Canopy effects of the invasive shrub Pyracantha angustifolia on seed bank composition,richness and density in a montane shrubland (Córdoba,Argentina)

Abstract Invasive woody species frequently change the composition of the established vegetation and the properties of the soil under their canopies. Accordingly, invasion may well affect regenerative phases of the community, especially at the seed bank level, likely influencing community restoration. Pyracantha angustifolia (Rosaceae) is an invasive shrub in central Argentina that affects woody recruitment, particularly enhancing the recruitment of other exotic woody species. There is though no information regarding its effect on the soil seed bank within the invaded community. The present study was set up to gain further insight into the canopy effects of P. angustifolia. We aimed to assess whether the invasive shrub affects seed bank composition, richness and seed density as compared with the dominant native shrub Condalia montana (Rhamnaceae), and to relate the observed seed bank patterns with those of the established vegetation. We evaluated the composition of the germinable seed bank and the established vegetation under the canopy of 16 shrubs of P. angustifolia, 16 shrubs of C. montana, and in 16 control plots (10 m²) without shrub cover. The floristic composition of the seed bank differed among canopy treatments. However, seed bank richness did not differ significantly. There was an overall high seed density of exotic species throughout the study site, though exotic forbs showed significantly lower seed densities under the invasive shrub. Pyracantha angustifolia would not promote the incorporation of new species into the seed bank of the invaded community but rather favour the establishment of woody species that do not depend on seed banks. The absence of dominant woody species in the seed bank, the dominance of exotic forbs, and the high similarity between established exotic species and those present in the seed bank may surely affect community restoration following the main disturbances events observed in the region.     

Does invasion by an alien plant species affect the soil seed bank?

Questions: How does invasion affect old‐field seed bank species richness, composition and density? How consistent are these effects across sites? Does the soil seed bank match vegetation structure in old‐fields? Location: Menorca, Balearic Islands, Spain, western Mediterranean basin. Methods: We monitored seed germination in soils from old‐fields that were both uninvaded and invaded (legacy effect) by the annual geophyte Oxalis pes‐caprae. We also added O. pes‐caprae bulbs to uninvaded soils to test O. pes‐caprae interference with seedling emergence (competitive effect). We compared species composition in the seed bank with that of the vegetation. Results: Species richness in the seed bank and in the vegetation was not significantly different between invaded and uninvaded areas. Uninvaded areas did not have larger seed banks than invaded areas. More seedlings, especially of geophytes, emerged when O. pes‐caprae bulbs were added to the soil. Species similarity between invaded and uninvaded areas was higher in the seed bank (74%) than in the vegetation (49%). Differences in species composition were as important as differences among sites. The degree of species similarity between the seed bank and the vegetation was very low (17%). Conclusions: Despite invasion by O. pes‐caprae not affecting species richness, the variation in the seed bank species composition in invaded and uninvaded areas, and the differences between the seed bank and the mature vegetation, highlights that even if the invader could be eradicated the vegetation could not be restored back to the exact composition as found in uninvaded areas.     

Community‐level changes in Australian subalpine vegetation following invasion by the non‐native shrub Cytisus scoparius

Question: What are the changes associated with the recent invasion by the non‐native legume, Cytisus scoparius? Location: Subalpine vegetation (1500 m a.s.l.) in Australia. Methods: We used multivariate techniques and regression analyses to assess vegetation and environmental changes across six study sites. Vegetation and environmental variables were investigated at three different stages of invasion: (1) recent invasion (8–10 yr), (2) mature invasion (15–16 yr) and (3) long‐term invasion (25 yr). Results: Substantial changes in floristic composition and species richness were evident after 15 yr and these changes became more pronounced after 25 yr. Changes due to invasion were associated with a dramatic loss of native species or a reduction in their abundance. No ‘new species’ were evident under invaded stands. Forbs were most affected by the establishment of C. scoparius, although all growth forms responded negatively. Dense canopy shading and an increasingly dense, homogeneous litter layer in the understorey as a result of C. scoparius were strong environmental drivers of vegetation change. Greenhouse studies confirmed the importance of these processes on the germination and growth of two native species. Conclusions: This study highlights the potential for C. scoparius to alter both vegetation and environmental processes in the subalpine region.     

