Biomass consumption by surface fires across Earth's most fire prone continent.

Landscape fire is a key but poorly understood component of the global carbon cycle. Predicting biomass consumption by fire at large spatial scales is essential to understanding carbon dynamics and hence how fire management can reduce greenhouse gas emissions and increase ecosystem carbon storage. An Australia-wide field-based survey (at 113 locations) across large-scale macroecological gradients (climate, productivity and fire regimes) enabled estimation of how biomass combustion by surface fire directly affects continental-scale carbon budgets. In terms of biomass consumption, we found clear trade-offs between the frequency and severity of surface fires. In temperate southern Australia, characterised by less frequent and more severe fires, biomass consumed per fire was typically very high. In contrast, surface fires in the tropical savannas of northern Australia were very frequent but less severe, with much lower consumption of biomass per fire (about a quarter of that in the far south). When biomass consumption was expressed on an annual basis, biomass consumed was far greater in the tropical savannas (>20 times that of the far south). This trade-off is also apparent in the ratio of annual carbon consumption to net primary production (NPP). Across Australia's naturally vegetated land area, annual carbon consumption by surface fire is equivalent to about 11% of NPP, with a sharp contrast between temperate southern Australia (6%) and tropical northern Australia (46%). Our results emphasise that fire management to reduce greenhouse gas emissions should focus on fire prone tropical savanna landscapes, where the vast bulk of biomass consumption occurs globally. In these landscapes, grass biomass is a key driver of frequency, intensity and combustion completeness of surface fires, and management actions that increase grass biomass are likely to lead to increases in greenhouse gas emissions from savanna fires.

Big eucalypts grow more slowly in a warm climate: evidence of an interaction between tree size and temperature.

Large trees are critical components of forest ecosystems, but are declining in many forests worldwide. We predicted that growth of large trees is more vulnerable than that of small trees to high temperatures, because respiration and tissue maintenance costs increase with temperature more rapidly than does photosynthesis and these costs may be disproportionately greater in large trees. Using 5 00 000 measurements of eucalypt growth across temperate Australia, we found that high temperatures do appear to impose a larger growth penalty on large trees than on small ones. Average stem diameter growth rates at 21 °C compared with 11 °C mean annual temperature were 57% lower for large trees (58 cm stem diameter), but only 29% lower for small trees (18 cm diameter). While our results are consistent with an impaired carbon budget for large trees at warmer sites, we cannot discount causes such as hydraulic stress. We conclude that slower growth rates will impede recovery from extreme events, exacerbating the effects of higher temperatures, increased drought stress and more frequent fire on the tall eucalypt forests of southern Australia.

Abrupt fire regime change may cause landscape-wide loss of mature obligate seeder forests.

Obligate seeder trees requiring high-severity fires to regenerate may be vulnerable to population collapse if fire frequency increases abruptly. We tested this proposition using a long-lived obligate seeding forest tree, alpine ash (Eucalyptus delegatensis), in the Australian Alps. Since 2002, 85% of the Alps bioregion has been burnt by several very large fires, tracking the regional trend of more frequent extreme fire weather. High-severity fires removed 25% of aboveground tree biomass, and switched fuel arrays from low loads of herbaceous and litter fuels to high loads of flammable shrubs and juvenile trees, priming regenerating stands for subsequent fires. Single high-severity fires caused adult mortality and triggered mass regeneration, but a second fire in quick succession killed 97% of the regenerating alpine ash. Our results indicate that without interventions to reduce fire severity, interactions between flammability of regenerating stands and increased extreme fire weather will eliminate much of the remaining mature alpine ash forest.

Using a rainforest-flame forest mosaic to test the hypothesis that leaf and litter fuel flammability is under natural selection.

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.

Climate, not Aboriginal landscape burning, controlled the historical demography and distribution of fire-sensitive conifer populations across Australia.

