Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north‐eastern Australia

Eucalypts (Eucalyptus spp. and Corymbia spp.) dominate many communities across Australia, including frequently burnt tropical savannas and temperate forests, which receive less frequent but more intense fires. Understanding the demographic characteristics that allow related trees to persist in tropical savannas and temperate forest ecosystems can provide insight into how savannas and forests function, including grass–tree coexistence. This study reviews differences in critical stages in the life cycle of savanna and temperate forest eucalypts, especially in relation to fire. It adds to the limited data on tropical eucalypts, by evaluating the effect of fire regimes on the population biology of Corymbia clarksoniana, a tree that dominates some tropical savannas of north‐eastern Australia. Corymbia clarksoniana displays similar demographic characteristics to other tropical savanna species, except that seedling emergence is enhanced when seed falls onto recently burnt ground during a high rainfall period. In contrast to many temperate forest eucalypts, tropical savanna eucalypts lack canopy‐stored seed banks; time annual seed fall to coincide with the onset of predictable wet season rain; have very rare seedling emergence events, including a lack of mass germination after each fire; possess an abundant sapling bank; and every tropical eucalypt species has the ability to maintain canopy structure by epicormically resprouting after all but the most intense fires. The combination of poor seedling recruitment strategies, coupled with characteristics allowing long‐term persistence of established plants, indicate tropical savanna eucalypts function through the persistence niche rather than the regeneration niche. The high rainfall‐promoted seedling emergence of C. clarksoniana and the reduction of seedling survival and sapling growth by fire, support the predictions that grass–tree coexistence in savannas is governed by rainfall limiting tree seedling recruitment and regular fires limiting the growth of juvenile trees to the canopy.     

Fire in Australian savannas: from leaf to landscape

Savanna ecosystems comprise 22% of the global terrestrial surface and 25% of Australia (almost 1.9 million km²) and provide significant ecosystem services through carbon and water cycles and the maintenance of biodiversity. The current structure, composition and distribution of Australian savannas have coevolved with fire, yet remain driven by the dynamic constraints of their bioclimatic niche. Fire in Australian savannas influences both the biophysical and biogeochemical processes at multiple scales from leaf to landscape. Here, we present the latest emission estimates from Australian savanna biomass burning and their contribution to global greenhouse gas budgets. We then review our understanding of the impacts of fire on ecosystem function and local surface water and heat balances, which in turn influence regional climate. We show how savanna fires are coupled to the global climate through the carbon cycle and fire regimes. We present new research that climate change is likely to alter the structure and function of savannas through shifts in moisture availability and increases in atmospheric carbon dioxide, in turn altering fire regimes with further feedbacks to climate. We explore opportunities to reduce net greenhouse gas emissions from savanna ecosystems through changes in savanna fire management.     

Fire frequency and biodiversity conservation in Australian tropical savannas: implications from the Kapalga fire experiment

Abstract Every year large proportions of northern Australia's tropical savanna landscapes are burnt, resulting in high fire frequencies and short intervals between fires. The dominant fire management paradigm in these regions is the use of low‐intensity prescribed fire early in the dry season, to reduce the incidence of higher‐intensity, more extensive wildfire later in the year. This use of frequent prescribed fire to mitigate against high‐intensity wildfire has parallels with fire management in temperate forests of southern Australia. However, unlike in southern Australia, the ecological implications of high fire frequency have received little attention in the north. CSIRO and collaborators recently completed a landscape‐scale fire experiment at Kapalga in Kakadu National Park, Northern Territory, Australia, and here we provide a synthesis of the effects of experimental fire regimes on biodiversity, with particular consideration of fire frequency and, more specifically, time‐since‐fire. Two recurring themes emerged from Kapalga. First, much of the savanna biota is remarkably resilient to fire, even of high intensity. Over the 5‐year experimental period, the abundance of most invertebrate groups remained unaffected by fire treatment, as did the abundance of most vertebrate species, and we were unable to detect any effect of fire on floristic composition of the grass‐layer. Riparian vegetation and associated stream biota, as well as small mammals, were notable exceptions to this general resilience. Second, the occurrence of fire, independent of its intensity, was often the major factor influencing fire‐sensitive species. This was especially the case for extinction‐prone small mammals, which have suffered serious population declines across northern Australia in recent decades. Results from Kapalga indicate that key components of the savanna biota of northern Australia favour habitat that has remained unburnt for at least several years. This raises a serious conservation concern, given that very little relatively long unburnt habitat currently occurs in conservation reserves, with most sites being burnt at least once every 2 years. We propose a conservation objective of increasing the area that remains relatively long unburnt. This could be achieved either by reducing the proportion of the landscape burnt each year, or by setting prescribed fires more strategically. The provision of appropriately long unburnt habitat is a conservation challenge for Australia's tropical savanna landscapes, just as it is for its temperate forests.     

