Evolutionary transition from resprouter to seeder life history in two Erica (Ericaceae) species: insights from seedling axillary buds

BACKGROUND AND AIMS: The genus Erica represents the epitome of plant biodiversity in the South African Cape region. It includes seeder and resprouter species, but both species diversity and narrow endemism are tightly associated with the seeder habit. It also includes 'mixed' species, in which both seeder and resprouter life histories are found. This intraspecific variation in life history is genetically based. METHODS: The cotyledonary region and basal stem of seeder and resprouter seedlings of two 'mixed' species, Erica calycina and E. coccinea, were examined to detect morphological and anatomical differences in axillary bud development between regeneration forms. KEY RESULTS: While at least some bud activity was observed for resprouter seedlings, none was detected for seeder seedlings. A closer examination allowed the detection of some axillary buds in seeder seedlings of the two species, but they appeared in an unequivocally atrophied state. CONCLUSIONS: The seeder and resprouter life histories are two character states and the seeder one (i.e. loss of resprouting) is derived within these two Erica species. Results allow the hypothesis that the loss of resprouting in a fire-prone scenario such as the Cape fynbos has promoted high diversification rates in seeder Erica lineages.     

Root starch storage and allocation patterns in seeder and resprouter seedlings of two Cape Erica (Ericaceae) species

Post-fire sprouting of dormant buds in resprouter plants is facilitated by stored carbohydrate reserves, with starch being the critical reserve. Starch is mainly stored in xylem parenchyma ray tissue of woody underground organs, such as burls, lignotubers, and roots. We carried out a comparative analysis of the pattern of starch storage and the proportion of parenchymatic ray tissue in the upper root or cotyledonary region of seedlings from seeder and resprouter forms within two Cape Erica (Ericaceae) species: E. coccinea L. and E. calycina L., which were raised in the greenhouse under controlled irrigation. We also explored the root-to-shoot allocation patterns of seeder and resprouter seedlings in these two species. Resprouter seedlings of both species showed higher relative amounts of upper-root starch and upper-root storage tissue as well as a higher root-to-shoot allocation than their seeder counterparts. Pronounced swelling of the upper root region suggests ontogenetic development of a lignotuber in the resprouter forms of the two Erica species. The distinct allocation of starch in roots seems to be genetically determined and would account for the apparent differences in the root-to-shoot allocation patterns between both regeneration forms from the early seedling stage.     

Underground starch storage in Erica species of the Cape Floristic Region – differences between seeders and resprouters

Concentrations of starch in roots of seeder species of Erica from the Cape Floristic Region, South Africa were found to be considerably less than in resprouters. Shoot starch was highly variable but mean values were similar in both seeder and resprouter species of Erica . Three distinct patterns of starch storage in roots were recognized. All seeder species fell within definitions of Categories 1 (narrow major and minor parenchymatous rays, one to two cells wide with no inter-ray storage) or 2 (thick major rays up to seven cells wide and thin minor rays with small amounts of inter-ray storage) whereas resprouter species were consistently within Categories 2 or 3 (broad major and minor rays, up to eight cells wide and conspicuous inter-ray starch storage). Results are discussed in light of similar studies of the related Epacridaceae. 'Mixed' species (i.e. with seeder or resprouter individuals present, often in distinct populations) were always classified as belonging to Category 2. Studies of populations of three 'mixed' species confirmed that seeder forms had consistently lower amounts of root starch than resprouters. Rays of xylem parenchyma were the main sites for starch storage in roots of both seeders and resprouters and greater proportions of cross-sectional area of roots were consistently devoted to such storage tissues in resprouter forms of the three 'mixed' species. Analyses of a number of seeder and resprouter Erica coccinea populations showed that differences in amounts of realised and potential root starch storage are best explained by the effect of regeneration behaviour rather than by among-population variability.     

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.     

