Fire influences ant diversity by modifying vegetation structure in an Australian tropical savanna.

Fire is a dominant ecological force shaping many faunal communities globally. Fire affects fauna either directly, such as by killing individuals, or indirectly, such as by modifying vegetation structure. Vegetation structure itself also modulates fire frequency and intensity. As such, faunal responses to fire need to be seen through the lens of variable fire activity and vegetation structure. Here, we incorporate information on fire activity and vegetation structure to enhance an understanding of the response of ants to long-term (17-year) experimental fire treatments in an extremely fire-prone tropical savanna in northern Australia. Previous analysis revealed limited divergence in ant communities after 5 years of experimental fire treatment. Hence, we first investigated the extent to which ant communities diverged over a subsequent 12 years of treatment. We then assessed the relative contribution of fire treatment, cumulative fire intensity (fire activity), and woody cover to responses of ant species frequency of occurrence, richness, and composition. We found that, even after 17 years, fire treatments explained little variation in any ant response variable. In contrast, woody cover was a strong predictor for all of them, while fire activity was a moderate predictor for abundance and richness. Ant species occurrence and richness increased in open habitats receiving higher levels of fire activity, compared with plots with higher vegetation cover experiencing low (or no) fire activity. Moreover, species composition differed between plots with high and low vegetation cover. Our findings provide experimental support to the principle that the effects of fire on fauna are primarily indirect, via its effect on vegetation structure. Furthermore, our results show that a "uniform" fire regime does not have uniform impacts on the ant fauna, because of variability imposed by interactions between vegetation structure and fire activity. This helps to explain why there is often a weak relationship between pyrodiversity and biodiversity, and it lessens the need for active management of pyrodiversity to maintain biodiversity.

Population genomics and conservation management of the threatened black-footed tree-rat (Mesembriomys gouldii) in northern Australia.

Genomic diversity is a fundamental component of Earth's total biodiversity, and requires explicit consideration in efforts to conserve biodiversity. To conserve genomic diversity, it is necessary to measure its spatial distribution, and quantify the contribution that any intraspecific evolutionary lineages make to overall genomic diversity. Here, we describe the range-wide population genomic structure of a threatened Australian rodent, the black-footed tree-rat (Mesembriomys gouldii), aiming to provide insight into the timing and extent of population declines across a large region with a dearth of long-term monitoring data. By estimating recent trajectories in effective population sizes at four localities, we confirm widespread population decline across the species' range, but find that the population in the peri-urban area of the Darwin region has been more stable. Based on current sampling, the Melville Island population made the greatest contribution to overall allelic richness of the species, and the prioritisation analysis suggested that conservation of the Darwin and Cobourg Peninsula populations would be the most cost-effective scenario to retain more than 90% of all alleles. Our results broadly confirm current sub-specific taxonomy, and provide crucial data on the spatial distribution of genomic diversity to help prioritise limited conservation resources. Along with additional sampling and genomic analysis from the far eastern and western edges of the black-footed tree-rat distribution, we suggest a range of conservation and research priorities that could help improve black-footed tree-rat population trajectories at large and fine spatial scales, including the retention and expansion of structurally complex habitat patches.

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.

Population genomics and conservation management of a declining tropical rodent.

Conservation management is improved by incorporating information about the spatial distribution of population genetic diversity into planning strategies. Northern Australia is the location of some of the world's most severe ongoing declines of endemic mammal species, yet we have little genetic information from this regional mammal assemblage to inform a genetic perspective on conservation assessment and planning. We used next-generation sequencing data from remnant populations of the threatened brush-tailed rabbit-rat (Conilurus penicillatus) to compare patterns of genomic diversity and differentiation across the landscape and investigate standardised hierarchical genomic diversity metrics to better understand brush-tailed rabbit-rat population genomic structure. We found strong population structuring, with high levels of differentiation between populations (FST = 0.21-0.78). Two distinct genomic lineages between the Tiwi Islands and mainland are also present. Prioritisation analysis showed that one population in both lineages would need to be conserved to retain at least ~80% of alleles for the species. Analysis of standardised genomic diversity metrics showed that approximately half of the total diversity occurs among lineages (δ = 0.091 from grand total γ = 0.184). We suggest that a focus on conserving remnant island populations may not be appropriate for the preservation of species-level genomic diversity and adaptive potential, as these populations represent a small component of the total diversity and a narrow subset of the environmental conditions in which the species occurs. We also highlight the importance of considering both genomic and ecological differentiation between source and receiving populations when considering translocations for conservation purposes.