Optimal Fire Regimes for Soil Carbon Storage in Tropical Savannas of Northern Australia

Modification of fire regimes in tropical savannas can have significant impacts on the global carbon (C) cycle, and therefore, on the climate system. In Australian tropical savannas, there has been recent, large-scale implementation of fire management that aims to decrease Kyoto-compliant non-CO₂ greenhouse gas emissions by reducing late dry season intense fires through strategic early dry season burning. However, there is no accounting for changes to soil C stocks resulting from changes to savanna fire management, although impacts on these pools may be considerable. We present a hypothesis that soil C storage is greatest under low intensity fires with an intermediate fire return interval. Simulations using the CENTURY Soil Organic Matter Model confirmed this hypothesis with greatest soil C storage under a fire regime of one low intensity fire every 5 years. Key areas of uncertainty for CENTURY model simulations include fine root dynamics, charcoal production and nitrogen (N) cycling, and better understanding of these processes could improve model predictions. Soil C stocks measured in the field after 5 years of annual, 3 year and unburned fire treatments were not significantly different (range 41–58 t ha⁻¹), but further CENTURY modelling suggests that changes in fire management will take up to 100 years to have a detectable impact (+4 t ha⁻¹) on soil C stocks. However, implementation of fire management that reduces fire frequency and intensity within the large area of intact savanna landscapes in northern Australia could result in emissions savings of 0.17 t CO₂-e ha⁻¹ y⁻¹, four times greater than reductions of non-CO₂ emissions.     

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.     

Arthropod responses to experimental fire regimes in an Australian tropical savannah: ordinal‐level analysis

Fire is widely used for conservation management in the savannah landscapes of northern Australia, yet there is considerable uncertainty over the ecological effects of different fire regimes. The responses of insects and other arthropods to fire are especially poorly known, despite their dominant roles in the functioning of savannah ecosystems. Fire often appears to have little long‐term effect on ordinal‐level abundance of arthropods in temperate woodlands and open forests of southern Australia, and this paper addresses the extent to which such ordinal‐level resilience also occurs in Australia’s tropical savannahs. The data are from a multidisciplinary, landscape‐scale fire experiment at Kapalga in Kakadu National Park. Arthropods were sampled in the two major savannah habitats (woodland and open forest) using pitfall traps and sweep nets, in 15–20 km² compartments subjected to one of three fire regimes, each with three replicates: ‘early’ (annual fires lit early in the dry season), ‘late’ (annual fires lit late in the dry season), and ‘unburnt’ (fires absent during the five‐year experimental period 1990–94). Floristic cover, richness and composition were also measured in each sampling plot, using point quadrats. There were substantial habitat differences in floristic composition, but fire had no measured effect on plant richness, overall composition, or cover of three of the four dominant species. Of the 11 ordinal arthropod taxa considered from pitfall traps, only four were significantly affected by fire according to repeated‐measures ANOVA . There was a marked reduction in ant abundance in the absence of fire, and declines in spiders, homopterans and silverfish under late fires. Similarly, the abundances of only four of the 10 ordinal taxa from sweep catches were affected by fire, with crickets and beetles declining in the absence of fire, and caterpillars declining under late fires. Therefore, most of the ordinal taxa from the ground and grass‐layer were unaffected by the fire treatments, despite the treatments representing the most extreme fire regimes possible in the region. This indicates that the considerable ordinal‐level resilience to fire of arthropod assemblages that has previously been demonstrated in temperate woodlands and open forests of southern Australia, also occurs in tropical savannah woodlands and open forests of northern Australia.     

