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.     

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.     

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.     

Fire, rain and the selection of seeder and resprouter life-histories in fire-recruiting, woody plants

Several Cape species of the genus Erica are known to present seeder and resprouter phenotypes, and this variation seems to have a genetic basis. Therefore, this genus provides ideal model systems for using to elucidate the evolution of nonsprouting or seeder and resprouter life-histories in woody, fire-recruiting plants. A simple simulation model was developed to identify, under life-history optimality, the ecological conditions (viz. rainfall conditions and fire frequency) that conferred a selective advantage to the seeder phenotype over the resprouter in a given Cape Erica species. The model illustrated that the seeder life-history was able to invade and replace a resprouter population only under a mild mediterranean climate, with short, moderate summer droughts. This simulation approach will contribute to a better understanding of the biogeographical pattern of seeder and resprouter lineages of one of the paradigmatic fynbos woody taxa throughout the Cape floristic region.     

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.     

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.     

Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

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.     

Resprouting Ability of Six Atlantic Shrub Species

Resprouting from subterranean structures is a principal method of vegetative regeneration that many shrub species show after a disturbance. This study, therefore, aims to determine the resprouting capacity and intensity of six dominant species in an Atlantic shrubland area located in the NW of the Iberian Peninsula and compare their resprouting and germinating strategies. Resprouting intensity is measured using three variables: individual probability of resprouting, number of resprouts and length of these sprouts in three plant-age classes. The intensity for each species was calculated using a simple index (IRI) that included the three measured variables. All studied species, excepting Erica umbellata, could resprout. According to the IRI values, there are three groups of species: strong resprouters (Ulex europaeus, Ulex minor and Pterospartum tridentatum), weak resprouters (Ulex micranthus and Genista triacanthos) and non-resprouters (E. umbellata). The germination of strong resprouters is highly stimulated by fire. Frequent disturbances remove the non- and weak resprouter populations and promote the strong resprouter ones.     

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.     

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.     

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.     

Population biology and regeneration of forbs and shrubs after fire in Brazilian <Emphasis Type="Italic">Campos</Emphasis> grasslands

Fire is an important factor in several ecosystems, affecting plant population biology. Campos grasslands are under constant influence of disturbance, mostly grazing and fire. However, few studies evaluated the effect of fire on plant population biology of grassland species. Therefore, we aim to analyze the effect of fire on the population biology of four species, from different functional groups and regeneration strategies: Chaptalia runcinata (forb, resprouter, absence of belowground organ), Vernonia flexuosa (forb, resprouter, presence of rhizophore), Eupatorium ligulaefolium (shrub, resprouter, presence of xylopodium) and Heterothalamus psiadioides (shrub, obligate seeder). Seven plots were established in different sites in southern Brazil: frequently burned (FB) and excluded from fire since 6 years (E). All plots were subjected to controlled burns during summer. Before experiments, populations were sampled. Further observations were carried out after 90 and after 360 days of fire experiments. In addition, we counted the number of seedlings and resprouters recruited after fire. Heat shock experiments were conducted with two species (H. psiadioides and V. flexuosa), as well as the study of the bud bank of the following species: E. ligulaefolium and V. flexuosa. The obligate seeder species had all individuals killed by fire and established only after 1 year. Resprouters, however, showed new stems immediately after fire. E. ligulaefolium and V. flexuosa showed only vegetative regeneration from belowground organs and more individuals in excluded sites 1 year after the fire. The bud bank of E. ligulaefolium tended to be larger in excluded sites, whilst V. flexuosa showed an opposite result. High temperatures did not enhance nor kill seeds from both studied species. Vegetative regeneration was the most important strategy for all studied species, except for H. psiadioides, the obligate seeder species. Fire thus, plays an important role on population structure and demography, being also important for plant recruitment.     

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.     

