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.     

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.     

Patterns of storage tissue and starch distribution in the young taproot of obligate seeders and resprouters of Australian Proteaceae (Juss.): Possible evidence of homoplastic evolution

The ancient Gondwanan family Proteaceae has its greatest speciation in fire‐prone environments of Australia. Fire response is either by seedling recruitment from parent plants that succumb to fire (obligate seeders), or survival and resprouting from protected buds (resprouters). Starch is the main source of energy for resprouting and in roots is restricted to parenchyma tissue. This study compared the size and distribution of storage parenchyma and the magnitude of starch reserves in roots of several proteaceous species from different genera in relation to their fire response and taxonomy. Cross‐sections (2 μm) of roots of 51 resprouter and 42 seeder species from 12 genera were stained for starch. Areas of cortex and ray parenchyma along with starch grain density were measured using image analysis software (Assess 2.0) and comparable samples of root tissue were assayed chemically for starch. Starch, where present, predominated in ray and cortex tissue with a greater percentage in resprouters (13.4 ± 1.03) than seeders (1.8 ± 0.26); these results correlated significantly with the chemical assay for starch (r = 0.93, P < 0.0001). Resprouters also had more storage parenchyma (56.9 ± 1.72%) than seeders (41.9 ± 1.91%) mostly due to broader rays (17.5 ± 1.22%) compared with seeders (8.2 ± 0.16%). Percentage of cortex tissue was similar for seeders and resprouters (39.4 ± 2.24 and 33.7 ± 2.04 respectively). Anatomical preferences for storage site were consistent within genera and broad suprageneric groupings. This study shows that histological analysis of root starch is a reliable predictor of resprouting capacity in Proteaceae and that patterns of storage tissue within genera, together with the persistence of parenchyma devoid of starch in seeders, are consistent with response to fire and suggests homoplastic evolution of this response within the family.     

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.     

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.     

Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest

Aim This study documents the effects of multiple fires and drought on the woody structure of a north Australian savanna never grazed by domestic stock. Location The study was conducted in a 500 ha pocket of Eucalyptus‐dominated savanna surrounded by a late Quaternary lava flow. The flow is known as the Great Basalt Wall, located c. 50 km northeast of Charters Towers in semi‐arid north‐eastern Australia. This region was exposed to the largest 5‐year rainfall deficit on record between 1992 and 1996. Methods All individual woody plants were tagged within a 1.56 ha plot. Species were segregated into their habitat affinities (rain forest, ecotone, savanna) and regeneration strategy (resprouter, seeder). The survivorship of plants within these categories was analysed in relation to fire intensity from the first fire, and to each of four fires lit between 1996 and 2001. Results Before the first fire, the plot contained thirty‐one tree species including twenty‐one typical of the surrounding dry rain forest. These rain forest species were represented by small individuals and constituted <1% of the total basal area of woody plants. The basal area of savanna trees was 7.5 m² ha^?1 at the commencement of monitoring, although 31% had recently died and others had major crown damage. Further death of the drought debilitated savanna trees was substantial during the first year of monitoring and the basal area of live savanna trees declined to 1.1 m² ha^?1 after 5 years. Most species from both rain forest and savanna were classified as resprouters and are capable of regenerating from underground organs after fire. Species without this ability (rain forest seeders and ecotone seeders) were mostly eliminated after the first two consecutive fires. Among resprouters, survivorship declined as fire intensity increased and this was more pronounced for rain forest than for savanna species. Repeated burning produced a cumulative effect of decreasing survivorship for rain forest resprouters relative to savanna resprouters. Main conclusions The study provides evidence that savanna and rain forest trees differ in fire susceptibility and that recurrent fire can explain the restricted distribution of rain forest in the seasonally arid Australian tropics. The time of death of the savanna trees is consistent with the regional pattern after severe drought, and highlights the importance of medium term climate cycles for the population dynamics of savanna tree species and structure of Australian savannas.     

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.     

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.     

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.     