Root dynamics influence tree–grass coexistence in an Australian savanna

Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C₃ (trees) and C₄ (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.     

Carbon emissions from fires in tropical and subtropical ecosystems

Global carbon emissions from fires are difficult to quantify and have the potential to influence interannual variability and long‐term trends in atmospheric CO₂ concentrations. We used 4 years of Tropical Rainfall Measuring Mission (TRMM) Visible and Infrared Scanner (VIRS) satellite data and a biogeochemical model to assess spatial and temporal variability of carbon emissions from tropical fires. The TRMM satellite data extended between 38°N and 38°S and covered the period from 1998 to 2001. A relationship between TRMM fire counts and burned area was derived using estimates of burned area from other satellite fire products in Africa and Australia and reported burned areas from the United States. We modified the Carnegie‐Ames‐Stanford‐Approach (CASA) biogeochemical model to account for both direct combustion losses and the decomposition from fire‐induced mortality, using both TRMM and Sea‐viewing Wide Field of view Sensor (SeaWiFS) satellite data as model drivers. Over the 1998–2001 period, we estimated that the sum of carbon emissions from tropical fires and fuel wood use was 2.6 Pg C yr^?1. An additional flux of 1.2 Pg C yr^?1 was released indirectly, as a result of decomposition of vegetation killed by fire but not combusted. The sum of direct and indirect carbon losses from fires represented 9% of tropical and subtropical net primary production (NPP). We found that including fire processes in the tropics substantially alters the seasonal cycle of net biome production by shifting carbon losses to months with low soil moisture and low rates of soil microbial respiration. Consequently, accounting for fires increases growing season net flux by ～12% between 38°N and 38°S, with the greatest effect occurring in highly productive savanna regions.     

Stocks and dynamics of carbon in trees across a rainfall gradient in a tropical savanna

In this study, systematic variation in tree morphology across a rainfall gradient in Australia's tropical savanna biome and its implications for carbon stocks and dynamics were quantified. The aim was to support efforts to manage fire regimes to increase vegetative carbon stocks as a greenhouse gas mitigation strategy. The height of trees for a given trunk diameter declines with decreasing rainfall from 2000 to 300 mm and increasing dry season length across the Australian savanna biome. It is likely that increasing dry season length is the main driver of this decline rather declining rainfall per se. By taking account of the response of total basal area to rainfall and soil type, stand structure, and tree height and diameter relationships, the carbon stocks in live trees were estimated to decline from about 34 t ha^?1 in the wetter savannas to 6 t ha^?1 in the drier savannas. These values are broadly consistent with field‐based estimates. Because of the declining ratio of height to trunk diameter, trees of a given diameter in drier regions will be more likely to be killed by fires of a given intensity than trees in wetter regions. Thus single fires of given intensity are likely to have a greater proportionate impact on live tree carbon stock in drier savannas, but a much greater absolute impact in wetter savannas due to the greater total carbon stock. Projected decreases in early wet season rainfall under climate change scenarios, despite projections of little change in total precipitation in northern Australia, may lead to decreased carbon stock in live trees through two mechanisms: a reduction in total basal area and decreases in tree height for given trunk diameters.     

BIODIVERSITY RESEARCH: Turning up the heat: the impacts of Andropogon gayanus (gamba grass) invasion on fire behaviour in northern Australian savannas

Aim This study aimed to quantify changes in fire severity resulting from the invasion of Australia’s tropical savannas by the African grass Andropogon gayanus Kunth. (gamba grass). Location Mesic savannas of the Northern Territory, Australia. Method Byram’s fire‐line intensity (I_f), fuel load and architecture, and two post‐fire indicators of fire intensity – scorch height (SH) and char height (CH) of woody vegetation – were determined for fires in native grass savanna and A. gayanus invaded savanna. Leaf scorch is the height at which the fire’s radiant heat browns leaf tissue, and leaf char is the height that radiant heat blackens or consumes leaf tissue and provides an indirect measure of flame height. These data, and 5 years of similar data collected from the Kapalga Fire Project in Kakadu National Park, were used to develop empirical relationships between I_f and the post‐fire indices of fire intensity. Results A relationship between A. gayanus I_f and SH could not be developed because complete canopy scorch occurred in most A. gayanus fires, even at low I_f. In contrast, A. gayanus I_f was strongly correlated with CH. This empirical relationship was substantially different from that for native grass fires. For a given I_f, there was a significantly greater CH in invaded sites. This increase in radiant heat is attributable to the increased biomass (mean 3.6 t ha^?1 in native grasses compared to 11.6 t ha^?1 in A. gayanus) and height (approximately 0.5 m in native grasses compared to 4 m in A. gayanus) of the standing fine fuel. Main conclusion Andropogon gayanus invasion resulted in substantial changes in fire behaviour. This has important regional implications owing to the current (10,000–15,000 km²) and predicted (380,000 km²) area of invasion and the negative consequences for the native savanna biota that has evolved with frequent but relatively low‐intensity fire.     

