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.     

Is there a cost to resprouting? Seedling growth rate and drought tolerance in sprouting and nonsprouting Ceanothus (Rhamnaceae)

Many woody plant species that depend upon fire-cued seed germination lack the ability to resprout. As the ability to resprout is widely assumed to be the ancestral condition in most plant groups, the failure to sprout is an evolutionary derived trait. Models for the evolutionary loss of sprouting assume a trade-off between seedling success and vegetative resprouting ability of adults. Such models require higher seedling success rates in nonsprouters than in sprouters. On the other hand, there seem to be few a priori reasons why a strong sprouter might not also have highly competitive post-fire seedlings. To test the hypothesis that nonsprouting plants have higher growth rates and/or drought survival, we grew seedlings of Ceanothus tomentosus from sprouting and nonsprouting populations in a common garden experiment. Each of these C. tomentosus populations was paired with a sympatric Ceanothus species that differed in resprouting ability. Sprouters exhibited greater allocation to root carbohydrate storage than did nonsprouters, but overall relative growth rates did not differ. Nonsprouters had earlier onset of flowering. These results provide mixed support for models of a sprouting/nonsprouting allocation trade-off.     

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.     

How to tell a shrub from a tree: A life‐history perspective from a South African savanna

The ecological differences between ‘shrubs’ and ‘trees’ are surprisingly poorly understood and clear ecological definitions of these two constructs do not exist. It is not clear whether a shrub is simply a small tree or whether shrubs represent a distinct life‐history strategy. This question is of special interest in African savannas, where shrubs and trees often co‐dominate, but are often treated uniformly as ‘woody plants’ even though the tree to shrub ratio is an important determinant of ecosystem functioning. In this study we use data from a long‐term fire experiment, together with a trait‐based approach to test (i) if woody species usually classified as shrubs or trees in African savanna differ in key traits related to disturbance and resource use; and (ii) if these differences justify the interpretation of the two growth forms as distinct life‐history strategies. We measured for 22 of the most common woody plant species of a South African savanna 27 plant traits related to plant architecture, life‐history, leaf characteristics, photosynthesis and resprouting capacity. Furthermore we evaluated their performance during a long‐term fire experiment. We found that woody plants authors call (i) shrubs; (ii) shrubs sometimes small trees; and (3) trees responded differently to long‐term fire treatments. We additionally found significant differences in architecture, diameter‐height‐allometry, foliage density, resprouting vigour after fire, minimum fruiting height and foliar δ¹³C between these three woody plant types. We interpret these findings as evidence for at least two different life‐history‐strategies: an avoidance/adaptation strategy for shrubs (early reproduction + adaptation to minor disturbance) and an escape strategy for trees (promoted investment in height growth + delayed reproduction).     

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.     

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.     

Resprouting as a life history strategy in woody plant communities

Resprouting is an efficient means by which woody plants regain biomass lost during disturbance, but there is a life history trade-off that occurs in all disturbance regimes between investment in the current generation through resprouting vs investment in future generations at the same or more distant sites. The relative allocation to resprouting vs seeding in woody plant communities is dictated by the nature of disturbance regimes. Resprouting is the predominant response to the least severe disturbance regimes, but is also a common response in disturbance regimes of high severity, those that destroy most or all above-ground biomass, and which occur at medium to high frequency. The response to disturbance either by resprouting or seeding is dictated by the site's productivity. We present a comprehensive model for relative allocation to resprouting vs seeding across a range of disturbance regimes. Competition between plants that mostly seed vs those that mostly resprout should accentuate differences in allocation along a gradient of disturbance frequency. However the patchy nature of disturbance in time and space, coupled with gene flow among populations undergoing different disturbance regimes, ensures that it is unlikely that either resprouting or seeding will be the sole response in most plant communities at most disturbance frequencies. Additional influences on resprouting in woody plant communities include changes in allocation during the lifespan of individual plants and phylogenetic constraints that are expressed as biogeographic patterns.     

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.     

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     

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.     

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.     

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.     

How do drought and fire influence the patterns of resprouting in Australian deserts?

Rainfall is the key driver of woody cover and life-history attributes in arid grassy biomes where disturbance is mostly rare and of low intensity. However, relatively little is known about the causes of woody community assembly in arid systems that are subject to periodic intense fire disturbance. In the central Australian desert region, grassland and shrubland fire can occur following above average rainfall. Patterns of species regeneration response (resprouting vs. reseeding) are poorly documented in this region. We tested the effects of rainfall and fire on species’ resprouting response across the latitudinal rainfall–fire gradient using constrained ordination of 385 sites and general linear models. A resprouting response was significantly greater in grassland habitat as well as at the high end of the rainfall–fire gradient. The frequency of epicormic stem resprouting also increased along the rainfall–fire gradient. We attribute this pattern to the combined effects of frequent fire and rapid gap closure on seedlings of slow-growing, fire-killed woody species in higher rainfall grasslands. In addition, we also demonstrated that rapidly maturing fire-recruiting species are similarly favoured by high fire disturbance. In arid grassy ecosystems, unlike in mesic savanna, flammable grassland supports a mix of resprouting and recruitment functional types, and habitat membership cannot be predicted by resprouting capacity. Regions, such as central Australia, that are characterised by grassland–shrubland mosaics of high and low fuel biomass, respectively, pose specific challenges to fire ecology research that are possibly best dealt with by focussing modelling at the habitat scale.     

