EFFECTS OF A FIRE RESPONSE TRAIT ON DIVERSIFICATION IN REPLICATED RADIATIONS

Fire has been proposed as a factor explaining the exceptional plant species richness found in Mediterranean regions. A fire response trait that allows plants to cope with frequent fire by either reseeding or resprouting could differentially affect rates of species diversification. However, little is known about the generality of the effects of differing fire response on species evolution. We study this question in the Restionaceae, a family that radiated in Southern Africa and Australia. These radiations occurred independently and represent evolutionary replicates. We apply Bayesian approaches to estimate trait‐specific diversification rates and patterns of climatic niche evolution. We also compare the climatic heterogeneity of South Africa and Australia. Reseeders diversify faster than resprouters in South Africa, but not in Australia. We show that climatic preferences evolve more rapidly in reseeder lineages than in resprouters and that the optima of these climatic preferences differ between the two strategies. We find that South Africa is more climatically heterogeneous than Australia, independent of the spatial scale we consider. We propose that rapid shifts between states of the fire response trait promote speciation by separating species ecologically, but this only happens when the landscape is sufficiently heterogeneous.     

Early post-fire plant succession in Peninsula Sandstone Fynbos: The first three years after disturbance

The early post-fire plant succession in fynbos vegetation in the Mediterranean-type climate area of South Africa was studied. Relatively little has been published on this early stage of plant succession in fynbos. Annual sampling over the first three post-fire years confirmed a steady, but relatively slow increase in plant canopy cover of shrubs and graminoids (mainly Restionaceae), whereas cover of geophytes and other herbs peaked in the first year and declined significantly, thereafter. Cover of annual plants increased each year, which may relate to the persistence of a relatively open vegetation cover by the third year. The responses of reseeder and resprouter species of the Restionaceae to the post-fire environment appeared to be habitat dependent. Cover of the reseeders increased rapidly in seep areas, but their recovery was distinctly delayed in dryland areas outside the seeps. Re-establishment of the many reseeder Erica species appeared to be delayed until the second post-fire year. Seed banks of these species were possibly negatively impacted by the fire, and required dispersal of seed from unburnt areas for recruitment. In contrast to some current generalisations, species richness appeared to increase after the fire; less certainly from the first to the second year, but more certainly from the second to the third year. Therefore, this study does not support a short-term monotonic decline in species richness after fire in fynbos.     

The influence of resprouting forest canopy species on richness in Southern Cape forests, South Africa

We investigated the relationship between species richness and numbers and types of individuals and species present in forests with different physiognomies in the southern Cape Province, South Africa. Data were collected from three different ‘plot’ types: 400 m², canopy‐scaled (plot length is directly proportional to canopy height) and per 100 individuals closest to a point. Plots were designed to control for the effect of scale on local richness. Canopy species richness was inversely proportional to the abundance of resprouting species. The strength of the relationship between the abundance of resprouters and canopy species richness increased progressively from the 400 m² plots to the canopy‐scaled plots and finally to the plots of 100 individuals. Resprouter abundance decreased, while canopy species richness increased, with increasing canopy height. Resprouters are able to retain their in situ position in the forests for longer periods of time than do reseeders. This reduces individual and species turnover, thus reducing species richness in resprouter‐dominated forests.     

Habitat and fire heterogeneity explain the co-occurrence of congeneric resprouter and reseeder Hypericum spp. along a Florida pine savanna ecocline

The distribution of resprouting and reseeding woody plants may be limited by the frequency of disturbances. Such species have a high probability of persisting in frequently and rarely disturbed habitats and may co-occur at intermediate disturbance frequencies. Nonetheless, resprouters and reseeders of the genus Hypericum co-occur in frequently burned pine savannas of southeastern North America. We predicted that these congeners would sort along a fire frequency gradient resulting from fine scale variation in topography and soil moisture. We examined habitat associations of a resprouter (H. microsepalum), facultative reseeder/resprouter (H. brachyphyllum), and reseeder (H. chapmanii) that occur along Northern Florida pine savanna ecoclines. We sampled five belt transects of 50 continuous 1?×?1?m² plots for edaphic characteristics, fire spread, and densities of each species. Hypericum microsepalum was associated with upland, drier pine savannas where fires are frequent and typically burn uniformly across landscapes (2?C3?year fire frequency). In contrast, H. brachyphyllum and H. chapmanii were associated with intermediate mesic areas where fires burn increasingly patchily downslope along ecoclines from upland flatwoods to lowland wet depressions (10?C30?year fire frequency). Hypericum species of all life histories co-occur in intermediate areas where small changes in topography and edaphic characteristics generate a fire frequency gradient on a local scale. In pine savannas, fires vary from frequent to infrequent on a local within-landscape level as a function of elevation gradients. We conclude that the occurrence of such fire gradients along ecoclines should facilitate co-occurrence of plants with different life histories and thereby increase overall biodiversity.     