Effect of fire regime on plant abundance in a tropical eucalypt savanna of north‐eastern Australia

Abstract Changes in plant abundance within a eucalypt savanna of north‐eastern Australia were studied using a manipulative fire experiment. Three fire regimes were compared between 1997 and 2001: (i) control, savanna burnt in the mid‐dry season (July) 1997 only; (ii) early burnt, savanna burnt in the mid‐dry season 1997 and early dry season (May) 1999; and (iii) late burnt, savanna burnt in the mid‐dry season 1997 and late dry season (October) 1999. Five annual surveys of permanent plots detected stability in the abundance of most species, irrespective of fire regime. However, a significant increase in the abundance of several subshrubs, ephemeral and twining perennial forbs, and grasses occurred in the first year after fire, particularly after late dry season fires. The abundance of these species declined toward prefire levels in the second year after fire. The dominant grass Heteropogon triticeus significantly declined in abundance with fire intervals of 4 years. The density of trees (>2 m tall) significantly increased in the absence of fire for 4 years, because of the growth of saplings; and the basal area of the dominant tree Corymbia clarksoniana significantly increased over the 5‐year study, irrespective of fire regime. Conservation management of these savannas will need to balance the role of regular fires in maintaining the diversity of herbaceous species with the requirement of fire intervals of at least 4‐years for allowing the growth of saplings >2 m in height. Whereas late dry season fires may cause some tree mortality, the use of occasional late fires may help maintain sustainable populations of many grasses and forbs.     

Fire history and the global carbon budget: a 1°× 1° fire history reconstruction for the 20th century

A yearly global fire history is a prerequisite for quantifying the contribution of previous fires to the past and present global carbon budget. Vegetation fires can have both direct (combustion) and long‐term indirect effects on the carbon cycle. Every fire influences the ecosystem carbon budget for many years, as a consequence of internal reorganization, decomposition of dead biomass, and regrowth. We used a two‐step process to estimate these effects. First we synthesized the available data available for the 1980s or 1990s to produce a global fire map. For regions with no data, we developed estimates based on vegetation type and history. Second, we then worked backwards to reconstruct the fire history. This reconstruction was based on published data when available. Where it was not, we extrapolated from land use practices, qualitative reports and local studies, such as tree ring analysis. The resulting product is intended as a first approximation for questions about consequences of historical changes in fire for the global carbon budget. We estimate that an average of 608 Mha yr^?1 burned (not including agricultural fires) at the end of the 20th century. 86% of this occurred in tropical savannas. Fires in forests with higher carbon stocks consumed 70.7 Mha yr^?1 at the beginning of the century, mostly in the boreal and temperate forests of the Northern Hemisphere. This decreased to 15.2 Mha yr^?1 in the 1960s as a consequence of fire suppression policies and the development of efficient fire fighting equipment. Since then, fires in temperate and boreal forests have decreased to 11.2 Mha yr^?1. At the same time, burned areas increased exponentially in tropical forests, reaching 54 Mha yr^?1 in the 1990s, reflecting the use of fire in deforestation for expansion of agriculture. There is some evidence for an increase in area burned in temperate and boreal forests in the closing years of the 20th century.     

