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.     

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.     

Legume seeders of the Cape Floristic Region inhabit more fertile soils than congeneric resprouters—sometimes

In fire-prone ecosystems, plants for the most part persist via either soil-stored seed banks (seeders) or below-ground storage structures (resprouters). Given their greater allocation of resources above ground to growth and reproduction, seeders are likely to have a higher nutrient requirement than resprouters. This may result in discernable differences in habitat nutrition and leaf morphology. These differences are probably accentuated in Cape legumes given their poor adaptation to low-P soils. It was hypothesized that legume seeders occupy habitats with greater fertility and possess larger, less sclerophyllous leaves than resprouters. Site nutrition and leaf morphologies were compared between seeders and resprouters in the genera Otholobium and Psoralea. There were no differences in leaf morphology between seeders and resprouters. Seeders sites had a higher total [N], exchangeable [Ca] and [Mg], and CEC, but lower [Fe] than resprouters. Only within Otholobium, did seeder sites have a higher Bray II [P]. This genus-specific variation in available P is probably a consequence of greater variation in soil type and precipitation between seeders and resprouters. Conversely, niche construction may contribute to the differences in soil fertility between seeders and resprouters in Psoralea. Thus, our data showed a general tendency for seeders to inhabit more fertile sites than resprouters. Caution is required, however, in generalizing these results, as our data indicate a difference in factors affecting soil nutrient availability between legume genera. Changes in soil fertility post-fire may limit legume persistence beyond the early stages of succession.     

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

Background and Aims

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

Methods

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

Key Results

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

Conclusions

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

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

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

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

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

Resprouters,assisted by somatic mutations,are as genetically diverse as nonsprouters in the world's fire-prone ecosystems

In fire-prone environments worldwide, resprouters mostly regenerate vegetatively after fire, whereas non (re)sprouters are killed by fire and rely entirely on stored seeds (soil or canopy storage) for regeneration. This dichotomy in post-fire regeneration strategies is a key mechanism for controlling the reproductive characteristics, demography, and population genetic structure of plant species in fire-prone ecosystems. Nonsprouters are considered to have higher within-population genetic variation than resprouters due to greater opportunities for recombination via their much greater seed production and frequent generation turnover. Empirical studies that explore this hypothesis are rare and the results are mixed. We collated published studies reporting genetic diversity measures of plant species in fire-prone ecosystems of four Mediterranean-climate regions. Ninety-two species were identified with unambiguous information on postfire regeneration type and with adequate sample sizes. We found no significant differences in population genetic diversity and structure between resprouters and nonresprouters across diversity parameters and genetic marker types, taxon groups or geographic regions. We conclude that resprouters are at least as genetically diverse as nonsprouters. We tested the hypothesis that accumulation of somatic mutations is a possible mechanism for maintaining genetic diversity among resprouters, by screening for microsatellite DNA mutations in the resprouter, Banksia attenuata. Genotyping of leaf material collected from disparate parts of individual plants demonstrated the existence of 2–5 somatic mutations among eleven microsatellite loci. The buildup of somatic mutations in meristematic buds during a resprouter's long lifespan and the considerable potential for interpopulation gene dispersal among resprouters may be key factors that enable their genetic diversity to keep pace with that of nonsprouters. Thus, resprouters and nonsprouters are equally capable of responding to natural selection, and therefore possess a similar potential to adapt to changing environmental conditions.     

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.     

Adult root structure of mediterranean shrubs: relationship with post‐fire regenerative syndrome

Life‐history attributes can impose differences on root system structures and properties related to nutrient and water uptake. Here, we assess whether plants with different post‐fire regenerative strategies (resprouters, seeders and seeder–resprouters) differ in the topological and morphological properties of their root systems (external path, altitude, magnitude, topological index, specific root length, root length, root‐to‐shoot biomass ratio, length of the main axis of the root system and link length). To achieve these objectives, we sampled individuals from eight woody species in a shrubland located in the western Mediterranean Basin. We sampled the adult root systems using manual field excavation with the aid of an air compressor. The results indicate that resprouters have a higher root‐to‐shoot ratio, confirming their higher ability to store water, starch and nutrients and to invest in the belowground biomass. Moreover, this pattern would allow them to explore deeper parts of the soil layers. Seeder species would benefit from a higher specific root length, pointing to increased relative root growth and water uptake rates. This study confirms that seeders and resprouters may differ in nutrient and water uptake ability according to the characteristics of their root system. Species that can both resprout and establish seedlings after fire had different patterns of root system structure; in particular, root:shoot ratio was more similar to resprouters and specific root length was closer to seeders, supporting the distinct functional performance of this type of species.     