普洱市周边地区4种土地利用类型土壤种子库特征

通过对云南普洱市周边地区次生季风常绿阔叶林、针阔混交林、人工更新形成的针叶林及茶园等4种土地利用类型的野外调查及土壤种子库的萌发实验,探讨其土壤种子库的密度大小、物种丰富度和组成及与地上植被的关系。结果表明:干扰强度与频度不同导致土地利用类型之间土壤种子库密度与物种丰富度存在较大差异,土壤种子库密度大小顺序为:针叶林(248.67±116.86)粒·m-2>针阔混交林(186.00±43.27)粒·m-2>次生季风常绿阔叶林(107.33±16.48)粒·m-2>茶园(51.67±10.17)粒·m-2;茶园土壤种子库物种丰富度要显著低于其他类型。4种土地利用类型土壤种子库生活型组成差异极显著,主要以草本植物组成,以菊科与禾本科占优势;针阔混交林的草本植物种子密度最多,非森林的原生物种是草本植物的主要组成;针叶林外来物种的种子密度要显著高于其他类型,紫茎泽兰(Eupatorium adenophorum)是其主要组成。土壤种子库与地上植被的相似性系数较低,其大小顺序为:次生季风常绿阔叶林(0.175)<针阔混交林(0.176)<针叶林(0.215)。     

Community changes following shrub invasion of grassland

Summary We studied the development of the shrub Baccharis pilularis ssp consanguinea and its effects on herbs of the annual grassland in Northern California. A series of shands of Baccharis was sampled of ages ranging from 1 yr to>9 yr, representing most of the life cycle of the shrub. In each stand we examined shrub biomass, structure and litterfall. We also determined cover and biomass of all herbaceous species and estimated seed production, seed rain and storage of seed in the soil. Abundances of all herbaceous species declined greatly after Baccharis formed a closed canopy at 2–3 yr, and little seed of herbaceous species was either dispersed into shrub stands or stored in the soil. Exclosures suggested that herbivory by small mammals in the closed shrub stands may be important in reducing the abundance of herbaceous species following shrub invasion of grassland.     

Variation in exotic and native seed arrival and recruitment of bird dispersed species in subtropical forest restoration and regrowth

Invasive bird-dispersed plants often share the same suite of dispersers as co-occurring native species, resulting in a complex management issue. Integrated management strategies could incorporate manipulation of dispersal or establishment processes. To improve our understanding of these processes, we quantified seed rain, recruit and seed bank density, and species richness for bird-dispersed invasive and native species in three early successional subtropical habitats in eastern Australia: tree regrowth, shrub regrowth and native restoration plantings. We investigated the effects of environmental factors (leaf area index (LAI), distance to edge, herbaceous ground cover and distance to nearest neighbour) on seed rain, seed bank and recruit abundance. Propagule availability was not always a good predictor of recruitment. For instance, although native tree seed rain density was similar, and species richness was higher, in native plantings, compared with tree regrowth, recruit density and species richness were lower. Native plantings also received lower densities of invasive tree seed rain than did tree regrowth habitats, but supported a similar density of invasive tree recruits. Invasive shrub seed rain was recorded in highest densities in shrub regrowth sites, but recruit density was similar between habitats. We discuss the role of microsite characteristics in influencing post-dispersal processes and recruit composition, and suggest ways of manipulating these processes as part of an integrated management strategy for bird-dispersed weeds in natural areas.     

Diversity,composition and guild structure relationships between soil-stored seed banks and mature vegetation in alien plant-invaded South African fynbos shrublands

We investigated vegetation-seed bank relationships at three fynbos sites on the Cape Peninsula, South Africa, and the impacts to these sites of invasion by the alien tree Acacia saligna. Soil-stored seed banks in uninvaded fynbos were of a similar density to those previously measured in fynbos (ca. 1100–1500 seeds m^-2) and were dominated by mostly short-lived species. Lack of similarity between mature vegetation and seed banks, suggests that seed banks are poor predictors of mature vegetation composition and structure in fynbos. This lack of correspondence was attributed to the ephemerals (present only in the soil seed bank) and the dominance of serotinous (aerial seed bank) and sprouting (soil seed bank low to absent) species, in mature vegetation. Long-lived seeders were among the 10 most abundant species in the seed banks at all sites and at two sites shrub species contributed more to seed bank richness than any other growth form. Soil-stored seed banks, therefore, boost species richness and diversity both in early post-fire and later seral stages.There was a decline in fynbos species richness, diversity and abundance both in the standing vegetation and seed banks with increasing duration of invasion by the alien tree, Acacia saligna. However, the rate of decline was higher for the vegetation than the seed banks, suggesting that many fynbos species have long-term persistent seed banks. At two sites, there was no obvious shift in community composition associated with Acacia invasion: invaded sites were depauperate versions of the uninvaded site. However, at a third site, the vegetation composition shifted towards a community dominated by bird-dispersed thicket species and its seed bank shifted towards a community dominated by wind-dispersed perennials. Community composition of the soil seed banks under dense, recent Acacia was very similar to that of the corresponding uninvaded fynbos at all sites, indicating that there is good potential to return to species-rich fynbos vegetation after removal of the alien Acacia. Most seed bank species persisted in the soil seed bank of the long-invaded fynbos at low frequency and density, indicating high seed longevity in many species. We suggest that either a thick Acacia litter layer or a deep (>5 cm) burial moderated the fire and ambient temperature effects, preventing these seeds from germinating after fire and thus preventing loss from the seed bank.     