Overlapping den tree selection by three declining arboreal mammal species in an Australian tropical savanna.

Tree cavities are important denning sites for many arboreal mammals. Knowledge of cavity requirements of individual species, as well as potential den overlap among species, is integral to their conservation. In Australia's tropical savannas, development of tree cavities is enhanced by high termite activity, and, conversely, reduced by frequent fires. However, it is poorly understood how the availability of tree cavities in the tropical savannas impacts tree cavity use and selection by cavity-dependent fauna. There has been a severe decline among arboreal mammal species in northern Australia over recent decades. Investigation of their cavity requirements may illuminate why these species have declined drastically in some areas but are persisting in others. Here we examined this issue in three species of arboreal mammals (Trichosurus vulpecula, Mesembriomys gouldii, Conilurus penicillatus) on Melville Island, northern Australia. We radiotracked individuals to their den sites to evaluate whether the species differ in their den tree and tree-cavity selection. The strongest influence on den tree selection was the presence of large cavities (> 10 cm entrance diameter), with all three species using larger cavities most frequently. Conilurus penicillatus, the smallest species, differed the most from the other species: it frequently was found in smaller, dead trees and its den sites were closer to the ground, including in hollow logs. The two larger species had broader den tree use, using larger live trees and dens higher up in the canopy. Dens of C. penicillatus are likely to be more susceptible to predation and destruction by high-intensity savanna fires. This may have contributed to this species' rapid decline, both on Melville Island and on the mainland. However, the apparent preference for larger tree cavities by all three arboreal species is concerning due to the limited availability of large trees across Australian savannas, which are subject to frequent, high-intensity fires.

Does rapid utilization of elevated nutrient availability allow eucalypts to dominate in the tropical savannas of Australia?

Northern Australia's savannas are among the most fire-prone biomes on Earth and are dominated by eucalypts (Eucalyptus and Corymbia spp.). It is not clear what processes allow this group to dominate under such extreme fire frequencies and whether a superior ability to compete for nutrients and water might play a role. There is evidence that eucalypts are adapted to frequent fires; juvenile eucalypts escape the fire trap by growing rapidly in height between fires. However, non-eucalypts are less able to escape the fire trap and tend to have stand structures strongly skewed toward suppressed juveniles. The mechanisms that drive these contrasting fire responses are not well understood. Here, we describe the results of a controlled glasshouse seedling experiment that evaluated the relative importance of nutrient and water availability in determining height growth and biomass growth of two eucalypt and one noneucalypt tree species, common in northern Australian savannas. We demonstrate that growth of eucalypt seedlings is particularly responsive to nutrient addition. Eucalypt seedlings are able to rapidly utilize soil nutrients and accumulate biomass at a much greater rate than noneucalypt seedlings. We suggest that a seasonal spike in nutrient availability creates a nutrient-rich microsite that allows eucalypt seedlings to rapidly gain height and biomass, increasing their likelihood of establishing successfully and reaching a fire-resistant size. Our results extend our understanding of how eucalypts dominate northern Australian savannas under extremely high fire frequencies.

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.

Facultative and Obligate Trees in a Mesic Savanna: Fire Effects on Savanna Structure Imply Contrasting Strategies of Eco-Taxonomic Groups.

