a polycarpic perennial thistle invasive in coastal grasslands in California that produces large numbers of wind-dispersed seeds. This study quantified and compared C. cardunculus dispersal patterns in an exotic grassland (vegetated site) and an agricultural field (non-vegetated site). Seed size variation in C. cardunculus was also quantified within inflorescences, between inflorescences, and between years of production, and compared across dispersal distances. Results indicate that dispersal distance increased dramatically from less than 20 m in the vegetated site to more than 40 m in the non-vegetated site. Plants producing fewer seeds produced heavier seeds and dispersal distance decreased over time, but seed size was not related to dispersal distance, which may serve to spread the risk to seeds or seedlings across the environment. C. cardunculus has great potential for dispersal in open areas such as agricultural fields or disturbed sites, but may be limited in highly structured natural communities. Management of C. cardunculus and other wind-dispersed plant invasions may be improved by prioritizing populations with open or disturbed areas downwind and minimizing the removal of vegetation during dispersal.     

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

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

POPULATION BIOLOGY OF BONAMIA GRANDIFLORA (CONVOLVULACEAE): EFFECTS OF FIRE ON PLANT AND SEED BANK DYNAMICS

The effects of disturbance on reproduction and plant and seed bank dynamics in the perennial herb Bonamia grandiflora were studied by comparing populations in recently burned, mechanically disturbed, and undisturbed habitats in central Florida over a 3-year period. Plant densities, seed production, and the occurrence of herbivory and predispersal seed predation varied considerably between sites and between years, with recently disturbed sites supporting the densest and most dynamic populations. Death of established plants was rare in all sites. In each site, the soil seed bank was several-fold larger than single season seed rains suggesting that B. grandiflora seeds are long-lived and accumulate in the soil. There was no evidence that postdispersal predation or pathogens have any significant influence on the seed bank dynamics. Fire resulted in large increases in stem densities due to both increased clonal stem production and new genet recruitment from seed. Burning also caused significant increases in the percentage of flowers producing seed and the numbers of capsules and seeds per plant. The seed rain was ten to thirty times greater in the burned site relative to adjacent unburned site during the 3 years after burning. However, additions to the seed bank from the postfire seed rain were balanced by equivalent losses due primarily to seed mortality during fire, and to a much lesser extent due to germination and new genet establishment. As a result, the subsequent densities of seeds stored in the soil in these two sites were similar, indicating that fire results in a significant turnover in the seed bank population but no immediate change in its size. These effects on seed bank dynamics, in addition to new genet recruitment, suggest that periodic fires may play an important role in the maintenance of genetic variability as well as the size of these populations.     

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.     

Limits and effects of invasion by the nonindigenous wetland plant <Emphasis Type="Italic">Lythrum salicaria</Emphasis> (purple loosestrife): a seed bank analysis

We used seed bank analyses to investigate the role of dispersal in limiting invasion by Eurasian Lythrum salicaria within and among North American wetlands, and the changes in seed bank diversity associated with this invader. We compared the number and species composition of seedlings emerging from soil sampled in 11 uninvaded wetlands and paired uninvaded and invaded sites within 10 invaded wetlands under both seedling competition and noncompetitive conditions. Almost no L. salicaria emerged in samples from uninvaded wetlands, indicating dispersal limitation despite prodigious seed production in nearby wetlands. However L. salicaria emerged in all samples from uninvaded sites in invaded wetlands, suggesting environmental limits on establishment within invaded wetlands. Conditions that provided opportunities for seedlings to compete reduced survival of Typha spp. but not L. salicaria seedlings. However, this was due to species-specific differences in post-emergence mortality rather than response to competition. Competition did reduce seedling mass, but this effect did not differ among species. Species richness of emerging seedlings was lower for invaded than uninvaded wetlands. Lower seed bank richness may be a cause or consequence of L. salicaria invasion. Efforts to reduce seed dispersal to uninvaded wetlands would likely slow the spread of this invader.     

Scale-dependent diversity effects of seed dispersal by a wild herbivore in fragmented grasslands

Dispersal limitation between habitat fragments is a known driver of landscape-scale biodiversity loss. In Europe, agricultural intensification during the twentieth century resulted in losses of both grassland habitat and traditional grassland seed dispersal vectors such as livestock. During the same period, populations of large wild herbivores have increased in the landscape. Usually studied in woodland ecosystems, these animals are found to disperse seeds from grasslands and other open habitats. We studied endozoochorous seed dispersal by roe deer (Capreolus capreolus) in fragmented grasslands and grassland remnants, comparing dispersed subcommunities of plant species to those in the established vegetation and the seed bank. A total of 652 seedlings of 67 species emerged from 219 samples of roe deer dung. This included many grassland species, and several local grassland specialists. Dispersal had potentially different effects on diversity at different spatial scales. Almost all sites received seeds of species not observed in the vegetation or seed bank at that site, suggesting that local diversity might not be dispersal limited. This pattern was less evident at the landscape scale, where fewer new species were introduced. Nonetheless, long-distance dispersal by large wild herbivores might still provide connectivity between fragmented habitats within a landscape in the areas in which they are active. Finally, as only a subset of the available species were found to disperse in space as well as time, the danger of future biodiversity loss might still exist in many isolated grassland habitats.