Isolation and characterization of microsatellite loci in the Cape fynbos heath Erica coccinea (Ericaceae)

Enriched genomic libraries were used to isolate and characterize dinucleotide microsatellite loci in Erica coccinea, a South African Cape fynbos heath species with distinct resprouter and seeder populations. Microsatellites were required to investigate the effect of the contrasting demographic pattern driven by these two post-fire responses in the population genetic structure of seeder and resprouter forms within this species. Eight microsatellite loci were characterised and amplified a total of 106 alleles in 2 samples each of 30 individuals from 1 resprouter and 1 seeder population. Mean allele numbers were 7.88 and 11.0 for the resprouter and seeder population, respectively. Both populations showed similar average observed and expected heterozygosity levels, H _O(resprouter) = 0.683, H _O(seeder) = 0.696; H _E(resprouter) = 0.726, H _E(seeder) = 0.756, and average positive inbreeding coefficients F _IS(resprouter) = 0.058, F _IS(seeder) = 0.080. This set of microsatellite loci will be used to conduct a population genetic survey of seeder and resprouter populations throughout the range of the species. Cross-species transferability was also assayed in four other South African and four European species of the genus Erica, supporting their potential use for population genetic analyses.     

Are woody seeder plants more prone than resprouter to population genetic differentiation in Mediterranean-type ecosystems?

High diversification of woody seeder lineages is characteristic of the south-western cape floristic region (CFR), South Africa, which has been explained as a consequence of its mild Mediterranean climate and reliable winter rainfall. Such climatic regime reduces the risk of post-fire recruitment failure, acting as an ecological filter that favours seeder populations, thus promoting genetic differentiation and diversification in seeder populations, as previously seen in the South African heath Erica coccinea. To explore this hypothesis further, genetic population structure was investigated in two Mediterranean Erica species, one seeder (Erica umbellata) and the other resprouter (Erica australis), using nuclear microsatellites. These two species are endemic to the western Mediterranean Basin and co-occur in heathlands of the Strait of Gibraltar region. Mean annual rainfall in this region is similar to that from the south-western CFR, but summer stress is more marked and winter rainfall is much less reliable. Contrary to what was found in E. coccinea, average genetic diversity levels were considerably lower in seeder populations (E. umbellata), regardless of an apparently higher gene flow among them. No differences in genetic differentiation among populations were found between the two species. The occurrence of less favourable climatic conditions for post-fire recruitment in the western Mediterranean compared to the south-western CFR may affect seeder populations more strongly than resprouter and may thus account for lower levels of within-population genetic diversity in the seeder E. umbellata. In addition, putatively higher migration rates in the seeder E. umbellata, may contribute to reduce its potential for genetic differentiation. This study provides evidence that high divergence of seeder populations is not a general rule in fire-prone, Mediterranean-type ecosystems.     

POSTFIRE RESPONSE AND GENETIC DIVERSITY IN ERICA COCCINEA: CONNECTING POPULATION DYNAMICS AND DIVERSIFICATION IN A BIODIVERSITY HOTSPOT

Understanding the processes of biological diversification is a central topic in evolutionary biology. The South African Cape fynbos, one of the major plant biodiversity hotspots out of the tropics, has prompted several hypotheses about the causes of generation and maintenance of biodiversity. Fire has been traditionally invoked as a key element to explain high levels of biodiversity in highly speciose fynbos taxa, such as the genus Erica. In this study, we have implemented a microevolutionary approach to elucidate how plant‐response to fire may contribute to explain high levels of diversification in Erica. By using microsatellite markers, we investigated the genetic background of seeder (fire‐sensitive) and resprouter (fire‐resistant) populations of the fynbos species Erica coccinea. We found higher within‐population genetic diversity and higher among‐population differentiation in seeder populations and interpreted these higher levels of genetic diversification as a consequence of the comparatively shorter generation times and faster population turnover in the seeder form of this species. Considering that genetic divergence among populations may be seen as the initial step to speciation, the parallelism between these results and the pattern of biodiversity at the genus level offers stimulating insights into understanding causes of speciation of the genus Erica in the Cape fynbos.     

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.     

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.     

