Article

Fire responses of flora in a sclerophyll–rainforest vegetation complex in the Nightcap Range, North Coast, New South Wales

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Abstract

Context Species fire responses were investigated in a mixed sclerophyll–rainforest ecosystem in the Nightcap Range, North Coast, New South Wales. Aims To examine rates of seedling recruitment and resprouting in functional and phytogeographic components of wet sclerophyll forest (WSRf), and adjacent open forest (OF) and rock outcrop shrubland (RO). Methods Species resprouting and seedling recruitment traits (fire responses) were recorded in 45 stem plots and 225 seedling subplots in WSRf, OF and RO. Species fire responses were classified, community fire-response spectra compiled and rates of seedling recruitment and resprouting in WSRf examined in relation to primary fire response, growth-form, habitat and broad functional and phytogeographic species groupings. Species size-regenerative class distribution was used to analyse population structure, fire impact, regeneration and recruitment in resprouter species that comprised most of the mesic-Gondwanan element of the WSRf flora. Key results WSRf, OF and RO habitats had distinctively different fire-response spectra. In WSRf, there was a high proportion of mesophyll resprouter species of Gondwanan origin with nil or very low seedling recruitment, a distinct component of mesophyll seeders of Indo-Malayan origin, as well as sclerophyll seeders and resprouters that also comprised most of the OF and RO floras. Resprouters comprised 75% of the WSRf flora, 50% OF and 10% pavement shrubland. Continuous size-class distributions indicated recruitment between fire events in the majority of mesophyll resprouters in WSRf. Lower total seedling density appeared to reflect inherent species traits and less canopy disturbance by fire. Large sclerophyll species forming the unburnt canopy of WSRf had very low seedling recruitment. Conclusions Different habitats (WSRf, OF and RO) and functional and phytogeographic clades in WSRf display distinctive patterns of resprouting and seedling-recruitment fire response. Fire responses of species that maintain species population and community composition are governed by fire regime, habitat variables and inherent species traits. Implications The distinctive fire-response spectrum of WSRf appears to be a direct consequence of the overlap of ‘new’ and ‘old’ floras in this broad vegetation type.

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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.
Book
Until recently, few people would have associated rainforest with Australia. The present-day distribution of Australian rainforest is limited but it was formerly more widespread and was the dominant vegetation type of the entire continent. The contraction to its present limits reflects environmental changes that have taken place over many millions of years. In this book Dr Adam gives a general account of the Australian rainforests: what they are, where they are, and how their present distribution has evolved. His aim is to provide a broad framework that will enable new results to be placed in context. The book concludes with a historical account of human interaction with the rainforest from late Pleistocene times to the 1980s. The history of the exploitation of the Australian rainforests and the development of a conservation ethic have important lessons for the conservation of rainforests world-wide.
Article
The vulnerability to repeated high severity fires of tall wet eucalypt forests (TWEF) dominated by obligate seeder species is widely understood. However, 80% of Tasmania’s TWEF are dominated by resprouter species, and while these forests are more resilient to wildfire than those dominated by obligate seeders, the degree of their resilience is not well documented. Recently, two wildfires affected five 1-ha forest plots that had been previously measured, providing the opportunity to assess fire effects on these forest stands. We characterised these wildfires using remote sensing and field measurements of char height (a proxy for fire intensity) and canopy scorch (fire severity). We assessed the influence of tree diameter, fire intensity and seedling densities on the survival and resprouting responses of eucalypt overstorey trees and mixed species understorey trees. Our results showed that these fires were predominantly of low to moderate intensity and severity. Our results showed that mature TWEF were resilient to these wildfires, with overall eucalypt survival of 75%. The major eucalypt species were either mostly not defoliated (E. regnans), or could recover rapidly by epicormic resprouting (E. delegatensis and E. obliqua). Results from this and other studies show recovery from topkill caused by high severity fires occurs but is slower, by basal resprouting (E. obliqua) or from seed (all species). By contrast, understorey species suffered high mortality (85% overall), with few species showing substantial resprouting. Fire resistance in both groups increased markedly with tree diameter. Our results have implications for forest management. The high survival of overstorey eucalypts leads to a multi-aged stand structure in most unlogged old growth TWEF in Tasmania. On the other hand, clearfell logging, originally designed to mimic stand replacing wildfire in obligate seeder systems, creates a vulnerable, even aged stand of young regrowth, and at a landscape scale, it also reduces average tree size, reducing overall resistance to fire. Adopting alternatives to clearfelling, such as partial logging systems, will increase landscape resilience to fire, as well as providing other previously shown benefits.