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.     

Resprouters vs reseeders in South African forest trees; a model based on forest canopy height

1. We expect tree species that regenerate primarily by sprouting to produce fewer seedlings than co-occurring species that regenerate mainly from seedlings, because of the trade-off between allocating resources either to ensuring vegetative reproduction (e.g. protective bark/latent buds) or to sexual reproduction (e.g. seeds).
2. Furthermore, resprouting species, because of their multi-stemmed nature, should be at a relative disadvantage, and therefore relatively infrequent, in tall forests. This is because a resprouting individual allocates resources to a number of basal branches/stems and buds rather than maximizing vertical extension of a single leader, as is the case in a seeder. Also, many tall stems arising from the same multi-stemmed base, as is the case in resprouters, will be relatively poorly supported in comparison to the single stem of a reseeder.
3. To test these two ideas we surveyed a number of plots in a range of South African forests and thicket communities. We noted the numbers of seedlings and resprouts for each species and determined a mean for each site.
4. Short forests and thickets were dominated by multi-stemmed species and there was an almost total absence of seedlings. In contrast, tall forests were dominated by single-stemmed reseeding species and were accompanied by seedlings.     

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.     

Ecological and evolutionary differences between Mediterranean seeders and resprouters

Abstract. Differences in allocation patterns between seeders and resprouters in several Mediterranean plant communities (Australia, California and South Africa) have led to the prediction that seedlings of seeders grow faster than those of resprouters. In the Mediterranean Basin, it has also been hypothesized that regeneration strategy of plants after fire is associated with several other life history traits. This paper tests both hypotheses for the dominant plants in the Mediterranean Basin from literature data. Results show that seeders from the Mediterranean Basin grow significantly faster and allocate more biomass to leaf plus paracotyledons than resprouters. Seeders are mainly non‐sclerophyllous, anemochorous, dry‐fruited, small‐seeded species that evolved in the Quaternary (post‐Pliocene) and are associated with earlier successional stages. Resprouters are mainly sclero‐phyllous, vertebrate‐dispersed, fleshy‐fruited, large‐seeded species that evolved in the Tertiary (pre‐Pliocene) and are associated with late successional stages.     

Tradeoffs between height growth rate, stem persistence and maximum height among plant species in a post-fire succession

One way species of low maximum height can accrue sufficient light income to persist in vegetation is via rapid height growth immediately following disturbance. By surveying patches of known time since fire, we reconstructed height–growth trajectories for 19 post-fire recruiting species from fire-prone vegetation in south-eastern Australia. Cross-species patterns of height growth were compared to several plant traits thought to influence height strategy, including leaf mass per area, stem tissue density, stem diameter and capacity to resprout. Shorter species were found to temporarily outpace taller species, both as resprouters and within reseeders. Among reseeders, a single axis of variation summarised patterns of height–growth, time to onset of reproduction and longevity. This axis was tightly correlated with maximum height, leaf mass per area and stem diameter at a given height. These results illustrate how a range of height strategies can coexist in fire-prone vegetation, via the time-process initiated by disturbance.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Seed germination ecology in southwestern Western Australia

Germination responses of species from the native plant communities of southwestern Western Australia can be related to syndromes of life history, fire response, and seed storage, and also to factors related to environmental stress. The Mediterranean-type climate of the region with periodic drought and recurrent fires affects the production of viable seeds in plants of limited stature and rooting depth. Fire response ephemerals and species cued to flower by fire tend to produce viable, readily germinable seeds, but there are instances where seed production is aborted in these predominantly herbaceous life forms. Clonal, rhizomatous species often produce mainly inviable seeds. Production of viable seeds in woody species of these highly diverse communities may also be restricted by limitations to cross pollination. Obligate post-fire seeding species tend to produce a greater proportion of viable seeds than species which are capable of resprouting following fire. Serotinous species, whether post-fire re-seeders or post-fire resprouting species, produce mainly viable seeds, which germinate readily once freed from protective fruits. Species of the legume families and a few others of the soil seed bank produce innately dormant seeds which can be germinated following heat shock treatments which simulate the effects of fire. Heat shock in these species appears mainly as a mechanism to crack the hard seed coats, but the effect of heat to denature seed coat inhibitors has not been eliminated. Western Australian species do not seem to break dormancy when exposed to leachates from burned wood as has been observed in comparable habitats in California and South Africa, but further research is advised. Germination in many native southwestern Australian species is cued by temperatures that correspond to the winter rainfall period. There are also indications that an after-ripening period of warm, dry storage increases percentage of germinable seeds. Stimulation of germination by hormones is almost unresearched in Western Australia, but germination percentages have been increased in a small number of species of horticultural potential. Stimulation of germination by soil nutrient concentrations is almost unresearched in Western Australia, except for the inhibitory effect of excess sodium chloride levels inEucalyptus andMelaleuca. These species only germinate when osmotic effects are reduced to lower levels as would occur when winter rains dilute soil salts. Application of research on seed germination has already enhanced the establishment of seedlings in the restoration of mine sites and is becoming important in aspects of the breeding and selection of native plants for the cut flower, bedding plant and essential oil industries.     