Climate and fire are the key environmental factors that shape the distribution and demography of plant populations in Australia. Because of limited palaeoecological records in this arid continent, however, it is unclear as to which factor impacted vegetation more strongly, and what were the roles of fire regime changes owing to human activity and megafaunal extinction (since ca 50 kya). To address these questions, we analysed historical genetic, demographic and distributional changes in a widespread conifer species complex that paradoxically grows in fire-prone regions, yet is very sensitive to fire. Genetic demographic analysis showed that the arid populations experienced strong bottlenecks, consistent with range contractions during the Last Glacial Maximum (ca 20 kya) predicted by species distribution models. In southern temperate regions, the population sizes were estimated to have been mostly stable, followed by some expansion coinciding with climate amelioration at the end of the last glacial period. By contrast, in the flammable tropical savannahs, where fire risk is the highest, demographic analysis failed to detect significant population bottlenecks. Collectively, these results suggest that the impact of climate change overwhelmed any modifications to fire regimes by Aboriginal landscape burning and megafaunal extinction, a finding that probably also applies to other fire-prone vegetation across Australia.

Population collapse of a Gondwanan conifer follows the loss of Indigenous fire regimes in a northern Australian savanna.

Colonialism has disrupted Indigenous socioecological systems around the globe, including those supported by intentional landscape burning. Because most disruptions happened centuries ago, our understanding of Indigenous fire management is largely inferential and open to debate. Here, we investigate the ecological consequences of the loss of traditional Aboriginal fire management on fire-exposed savannas on the Arnhem Plateau, northern Australia, using the fire-sensitive conifer Callitris intratropica as a bio-indicator. We contrast Kakadu National Park, where traditional Aboriginal fire management was severely disrupted during the early twentieth century following Aboriginal relocation to surrounding settlements, and an adjacent Aboriginal estate where traditional Aboriginal fire management endures. Since 2006, traditional Aboriginal fire management at this site has been overlaid by a program of broad-scale institutionalized burning in the early dry season, designed to reduce greenhouse emissions. Using remote sensing, field survey, and dendrochronology, we show that on the Aboriginal estate, C. intratropica populations depend on the creation of a shifting patch mosaic of long unburned areas necessary for the recruitment of C. intratropica. However, the imposition of broad-scale fire management is disrupting this population patch dynamic. In Kakadu, there have been extreme declines of C. intratropica associated with widespread fires since the mid twentieth century and consequent proliferation of grass fuels. Fire management in Kakadu since 2007, designed to increase the size and abundance of patches of unburned vegetation, has not been able to reverse the population collapse of C. intratropica. Our study demonstrates that colonial processes including relocation of Indigenous people and institutional fire management can have deleterious consequences that are nearly irreversible because of hysteresis in C. intratropica population dynamics.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Isolation and characterization of 52 polymorphic EST-SSR markers for Callitris columellaris (Cupressaceae).

PREMISE OF THE STUDY: We developed simple sequence repeat (SSR) markers from expressed sequence tags (ESTs) for Callitris columellaris sensu lato (s.l.) to elucidate population genetic structure and detect outlier loci by genome scan. METHODS AND RESULTS: mRNA from an individual seedling was subjected to cDNA synthesis and then de novo pyrosequencing. Two hundred and nineteen primer pairs bordering sequence regions were designed from the obtained sequence data. In total, 52 showed polymorphism within 16 individuals representative of the species' entire range, with the number of alleles per locus and expected heterozygosity ranging from two to 10 and 0.06 to 0.84, respectively. CONCLUSIONS: The EST-SSR markers developed in this study will be useful for evaluating the range-wide genetic structure of C. columellaris s.l. and detecting outlier loci under selection, as well as providing useful markers to investigate the conservation genetics and reproductive ecology of the species.

The worldwide leaf economics spectrum.

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

Fire controls population structure in four dominant tree species in a tropical savanna.

The persistence of mesic savannas has been theorised as being dependent on disturbances that restrict the number of juveniles growing through the sapling size class to become fire-tolerant trees. We analysed the population structures of four dominant tropical savanna tree species from 30 locations in Kakadu National Park (KNP), northern Australia. We found that across KNP as a whole, the population size structures of these species do not exhibit recruitment bottlenecks. However, individual stands had multimodal size-class distributions and mixtures of tree species consistent with episodic and individualistic recruitment of co-occurring tree species. Using information theory and multimodel inference, we examined the relative importance of fire frequency, stand basal area and elevation difference between a site and permanent water in explaining variations in the proportion of sapling to adult stems in four dominant tree species. This showed that the proportion of the tree population made up of saplings was negatively related to both fire frequencies and stand basal area. Overall, fire frequency has density-dependent effects in the regulation of the transition of saplings to trees in this Australian savanna, due to interactions with stem size, regeneration strategies, growth rates and tree-tree competition. Although stable at the regional scale, the spatiotemporal variability of fire can result in structural and floristic diversity of savanna tree populations.