The role of environmental gradients in non-native plant invasion into burnt areas of Yosemite National Park, California

Fire is known to facilitate the invasion of many non-native plant species, but how invasion into burnt areas varies along environmental gradients is not well-understood. We used two pre-existing data sets to analyse patterns of invasion by non-native plant species into burnt areas along gradients of topography, soil and vegetation structure in Yosemite National Park, California, USA. A total of 46 non-native species (all herbaceous) were recorded in the two data sets. They occurred in all seven of the major plant formations in the park, but were least common in subalpine and upper montane conifer forests. There was no significant difference in species richness or cover of non-natives between burnt and unburnt areas for either data set, and environmental gradients had a stronger effect on patterns of non-native species distribution, abundance and species composition than burning. Cover and species richness of non-natives had significant positive correlations with slope (steepness) and herbaceous cover, while species richness had significant negative correlations with elevation, the number of years post-burn, and cover of woody vegetation. Non-native species comprised a relatively minor component of the vegetation in both burnt and unburnt areas in Yosemite (percentage species = 4%, mean cover < 6.0%), and those species that did occur in burnt areas tended not to persist over time. The results indicate that in many western montane ecosystems, fire alone will not necessarily result in increased rates of invasion into burnt areas. However, it would be premature to conclude that non-native species could not affect post-fire succession patterns in these systems. Short fire-return intervals and high fire severity coupled with increased propagule pressure from areas used heavily by humans could still lead to high rates of invasion, establishment and spread even in highly protected areas such as Yosemite.     

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.     

Importance of the soil seed bank of disturbed sites in Chilean matorral in early secondary succession

Abstract. The dynamics of the seed bank may provide clues to the process of recovery of the vegetation of disturbed sites. The role of the seed bank may be more important in areas with a seasonal climate than in areas where seedling recruitment is not limited to one season. We studied the seed bank and the seed rain in three sites of the Chilean mediterranean-climate region (33° 48'S) which differed in the degree of anthropic disturbance: a closed-canopy, second-growth forest; an open matorral; and an old-field. Additionally, we tested the germination of seeds from the soil and from the current-year seed crop. The seed bank varied considerably between the two years of study, although no change in the vegetation could be observed. Seed density and species richness were lower in 1989 than in 1988. The seed bank of the second-growth forest changed less between years, while the old-field showed the largest change. The highest seed rain occurred under shrub patches in the open matorral, while few seeds fell in the spaces between shrub clumps or in the old-field. In the forest, seed rain was low and correlated with species cover. Germination was low (0 - 15%) in tests using either soil samples or fresh seeds. These results indicate that matorral succession is a very slow process, limited mainly by low germination and low arrival of propagules to open areas. Most woody species have animal-disseminated fleshy propagules. The presence of established shrubs which may serve as perches or refuges for animals increases species richness in the seed rain and the seed bank.     

Soil seed banks in Pinus ponderosa forests in Arizona: Clues to site history and restoration potential

Question: How does the relationship between the viable soil seed bank species composition and the above‐ground vegetation in northern Arizona Pinus ponderosa forests differ under varying historical land use disturbances (low, intermediate, high)? Is above‐ground vegetation correlated to the viable soil seed bank immediately following soil disturbance from restoration thinning treatments? Location: Northern Arizona, USA. Methods: Soil seed bank samples were taken along replicated transects and collected with a 5‐cm diameter bulk density hammer. Samples included a 5 ‐cm diameter O‐horizon sample (at varying depths) plus the underlying mineral soil to a depth of 5 cm. The seedling emergent method was used to quantify seed bank species composition and density. The herbaceous and shrub plant community was quantified along the same transects using the point intercept method. Results: Early‐successional or ruderal species were common in the soil seed bank at all three disturbance sites. Non‐native species, notably Verbascum thapsus, were more numerous (up to 940 seeds/m²) under high disturbance with overgrazing and logging, and less common or absent under low disturbance. Most viable seeds were found in the O‐horizon and the upper 5 cm of mineral soil; there was little correlation between species in the soil seed bank and the above‐ground vegetation. Conclusions: We recommend that restoration plans be geared toward minimizing activities, such as severe soil disturbance, that may promote the spread of non‐native invasive species, and that manual seeding be explored as an option to restore plant species diversity and abundance.     