Chapter
Wet sclerophyll forests (WSFs) are dominated by sclerophyllous trees (Eucalyptus and closely allied genera; family Myrtaceae) usually taller than 30 m at maturity in areas of greater than 1 000 mm of annual rainfall. They include the tallest angiosperms, and exhibit biomass amongst the highest in terrestrial ecosystems. Although occupying only 0.75% of Australia’s land-cover (4.83 million ha), the environmental, economic and cultural significance of WSF is far out of proportion to their distributional area. They occur discontinuously in a zone of high rainfall from north-eastern Queensland to southern Tasmania, with an outlier in south-western Australia; and also occur in east Malesia. In moist or protected margins in eastern Australia (and Malesia), and more extensively in Tasmania, mature eucalypts may occur above a well-developed rainforest; and are sometimes classified separately as mixed forest. This contrasts with WSF in south-western Australia, which includes no rainforest trees. Wet sclerophyll forest grades into rainforest in wetter, more fertile, or less frequently burnt margins; and into dry sclerophyll forests and woodlands in drier, more seasonal, less fertile, more frequently burnt, or cooler sites. Boundaries to adjacent forest types may be sharp, but sometimes unstable in eastern Australia, depending on disturbance history. These forests include important refugia under climate change, harbouring a diverse biota, featuring many endemic taxa. However, they are impacted by climate disruption and associated changes in fire regime, and distributions of weed and pest taxa. As a result, there are conservation concerns for the old-growth elements that are unique to these systems.
Article
Provides a synoptic description of the forests, with particular reference to the various eucalyptus species, and key environmental factors are identified. Fires in the upper canopy, undergrowth, litter, peat and soil-humus are described, and the effects of burning indicated. Pyric succession is noted, indicating the rich herb vegetation characteristic of the initial post-burn years, and the wet schlerophyll strategy is outlined. It seems paradoxical that forests producing vast amounts of fuel and generating the fiercest fires consist of fire-sensitive species, although the fact that they need to be burned at some time in their seed-bearing life is axiomatic. -P.J.Jarvis
Article
Plant traits considered adaptive to fire may be distinguished by: their vegetative survival; enhanced reproduction through fire-stimulated disperal; or enhanced reproduction facilitated by in-soil storage with fire-stimulated germination. A single trait, eg. vegetative survival and resprouting, or hard-seededness, may be adaptive not only to fires but also to other variables. Different species in the same environment may have different adaptive traits but persist with equal success. Adaption must be considered in terms of the fire regime: type, season, frequency and intensity. Traits enabling survival and reproduction during a succession of fires must also enable survival and reproduction to take place during stresses imposed by other selective agents, eg. drought. -from Author
Article
The importance of dormant buried seeds of understorey species of the wet sclerophyll forests of northern N.S.W. is discussed in relation to their germination ability following fire or mechanical or chemical abrasion.
Article
Estimation of tree age in various species along the east coast of Australia is made difficult by the lack of identifiable annual growth rings in stem wood and this creates problems in studying long term forest dynamics. It is possible to use radiocarbon dating to obtain general information for this purpose. In an area on the north coast of New South Wales where three forest types, were in close proximity, individuals of Eucalyptus pilularis (blackbutt) were found to be about 100 years old, Tristania conferta (brushbox) between 100 and 1340 years old and subtropical rainforest was between 100 and 880 years old. From these data, together with that from dating of soil charcoal, the frequency of fire (with sufficient severity to result in the production of charcoal) was estimated to be 300 years in the blackbutt forest, 300–400 years in the brushbox forest and in excess of 1000 years in the rainforest. Although only a few results are available, they indicate the potential of radiocarbon dating for use in studies on long term dynamics of forest ecosystems.