Influence of summer marine fog and low cloud stratus on water relations of evergreen woody shrubs (Arctostaphylos: Ericaceae) in the chaparral of central California

Mediterranean-type climate (MTC) regions around the world are notable for cool, wet winters and hot, dry summers. A dominant vegetation type in all five MTC regions is evergreen, sclerophyllous shrubland, called chaparral in California. The extreme summer dry season in California is moderated by a persistent low-elevation layer of marine fog and cloud cover along the margin of the Pacific coast. We tested whether late dry season water potentials (Ψ(min)) of chaparral shrubs, such as Arctostaphylos species in central California, are influenced by this coast-to-interior climate gradient. Lowland coastal (maritime) shrubs were found to have significantly less negative Ψ(min) than upland interior shrubs (interior), and stable isotope (δ(13)C) values exhibited greater water use efficiency in the interior. Post-fire resprouter shrubs (resprouters) had significantly less negative Ψ(min) than co-occurring obligate seeder shrubs (seeders) in interior and transitional chaparral, possibly because resprouters have deeper root systems with better access to subsurface water than shallow-rooted seeders. Unexpectedly, maritime resprouters and seeders did not differ significantly in their Ψ(min), possibly reflecting more favorable water availability for shrubs influenced by the summer marine layer. Microclimate and soil data also suggest that maritime habitats have more favorable water availability than the interior. While maritime seeders constitute the majority of local Arctostaphylos endemics, they exhibited significantly greater vulnerability to xylem cavitation than interior seeders. Because rare seeders in maritime chaparral are more vulnerable to xylem cavitation than interior seeders, the potential breakdown of the summer marine layer along the coast is of potential conservation concern.     

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

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

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.     

Fire and resprouting in Mediterranean ecosystems: insights from an external biogeographical region, the mexical shrubland

We investigated modes of regeneration of dominant species of the mexical vegetation after fire. The mexical shrubland shows a remarkable structural, morphological, and floristic similarity to Mediterranean-type vegetation and is considered a relict of the Madro-Tertiary Geoflora under a non-Mediterranean climate. This vegetation provides an ideal scenario to test the role of fire in Mediterranean ecosystems because historical fire occurrence is absent and the species assembly is constituted mostly by Madro-Tertiary elements and Neotropical species (some of them, endemic species from Mexico). The existence of congeneric species of the California chaparral allows us to determine the regeneration ability of these communities after fire in relation to resprouting and seeding strategies, which are widespread modes reported in the Mediterranean-type vegetation. By the experimental application of fire in the two biogeographical groups of species, we tested the hypothesis that low resprouting ability of California congeneric species (Madro-Tertiary species) after fire would indicate that fire has played an important selective force in the resprouting habit. A low resprouting ability in the Neotropical group of species would suggest that fire has molded the set of species dominating fire-prone environments.Our results indicated that resprouting is a widespread trait in the mexical species characterized by the presence of lignotubers and burls. Resprouting can be considered an ancient trait, probably linked to losses of aboveground biomass, that became a pre-adaptation in Mediterranean fire-prone communities. The Neotropical group of species showed less ability to regenerate after fire, and small plants were more likely to die after disturbance in this group than in the Madro-Tertiary group. The resprouting feature and the seeder strategy of other species after a fire in the mexical shrubland are similar to Mediterranean-type ecosystems, emphasizing their common origin and the relevance of phylogenetic and biogeographical studies to explain current patterns of vegetation.     

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

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

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.     

Demography of woody plants in relation to fire: Banksia serrata Lf. and Isopogon anemonifolius (Salisb.) Knight

Banksia serrata and Isopogon anemonifolius are serotinous resprouters (single-stemmed tree, multi-stemmed shrub, respectively) found in forests within the Sydney region. Studies were conducted to predict the population dynamics of these species. Seed production and survival and the accumulation of seed-bank within cones were estimated in relation to time since fire. Emergence, survival and development of lignotubers were measured in young juveniles and the time taken to reach adulthood was estimated. This information and published data on survival were used to estimate the amount of recruitment of adults and juveniles necessary for stands to remain in a stable state under frequent (<16 years), high and low intensity fire regimes. The effects of longer intervals (up to 80 years) between fires were also estimated. It was predicted that B. serrata populations will decline in numbers when the interval between high intensity fires is <9 years, while under low intensity fires the critical interval was 12–13 years. In I. anemonifolius the predicted intervals were 14 and 16 years, respectively. When fires are timed so that maximum seed-bank is available (about 30 year interval), it is unlikely that resprouters will dominate communities because the seed-banks and rates of growth of seedlings of obligate seeder shrubs are greater than these resprouters. Populations of these resprouters may be more able to persist than obligate seeders when the fire frequency is either very high (<6 years) or low (>50years), though the density of resprouter populations may slowly decline under such fire regimes.     

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.     

Fitness and evolution of resprouters in relation to fire

There are many ways that plants may recover vegetatively from dieback caused by fires. Compared with fire-killed species, the presence of woody resprouters in fire-prone floras increases with fire frequency, though this is affected by site productivity that may have opposing correlates along different gradients. Population recovery is enhanced by resprouting when fecundity is low and/or seedling recruitment is not guaranteed. There is resource cycling between vegetative growth and storage but no clear trade-off between fecundity and storage, and more attention needs to be given to the role of somatic mutations in reducing fecundity. Seven fitness benefits of post-fire resprouting are noted that centre around the rapid return to adult growth rates and early flowering and seed set without the risks of recruitment failure. The extent of resprouting at the individual, population, and species levels varies greatly but it is under genetic control. Recent studies on the evolution of resprouting in fire-prone systems have shown that types of resprouting (clonality, rootstocks, epicormic bud strands) are derived from surrounding parent lineages lacking these traits and confined to non-fire-prone environments. The oldest lineages with fire-related resprouting date to at least 61 million years ago, indicating that seed plants have had a long evolutionary relationship with fire. Various genetic mechanisms, including beneficial somatic mutations, have been invoked to explain how speciation of resprouters may keep pace with non-sprouters.     

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.     

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