Root traits explain different foraging strategies between resprouting life histories

Drought and fire are prevalent disturbances in Mediterranean ecosystems. Plant species able to regrow after severe disturbances (i.e. resprouter life history) have higher allocation to roots and higher water potential during the dry season than coexisting non-resprouting species. However, seedlings of non-resprouters have a higher survival rate after summer drought. We predict that, to counteract their shallow-rooting systems and to maximize seedling survival, non-resprouters have root traits that confer higher efficiency in soil resource acquisition than resprouters. We tested this prediction in seedlings of less than 1.5 months old. We select 13 coexisting woody species (including both resprouters and non-resprouters), grew them in a common garden and measured the following root traits: length, surface, average diameter, root tissue density (RTD), specific root length (SRL), surface:volume ratio (SVR), specific tip density (STD), tip distribution in depth, internal links ratio (ILR), and degree of branching. These root traits were compared between the two resprouting life histories using both standard cross-species and phylogenetic-informed analysis. Non-resprouters showed higher SRL and longer, thinner and more branched laterals, especially in the upper soil layers. The external links (i.e. the most absorptive root region) were also more abundant, longer, thinner and with higher SVR for non-resprouters. The results were supported by the phylogenetic-informed analysis for the root traits most strongly related to soil resource acquisition (SRL, SVR and branching pattern). The seedling root structure of non-resprouters species allows them to more efficiently explore the upper soil layer, whereas seedling roots of resprouters will permit both carbon storage and deep soil penetration.     

Herbivory, serotiny and seedling defence in Western Australian Proteaceae

We examined how acceptability characteristics displayed by 28-day-old seedlings of 12 species of Western Australian Proteaceae affect the likelihood of seedling herbivory in the field. The seedling attributes quantified were cotyledon phenolic, cyanide and nitrogen concentrations, and cotyledon area, thickness and specific leaf area. Only phenolic content was significantly correlated (negatively) with field rates of herbivore attack. This finding shows that the phenomenon of selective herbivore attack on seedlings may be influenced by a specific plant life-history trait, (in this case cotyledon phenolic concentration). In addition, we also studied the interaction between fire, serotiny and herbivory in matched burned and unburned plots. Although herbivore activity was greater in unburned plots, weakly serotinous species were as prone to defoliation as congeneric, strongly serotinous species, even though their seedlings recruit successfully in the absence of fire. This result suggests that seedlings of species able to establish between fires are not better defended against the higher levels of herbivory normally associated with unburned vegetation.     

Biogeography of seeder and resprouter Erica species in the Cape Floristic Region—Where are the resprouters?

The genus Erica L., with more than 600 species, and a high number of endemics, represents the most remarkable example of floristic diversity in the Cape Floristic Region (CFR). It is largely confined to nutrient-poor, acidic, sandy soils, being one of the most characteristic element of fynbos. The ability to survive fires, resprouting from a lignotuber, is a common trait among Euro-mediterranean Erica species. In contrast, resprouting is fairly uncommon among ericas in the CFR (less than 10%). Most of them are killed by fire, regenerating only but readily by seed germination. An extensive survey on the resprouting ability of South African Erica species was carried out and the pattern of geographical distribution of resprouters and seeders in the CFR was determined. The geographical distribution of these two regeneration classes was related to a climatic gradient of seasonality along the CFR. A pattern of higher proportions of resprouter species towards the mediterranean, strongly seasonal northwestern CFR and the non-seasonal eastern CFR and summer rainfall area outside the CFR was identified. The number of resprouter species reaches a maximum in the eastern CFR and is lower in the southwestern CFR despite the overall higher concentration of species in this subregion. Summer drought strongly influences the effectiveness of post-fire regeneration and growth (i.e. new recruits plus survivors) of Erica species, and is the major selective force accounting for the pattern of distribution of seeders and resprouters in the CFR. A mild mediterranean climate with reliable autumn-winter rains and a short summer drought, typical of the mountain areas of the southwestern CFR, favours recruitment of seeders but hampers recruitment of resprouters. Resprouter species persist and become dominant under harsh conditions for recruitment (severe summer drought) and would coexist with seeders under situations of no summer stress. Diversification is associated with seeder lineages. Hence, number of seeder species will be higher than number of resprouters, especially in the southwestern CFR, where favourable conditions for recruitment allow a massive concentration of seeder species, many of them narrow endemics.     