Strong signature of selection in seeder populations but not in resprouters of the fynbos heath Erica coccinea (Ericaceae)

A higher frequency of natural selection is expected in populations of organisms with shorter generation times. In fire‐prone ecosystems, populations of seeder plants behave as functionally semelparous populations, with short generation times compared to populations of resprouter plants, which are truly iteroparous. Therefore, a stronger signature of natural selection should be detected in seeder populations, favoured by their shorter generation times and higher rates of population turnover. Here we test this idea in Erica coccinea from the Cape Floristic Region, which is dimorphic for post‐fire regeneration mode. We measured three floral traits supposedly subject to natural selection in seeder and resprouter populations. We then compared phenotypic trait variation with neutral genetic variation in each group of populations using P_ST–F_ST comparisons to detect signatures of natural selection in seeders and resprouters. We found a strong signature of selection in seeder populations, but not in resprouters. Furthermore, anthers of seeders were more exserted (and larger) than those of resprouters. These differences were maintained at sites where seeders and resprouters co‐occurred, suggesting that phenotypic plasticity or adaptation to different growth environments are unlikely explanations for trait variation. These results provide empirical support for the hypothesis that the genetic signature of natural selection is certainly more intense in seeder than in resprouter populations, favoured by their comparatively faster generation turnovers. Increased frequency of natural selection would increase differentiation among populations, thus promoting speciation in pyrophytic seeder lineages of the Cape flora.     

Fire regenerative syndromes of forest woody species across fire and climatic gradients

Community resilience after fire is determined by species’ ability to regenerate through two main mechanisms growth of new sprouts (resprouter species) and germination from surviving seed banks or from seeds arriving from neighbouring populations (seeder species). Both the mechanisms are present in Mediterranean communities. The occurrence of both the types in a community depends on fire history and the bio-geographical history determining the available species pool. Regenerative traits also covary with other functional attributes associated with resource acquisition and stress tolerance. As post-fire regenerative responses can be related to various ecological factors other than fire, we tested the hypothesis of a different proportional representation of post-fire regenerative syndromes in forest woody species along a climatic gradient in Catalonia (NE Spain) ranging from Mediterranean to temperate-boreal climates. Specifically, we expected seeder species to become less common with colder and moister conditions while resprouters would not be so influenced by the climatic gradient. We also tested the hypothesis of change in the relative abundance of regenerative syndromes in relation to recent fire history. We analysed a large database obtained from extensive forestry surveys and remote sensing fire records. After correction for spatial autocorrelation, we found an increase in the proportion of seeder species under more Mediterranean conditions and a decrease in fire-sensitive species (with no efficient mechanisms of post-fire recovery) in moister conditions. Resprouter species were similarly present across the whole gradient. A similar pattern was observed after excluding recently burnt plots. Therefore, post-fire regenerative syndromes segregate along the climatic gradient. Recent fires reduced the occurrence of fire-sensitive species and increased the proportion of seeder species. No significant effect was observed on resprouter species. Fire has a sorting effect, shaping the occurrence of species with different regenerative traits. Overall, fire seems to explain better the variability of the proportion of fire-sensitive species and climate the variability of seeder species. In addition, other factors (forestry practices and the covariation between regenerative and functional attributes) are likely to contribute to the regional pattern of regenerative syndromes.     

Linking post-fire regenerative strategy and leaf nutrient content in Mediterranean woody plants

We explored differences in leaf nutrient contents between species from Mediterranean shrublands with the ability to resprout after disturbances such as fire (resprouters) and others without this capacity (non-resprouters). Since it is to be expected that resprouting capacity is related to a more conservative use of nutrients, we hypothesize that resprouter and non-resprouter species will differ in their leaf nutrient concentrations.We measured the following leaf traits: leaf carbon content (LCC), leaf nitrogen content, leaf phosphorous content (LPC), leaf potassium content, leaf calcium content (LCaC), leaf magnesium content and leaf sodium content, in 30 woody species coexisting in a coastal shrubland. We also considered the influence of species’ taxonomic affiliation in our analysis.Non-resprouters had higher LPC and LCaC than resprouter species, and lower LCC, which could be related to their cell and life-history properties. This study also suggests that non-resprouter species have more P in their leaves and are less limited by P than resprouter species.Overall, the differences in leaf nutrient contents suggest that shifts in the proportion of resprouter and non-resprouter species resulting from changes in the fire regime may have effects on the functional properties of the ecosystem.     

