What predicts the richness of seeder and resprouter species in fire‐prone Cape fynbos: Rainfall reliability or vegetation density?

In ecosystems subject to regular canopy fires, woody species have evolved two general strategies of post‐fire regeneration. Seeder species are killed by fire and populations regenerate solely by post‐fire recruitment from a seed bank. Resprouter species survive fire and regenerate by vegetative regrowth from protected organs. Interestingly, the abundance of these strategies varies along environmental gradients and across regions. Two main hypotheses have been proposed to explain this spatial variation: the gap dependence and the environmental‐variability hypotheses. The gap‐dependence model predicts that seeders are favoured in sparse vegetation (vegetation gaps allowing effective post‐fire recruitment of seedlings), while resprouters are favoured in densely vegetated sites (seedlings being outcompeted by the rapid crown regrowth of resprouters). The environmental‐variability model predicts that seeders would prevail in reliable rainfall areas, whereas resprouters would be favoured in areas under highly variable rainfall that are prone to severe dry events (leading to high post‐fire seedling mortality). We tested these two models using distribution data, captured at the scale of quarter‐degree cells, for seeder and resprouter species of two speciose shrub genera (Aspalathus and Erica) common in fire‐prone fynbos ecosystems of the mediterranean‐climate part of the Cape Floristic Region. Contrary to the predictions of the gap‐dependence model, species number of both resprouters and seeders increased with values of the Normalized Difference Vegetation Index (a widely used surrogate for vegetation density), with a more marked increase for seeders. The predictions of the environmental‐variability hypothesis, by contrast, were not refuted by this study. Seeder and resprouter species of both genera showed highest richness in environments with high rainfall reliability. However, with decreasing reliability, seeder numbers dropped more quickly than those of resprouters. We conclude that the environmental‐variability model is better able to explain the abundance of woody seeder and resprouter species in Southern Hemisphere fire‐prone shrublands (fynbos and kwongan) than the gap‐dependence model.     

Seed bank dynamics of two obligate seeders, <Emphasis Type="Italic">Cistus monspeliensis</Emphasis> and <Emphasis Type="Italic">Rosmarinus officinalis</Emphasis>, in relation to time since fire

Many species in Mediterranean-type ecosystems regenerate after fire by seed germination from soil seed banks. Seed bank dynamics of two of those obligate seeders, Cistus monspeliensis and Rosmarinus officinalis, were investigated in relation to stand age since fire in southwestern Portugal. Soil seed density, annual seed input, annual seed losses through germination and seed persistence were compared between species at stands differing in age since fire (5, 10 and 35 years). Soil seed density and seed input increased over the first decade after fire and were lowest at 35-year-old stands for C. monspeliensis. In R. officinalis, few seeds were produced and found in the soil at early stages, and maximum seed input and soil seed density were attained at 35-year-old stands. Soil seed density was mostly driven by seed production in both species, which is largely dependent on plant traits and population dynamics related to fire. Overall, stand age since fire had a negligible effect on seed germination, seed persistence and viability. Ten to 39% of buried seeds were not recovered after 1 year, and viability of seeds recovered was 97–100% for C. monspeliensis and only 0–3% for R. officinalis. Variation in plant traits within the seeder syndrome was evidenced by this study. R. officinalis evidenced lower seed persistence, lower proportion of viable seed produced and lower density of viable soil seed than C. monspeliensis at any stage after fire. R. officinalis is expected to depend largely on previous year seed production for population replacement after fire.     

Relationship between leaf traits and fire-response strategies in shrub species of a mountainous region of south-eastern Australia

Background and Aims

Resprouting and seed recruitment are important ways in which plants respond to fire. However, the investments a plant makes into ensuring the success of post-fire resprouting or seedling recruitment can result in trade-offs that are manifested in a range of co-occurring morphological, life history and physiological traits. Relationships between fire-response strategies and other traits have been widely examined in fire-prone Mediterranean-type climates. In this paper, we aim to determine whether shrubs growing in a non-Mediterranean climate region exhibit relationships between their fire-response strategy and leaf traits.

Methods

Field surveys were used to classify species into fire-response types. We then compared specific leaf area, leaf dry-matter content, leaf width, leaf nitrogen and carbon to nitrogen ratios between (a) obligate seeders and all other resprouters, and (b) obligate seeders, facultative resprouters and obligate resprouters.