Fire is a major determinant of savanna tree communities and, as such, manipulation of fire frequency is an important management tool. Resolving the effects of fire management on tree size class distributions can help managers predict and plan for short-term ecological and economic outcomes, reveal different strategies by which woody plants cope with frequent fire, and help us predict vegetation changes under future fire scenarios. Savanna structure and size class distribution are strongly influenced by the ability of suppressed tree resprouts to escape stem death by frequent fire. A widespread assumption is that resprouts have an imperative to escape fire to reach sexual maturity in the canopy and thereby ensure long-term species viability. We use a census of Australian mesic savanna tree communities subjected to annual, triennial, and fire exclusion (unburnt) fire treatments to ask how fire frequency affects size class distributions within and between eco-taxonomic groups of species. Total tree densities did not significantly differ, but were highest in the triennial (7,610 ± se 1,162 trees ha-1) and unburnt fire treatments (7,051 ± se 578 trees ha-1) and lowest in the annual fire treatment (6,168 ± se 523 trees ha-1). This was caused by increased sapling densities in the triennial and unburnt fire treatments, predominantly of Acacia and pantropical genera. Eucalypts (Eucalyptus and Corymbia spp.) dominated the canopy across all fire treatments indicating relatively greater success in recruiting to larger sizes than other species groups. However, in the sub-canopy size classes eucalypts co-dominated with, and in some size classes were outnumbered by, pantropicals and Acacia, regardless of fire treatment. We hypothesize that such results are caused by fundamental differences in woody plant strategies, in particular sexual reproduction, that have not been widely recognized in Australian savannas.

Savanna woody encroachment is widespread across three continents.

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta-analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO2 , changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N-fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.

The underestimated biodiversity of tropical grassy biomes.

For decades, there has been enormous scientific interest in tropical savannahs and grasslands, fuelled by the recognition that they are a dynamic and potentially unstable biome, requiring periodic disturbance for their maintenance. However, that scientific interest has not translated into widespread appreciation of, and concern about threats to, their biodiversity. In terms of biodiversity, grassy biomes are considered poor cousins of the other dominant biome of the tropics-forests. Simple notions of grassy biomes being species-poor cannot be supported; for some key taxa, such as vascular plants, this may be valid, but for others it is not. Here, we use an analysis of existing data to demonstrate that high-rainfall tropical grassy biomes (TGBs) have vertebrate species richness comparable with that of forests, despite having lower plant diversity. The Neotropics stand out in terms of both overall vertebrate species richness and number of range-restricted vertebrate species in TGBs. Given high rates of land-cover conversion in Neotropical grassy biomes, they should be a high priority for conservation and greater inclusion in protected areas. Fire needs to be actively maintained in these systems, and in many cases re-introduced after decades of inappropriate fire exclusion. The relative intactness of TGBs in Africa and Australia make them the least vulnerable to biodiversity loss in the immediate future. We argue that, like forests, TGBs should be recognized as a critical-but increasingly threatened-store of global biodiversity.This article is part of the themed issue 'Tropical grassy biomes: linking ecology, human use and conservation'.

Pyrodiversity is the coupling of biodiversity and fire regimes in food webs.

Fire positively and negatively affects food webs across all trophic levels and guilds and influences a range of ecological processes that reinforce fire regimes, such as nutrient cycling and soil development, plant regeneration and growth, plant community assembly and dynamics, herbivory and predation. Thus we argue that rather than merely describing spatio-temporal patterns of fire regimes, pyrodiversity must be understood in terms of feedbacks between fire regimes, biodiversity and ecological processes. Humans shape pyrodiversity both directly, by manipulating the intensity, severity, frequency and extent of fires, and indirectly, by influencing the abundance and distribution of various trophic guilds through hunting and husbandry of animals, and introduction and cultivation of plant species. Conceptualizing landscape fire as deeply embedded in food webs suggests that the restoration of degraded ecosystems requires the simultaneous careful management of fire regimes and native and invasive plants and animals, and may include introducing new vertebrates to compensate for extinctions that occurred in the recent and more distant past.This article is part of the themed issue 'The interaction of fire and mankind'.