Article
We used a mosaic of infrequently burnt temperate rainforest and adjacent, frequently burnt eucalypt forests in temperate eastern Australia to test whether: (1) there were differences in flammability of fresh and dried foliage amongst congeners from contrasting habitats, (2) habitat flammability was related to regeneration strategy, (3) litter fuels were more flammable in frequently burnt forests, (4) the severity of a recent fire influenced the flammability of litter (as this would suggest fire feedbacks), and (5) microclimate contributed to differences in fire hazard amongst habitats. Leaf-level comparisons were made among 11 congeneric pairs from rainforest and eucalypt forests. Leaf-level ignitability, combustibility and sustainability were not consistently higher for taxa from frequently burnt eucalypt forests, nor were they higher for species with fire-driven recruitment. The bulk density of litter-bed fuels strongly influenced flammability, but eucalypt forest litter was not less dense than rainforest litter. Ignitability, combustibility and flame sustainability of community surface fuels (litter) were compared using fuel arrays with standardized fuel mass and moisture content. Forests previously burned at high fire severity did not have consistently higher litter flammability than those burned at lower severity or long unburned. Thus, contrary to the Mutch hypothesis, there was no evidence of higher flammability of litter fuels or leaves from frequently burnt eucalypt forests compared with infrequently burnt rainforests. We suggest the manifest pyrogenicity of eucalypt forests is not due to natural selection for more flammable foliage, but better explained by differences in crown openness and associated microclimatic differences.
Article
Alternative stable state theory is often used to explain the occurrence of flammable vegetation adjacent to less flammable vegetation where fire regimes mediate the shift between states. In 2002 an extreme landscape scale fire burnt extensive areas of forests in eastern Australia, including rainforests that are rarely severely burnt. This unique event allowed us to test long-held assumptions that predict landscape scale change after major disturbance. We tested three assumptions for detecting alternative community states; (1) that the scale of the event was large enough to remove canopy dominants, (2) fire feedbacks, both positive and negative are present, and (3) shifts in the floristic composition of communities are detected. We also examined whether high severity fires resulted in a community shift from less flammable to more flammable vegetation (e.g. from rainforest to wet sclerophyll vegetation), by examining floristic composition of vegetation communities (rainforest, wet sclerophyll forest, and dry sclerophyll forest) when burnt at different fire severities (high and low). Conversely, we tested whether there was a state shift from flammable to less flammable vegetation communities in sclerophyll forests long unburnt. In addition, we determined if there was any evidence that antecedent fire regimes and fire severity influenced flammability feedbacks.
Article
Disturbance is a dominant factor in many ecosystems, and the disturbance regime is likely to change over the next decades in response to land-use changes and global warming. We assume that predictions of vegetation dynamics can be made on the basis of a set of life-history traits that characterize the response of a species to disturbance. For crown-fire ecosystems, the main plant traits related to postfire persistence are the ability to resprout (persistence of individuals) and the ability to retain a persistent seed bank (persistence of populations). In this context, we asked (1) to what extent do different life- history traits co-occur with the ability to resprout and/or the ability to retain a persistent seed bank among differing ecosystems and (2) to what extent do combinations of fire- related traits (fire syndromes) change in a fire regime gradient? We explored these questions by reviewing the literature and analyzing databases compiled from different crown-fire ecosystems (mainly eastern Australia, California, and the Mediterranean basin). The review suggests that the pattern of correlation between the two basic postfire persistent traits and other plant traits varies between continents and ecosystems. From these results we predict, for instance, that not all resprouters respond in a similar way everywhere because the associated plant traits of resprouter species vary in different places. Thus, attempts to generalize predictions on the basis of the resprouting capacity may have limited power at a global scale. An example is presented for Australian heathlands. Considering the com- bination of persistence at individual (resprouting) and at population (seed bank) level, the predictive power at local scale was significantly increased.