Key Results

Leaf traits only varied between fire-response types when we considered facultative resprouters as a separate group to obligate resprouters, as observed after a large landscape-scale fire. We found no differences between obligate seeders and obligate resprouters, nor between obligate seeders and resprouters considered as one group.

Conclusions

The results suggest that facultative resprouters may require a strategy of rapid resource acquisition and fast growth in order to compete with species that either resprout, or recruit from seed. However, the overall lack of difference between obligate seeders and obligate resprouters suggests that environmental factors are exerting similar effects on species'' ecological strategies, irrespective of the constraints and trade-offs that may be associated with obligate seeding and obligate resprouting. These results highlight the limits to trait co-occurrences across different ecosystems and the difficulty in identifying global-scale relationships amongst traits.     

Long-term post-fire dynamics of co-occurring woody species in Pinus brutia forests: the role of regeneration mode

Regeneration mode is one of the key attributes determining population structure and dynamics of plant species. We investigated long-term patterns after fire in the cover of plant species in a 100-year chronosequence of burned Pinus brutia forests in a humid Mediterranean climate region in Turkey. Significant trends were present in the change of cover in major species through post-fire chronosequence, and species with similar trends were clustered in relation to their regeneration modes. Obligate resprouters increased their cover from the early post-fire years to the later stages, while cover of obligate seeders with a soil seed bank increased in the early years, but then decreased through time. Facultative resprouters were at an intermediate position, with an increase in cover until mid-successional stages and then a decrease through time. The cover of the only obligate seeder with a canopy seed bank (P. brutia) followed a linear increasing trend during the succession. When species with the same regeneration mode were grouped, the same trends were observed with more explained variances. A few life-history traits were enough to explain the observed trends. Our study shows that regeneration mode is an explanatory functional grouping system for describing long-term post-fire dynamics of Mediterranean Basin woody species. We suggest that regeneration mode must be a major component of any vegetation or forest stand dynamics model in the Mediterranean Basin. This result has important implications for the management of Mediterranean Basin ecosystems, and can potentially be extrapolated to other Mediterranean-type fire-prone ecosystems.     

Post‐fire regeneration of Mediterranean plant communities at a regional scale is dependent on vegetation type and dryness

Question: We tested whether (1) the change in composition and structure of whole plant communities after fire is directly related to regeneration of the dominant tree species in the canopy; (2) the change in structure and composition of plant communities several years after fire decreases with the proportion of obligate seeders and (3) the proportion of obligate seeders in plant communities increases with the dryness gradient. Location: Catalonia (NE Spain) Methods: We measured floristic differences between burned and long‐since burned sites in eight vegetation types across a climate gradient. We compared 22 sites burnt in 1994 in paired plots with 22 sites that had not been burnt since the 1940s. In each site we placed plots in burned and long‐since burned areas, where we identified the presence and abundance of all plant species. Results: When the tree canopy recovers, structure and composition of the vegetation also return to the long‐since burned community; when tree canopy does not recover, composition of the post‐fire community varies compared to the long‐since burned one. A higher proportion of obligate seeders in the pre‐fire community promotes quicker regeneration of the original community. The proportion of obligate seeders increased along the dryness gradient. Conclusions: Regeneration of plant communities after fire depends on the vegetation type before the fire. Regeneration increases when the dominant tree or shrub species persists after fire and with a higher proportion of obligate seeders in the pre‐fire community. The proportion of obligate seeders varies along the dryness gradient, which suggests that vegetation in drier areas (when seeders are more abundant) recovers earlier than in moister areas.     

The relationship between soil seed bank,above‐ground vegetation and disturbance intensity on old‐field successional permanent plots

Questions: How does disturbance and successional age influence richness, size and composition of the soil seed bank? What is the potential contribution of the soil seed bank to the plant community composition on sites differing in their successional age or disturbance intensity? Location: Experimental Botanical Garden of Göttingen University, central Germany. Methods: Above‐ground vegetation and soil seed bank were studied on formerly arable fields in a 36‐year‐old permanent plot study with five disturbance intensities, ranging from yearly ploughing via mowing to long‐term uninterrupted succession. We compared species compositions, seed densities and functional features of the seed bank and above‐ground vegetation by using several methods in parallel. Results: The seed bank was mainly composed of early successional species typical of strongly disturbed habitats. The difference between seed bank composition and above‐ground vegetation decreased with increasing disturbance intensity. The species of greatest quantitative importance in the seed bank was the non‐native forb Solidago canadensis. Conclusions: The ability of a plant community to regenerate from the soil seed bank dramatically decreases with increasing time since abandonment (successional age) and with decreasing disturbance intensity. The present study underlines that plant species typical of grasslands and woodlands are limited by dispersal capacity, owing to low capacity for accumulation of seeds in the soil and the fact that most species do not build up persistent seed banks. Rare and target species were almost absent from the seed bank and will, after local elimination, depend on reintroduction for continuation of their presence.     