Deriving Multiple Benefits from Carbon Market-Based Savanna Fire Management: An Australian Example.

Carbon markets afford potentially useful opportunities for supporting socially and environmentally sustainable land management programs but, to date, have been little applied in globally significant fire-prone savanna settings. While fire is intrinsic to regulating the composition, structure and dynamics of savanna systems, in north Australian savannas frequent and extensive late dry season wildfires incur significant environmental, production and social impacts. Here we assess the potential of market-based savanna burning greenhouse gas emissions abatement and allied carbon biosequestration projects to deliver compatible environmental and broader socio-economic benefits in a highly biodiverse north Australian setting. Drawing on extensive regional ecological knowledge of fire regime effects on fire-vulnerable taxa and communities, we compare three fire regime metrics (seasonal fire frequency, proportion of long-unburnt vegetation, fire patch-size distribution) over a 15-year period for three national parks with an indigenously (Aboriginal) owned and managed market-based emissions abatement enterprise. Our assessment indicates improved fire management outcomes under the emissions abatement program, and mostly little change or declining outcomes on the parks. We attribute improved outcomes and putative biodiversity benefits under the abatement program to enhanced strategic management made possible by the market-based mitigation arrangement. For these same sites we estimate quanta of carbon credits that could be delivered under realistic enhanced fire management practice, using currently available and developing accredited Australian savanna burning accounting methods. We conclude that, in appropriate situations, market-based savanna burning activities can provide transformative climate change mitigation, ecosystem health, and community benefits in northern Australia, and, despite significant challenges, potentially in other fire-prone savanna settings.

Local and global pyrogeographic evidence that indigenous fire management creates pyrodiversity.

Despite the challenges wildland fire poses to contemporary resource management, many fire-prone ecosystems have adapted over centuries to millennia to intentional landscape burning by people to maintain resources. We combine fieldwork, modeling, and a literature survey to examine the extent and mechanism by which anthropogenic burning alters the spatial grain of habitat mosaics in fire-prone ecosystems. We survey the distribution of Callitris intratropica, a conifer requiring long fire-free intervals for establishment, as an indicator of long-unburned habitat availability under Aboriginal burning in the savannas of Arnhem Land. We then use cellular automata to simulate the effects of burning identical proportions of the landscape under different fire sizes on the emergent patterns of habitat heterogeneity. Finally, we examine the global extent of intentional burning and diversity of objectives using the scientific literature. The current distribution of Callitris across multiple field sites suggested long-unburnt patches are common and occur at fine scales (<0.5 ha), while modeling revealed smaller, patchy disturbances maximize patch age diversity, creating a favorable habitat matrix for Callitris. The literature search provided evidence for intentional landscape burning across multiple ecosystems on six continents, with the number of identified objectives ranging from two to thirteen per study. The fieldwork and modeling results imply that the occurrence of long-unburnt habitat in fire-prone ecosystems may be an emergent property of patch scaling under fire regimes dominated by smaller fires. These findings provide a model for understanding how anthropogenic burning alters spatial and temporal aspects of habitat heterogeneity, which, as the literature survey strongly suggests, warrant consideration across a diversity of geographies and cultures. Our results clarify how traditional fire management shapes fire-prone ecosystems, which despite diverse objectives, has allowed human societies to cope with fire as a recurrent disturbance.

A synthesis of postfire recovery traits of woody plants in Australian ecosystems.