Article
Woody plants may be killed by severe disturbance or resprout from vegetative tissue. Sprouters can persist at a site through several generations of nonsprouters. Differences in sprouting behavior are therefore important for understanding vegetation dynamics, extinction risks, and woody plant management. Although sprouting appears not to be uniquely correlated with many other intrinsic attributes, such as specific leaf area or breeding systems, a clear correlate is reduced seedling aboveground growth rates from sprouters allocating more to belowground structures. Consequently, sprouters tend to have low seedling recruitment rates, and saplings take longer to reach maturity. Sprouters also tend to have lower seed output than nonsprouters, but comparative studies have seldom taken other trait differences such as plant size into account. Added to these trade-offs between persistence and recruitment, sprouters are often multistemmed and shorter than related nonsprouters and may be outcompeted by them when disturbances are rare. Since sprouters tend to have long generation times, damped demographic trends, and gene flow across generations, it has been suggested that their speciation rates would be low. The available data, primarily from fire-prone Gondwanan shrublands in South Africa, show no strong differences in speciation rates of related sprouters versus seeders. This indicates that ecological factors are important determinants of the evolution of fire life histories. Analysis of disturbance regimes indicates a fundamental ecological correlate: sprouters are favored where disturbance regimes are frequent and severe in comparison to regrowth rates. To bridge the gap between ecology and evolution, data are needed on the genetical differences between related species with contrasting sprouter life histories.
Article
Following an introduction of the Australian environment, the remaining chapters examine: definition and classification of rain forest; an overview of Australian rain forests; rain forest boundaries and the problem of mixed forests; regeneration and responses to disturbance; origins and history of Australian rain forest; the rain forest fauna; mangroves; and the human influence. Rainforest is currently found along the eastern seaboard, from the NE tip of Queensland south to Tasmania; there are also patches in N Northern Territory. The vegetation type was once more widespread in Australia, and its contraction reflects millennia of environmental change together with human activity. Distinctions are drawn between subtropical, dry, warm temperate and cool temperate rainforests in terms of flora and physiognomy as well as environment. Threats to Australian rain forests come from recent exploitation, especially for logging, and conservation and management issues are raised. -P.J.Jarvis
Article
Transects across the margins of rainforests with eucalypt forests at two sites in New South Wales are described. At Girard State Forest along ten transects it was consistently found that pure rainforest occurs where there is no evidence of past burning; and that rainforest tree species appear to be invading and replacing eucalypt forest through a formerly burned ecotone area. A similar pattern was found at Barrington Tops. It is suggested that adaptations shown by species of tall open forests which enable them to regenerate rapidly after burning may have evolved initially in a fire free, prehuman rainforest environment in response to other types of disturbance.
Article
Eidothea hardeniana (Proteaceae) is a narrow endemic representative of an ancient lineage restricted to a single site in northern New South Wales (Australia). This study aims to identify the life‐history traits most likely to have influenced the current distribution pattern of this rain forest tree. Using ecological and molecular analyses we found that its range is limited by the absence of efficient dispersal mechanisms rather than by habitat availability, or as a result of a recent bottleneck or of reproductive failure. Resprouts can take over the position of senesced plants, even in the absence of disturbance, allowing local persistence and the maintenance of genetic diversity. Long‐term persistence and preferential outcrossing further favour relatively high levels of diversity ( H E = 0.542) despite a small effective population size (N e = 21.6). We used eight life‐history traits to assess if the E. hardeniana findings were valid across 258 local taxa. Current distribution patterns of rain forest species within a species‐rich community were accounted for by linking two important components of community assembly theory: dispersal and niche assembly. The interaction of the seed‐based dispersal dimension and resprouting potential best explains the current distribution of rare local taxa. Major dimensions of life‐history trait variation were identified among local plant species, suggesting that a range of interacting traits contribute to a species’ response to environmental variables and mitigate the influence of potentially adverse circumstances. The benefits of merging ecological and genetic approaches to interpret species distribution and population structure are applicable across a broader range of studies. Our findings highlight how currently constrained palaeo‐endemic species with small populations in refugial habitats may retain the capacity to both persist and expand in response to changing circumstances and opportunities. This has important implications for species conservation, habitat management and reserve design.