Response to recurrent disturbance in two co-occurring resprouter heath species: the ecological consequences of withstanding herbivores

Resprouting woody plants are vulnerable to large mammal herbivores in the early stages of regeneration after disturbance (e.g., fire, slashing), because herbivory reduces the storage of below-ground starch reserves and may thus compromise regeneration success. Defence from herbivory may incur a cost to resprouter plants in terms of lengthening the time to recovery of pre-disturbance starch levels. Erica scoparia and E. australis (Ericaceae), two resprouter woody species that co-occur in Mediterranean heath-communities of the Strait of Gibraltar, differ in the way they cope with herbivory: E. scoparia is browsing-tolerant and E. australis is a browsing-avoider. Our experimental field study demonstrated that exposure to a moderate density of ungulate herbivores markedly lowered starch re-storage after resprouting in E. scoparia, but not in E. australis. Nevertheless, time to recover pre-disturbance starch levels was shorter in the tolerant E. scoparia, even under exposure to herbivores. The slower starch recovery rate of E. australis is a likely outcome of its resource-demanding browsing-avoidance mechanism. Differences in starch recovery rates after disturbance between the two Erica species partly explains their ecological segregation at the landscape scale.     

Burning phylogenies: fire, molecular evolutionary rates, and diversification

Mediterranean-type ecosystems are among the most remarkable plant biodiversity "hot spots" on the earth, and fire has traditionally been invoked as one of the evolutionary forces explaining this exceptional diversity. In these ecosystems, adult plants of some species are able to survive after fire (resprouters), whereas in other species fire kills the adults and populations are only maintained by an effective post-fire recruitment (seeders). Seeders tend to have shorter generation times than resprouters, particularly under short fire return intervals, thus potentially increasing their molecular evolutionary rates and, ultimately, their diversification. We explored whether seeder lineages actually have higher rates of molecular evolution and diversification than resprouters. Molecular evolutionary rates in different DNA regions were compared in 45 phylogenetically paired congeneric taxa from fire-prone Mediterranean-type ecosystems with contrasting seeder and resprouter life histories. Differential diversification was analyzed with both topological and chronological approaches in five genera (Banksia, Daviesia, Lachnaea, Leucadendron, and Thamnochortus) from two fire-prone regions (Australia and South Africa). We found that seeders had neither higher molecular rates nor higher diversification than resprouters. Such lack of differences in molecular rates between seeders and resprouters-which did not agree with theoretical predictions-may occur if (1) the timing of the switch from seeding to resprouting (or vice versa) occurs near the branch tip, so that most of the branch length evolves under the opposite life-history form; (2) resprouters suffer more somatic mutations and therefore counterbalancing the replication-induced mutations of seeders; and (3) the rate of mutations is not related to shorter generation times because plants do not undergo determinate germ-line replication. The absence of differential diversification is to be expected if seeders and resprouters do not differ from each other in their molecular evolutionary rate, which is the fuel for speciation. Although other factors such as the formation of isolated populations may trigger diversification, we can conclude that fire acting as a throttle for diversification is by no means the rule in fire-prone ecosystems.     

Relationships Between Fire Response, Morphology, Root Anatomy and Starch Distribution in South-west Australian Epacridaceae

Selected epacrids (92 species in 15 genera) were examined withrespect to fire response type, morphology, root anatomy andstarch storage. Seeders, 75% of the species investigated, possesseda single main stem and a small root system with lateral rootswhich in most cases did not spread beyond the shoot canopy.Resprouter species were generally multi-stemmed with large lignotuberousroot stocks. Certain seeder and resprouter species were intermediatein form and showed small root systems and basally branched mainstems. Amounts of starch in roots of seeders (1.9±0.5mgstarch gd.wt per root) were much less than in resprouters (14.1±3.3)whereas amounts in shoots were similar (1.9±0.5 and 1.6±0.6mgstarch gd.wt per shoot, respectively). Starch storage in rootswas mostly confined to rays of xylem parenchyma and inter-rayxylem parenchyma and the greater storage capacity of resprouterswas generally due to broader rays. Growth zones in root xylemranged from clear, verifiable annual rings, as in many seederspecies, to indistinct growth zones, typical of many resprouterspecies. Shoot:root dry weight ratios were higher in seedersthan resprouters. The study suggests that speciation withinthe Epacridaceae into seeder and resprouter forms involved divergentdifferentiation in terms of morphology, shoot:root dry weightratio root storage of starch. Seeder; starch storage; growth rings; growth zones; south-west Australia; resprouter; Epacridaceae     

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.     