Vegetation Management for Promoting Ecosystem Resilience in Fire-Prone Mediterranean Shrublands

Dense shrublands constitute highly hazardous fuels in Mediterranean countries. The combination of agricultural land abandonment and fire occurrence in many Mediterranean areas has led to a landscape dominated by shrublands where resprouter species are scarce or absent. Major goals in the management of these areas are to reduce both the fuel loads and their continuity and increase the resilience of the ecosystems by introducing resprouter species. We investigated the performance of one-year-old seedlings of native resprouters, that is, Pistacia lentiscus , Quercus ilex , and Rhamnus alaternus , in a combination of vegetation clearing and mulching with brush chipping in three sites with high amount of fuel load. The planting holes inside the shrublands showed lower radiation and soil surface temperature than those in the cleared areas, whereas soil water content was higher in the mulched holes than in the unmulched ones, especially when water availability was lowest. Seedling survival of Q. ilex and R. alaternus significantly increased within the shrubland; P. lentiscus showed the opposite effect, but its survival was enhanced by the mulching treatment. The three species grew faster in the cleared plots although, unexpectedly, mulching showed a negative effect on Q. ilex relative growth rate in diameter. Our results suggest that the combination of fuel control and reforestation as techniques for shrubland management is an appropriate option, at least in the short term, to redirect vegetation dynamics toward later successional stages, improving ecosystem resilience by the introduction of woody resprouter species.     

Post‐fire recovery of revegetated woodland communities in south‐eastern Australia

The primary goal of restoration is to create self‐sustaining ecological communities that are resilient to periodic disturbance. Currently, little is known about how restored communities respond to disturbance events such as fire and how this response compares to remnant vegetation. Following the 2003 fires in south‐eastern Australia we examined the post‐fire response of revegetation plantings and compared this to remnant vegetation. Ten burnt and 10 unburnt (control) sites were assessed for each of three types of vegetation (direct seeding revegetation, revegetation using nursery seedlings (tubestock) and remnant woodland). Sixty sampling sites were surveyed 6 months after fire to quantify the initial survival of mid‐ and overstorey plant species in each type of vegetation. Three and 5 years after fire all sites were resurveyed to assess vegetation structure, species diversity and vigour, as well as indicators of soil function. Overall, revegetation showed high (>60%) post‐fire survival, but this varied among species depending on regeneration strategy (obligate seeder or resprouter). The native ground cover, mid‐ and overstorey in both types of plantings showed rapid recovery of vegetation structure and cover within 3 years of fire. This recovery was similar to the burnt remnant woodlands. Non‐native (exotic) ground cover initially increased after fire, but was no different in burnt and unburnt sites 5 years after fire. Fire had no effect on species richness, but burnt direct seeding sites had reduced species diversity (Simpson's Diversity Index) while diversity was higher in burnt remnant woodlands. Indices of soil function in all types of vegetation had recovered to levels found in unburnt sites 5 years after fire. These results indicate that even young revegetation (stands <10 years old) showed substantial recovery from disturbance by fire. This suggests that revegetation can provide an important basis for restoring woodland communities in the fire‐prone Australian environment.     

Vegetation dynamics of Mediterranean shrublands in former cultural landscape at Grazalema Mountains,South Spain

Plant community dynamics in Mediterranean basin ecosystems are mainly driven by an alternation of episodes of human intervention and land abandonment. As a result, a mosaic of plant communities has evolved following different stages of degradation and regeneration. Some authors has relate secondary succession to abandoned culture lands and regeneration to natural systems with abandonment of livestock or forestry exploitation. In this paper, the dynamics of shrublands in mid-mountain areas in the South of Spain after disturbance and land abandonment has been studied. The plant cover and 13 environmental variables of 137 selected sites on the Grazalema mountains was analysed to determine the vegetation pattern in relation to environmental factors and the succession types, either regenerative or secondary succession. The results show that today the Grazalema mountains have a heterogeneous vegetation pattern. Besides physical factors such as altitude or soil , human disturbance has modulated current vegetation patterns and dynamics. Two main types of vegetation dynamics can be distinguished in the study area. In areas affected by cutting, regeneration results in rich and dense shrub land, with resprouters as dominant species. In areas affected by recurrent wildfires or agriculture, secondary succession became dominant, resulting in less diverse shrubland, due to the dominance of seeders and decrease in resprouter species richness and cover.     