Land‐use and fire history effects on post‐fire vegetation dynamics in eastern Spain

Question: Is post‐fire, medium‐term vegetation dynamics determined by land‐use or fire history prior to fire? Location: South‐facing slope in the Gallinera valley, Alicante province, eastern Spain. Methods: After mapping the land‐use and fire history of the study site using photo‐interpretation, we sampled vegetation structure on a set of plots representing the most frequent land‐use and fire history combinations on an area burned six years before sampling. We studied the effects of land‐use history, comparing the one‐fire land‐use trajectories. We analysed the effects of fire history; comparing one‐ and two‐fire plots for both previously cropped and uncropped areas. Results: Most variables were not significantly different between the earliest abandoned plots (abandoned at least 38 years before the fire) and the uncropped plots. On the most recently abandoned plots (abandoned between one and four years before the fire), the therophyte richness and the ratio of seeder: resprouter richness were significantly greatest. Different fire recurrences did not determine different post‐fire vegetation on either the uncropped or the early abandoned plots (all dominated by fire‐recruited seeder shrubs). The most recently abandoned plots had a lower resilience to fire. Conclusions: Land‐use history and recent pre‐fire land use, in particular, determined the post‐fire vegetation in the medium term. The vegetation composition converged during secondary succession among land‐use histories. Increasing fire recurrence had a small effect on mature plant communities, due to the combination of life‐history traits determining the response to fire of the dominant species.     

Fire severity and nutrient availability do not constrain resprouting in forest shrubs

Plants often survive disturbances such as fire by resprouting which involves having both protection traits and carbohydrate storage capacity. Protection traits not only act directly to insulate meristems but also prevent combustion of carbohydrate stores. Rapid stem growth also allows replenishment of carbohydrate stores ensuring persistence through another event. Resource availability may, however, constrain the ability to develop resilience to high-severity fires through either nutrient limitation or light limitation. We tested whether fire severity influenced resprouting ability of woody plants in two contrasting environments, low nutrient dry sclerophyll forest and more fertile wet sclerophyll forest. We tested which fire protection and growth traits were associated with resprouting ability (27 species) and resprouting vigour (16 species). Fire severity did not limit the ability of most species to resprout in either forest type. There was no generalized protection syndrome for surviving top kill, but combinations of bud protection and growth together with storage capacity appear to drive resprouting ability. In nutrient-limited forests, low specific leaf area (SLA) may reduce stem growth in resprouters, causing more reliance on bud protection through bark thickness. Conversely, in the more fertile forests, where light becomes limiting with time-since-fire, high SLA appears to increase the capacity for rapid stem growth with less emphasis on developing thicker bark. These different syndromes appear to be adaptive as fire severity did not influence survival in either forest type.     

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.     

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.     

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.     

No allocation trade-offs between flowering and sproutingin the lignotuberous,Mediterranean shrub Erica australis

Trade-offs between allocation to sexual or vegetative regeneration capacity are well established as a driving force in the life history patterns of plants in fire-prone environments. However, it is not known whether such trade-offs exist in plants which after aboveground removing disturbances, such as fire, may regenerate by sexual (seeding) or asexual (sprouting) mechanisms. We evaluated whether in the fire-recruiting resprouter Erica australis, which after fire can regenerate by seedling establishment or resprouting, a larger investment in flowers and seeds prior to being disturbed by clipping its aboveground parts would decrease subsequent sprouting, that is, its vegetative regeneration capacity. We analysed the relationships between flower and seed production and the ensuing production and growth of sprouts of six plants from thirteen different sites in central-western Spain. We found no significant relationships between measures of sexual reproductive effort and resprout production and growth 6 months after clipping the aboveground parts of the plants. No evidence of trade-offs between sexual and asexual efforts was found. Furthermore, no significant relationship was found between lignotuber total non-structural carbohydrates and sexual reproductive effort. In addition, 2 years after the disturbance, resprout biomass was positively and significantly correlated with sexual reproductive effort prior to the disturbance. This indicates that growth of resprouts was higher at the sites where plants made a greater reproductive effort. The sites that were more favourable to producing flowers and seeds could also be more favourable to resprouting.