Experimental drought and warming decrease diversity and slow down post-fire succession in a Mediterranean shrubland

Plant community recovery (species richness, diversity and composition) of a post-fire Mediterranean shrubland was monitored over a seven year period (1998–2005) under experimental drought and warming that simulated the environmental conditions forecast for this area in the coming decades. Species richness and Shannon's index were positively correlated with accumulated precipitation in the growing season and both variables were negatively affected by reduced water availability in drought plots. The relative abundance of the different species in both treatments was linearly correlated with their relative abundance in control plots. Moreover, we found species-specific responses to treatments. Drought and warming treatment reduced the competitive ability of the obligate seeder tree Pinus halepensis against native resprouter shrubs and consequently, the transformation from shrub to pine tree dominated vegetation was slowed down. Conversely, the water use strategy of Globularia alypum may allow this species to maintain a dominant position in drought plots. Therefore, future drier and warmer conditions in Mediterranean areas may severely affect plant community recovery after a disturbance, due to the existence of both abundance-dependent and species-specific responses that may change inter-specific competitive relationships.     

Enhancement of photosynthesis in post-disturbance resprouts of two co-occurring Mediterranean <Emphasis Type="Italic">Erica</Emphasis> species

The higher growth rates of resprouting shoots compared with those of mature plants in resprouter woody species are supported by higher rates of photosynthesis and transpiration. In this contribution we hypothesize that species with higher resprouting vigour will show a larger enhancement of photosynthesis in resprouting shoots. We test this hypothesis by comparing gas exchange and leaf parameters between resprouting and mature plants in Erica scoparia and E. australis. These two Erica species co-occur in Mediterranean heathlands of the Strait of Gibraltar. Erica scoparia has a higher rate of post-disturbance starch recovery than E. australis, which makes it more resistant to recurrent disturbance. We tested the hypothesis that enhancement of photosynthesis and water use characteristics of resprouting shoots compared with mature plants should be more pronounced in E. scoparia. In both species, resprouts had higher efficiency in the use of light and higher maximum net photosynthesis than mature shoots. However, contrary to expectations, differences in the photosynthetic performance between resprouts and mature plant shoots were larger in E. australis. Higher root to shoot ratios in resprouting E. australis plants, determined by their slower above-ground recovery, together with stronger demand from carbon sinks might explain this result.     

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.     

Chloroplast DNA evidence for the roles of island colonization and extinction in Tolpis (Asteraceae: Lactuceae)

Tolpis consists of ～13 species native to Africa, Europe, and Macaronesia, with at least one species endemic to each of the four major archipelagos of the Azores, Madeira Islands, Canary Islands, and Cape Verde Islands. All but two of these species develop woody stems by maturity. Chloroplast DNA restriction site variation was analyzed for all species of Tolpis and four outgroups in order to understand the patterns of island colonization and evolution of woodiness in this genus. Parsimony analyses revealed a strongly supported monophyletic Tolpis. Within the genus, the following three well-supported groups were detected: all species from the Canary Islands and Cape Verde Islands, both Azorean species, and both continental species. The Canary Island/Cape Verde clade was sister to the two continental species, and the Azorean clade was sister to this group. The two Madeiran species of Tolpis occupied the basalmost positions within the genus. When biogeography was mapped onto this phylogeny, nine equally parsimonious reconstructions (five steps each) of dispersal history were detected, which fell into two groups: eight reconstructions implied that Tolpis colonized Madeira from the continent, followed by continental extinction and subsequent continental recolonization, while one reconstruction implied that Tolpis colonized Macaronesia four times. Two of the reconstructions involving continental extinction required the least amount of overall dispersal distance. The cpDNA phylogeny also suggests that woodiness arose in the common ancestor of all extant Tolpis, followed by two independent reversals to an herbaceous habit. Assuming that one of the eight reconstructions favoring continental extinction and recolonization is true, our results suggest that Tolpis may represent the first documented example of a woody plant group in Macaronesia that has recolonized the mainland in herbaceous form.