Soil organic carbon content at a range of north Australian tropical savannas with contrasting site histories

Soils play an important role in the global carbon cycle, and can be major source or sink of CO₂ depending upon land use, vegetation type and soil management practices. Natural and human impact on soil carbon concentration and storage is poorly understood in native north Australian savanna, yet this represents the largest carbon store in the ecosystem. To gain understanding of possible management impacts on this carbon pool, soil organic carbon (SOC) of the top 1m of red earth sands and sandy loams common in the region was sampled at 5 sites with different vegetation cover and site history (fire regime and tree removal). SOC was high when compared to other published values for savannas and was more comparable with dry-deciduous tropical forests. Sites sampled in this study represent high rainfall savannas of northern Australia (> 1700 mm annual rainfall) that feature frequent burning (2 in 3 years or more frequent) and a cycle of annual re-growth of tall C₄ grasses that dominate the savanna understorey. These factors may be responsible for the higher than expected SOC levels of the surface soils, despite high respiration rates. Medium term fire exclusion (15–20 years) at one of the sampled sites (Wildlife Park) dramatically reduced the grassy biomass of the understorey. This site had lower SOC levels when compared to the grass dominated and frequently burnt sites, which may be due to a reduction in detrital input to surface (0–30 cm) soil carbon pools. Exclusion of trees also had a significant impact on both the total amount and distribution of soil organic carbon, with tree removal reducing observed SOC at depth (100 cm). Soil carbon content was higher in the wet season than that in the dry season, but this difference was not statistically significant. Our results indicated that annual cycle of grass growth and wildfire resulted in small carbon accumulation in the upper region of the soil, and removal of woody plants resulted in significant carbon losses to recalcitrant, deep soil horizons greater than 80 cm depth.     

Frequent fires reduce tree growth in northern Australian savannas: implications for tree demography and carbon sequestration

Tropical savannas are typically highly productive yet fire‐prone ecosystems, and it has been suggested that reducing fire frequency in savannas could substantially increase the size of the global carbon sink. However, the long‐term demographic consequences of modifying fire regimes in savannas are difficult to predict, with the effects of fire on many parameters, such as tree growth rates, poorly understood. Over 10 years, we examined the effects of fire frequency on the growth rates (annual increment of diameter at breast height) of 3075 tagged trees, at 137 locations throughout the mesic savannas of Kakadu, Nitmiluk and Litchfield National Parks, in northern Australia. Frequent fires substantially reduced tree growth rates, with the magnitude of the effect markedly increasing with fire severity. The highest observed frequencies of mild, moderate and severe fires (1.0, 0.8 and 0.4 fires yr^?1, respectively) reduced tree growth by 24%, 40% and 66% respectively, relative to unburnt areas. These reductions in tree growth imply reductions in the net primary productivity of trees by between 0.19 t C ha^?1 yr^?1, in the case of mild fires, and 0.51 t C ha^?1 yr^?1, in the case of severe fires. Such reductions are relatively large, given that net biome productivity (carbon sequestration potential) of these savannas is estimated to be just 1–2 t C ha^?1 yr^?1. Our results suggest that current models of savanna tree demography, that do not account for a relationship between severe fire frequency and tree growth rate, are likely to underestimate the long‐term negative effects of frequent severe fires on tree populations. Additionally, the negative impact of frequent severe fires on carbon sequestration rates may have been underestimated; reducing fire frequencies in savannas may increase carbon sequestration to a greater extent than previously thought.     

Nitrogen deposition in tropical forests from savanna and deforestation fires

We used satellite‐derived estimates of global fire emissions and a chemical transport model to estimate atmospheric nitrogen (N) fluxes from savanna and deforestation fires in tropical ecosystems. N emissions and reactive N deposition led to a net transport of N equatorward, from savannas and areas undergoing deforestation to tropical forests. Deposition of fire‐emitted N in savannas was only 26% of emissions – indicating a net export from this biome. On average, net N loss from fires (the sum of emissions and deposition) was equivalent to approximately 22% of biological N fixation (BNF) in savannas (4.0 kg N ha^?1 yr^?1) and 38% of BNF in ecosystems at the deforestation frontier (9.3 kg N ha^?1 yr^?1). Net N gains from fires occurred in interior tropical forests at a rate equivalent to 3% of their BNF (0.8 kg N ha^?1 yr^?1). This percentage was highest for African tropical forests in the Congo Basin (15%; 3.4 kg N ha^?1 yr^?1) owing to equatorward transport from frequently burning savannas north and south of the basin. These results provide evidence for cross‐biome atmospheric fluxes of N that may help to sustain productivity in some tropical forest ecosystems on millennial timescales. Anthropogenic fires associated with slash and burn agriculture and deforestation in the southern part of the Amazon Basin and across Southeast Asia have substantially increased N deposition in these regions in recent decades and may contribute to increased rates of carbon accumulation in secondary forests and other N‐limited ecosystems.     