Article
Fire has played a decisive role in Post-Glacial biological and cultural evolution in the Mediterranean Region. Its evolutionary impact on plants has been manifested by feedback responses, in which the fire and its after-effects selected plants for physiological and other mechanisms that enable direct fire tolerance or permit avoidance followed by vegetative and reproductive regeneration. The dominant, sclerophyll, drought-tolerant phanerophytes of the maqui are obligatory rootsprouters, whereas the subordinate, drought-evading chamaephytes, as well as herbaceous perennials, are both vegetative and reproductive regenerators and are well adapted to new, fire-denuded habitats. Annual and perennial grass fire-followers are also able to escape high surface fire temperatures with the aid of torsion devices on seeds. Evolutionary strategies to overcome fire are closely interwoven with those against other environmental stresses such as drought and grazing. These combinations of stresses have brought about convergence in plant form and function in mediterranean climates. Fire has thus been important in the evolution of more xeric grasslands and woodlands and more mesic maqui and shrubland communities. Contrary to the present view of fire as simply destructive, both fire and grazing have favored genetical as well as ecological diversity. They should be studied as integral components of Mediterranean ecosystems and their evolution.
Article
A widely‐used description of vegetation response to fire is that species can be clearly classified as sprouters or non‐sprouters. We aimed to assess: (1) how well this dichotomous classification (sprouter/non‐sprouter) described the responses of a semi‐arid flora to experimental disturbance; (2) how similar were sprouting responses to treatments mimicking intense herbivory and fire; (3) how well easily‐measured traits could predict sprouting. Sprouting was assessed for 45 species from a range of growth forms (grasses, forbs, sub‐shrubs, woody shrubs and trees) from semi‐arid south‐eastern Australia. We used two treatments: clipping at stem base, and clipping followed by burning with a blowtorch. A dichotomy accounted for >60% of deviance explained by species identity. Models with three or four groups were not substantially better. The dichotomy was not between 0% and 100% sprouting, rather between ‘weak’ and ‘strong’ sprouters. Probabilities of sprouting for weak sprouters were 23% after clipping and 6% after burning, while strong sprouters had sprouting probabilities of 90% after clipping and 79% after burning. While sprouting varied in space and time, the dichotomy was robust to this variation. Sprouting ability increased with size in most of the species with variable sprouting. Sprouting was partially related to growth form; grasses sprouted strongly, chenopods weakly, and forb and woody species covered the range of sprouting. Strong sprouters were likely to have more stems per plant, greater basal area, shorter potential height and deeper buds than weak sprouters. A hierarchical model that used growth form and then stems‐per‐plant provided a simple, robust predictor of sprouting. Four‐fifths of species responded consistently to clipping and burning while one‐fifth of species were strong sprouters after clipping but weak sprouters after burning. Burning reduced sprouting most in intermediate sprouters. Differences between sprouting after clipping and burning reflected increased intensity and were related to the depth of buds below ground.
Article
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.
Soil phosphate and the delimitation of plant communities in eastern Australia.
Beadle NCW (1954). Soil phosphate and the delimitation of plant communities in eastern Australia. Ecology 35(3) 370-375. doi:10.2307/ 1930100
Special Climate Statement 72 -dangerous bushfire weather in spring 2019
BOM (2019) 'Special Climate Statement 72 -dangerous bushfire weather in spring 2019.' (Bureau of Meteorology: Commonwealth of Australia)
Response of montane wet sclerophyll forest understorey species to fire: evidence from high and low intensity fires
  • M L Campbell
  • P J Clarke
Campbell ML, Clarke PJ (2006) Response of montane wet sclerophyll forest understorey species to fire: evidence from high and low intensity fires. Proceedings of the Linnean Society of New South Wales 127, 63-73.