The evolution of growth rate, resource allocation and competitive ability in seeder and resprouter tree seedlings

Many woody plant species in fire disturbed communities survive disturbance events by resprouting. The resprouting life history is predicted to be costly to plants as resources are diverted into storage for post-fire regrowth rather than allocated to current growth, and resprouting species typically grow more slowly than seeder species (species that do not resprout after disturbance events). Differences in allocation to current growth are also predicted to make resprouter species poorer competitors compared to seeder species. We tested the predictions that the evolution of a resprouter life history is associated with slow growth, increased allocation to storage, and low competitive ability in woody plant seedlings. We grew eight phylogenetically independent pairs of seeder and resprouter species in competition and no competition treatments in a field experiment near Sydney, Australia. The presence of competitors reduced plant growth rates across taxa and fire response life histories. However, relative to seeder species, resprouter species were not slower growing, they did not allocate more resources to storage, and they did not have lower competitive abilities. We propose that differences in resource allocation to storage are not responsible for differences in growth rate and competitive ability. Rather, growth rate and competitive ability in seedlings are associated with key aspects of plant life history such as life-span and body size at maturity. These traits that are sometimes, but not always, related to fire response life histories.     

Biodiversity responses to vegetation structure in a fragmented landscape: ant communities in a peri-urban coastal dune system

Habitat fragmentation often results in significant degradation of the structure and composition of remnant natural vegetation, leading to substantial biodiversity decline. Ants are an ecologically dominant faunal group known to be sensitive to vegetation degradation following fragmentation. We examined ant diversity and composition in relation to changes in vegetation structure in remnant coastal vegetation in the global biodiversity hotspot of southwestern Western Australia. The key features of vegetation structure driving the species and functional diversity and composition of ant communities were measures of cover of vegetation and bare ground. However, these effects were highly idiosyncratic at the species level. Cluster analyses based on plant species composition classified plots into two groups corresponding to relatively intact and degraded vegetation respectively. Although systematic changes in plant diversity and vegetation structure were observed between the two groups, key features from an ant perspective (native plant cover and bare ground) remained unchanged. Vegetation degradation consequently had little overall effect on ant species composition and functional diversity. The major disturbance–related impact on ant communities was through invasion by exotic ants, especially Pheidole megacephala; however, this occurred only in close proximity to development. Our results suggest that the priority for conserving ant diversity in our coastal dune system is the prevention of invasion by exotic species.     

Community Characteristics of Early Recovery Vegetation on Abandoned Lands of Shifting Cultivation in Bawangling of Hainan Island, South China

Shifting cultivation is a major form of agricultural practice in most parts of tropical regions worldwide. In places where the bush fallow period is excessively shortened or the period of cultivation is extended for too long, the rate of vegetation recovery and biodiversity on abandoned lands of shifting cultivation would decline. The recovery of the secondary plant communities could even be inhibited for a prolonged period because of grass occupancy. Because of the vital significance of the early recovery communities to secondary succession, we studied the community characteristics of early recovery vegetation on abandoned lands of shifting cultivation in Bawangling of Hainan Island. Measurements were made of the community composition and structure of early recovery vegetation. The sprouting abilities of different functional groups and different species in the same functional group, and the effect of the grass functional group on the composition and quantitative characteristics of tree and shrub functional groups were analyzed. Results indicated that only a few families, genera, or species apparently dominated in the early recovery vegetation on the abandoned lands of shifting cultivation and that deciduous species occurred with a rather high percentage in this early recovery community compared with the natural secondary or old growth forests. Smallsized individuals dominated the woody community. The abundance and basal area of sprouting stems for species in the tree functional group were greater than those of seeder stems, whereas the abundance and basal area of resprouters and seeders for species in the shrub functional group did not differ. The total abundance of stems for the community, stem abundances for species in tree or shrub functional groups, and for seeder or resprouter stems were all negatively correlated with coverage of the grass functional group. The mean sprouting ability in the tree functional group was greater than in the shrub functional group. The sprouting ability for different species in the same functional group was also significantly different.