Convergence of bark investment according to fire and climate structures ecosystem vulnerability to future change

Fire regimes in savannas and forests are changing over much of the world. Anticipating the impact of these changes requires understanding how plants are adapted to fire. In this study, we test whether fire imposes a broad selective force on a key fire‐tolerance trait, bark thickness, across 572 tree species distributed worldwide. We show that investment in thick bark is a pervasive adaptation in frequently burned areas across savannas and forests in both temperate and tropical regions where surface fires occur. Geographic variability in bark thickness is largely explained by annual burned area and precipitation seasonality. Combining environmental and species distribution data allowed us to assess vulnerability to future climate and fire conditions: tropical rainforests are especially vulnerable, whereas seasonal forests and savannas are more robust. The strong link between fire and bark thickness provides an avenue for assessing the vulnerability of tree communities to fire and demands inclusion in global models.     

Contrasting fire-related resilience of ecologically dominant ants in tropical savannas of northern Australia

This paper examines the role of fire in mediating the relative abundance of two of the world's major ecologically dominant ant genera, Iridomyrmex and Oecophylla, where they coexist across the tropical savanna landscapes of northern Australia. These taxa have contrasting biogeographical histories, which are predicted to lead to contrasting responses to fire. Iridomyrmex is an autochthonous Australian genus that has radiated primarily in the arid zone; as such, its abundance is predicted to be promoted by frequent fire because this maintains an open habitat. In contrast, Oecophylla is a genus of leaf‐nesting ants occurring in the canopies of Old World tropical rainforest, and is a recent arrival to Australia in geological time; the abundance of these ants is predicted to decline under frequent fire. We test these predictions using results from a landscape‐scale fire experiment, where three experimental fire regimes (including no fire) were applied to replicated subcatchments over a 5‐year period. Using sweep nets, ants were sampled in the grass layer (the habitat layer of greatest overlap between Iridomyrmex and Oecophylla) in eucalypt woodland (canopy cover < 30%) and open eucalypt forest (canopy cover about 50%) habitats. A total of 27 species from 11 genera were collected during the study; eight were common enough for statistical analysis, and the abundances of four of these were significantly affected by fire treatment. As predicted, the abundance of Iridomyrmex was promoted by fire, whereas that of Oecophylla declined. These changes occurred only under late‐season (relatively high intensity) fires, and for Oecophylla occurred only in open forest (not woodland) habitat. This fire‐mediated relationship between Iridomyrmex and Oecophylla mirrors the much broader, ecosystem‐wide dynamic between eucalypt‐dominated savanna and rainforest in tropical Australia, with savannas dominated by fire‐resistant sclerophyll elements of Australian origin, and rainforest dominated by fire‐sensitive mesophyll elements of South‐East Asian origin.     

Testing the grass-fire cycle: alien grass invasion in the tropical savannas of northern Australia

Abstract. Invasive alien grasses can increase fuel loads, leading to changes in fire regimes of invaded ecosystems by increasing the frequency, intensity and spatial extent of fires. Andropogon gayanus Kunth. (Gamba grass), a tall perennial grass from Africa, is invading ecosystems in the Top End of northern Australia. To determine whether A. gayanus alters savanna fire regimes, we compared fuel loads and fire intensities at invaded sites with those from native grass savannas. Savanna invaded by A. gayanus had fuel loads up to seven times higher than those dominated by native grasses. This higher fuel load supported a fire that was on average eight times more intense than those recorded in native grass savannas at the same time of year (means 15700 ± 6200 and 2100 ± 290 kW m⁻¹, respectively), and was the highest early dry season fire intensities ever recorded in the Northern Territory. These results suggest that A. gayanus is a serious threat to northern Australia's savannas, with the potential to alter vegetation structure and initiate a grass-fire cycle.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.