Impact of two wildfires on endemic granite outcrop vegetation in Western Australia

Abstract. In seasonally dry regions of the world fire is a recurring disturbance but little is known of how fire interacts with granite outcrop vegetation. We hypothesize that the floristic composition in granite vegetation, usually attributed to the edaphic environment, may also reflect the impact of disturbances such as fire. Dramatic differences in floristic composition and cover over 13 years and two fires were observed in vegetation on a Western Australian granite outcrop. This was very marked in the first year following the two fires, with annuals and geophytes showing the greatest turnover of species. Even among the perennial shrubs there was considerable turnover in a number of obligate seeders. After the first fire the number of species declined for woody perennials, herbaceous perennials and annuals, remained unchanged for perennial grasses and sedges, and varied with highest richness 4 yr after fire for geophytes. Demographic studies of two endemic woody obligate seeders and three endemic mallee eucalypt resprouters similarly showed dramatic differences within and between species in seedling recruitment following the two fires. Fire does have a significant impact on the floristic composition of semi‐arid granite outcrop vegetation communities. Studies on other granite outcrop systems are needed to test the generality of this conclusion.     

Seed bank patterns in Restionaceae and Epacridaceae after wildfire in kwongan in southwestern Australia

Abstract. Post-fire seed germination, seedling mortality and seed banks were investigated in scrub-heath (kwongan) in SW Australia. Study species included herbaceous and woody, obligate seeders and resprouters in two non-bradysporous but significant plant families (Restionaceae and Epacridaceae). In all species, seedlings were recruited only in the first autumn-spring after fire and occurred in similar densities as the estimated germinable annual seed input. Seedlings were absent from unburnt vegetation. Although most species retained some residual seeds after fire, tests (excised embryo culture) indicated that a negligible number of seeds were germinable. Regardless of fire response or species, there appeared to be a large loss of seeds each year and in most cases, only a small proportion of the annual seed production was used in post-fire recovery of plants. Based on seedling: parent ratios, all species had the capacity to reconstitute parent densities from germinants in the first year after fire, but high seedling mortality and no further recruitment resulted in less seedlings than replacements for four resprouter Restionaceae and three Epacridaceae (all obligate seeders) at the end of the third year after fire.     

Post-fire and post-quarry rehabilitation successions in Mediterranean-like ecosystems: Implications for ecological restoration

Resilience against sudden changes in the environment is a very desirable trait in plants used for ecosystem restoration. Mediterranean-like vegetation exhibits particularly strong fire resilience. There are two main functional groups of fire-prone species among Mediterranean-like vegetation: seeders and resprouters. Our aims were to describe how the theory of succession after fire relates to rehabilitation and to use this knowledge to improve the results of rehabilitation attempts in Mediterranean-like ecosystems. Eight post-fire (PF) sites, 14 post-rehabilitated (PR) quarry sites and two woodland sites were sampled. Detrended Canonical Correspondence Analysis (DCCA) showed that PF and PR successions were quite different. Both displayed an increasing abundance of resprouters over time, but seeder density increased throughout PR succession and decreased during PF succession. Nine species were common to both successions in all studied stages. The results showed that until 15-21 years of succession, the post-rehabilitation sites had not become as resilient to fires as sites populated by indigenous vegetation due to the lack of a seeder seed bank. However, after 21 years of PR succession, the exponentially increasing seeder population may allow for seed bank formation and thus eventually improve the fire resilience of the site.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.     

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.     

Biological significance of dead biomass retention trait in Mediterranean Basin species: an analysis between different successional niches and regeneration strategies as functional groups

Standing dead biomass retention is considered one of the most relevant fuel structural traits to affect plant flammability. However, very little is known about the biological significance of this trait and its distribution between different functional groups. Our aim was to analyse how the proportion of dead biomass produced in Mediterranean species is related to the successional niche of species (early‐, mid‐ and late‐successional stages) and the regeneration strategy of species (seeders and resprouters). We evaluated biomass distribution by size classes and standing dead biomass retention in nine dominant species from the Mediterranean Basin in different development stages (5, 9, 14 and 26 years since the last fire). The results revealed significant differences in the standing dead biomass retention of species that presented a distinct successional niche or regeneration strategy. These differences were restricted to the oldest ages studied (>9 years). Tree and small tree resprouters, typical in late‐successional stages, presented slight variations with age and a less marked trend to retain dead biomass, while seeder shrubs and dwarf shrubs, characteristic of early‐successional stages, showed high dead biomass loads. Our results suggest that the species that tend to retain more dead branches are colonising species that may promote fire in early‐successional stages.     