Postfire resprouting and recruitment from seed are key plant life-history traits that influence population dynamics, community composition and ecosystem function. Species can have one or both of these mechanisms. They confer resilience, which may determine community composition through differential species persistence after fire. To predict ecosystem level responses to changes in climate and fire conditions, we examined the proportions of these plant fire-adaptive traits among woody growth forms of 2880 taxa, in eight fire-prone ecosystems comprising ~87% of Australia's land area. Shrubs comprised 64% of the taxa. More tree (>84%) than shrub (~50%) taxa resprouted. Basal, epicormic and apical resprouting occurred in 71%, 22% and 3% of the taxa, respectively. Most rainforest taxa (91%) were basal resprouters. Many trees (59%) in frequently-burnt eucalypt forest and savanna resprouted epicormically. Although crown fire killed many mallee (62%) and heathland (48%) taxa, fire-cued seeding was common in these systems. Postfire seeding was uncommon in rainforest and in arid Acacia communities that burnt infrequently at low intensity. Resprouting was positively associated with ecosystem productivity, but resprouting type (e.g. basal or epicormic) was associated with local scale fire activity, especially fire frequency. Although rainforest trees can resprout they cannot recruit after intense fires and may decline under future fires. Semi-arid Acacia communities would be susceptible to increasing fire frequencies because they contain few postfire seeders. Ecosystems dominated by obligate seeders (mallee, heath) are also susceptible because predicted shorter inter-fire intervals will prevent seed bank accumulation. Savanna may be resilient to future fires because of the adaptive advantage of epicormic resprouting among the eucalypts. The substantial non-resprouting shrub component of shrublands may decline, but resilient Eucalyptus spp. will continue to dominate under future fire regimes. These patterns of resprouting and postfire seeding provide new insights to ecosystem assembly, resilience and vulnerability to changing fire regimes on this fire-prone continent.

A grass-fire cycle eliminates an obligate-seeding tree in a tropical savanna.

A grass-fire cycle in Australian tropical savannas has been postulated as driving the regional decline of the obligate-seeding conifer Callitris intratropica and other fire-sensitive components of the regional flora and fauna, due to proliferation of flammable native grasses. We tested the hypothesis that a high-biomass invasive savanna grass drives a positive feedback process where intense fires destroy fire-sensitive trees, and the reduction in canopy cover facilitates further invasion by grass. We undertook an observational and experimental study using, as a model system, a plantation of C. intratropica that has been invaded by an African grass, gamba (Andropogon gayanus) in the Northern Territory, Australia. We found that high grass biomass was associated with reduced canopy cover and restriction of foliage to the upper canopy of surviving stems, and mortality of adult trees was very high (>50%) even in areas with low fuel loads (1 t·ha(-1)). Experimental fires, with fuel loads >10 t·ha(-1), typical of the grass-invasion front, caused significant mortality due to complete crown scorch. Lower fuel loads cause reduced canopy cover through defoliation of the lower canopy. These results help explain how increases in grass biomass are coupled with the decline of C. intratropica throughout northern Australia by causing a switch from litter and sparse perennial grass fuels, and hence low-intensity surface fires, to heavy annual grass fuel loads that sustain fires that burn into the midstorey. This study demonstrates that changes in fuel type can alter fire regimes with substantial knock-on effects on the biota.

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.

Cultural legacies, fire ecology, and environmental change in the Stone Country of Arnhem Land and Kakadu National Park, Australia.