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.     

Ecosystem response to removal of exotic riparian shrubs and a transition to upland vegetation

Understanding plant community change over time is essential for managing important ecosystems such as riparian areas. This study analyzed historic vegetation using soil seed banks and the effects of riparian shrub removal treatments and channel incision on ecosystem and plant community dynamics in Canyon de Chelly National Monument, Arizona. We focused on how seeds, nutrients, and ground water influence the floristic composition of post-treatment vegetation and addressed three questions: (1) How does pre-treatment soil seed bank composition reflect post-treatment vegetation composition? (2) How does shrub removal affect post-treatment riparian vegetation composition, seed rain inputs, and ground water dynamics? and (3) Is available soil nitrogen increased near dead Russian olive plants following removal and does this influence post-treatment vegetation? We analyzed seed bank composition across the study area, analyzed differences in vegetation, ground water levels, and seed rain between control, cut-stump and whole-plant removal areas, and compared soil nitrogen and vegetation near removed Russian olive to areas lacking Russian olive. The soil seed bank contained more riparian plants, more native and fewer exotic plants than the extant vegetation. Both shrub removal methods decreased exotic plant cover, decreased tamarisk and Russian olive seed inputs, and increased native plant cover after 2 years. Neither method increased ground water levels. Soil near dead Russian olive trees indicated a short-term increase in soil nitrogen following plant removal but did not influence vegetation composition compared to areas without Russian olive. Following tamarisk and Russian olive removal, our study sites were colonized by upland plant species. Many western North American rivers have tamarisk and Russian olive on floodplains abandoned by channel incision, river regulation or both. Our results are widely applicable to sites where drying has occurred and vegetation establishment following shrub removal is likely to be by upland species.     

洮河上游紫果云杉群落土壤种子库特征及其与地上植被的关系

该研究以甘肃洮河国家级自然保护区大峪沟林区(TH)、冶力关林区(YLG)、尕海 则岔国家级自然保护区则岔林区(GZ)的紫果云杉(Picea purpurea)天然林群落为研究对象,通过样地调查与萌发实验,分析洮河上游紫果云杉群落土壤种子库特征,以揭示洮河上游紫果云杉群落土壤种子库特征及与地上植被的关系,为紫果云杉群落恢复和管理提供依据。结果表明：(1)3个林区土壤种子库共萌发27科45属50种植物,种子库密度在958~1 129粒/m²之间,草本植物是构成研究区土壤种子库的主体。(2)3个林区紫果云杉群落土壤种子库垂直结构明显,物种数及种子密度随土层的加深呈减少趋势。(3)3个林区地上植被物种数大于土壤种子库物种数,相似性系数在0.15~0.23之间,表现为极不相似性;土壤种子库的多样性指数除Pielou指数高于地上植被外,Margalef指数、Shannon Wiener指数、Simpson指数均低于地上植被。研究认为,洮河上游紫果云杉群落土壤种子库以草本植物为主,乔木和灌木的储量较小,特别是建群种紫果云杉的储量较小;地上植被和土壤种子库之间相互的贡献力较低,仅依靠紫果云杉林土壤种子库进行自然恢复,远不能缓解现阶段紫果云杉林退化的问题,因此需开展人工育苗、造林等措施来促进紫果云杉的更新与发展。     

Seed Stores for Restoration of Species-Rich Shrubland Vegetation Following Mining in Western Australia

The extreme species richness of native shrubland vegetation (kwongan) near Eneabba, Western Australia, presents a major problem in the restoration of sites following mineral sand mining. Seed sources available for post-mining restoration and those present in the native kwongan vegetation were quantified and compared. Canopy-borne seeds held in persistent woody fruits were the largest seed source of perennial species in the undisturbed native vegetation and also provided the most seeds for restoration. In undisturbed vegetation, the germinable soil seed store (140–174 seeds · m^?2) was only slightly less than the canopy-borne seed store (234–494 seeds · m^?2), but stockpiled topsoil provided only 9% of the germinable seeds applied to the post-mining habitat. The age of stockpiled soil was also important. In the three-year-old stockpiled topsoil, the seed bank was only 10.5 seeds · m^?2 in the surface 2.5 cm, compared to 56.1 to 127.6 seeds · m^?2 in fresh topsoil from undisturbed vegetation sites. In the stockpiled topsoil, most seeds were of annual species and 15–40% of the seeds were of non-native species. In the topsoil from undisturbed vegetation, over 80% of the seeds were of perennial species, and non-native species comprised only 2.7% of the seed bank. Additional seeds of native species were broadcast on restoration areas, and although this represented only 1% of the seed resources applied, the broadcast seed mix was an important resource for increasing post-mining species richness. Knowledge of the life-history characteristics of plant species may relate to seed germination patterns and assist in more accurate restoration where information on germination percentages of all species is not available.     