Long‐term seed bank dynamics in a temperate forest under conversion from coppice‐with‐standards to high forest management

Questions: How do changes in forest management, i.e. in disturbance type and frequency, influence species diversity, abundance and composition of the seed bank? How does the relationship between seed bank and vegetation change? What are the implications for seed bank dynamics? Location: An ancient Quercus petraea — Carpinus betulus forest in conversion from coppice‐with‐standards to regular Quercus high forest near Montargis, France. Methods: Seed bank and vegetation were sampled in six replicated stand types, forming a chronosequence along the conversion pathway. The stand types represented mid‐successional stages of stands in transition from coppice‐with‐standards (to high forest (16 plots) and early‐ and mid‐successional high forest stands (32 plots). Results: Seed bank density and species richness decreased with time since last disturbance. Adjusting for seed density effects obscured species richness differences between stand types, but species of later seres were nested subsets of earlier seres, implying concomitant shifts in species richness and composition with time since disturbance. Later seres were characterized by species with low seed weight and high seed longevity. Seed banks of early seres were more similar to vegetation than to later seres. Conclusions: Abandonment of the coppice‐with‐standards regime altered the seed bank characteristics, as well as its relationship with vegetation. Longer management cycles under high forest yield impoverished seed banks. For their persistence, seed bank species will increasingly rely on management of permanently open areas in the forest landscape. Thus, revegetation at the beginning of new high‐forest cycles may increasingly depend on inflow from seed sources.     

Fire persistence traits can be used to predict vegetation response to changing fire regimes at expansive landscape scales – an Australian example

Aim Building on a substantial literature addressing the fire responses of woody plants, particularly under mediterranean climates, we assess the extent to which fire persistence traits can be used to predict vegetation responses to fire regime changes in fire‐prone arid and savanna landscape settings. Location Australia, applying data from arid central to monsoonal northern regions (11–26° S, 129–138° E). Methods With reference to a substantial sub‐continental floristics dataset, we first assigned the fire response (obligate seeder, resprouter) and seedbank persistence (transient, dormant) of rapid and longer‐maturing (> 3 years) woody taxa. Using logistic regression, we then modelled the proportions of taxa possessing these traits as a function of mean annual rainfall (highly correlated with fire frequency) and terrain roughness (a measure of topographic variability) in 0.25° × 0.25° and 1° × 1° grid cells. Separate assessments were undertaken with datasets for 1264 sclerophyll and 236 rain forest taxa. Results This woody flora is characterized by taxa exhibiting mostly resprouting and dormant seedbank traits that promote site persistence. While numbers of obligate seeder and resprouter taxa were related positively to both rainfall and roughness, the relative abundance of both sclerophyll and rain forest obligate seeders decreased significantly with rainfall. The relative abundance of sclerophyll (especially long‐lived) obligate seeders alone increased with topographic roughness. The proportion of taxa with transient seedbanks increased with rainfall in resprouters generally, and in rain forest obligate seeders alone. Main conclusions We find that resprouters are favoured on more productive, fire‐prone sites, and obligate seeders are favoured in less productive, more fire‐protected settings. Seedbank persistence responses are more variable. These findings concur generally with theoretical constructs, and support comparable assessments in Australian and other fire‐prone systems ranging from mediterranean to boreal environments. Our observations illustrate that resprouting and obligate seeding syndromes, but not necessarily seedbank persistence, are useful predictors of vegetation responses to changing fire regime conditions at large landscape scales.     