Can trophic rewilding reduce the impact of fire in a more flammable world?

Large vertebrates affect fire regimes in several ways: by consuming plant matter that would otherwise accumulate as fuel; by controlling and varying the density of vegetation; and by engineering the soil and litter layer. These processes can regulate the frequency, intensity and extent of fire. The evidence for these effects is strongest in environments with intermediate rainfall, warm temperatures and graminoid-dominated ground vegetation. Probably, extinction of Quaternary megafauna triggered increased biomass burning in many such environments. Recent and continuing declines of large vertebrates are likely to be significant contributors to changes in fire regimes and vegetation that are currently being experienced in many parts of the world. To date, rewilding projects that aim to restore large herbivores have paid little attention to the value of large animals in moderating fire regimes. Rewilding potentially offers a powerful tool for managing the risks of wildfire and its impacts on natural and human values.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

Water, land, fire, and forest: Multi-scale determinants of rainforests in the Australian monsoon tropics.

The small rainforest fragments found in savanna landscapes are powerful, yet often overlooked, model systems to understand the controls of these contrasting ecosystems. We analyzed the relative effect of climatic variables on rainforest density at a subcontinental level, and employed high-resolution, regional-level analyses to assess the importance of landscape settings and fire activity in determining rainforest density in a frequently burnt Australian savanna landscape. Estimates of rainforest density (ha/km2) across the Northern Territory and Western Australia, derived from preexisting maps, were used to calculate the correlations between rainforest density and climatic variables. A detailed map of the northern Kimberley (Western Australia) rainforests was generated and analyzed to determine the importance of geology and topography in controlling rainforests, and to contrast rainforest density on frequently burnt mainland and nearby islands. In the northwestern Australian, tropics rainforest density was positively correlated with rainfall and moisture index, and negatively correlated with potential evapotranspiration. At a regional scale, rainforests showed preference for complex topographic positions and more fertile geology. Compared with mainland areas, islands had significantly lower fire activity, with no differences between terrain types. They also displayed substantially higher rainforest density, even on level terrain where geomorphological processes do not concentrate nutrients or water. Our multi-scale approach corroborates previous studies that suggest moist climate, infrequent fires, and geology are important stabilizing factors that allow rainforest fragments to persist in savanna landscapes. These factors need to be incorporated in models to predict the future extent of savannas and rainforests under climate change.

Macroecology of Australian Tall Eucalypt Forests: Baseline Data from a Continental-Scale Permanent Plot Network.

Tracking the response of forest ecosystems to climate change demands large (≥1 ha) monitoring plots that are repeatedly measured over long time frames and arranged across macro-ecological gradients. Continental scale networks of permanent forest plots have identified links between climate and carbon fluxes by monitoring trends in tree growth, mortality and recruitment. The relationship between tree growth and climate in Australia has been recently articulated through analysis of data from smaller forest plots, but conclusions were limited by (a) absence of data on recruitment and mortality, (b) exclusion of non-eucalypt species, and (c) lack of knowledge of stand age or disturbance histories. To remedy these gaps we established the Ausplots Forest Monitoring Network: a continental scale network of 48 1 ha permanent plots in highly productive tall eucalypt forests in the mature growth stage. These plots are distributed across cool temperate, Mediterranean, subtropical and tropical climates (mean annual precipitation 850 to 1900 mm per year; mean annual temperature 6 to 21°C). Aboveground carbon stocks (AGC) in these forests are dominated by eucalypts (90% of AGC) whilst non-eucalypts in the understorey dominated species diversity and tree abundance (84% of species; 60% of stems). Aboveground carbon stocks were negatively related to mean annual temperature, with forests at the warm end of the temperature range storing approximately half the amount of carbon as forests at the cool end of the temperature range. This may reflect thermal constraints on tree growth detected through other plot networks and physiological studies. Through common protocols and careful sampling design, the Ausplots Forest Monitoring Network will facilitate the integration of tall eucalypt forests into established global forest monitoring initiatives. In the context of projections of rapidly warming and drying climates in Australia, this plot network will enable detection of links between climate and growth, mortality and carbon dynamics of eucalypt forests.