Persistence on inselbergs: the role of obligate seeders and resprouters

Aim To investigate if the current theories associated with recruitment strategies are sufficient to explain differences in the proportions of obligate seeders and resprouters on inselbergs and in the matrix. Location The New England Batholith of eastern Australia. Methods The importance of life form and regenerative response of flora species are compared for twenty‐four archipelagos of inselbergs habitat islands and for their adjacent matrix. Forward stepwise multiple linear regression is used to correlate measured and modelled variables with differences in the proportional abundance of obligate seeders on inselbergs at three levels of sampling. Results A total of 216 inselbergs were surveyed and information on 826 vascular plant species was collated. Herbs were the most common life form both on and off inselbergs. There were generally more shrubs on inselbergs than in the matrix. Resprouters were more common everywhere. However, obligate seeders were proportionately more common on inselbergs. The degree of insularity of outcrop floras accounted for 23–37% of the variance in the proportional abundance of obligate seeders at three sampling scales. Main conclusions A model is proposed whereby the mechanisms that influence the abundances of obligate seeders and resprouters are different on inselbergs and in the matrix. Inselberg environments are harsh, arid, have limited safe recruitment sites and below ground resources, but are not light limited. In such habitats, obligate seeders with persistence life‐history traits are favoured. Within the matrix in eastern Australia frequent biomass removal disturbances occur (such as fire) and resprouting is advantageous. High turnover obligate seeders with no persistence life‐history traits can be favoured in the matrix with a reduced disturbance regime and where competition for light still occurs. Some of these matrix obligate seeders occur on inselbergs due to their infrequent disturbance regimes.     

Effects of inter‐fire intervals on the reproductive output of resprouters and obligate seeders in the Proteaceae

Abstract Fire is often used as a management tool in fire‐prone communities to reduce fuel loads with the intention of reducing the severity and extent of unplanned fires, often resulting in the increased occurrence of fire in the dry sclerophyll vegetation of Australia. This study examined the effects of fire frequency (length of the inter‐fire interval) on the reproductive output of seven plant species in the Proteaceae, including obligate seeding shrubs (Hakea teretifolia, Petrophile pulchella), resprouting shrubs (Banksia spinulosa, Isopogon anemonifolius, Lambertia formosa) and resprouting trees (Banksia serrata, Xylomelum pyriforme). Reproductive output (measured as either number of confructescences or follicles) and relative size were estimated for 100 individuals at each of five sample sites, covering a range of past fire frequencies over 26 years including repeated short inter‐fire intervals. Patterns in reproductive output (after standardizing for size) were related to the life‐history attributes of the species. In areas that had experienced short inter‐fire intervals, obligate seeders had greater reproductive output compared with longer intervals, and the reproductive output of resprouting shrubs was less. Fire frequency did not affect reproductive output of the resprouting trees. The decreased reproductive output of the resprouting shrubs could be due to the allocation of resources to regrowth following fire rather than to reproduction. It is less clear what process resulted in the increased reproductive output of obligate seeders in high fire frequency areas, but it could be due to the most recent fires being more patchy in the areas experiencing shorter inter‐fire intervals, or it may have resulted from the selection for early reproduction in the high fire frequency areas. These results highlight the need to take into account past fire frequency at a site, in addition to time since the last fire, when planning prescribed fires.     

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     

Fire persistence traits of plants along a productivity and disturbance gradient in mediterranean shrublands of south-east Australia

Aim To understand changes in fire persistence traits of plants along a latitudinal gradient, considering the interactions between productivity, community (fuel) structure and fire regime. Location A gradient in the south of Australia (latitude 33–37° S; longitude 140–143° E), including: Little Desert National Park (VIC), Big Desert Wilderness Park (VIC), Murray‐Sunset National Park (VIC), Danggali Conservation Park (SA) and Tarawi Nature Reserve (NSW). Methods We selected four areas along a latitudinal gradient for which information on fire history and vegetation was available. Then, we tested to what extent the four selected areas have different climate and different fire regimes. Plant cover values of different life forms provided an indication of the plant community structure and flammability, and the proportion of species with different fire persistence traits (resprouting, seedbank persistence) informed us on the trait selection. Results Precipitation decreases and temperature increases from south to north. Thus the selected sites represent a gradient from high productivity (low aridity) in the south to low productivity (high aridity) in the north. Fire statistics suggest that fire frequency parallels productivity. There is a tendency for life form dominance and community structure to shift in such a way that fuel connectivity is reduced towards the north. Resprouting species increase and obligate seeders decrease along the fire–productivity gradient. Main conclusions Changes in plant traits are difficult to understand without simultaneous consideration of both the disturbance and the productivity gradients. In our study area, fire regime and productivity interact in such a way that decreases in productivity imply changes in fuel structure that produce a reduction in fire frequency. Resprouting species are better represented at the high fire–productivity part of the gradient, while obligate seeders are better represented at the opposite end of the gradient. The results also emphasize the importance of considering not only climate changes but also changes in fuel structure to predict future fire regimes.     