We use the fire ecology and biogeographical patterns of Callitris intratropica, a fire-sensitive conifer, and the Asian water buffalo (Bubalus bubalis), an introduced mega-herbivore, to examine the hypothesis that the continuation of Aboriginal burning and cultural integration of buffalo contribute to greater savanna heterogeneity and diversity in central Arnhem Land (CAL) than Kakadu National Park (KNP). The 'Stone Country' of the Arnhem Plateau, extending from KNP to CAL, is a globally renowned social-ecological system, managed for millennia by Bininj-Kunwok Aboriginal clans. Regional species declines have been attributed to the cessation of patchy burning by Aborigines. Whereas the KNP Stone Country is a modern wilderness, managed through prescribed burning and buffalo eradication, CAL remains a stronghold for Aboriginal management where buffalo have been culturally integrated. We surveyed the plant community and the presence of buffalo tracks among intact and fire-damaged C. intratropica groves and the savanna matrix in KNP and CAL. Aerial surveys of C. intratropica grove condition were used to examine the composition of savanna vegetation across the Stone Country. The plant community in intact C. intratropica groves had higher stem counts of shrubs and small trees and higher proportions of fire-sensitive plant species than degraded groves and the savanna matrix. A higher proportion of intact C. intratropica groves in CAL therefore indicated greater gamma diversity and habitat heterogeneity than the KNP Stone Country. Interactions among buffalo, fire, and C. intratropica suggested that buffalo also contributed to these patterns. Our results suggest linkages between ecological and cultural integrity at broad spatial scales across a complex landscape. Buffalo may provide a tool for mitigating destructive fires; however, their interactions require further study. Sustainability in the Stone Country depends upon adaptive management that rehabilitates the coupling of indigenous culture, disturbance, and natural resources.

What controls the distribution of tropical forest and savanna?

Forest and savanna biomes dominate the tropics, yet factors controlling their distribution remain poorly understood. Climate is clearly important, but extensive savannas in some high rainfall areas suggest a decoupling of climate and vegetation. In some situations edaphic factors are important, with forest often associated with high nutrient availability. Fire also plays a key role in limiting forest, with fire exclusion often causing a switch from savanna to forest. These observations can be captured by a broad conceptual model with two components: (1) forest and savanna are alternative stable states, maintained by tree cover-fire feedbacks, (2) the interaction between tree growth rates and fire frequency limits forest development; any factor that increases growth (e.g. elevated availability of water, nutrients, CO(2)), or decreases fire frequency, will favour canopy closure. This model is consistent with the range of environmental variables correlated with forest distribution, and with the current trend of forest expansion, likely driven by increasing CO(2) concentrations. Resolving the drivers of forest and savanna distribution has moved beyond simple correlative studies that are unlikely to establish ultimate causation. Experiments using Dynamic Global Vegetation Models, parameterised with measurements from each continent, provide an important tool for understanding the controls of these systems.

Humid tropical rain forest has expanded into eucalypt forest and savanna over the last 50 years.

Tropical rain forest expansion and savanna woody vegetation thickening appear to be a global trend, but there remains uncertainty about whether there is a common set of global drivers. Using geographic information techniques, we analyzed aerial photography of five areas in the humid tropics of northeastern Queensland, Australia, taken in the 1950s and 2008, to determine if changes in rain forest extent match those reported for the Australian monsoon tropics using similar techniques. Mapping of the 1950s aerial photography showed that of the combined study area (64,430 ha), 63% was classified as eucalypt forests/woodland and 37% as rain forest. Our mapping revealed that although most boundaries remained stable, there was a net increase of 732 ha of the original rain forest area over the study period, and negligible conversion of rain forest to eucalypt forest/woodland. Statistical modeling, controlling for spatial autocorrelation, indicated distance from preexisting rain forest as the strongest determinant of rain forest expansion. Margin extension had a mean rate across the five sites of 0.6 m per decade. Expansion was greater in tall open forest types but also occurred in shorter, more flammable woodland vegetation types. No correlations were detected with other local variables (aspect, elevation, geology, topography, drainage). Using a geographically weighted mean rate of rain forest margin extension across the whole region, we predict that over 25% of tall open forest (a forest type of high conservation significance) would still remain after 2000 years of rain forest expansion. This slow replacement is due to the convoluted nature of the rain forest boundary and the irregular shape of the tall open forest patches. Our analyses point to the increased concentration of atmospheric CO(2) as the most likely global driver of indiscriminate rain forest expansion occurring in northeastern Australia, by increasing tree growth and thereby overriding the effects of fire disturbance.