Across a macro-ecological gradient forest competition is strongest at the most productive sites.

We tested the hypothesis that the effect of forest basal area on tree growth interacts with macro-ecological gradients of primary productivity, using a large dataset of eucalypt tree growth collected across temperate and sub- tropical mesic Australia. To do this, we derived an index of inter-tree competition based on stand basal area (stand BA) relative to the climatically determined potential basal area. Using linear mixed effects modeling, we found that the main effects of climatic productivity, tree size, and competition explained 26.5% of the deviance in individual tree growth, but adding interactions to the model could explain a further 8.9%. The effect of competition on growth interacts with the gradient of climatic productivity, with negligible effect of competition in low productivity environments, but marked negative effects at the most productive sites. We also found a positive interaction between tree size and stand BA, which was most pronounced in the most productive sites. We interpret these patterns as reflecting intense competition for light amongst maturing trees on more productive sites, and below ground moisture limitation at low productivity sites, which results in open stands with little competition for light. These trends are consistent with the life history and stand development of eucalypt forests: in cool moist environments, light is the most limiting resource, resulting in size-asymmetric competition, while in hot, low rainfall environments are open forests with little competition for light but where the amount of tree regeneration is limited by water availability.

Have plants evolved to self-immolate?

By definition fire prone ecosystems have highly combustible plants, leading to the hypothesis, first formally stated by Mutch in 1970, that community flammability is the product of natural selection of flammable traits. However, proving the "Mutch hypothesis" has presented an enormous challenge for fire ecologists given the difficulty in establishing cause and effect between landscape fire and flammable plant traits. Individual plant traits (such as leaf moisture content, retention of dead branches and foliage, oil rich foliage) are known to affect the flammability of plants but there is no evidence these characters evolved specifically to self-immolate, although some of these traits may have been secondarily modified to increase the propensity to burn. Demonstrating individual benefits from self-immolation is extraordinarily difficult, given the intersection of the physical environmental factors that control landscape fire (fuel production, dryness and ignitions) with community flammability properties that emerge from numerous traits of multiple species (canopy cover and litter bed bulk density). It is more parsimonious to conclude plants have evolved mechanisms to tolerate, but not promote, landscape fire.

Detecting trends in tree growth: not so simple.

Tree biomass influences biogeochemical cycles, climate, and biodiversity across local to global scales. Understanding the environmental control of tree biomass demands consideration of the drivers of individual tree growth over their lifespan. This can be achieved by studies of tree growth in permanent sample plots (prospective studies) and tree ring analyses (retrospective studies). However, identification of growth trends and attribution of their drivers demands statistical control of the axiomatic co-variation of tree size and age, and avoiding sampling biases at the stand, forest, and regional scales. Tracking and predicting the effects of environmental change on tree biomass requires well-designed studies that address the issues that we have reviewed.

Short Communication: Leaf trait relationships in Australian plant species.

Leaf trait data were compiled for 258 Australian plant species from several habitat types dominated by woody perennials. Specific leaf area (SLA), photosynthetic capacity, dark respiration rate and leaf nitrogen (N) and phosphorus (P) concentrations were positively correlated with one another and negatively correlated with average leaf lifespan. These trait relationships were consistent with previous results from global datasets. Together, these traits form a spectrum of variation running from species with cheap but frequently replaced leaves to those with strategies more attuned to a nutrient-conserving lifestyle. Australian species tended to have SLAs at the lower end of the spectrum, as expected in a dataset dominated by sclerophyllous species from low fertility or low rainfall sites. The existence of broad-scale, 'global' relationships does not imply that the same trait relationships will always be observed in small datasets. In particular, the probability of observing concordant patterns depends on the range of trait variation in a dataset, which, itself, may vary with sample size or species-sampling properties such as the range of growth forms, plant functional 'types', or taxa included in a particular study. The considerable scatter seen in these broad-scale trait relationships may be associated with climate, physiology and phylogeny.