Long-term post-fire vegetation dynamics in Pinus halepensis forests of Central Greece: A functional group approach

A hierarchical approach for plant functional classification was applied to describe long-term vegetation change in Pinus halepensis burned forests. Plant species were initially grouped according to their growth form and afterwards data on species modes of regeneration, persistence and dispersal, together with some other specific competitive advantages were explored, resulting in the identification of 29 different functional groups, 14 for woody and 15 for herbaceous species. Three types of Pinus halepensis forests were identified, according to the structure of the understorey. For each forest type, a post-fire chronosequence of communities was selected for sampling. Data sampling was performed for at least two consecutive years in each community, so as to reduce the shortcomings of the synchronic approach and to increase the age range of each chronosequence. Even though the vast majority of the functional groups proved to be persistent throughout the post-fire development of vegetation, their species richness and abundance did not remain stable. An increase of annual herb richness and abundance was recorded in the first years after the fire, with the leguminous species forming the dominant functional group. For perennial herbs, the most abundant group was of species with vivid lateral growth, while the group of species with subterranean resource organs included the highest number of species. Finally, as far as the woody species are concerned, the groups that played the most important role in defining vegetation structure were the mono-specific group of the pine, the group of resprouting sclerophyllous tall shrubs and the group of obligate seeder short shrubs (with Cistusspp., among others). A negative relationship between the abundance of woody obligate resprouters and the regeneration of woody obligate seeders was found. The advantage of the proposed functional group approach over classical floristic or structural approaches for the long-term study of communities is discussed, together with the applicability of this approach in studies of vegetation risk assessments due to fire regime alterations.     

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.     

Seed bank patterns in Restionaceae and Epacridaceae after wildfire in kwongan in southwestern Australia

Abstract. Post-fire seed germination, seedling mortality and seed banks were investigated in scrub-heath (kwongan) in SW Australia. Study species included herbaceous and woody, obligate seeders and resprouters in two non-bradysporous but significant plant families (Restionaceae and Epacridaceae). In all species, seedlings were recruited only in the first autumn-spring after fire and occurred in similar densities as the estimated germinable annual seed input. Seedlings were absent from unburnt vegetation. Although most species retained some residual seeds after fire, tests (excised embryo culture) indicated that a negligible number of seeds were germinable. Regardless of fire response or species, there appeared to be a large loss of seeds each year and in most cases, only a small proportion of the annual seed production was used in post-fire recovery of plants. Based on seedling: parent ratios, all species had the capacity to reconstitute parent densities from germinants in the first year after fire, but high seedling mortality and no further recruitment resulted in less seedlings than replacements for four resprouter Restionaceae and three Epacridaceae (all obligate seeders) at the end of the third year after fire.     

Tissue water relations of three chaparral shrub species after wildfire

Summary We compared the tissue water relations among resprouts and seedlings of three chaparral species during the first summer drought after wildfire. Two of the species, Rhus laurina and Ceanothus spinosus recover after fire by a combination of resprouting and seedling establishment (facultative resprouters), whereas a third species, Ceanothus megacarpus recovers by seedling establishment alone (obligate seeder). Our objectives were to document any differences in tissue water characteristics that might arise between resprouts and seedlings and to test the hypothesis that seedlings of obligate seeders develop more drought tolerant characteristics of their tissues than seedlings of facultative resprouters. We found that resprouts had much higher predawn values of water potential, osmotic potential, and turgor potentials than seedlings. Predawn turgor potentials of resprouts were 1.5 MPa through July and August when turgor potentials for seedlings remained near 0 MPa. During summer months, midday water potentials were 2 to 3 MPa higher for resprouts than seedlings and midday conductances of resprouts were two to five fold greater than those of seedlings. Even though resprouts did not experience severe water stress like seedlings, their tissue water characteristics, as determined by pressure-volume curve analyses, were similar by the peak of the drought in August. Further-more, the tissue water characteristics of seedlings from the obligate seeder, C. megacarpus, were similar to those of facultative resprouters — R. laurina, and C. spinosus. We attribute the observed differences in plant water status between resprouts and seedlings to differences in rooting depths and access to soil moisture reserves during summer drought. We conclude that the higher growth rates, photosynthetic performance, and survivorship of postfire resprouts are primarily a result of higher water availability to resprouting tissues during summer months. It appears that the greater seedling survivorship during summer drought observed for the obligate seeder, C. megacarpus, is not associated with more favorable tissue water characteristics.     

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.     

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.