Postfire regeneration of resprouting mountain fynbos shrubs: differentiating obligate resprouters and facultative seeders

Plant species in Mediterranean-type climate regions have a diversity of traits that facilitate their persistence under a given fire regime. Obligate resprouters (OR) are dependent on resprouting to persist through a burn episode, as their seeds are killed by fire. Facultative seeders (FS) combine strategies by resprouting and recruiting new seedlings after fire. We hypothesised that these life history differences would lead to differential resprout success and we predicted that OR would be more successful than FS. We performed a 2-year study to assess resprout success of co-occurring Western Cape mountain fynbos FS and OR species, and to determine predictors of resprout success following a wildfire. All the FS species recruited seedlings postfire, whereas the OR did not. OR demonstrated near complete (99 %) survival after 2 years and all resprouted within 4 months postfire. In contrast, only 81 % of FS resprouted with only 65 % surviving 2 years postfire. Numerous factors were linked to resprout success: decreased lignotuber exposure and pre-fire vigour (number of pre-fire shoots) were significantly associated with postfire resprouting, while early resprouting (days to first resprout) and growth rate were significant predictors of post-resprout survival. The difference in resprout survival between the FS and OR may also be partially due to phylogenetic differences. These findings confirm the heterogeneity and complexity of postfire resprouting and support the distinction of OS and FS life history types.     

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.     

Seedling Growth and Storage Characteristics of Seeder and Resprouter Species of Mediterranean-type Ecosystems of S. W. Australia

Juvenile (2–4 years old) plants of a taxonomically diverserange of dicotyledonous species were examined following recruitmentfrom seed in recently burnt habitats in S.W. Australia. Obligateseeder species (those succumbing to fire) had on average, analmost threefold greater total plant d. wt and more than a fourfoldgreater shoot: root d. wt ratio than comparably-aged, cohabiting,resprouter species (those capable of surviving fire). Starchwas generally much more concentrated in root dry matter of resproutersthan seeders, and both categories exhibited greater starch storagecapacity in roots than shoots. Members of the Myrtaccae wereexceptional in not showing a greater root starch reserve inresprouter than in seeder species. and in carrying as high,or higher, starch levels in shoots as in roots. Anatomical investigationson roots provided instances of zero starch storage, storage,only in rays or in cortex, in rays and in xylem parenchyma,in rays and in cortex, or in all three locations. High starchratings of resprouter roots related mostly to higher starchgrain packing density at storage sites, but in certain instancesthese also reflected proportionally greater areas of tissuespecifically devoted to storage. Dry matter of shoots of bothseeders and resprouters generally contained higher levels ofN, P, K, Ca and Mg than that of roots, but there was no significantevidence of elements being more concentrated in resproutersthan in seeders. Fire response, seedlings, resprouter, obligate seeder, shoot: root ratio, starch storage, mineral nutrition     

Drivers of post-fire resprouting success in restored Banksia woodlands

Disturbances can alter persistence trajectories of restored ecosystems. Resprouting is a common response of plants to disturbances such as fire or herbivory. Therefore, understanding a plant's resprouting response can inform successful restoration. We investigated patterns and drivers of resprouting following fire in fire-prone Banksia woodlands restored after sand mining in the Mediterranean-climate region of Western Australia. We applied experimental fire to samples of nine species with different resprouting types (rhizome, root crown, root sucker and lignotuber) across a 4- to 27-year-old restoration chronosequence. We investigated the influence of pre-fire plant size, restoration age and soil conditions on resprouting success, defined by: (i) the probability of resprouting (measured ~5 months after fire), (ii) the probability of surviving the first summer and, (iii) vigour (both measured ~12 months post-fire). We found that the probability of initial resprouting was high across most species, but summer survival was lower but comparable to that in other post-mining restored ecosystems following fire. Generally, pre-fire plant size did not influence probability of resprouting, while size and soil conditions were important for two species survival. Pre-fire plant size was a significant predictor of vigour for all species with soil conditions influencing four species. Restoration age significantly influenced survival of three species. However, as our models explained low amounts of variation in probabilities of resprouting and survival (R² = <0.11), other factors influencing resprouting success remain unidentified. Resprouting response to fire disturbance in restored Banksia woodlands are species and resprouter type specific, with plant size and soil conditions potentially more informative for understanding responses to disturbances than restoration age alone.