Scale-dependent segregation of seeders and resprouters in cork oak (<Emphasis Type="Italic">Quercus suber</Emphasis>) forests

Recent studies showed that disturbances and water availability determine the richness among plants with different post-fire strategies of Mediterranean-type ecosystems. The aim of this study was to determine whether or not the scale of analysis has an influence on the effects of these factors and, therefore, on the segregation of the dominant post-fire strategies, obligate seeders and obligate resprouters, and facultative species. We recorded all woody species and geographical features on 94 (75 m²) plots of cork oak woodlands in the southern Iberian Peninsula. For each regenerative type (resprouters, seeders and species with both traits—facultative species), we tested the relationship between the number of species and the predictors using a generalised linear mixed model. The fixed predictor considered at the large scale was altitude, and fixed predictors considered at the local scale were aspect (north/south) and disturbance (fire and clearing by heavy machinery; yes/no). The random predictor was the factor of site. When this factor did not have significant effect for some regenerative types, these relationships was tested using a generalised linear model. Resprouting species were most represented at lower altitudes and in undisturbed sites, while seeders were also at lower altitudes but mostly on south-facing slopes, especially south-facing disturbed sites. For facultative species, site is the most important variable. The proportion of seeders from the total species is not related to altitude, but it is related to disturbance and aspect. These results suggest that there is no segregation of the richness of seeders and resprouters at the large scale (altitudinal gradient). Differences appeared at the local scale (aspect and disturbance).     

Soil seed bank dynamics in alpine wetland succession on the Tibetan Plateau

The primary goal was to address several questions with regard to how soil seed banks change in a successional series. How does the composition of the viable seed bank change, and how does the relationship of the soil seed bank and vegetation change with succession? Can the seed bank be regarded as a potential as a source of seeds for wetland restoration? We collected soil seed bank samples and sampled the vegetation in four different successional stages and used the NMDS (nonmetric multidimensional scaling) to evaluate the relationship of species composition between the seed banks and vegetation. The difference of seed density and species richness in different habitats and soil depths also was compared. Viable seeds of half (37) the species in the early-successional stage were found in all the successional stages. Similarity between seed bank and vegetation increased with succession. Both seed density and species richness in the seed bank increased with successional age and decreased with soil depth. The majority of species from the early-successional stage produced long-lived seeds. Seed density and species richness increased with succession, mainly as a result of increasing seed production, and hypotheses predicting decreasing density of buried seeds and species richness were not confirmed. Seed banks play a minor role in contributing to the regeneration of vegetation, and managers cannot rely on soil-stored seed banks for restoration of wetlands.     

Can soil seed banks contribute to the restoration of dune slacks under conservation management?

Questions: Does the soil seed bank resemble the former early successional stages of a dune slack system more than the established later successional vegetation? Does it have the potential to contribute to the conservation of a highly endangered habitat? Location: Dune slacks at Newborough Warren, UK. Methods: The composition of the soil seed bank in two depth layers was determined using the seedling emergence method between March 2004 and April 2005. Long-term monitoring data on the floristic composition of the established vegetation were obtained from the national conservation agency, and additional monitoring was undertaken in 2003. Floristic composition, seed weights, seed longevity of component species and Ellenberg indicator values were used to compare the seed bank and established vegetation. Results: The soil seed bank was diverse and contained typical dune slack species, species of early successional stages and species of conservation interest. A comparison between the composition of the seed bank and historical data on the composition of the established vegetation showed that the seed bank reflects earlier successional stages more closely than the current aboveground vegetation. This study increases the scarce information currently available on the seed bank ecology of several species, including two orchid species. Conclusions: The soil seed bank can be expected to contribute to vegetation change after disturbance. Stimulation of germination from the seed bank through management may contribute to the conservation of both characteristic and threatened species typical of dune slacks.     

Composition,size and dynamics of the seed bank in a mediterranean shrubland of Chile

Abstract Analysis was performed of the richness and abundance of woody species, forbs, and annual grasses in the easily germinating soil seed bank (henceforth seed bank) in a mediterranean shrubland of central Chile. The effects of successional development after fire and by microsite type (underneath or outside shrubs) on the density of seeds in the soil, and the relationship of species abundance in the seed bank with its abundance in the above‐ground vegetation was examined. A total of 64 plant species were recorded in the seed bank, of which 44 were annual or biannual. Eight species were woody and another eight were perennial herbs. Four could not be identified to species level. The highest richness of established herbaceous species was recorded in late spring, with 31 species. The regeneration of the herbaceous vegetation was driven by the annual production of seeds and by a reserve of short‐lived propagules in the soil. Density of all germinating seeds was significantly higher during late spring and late summer. Density of grass seeds was greater during late spring, while that of all other species was greater during late summer. Annual grass seeds accumulated in higher proportion at exposed microsites rather than under woody canopy, and in young (< 5 years old) and intermediate‐age patches (10–20 years old) rather than in mature vegetation (30–50 years old). The abundance of established woody and herb species was uncorrelated with that of the seed bank.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.     

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.