Disturbance of suburban Fagus forests by recreational activities: Effects on soil characteristics,above‐ground vegetation and seed bank

Questions: How does recreational disturbance (human trampling) affect soil characteristics, the performance of the understorey vegetation, and the density and species composition of the soil seed bank in Fagus sylvatica forests? Location: Suburban forests near Basel, northwestern Switzerland. Methods: We compared various soil characteristics and the performance of the understorey vegetation in six beech forest areas frequently disturbed by recreational activities with those in six undisturbed control areas, in spring 2003. In the same forest areas, the soil seed bank was investigated using the seedling emergence method. Samples were obtained from soil cores in January 2003. Results: We found substantial changes in soil compaction, above‐ground vegetation and in the soil seed bank due to recreational activities. In frequently visited areas, soil compaction was enhanced which caused a decrease in cover, height and species richness of both herb and shrub layers. Compared with control areas, the number of trampling‐tolerant species of the seed bank was significantly higher in disturbed areas, and total species richness tended to be higher in disturbed than in control areas. Furthermore, the similarity in species composition between the above‐ground vegetation and seed bank was significant lower in disturbed than in control areas. Conclusions: The intensive use of suburban forests for recreational activities, mainly picnicking, affects the vegetation of natural beech forests. Our study indicates that a restoration of degraded forest areas from the soil seed bank would result in a substantial change of the vegetation composition.     

Depth distribution and composition of seed‐banks in alien‐invaded and uninvaded fynbos vegetation

South African fynbos vegetation is threatened on a large scale by invasive woody plants. A major task facing nature conservation managers is to restore invaded areas. The aim of this study was to determine the restoration potential of fynbos following dense invasion by the Australian tree Acacia saligna. The impacts of dense invasion on seed‐bank composition and depth distribution were investigated to determine which fynbos guilds and species have the most persistent seed‐banks. Soil samples were excavated at three different depths for invaded and uninvaded vegetation at two sand plain and mountain fynbos sites. Seed‐banks were determined using the seedling emergence approach. Invasion caused a significant reduction in seed‐bank density and richness at all sites. There was a significant, but smaller, reduction in seed‐bank density and richness with soil depth at three sites. Seed‐bank composition and guild structure changed following invasion. Low persistence of long‐lived obligate seeders in sand plain fynbos seed‐banks indicates that this vegetation type will be difficult to restore from the seed‐bank alone following alien clearance. The dominance of short‐lived species, especially graminoids, forbs and ephemeral geophytes, suggests that regenerating vegetation will develop into a herbland rather than a shrubland. It is recommended that seed collecting and sowing form part of the restoration plan for densely invaded sand plain sites. As seed density remained higher towards the soil surface following invasion, there is no general advantage in applying a mechanical soil disturbance treatment. However, if the shallow soil seed‐bank becomes depleted, for example following a hot fire through dense alien slash, a soil disturbance treatment should be given to exhume the deeper viable seed‐bank and promote recruitment.     

Non-native and native shrubs have differing impacts on species diversity and composition of associated plant communities

The spread of non-native plants has been depicted as a serious threat to biodiversity. However, it remains unclear whether the indigenousness of the invading plant plays a marked role for the ecological consequences of an invasion as few studies have compared the ecological impacts of non-native shrubs with structurally or functionally comparable native shrubs. We studied patches of introduced and native shrubs to assess whether there are general differences in plant species composition or biomass between patches formed by non-native versus native shrubs. The indigenousness of the shrub (non-native vs. native) did not explain the variation in soil nutrients, neither the production of shoot biomass or allocation of growth to different parts of the shoot. The amount of light reaching ground level did not differ between patches of a non-native and a native shrub. However, species richness and biomass of herbaceous plants were lower in patches of non-native than native shrubs and the amount of litter was higher below non-native than native shrubs. Our results suggest that the indigenousness of the patch-forming plant may be an important factor for the diversity and composition of associated herbaceous vegetation. Based on our results, resource availability (light and nutrients) is not a sufficient explanation for the negative effects of non-native shrubs on plant communities. Further research is needed to investigate whether alternative explanations, such as the novelty of the toxic compounds produced by non-native plants, can explain the differences we observed.     

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.     

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.