Fire resilience of ant assemblages in long‐unburnt savanna of northern Australia

Abstract Tropical savannas and rainforests contrast in their flammability and the fire resilience of their associated species. While savanna species generally exhibit high resilience to burning, there is much debate about the fire resilience of forest‐associated species, and the persistence of forest patches in a flammable savanna matrix. Where fire has been excluded, savanna tends on a trajectory towards forest, with an increase in forest‐associated plants and animal species. This study tested the idea that given the high proportion of forest‐associated taxa in long‐unburnt savanna, the fauna of these areas would be expected to exhibit less resilience to fire than the fauna in frequently burnt savannas. The study investigated the immediate and short‐term effects on ant assemblages of re‐introducing fire into long‐unburnt savanna in northern Australia. The ant fauna exhibited high resistance to fires, with no significant short‐term change in mean abundance or species richness; instead, seasonality had a far stronger influence on overall ant activity. Fire caused dramatic declines in dominance of the patchily distributed forest‐associated species Oecophylla smaragdina and Papyrius sp., but had no effect on overall dominance by open savanna species of Iridomyrmex. Dominance by Iridomyrmex pallidus declined, but this was compensated for by increases in I. reburrus, while two other species of Iridomyrmex showed no change. This indicates a high level of functional redundancy among dominant species of Iridomyrmex, which universally dominate open savanna communities, but not of dominant forest‐associated species. Overall, our findings demonstrate a high degree of fire‐resilience of the long‐unburnt savanna ant fauna. Despite the occurrence of forest‐associated species, the high proportion of savanna species persisting in this habitat means that long‐unburnt savanna retains the general response characteristics of frequently burnt savanna.     

The interdependence of fire, grass, kangaroos and Australian Aborigines: a case study from central Arnhem Land, northern Australia

Aim To describe the nexus between Aboriginal landscape burning and patterns of habitat use by kangaroos in a tropical savanna habitat mosaic, and to provide evidence to evaluate the claim that Aboriginal landscape burning is a game management tool. Location Central Arnhem Land, a stronghold of traditional Aboriginal culture, in the monsoon tropics of northern Australia. Methods The abundance of kangaroo scats was recorded throughout a landscape burnt by Aboriginal people, and used as a proxy for the intensity of habitat use by kangaroos. Scat abundance was assessed along field traverses totalling 112 km, at three time periods: (1) 1–4 weeks following mid‐dry season burning (July 2003); (2) in the late dry season (November 2003); and (3) in the following mid‐dry season (July 2004). We compared the intensity with which kangaroos used burnt vs. unburnt areas in various habitat types, with time since mid‐dry season burning. Scats were collected from areas that had been burnt to a varying extent and the abundance of carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) and carbon to nitrogen ratios (C : N) determined. Results There was clear evidence of an interaction between burning and habitat type on the abundance of kangaroo scats. Scats were much more abundant in burnt vs. unburnt areas in the moist habitats, but the opposite effect was observed in the dry rocky habitats, with higher scat abundance in unburnt areas. This interactive effect of burning and habitat type on scat abundance was observed immediately (< 4 weeks) following fire, and was still present one year later. High concentrations of nitrogen in resprouting grasses indicate that burnt areas may provide kangaroos with greater access to nutrients. The isotopic composition of scats indicates that kangaroos feeding in extensively burnt areas were consuming more grasses, and possibly sedges, than kangaroos feeding in unburnt areas. Main conclusions The fine‐scale mosaic of burnt and unburnt areas created by mid‐dry season Aboriginal landscape burning has clear effects on the distribution of kangaroos. Kangaroos move into burnt moist habitats and away from burnt dry, rocky habitats. Isotopic analysis of scats suggests that the mechanism driving this effect is the increased abundance of nitrogen rich grasses in burnt moist habitats.     

Long‐term fire exclusion and ant community structure in an Australian tropical savanna: congruence with vegetation succession

Aim To examine the extent to which succession from tropical savanna to rain forest in the long‐term absence of fire is matched by successional changes in ant communities. This is done by describing ant community responses to 23 years of fire exclusion in a northern Australian tropical savanna, with a particular focus on the extent of colonization by specialist rain forest taxa. Location Solar Village, near Darwin in Australia's Northern Territory. Methods Ants were sampled within 12 plots located inside (‘unburnt’– protected from fire for 23 years) and outside (burnt every 1–2 years) Solar Village in ridge and slope habitat dominated by Eucalyptus spp. The litter, ground‐foraging and arboreal faunas were sampled separately, using Berlese funnels, unbaited pitfall traps and baited pitfall traps attached to tree trunks, respectively. Each species was assigned a forest‐association score ranging from 0 (open savanna species) to 3 (specialist forest species) based on their known habitat preferences in the region. Results A total of 85 ant species from 35 genera were recorded, with multivariate analysis demonstrating distinct litter, ground and arboreal communities. Ant communities also varied substantially with topographic position, which interacted strongly with fire exclusion. A total of 72 species were recorded in burnt habitat, compared with only 45 in unburnt, and the number of ant species records was also about twice as high in burnt compared with unburnt habitat. Fire exclusion has resulted in a dramatic increase in forest‐associated taxa (those occurring in forest and denser, but rarely open, savanna), with such species representing 51% of species records in unburnt habitat compared with 19% in burnt. However, only five specialist forest species were recorded, representing < 1% of total ant records. Main conclusions Fire exclusion at Solar Village has markedly increased the prevalence of forest‐associated ant species, but has led to only very minor incursions by specialist rain forest ant taxa. These responses match very closely those of the vegetation.     

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.     

Effects of a short fire‐return interval on resources and assemblage structure of birds in a tropical savanna

Fire frequency is a key land management issue, particularly in tropical savannas where fire is widely used and fire recurrence times are often short. We used an extended Before‐After‐Control‐Impact design to examine the impacts of repeated wet‐season burning for weed control on bird assemblages in a tropical savanna in north Queensland, Australia. Experimentally replicated fire treatments (unburnt, singularly bunt, twice burnt), in two habitats (riparian and adjacent open woodland), were surveyed over 3 years (1 year before the second burn, 1 year post the second burn, 2 years post the second burn) to examine responses of birds to a rapid recurrence of fire. Following the second burn, species richness and overall bird abundance were lower in the twice‐burnt sites than either the unburnt or singularly burnt sites. Feeding group composition varied across year of survey, but within each year, feeding guilds grouped according to fire treatment. In particular, abundance of frugivores and insectivores was lower in twice‐burnt sites, probably because of the decline of a native shrub that produces fleshy fruits, Carissa ovata. Although broader climatic variability may ultimately determine overall bird assemblages, our results show that a short fire‐return interval will substantially influence bird responses at a local scale. Considering that fire is frequently used as a land management tool, our results emphasize the importance of determining appropriate fire‐free intervals.     

Threatened mammals become more predatory after small‐scale prescribed fires in a high‐rainfall rocky savanna

Understanding mechanisms underlying fire regime effects on savanna fauna is difficult because of a wide range of possible trophic interactions and feedbacks. Yet, understanding mechanisms underlying fauna dynamics is crucial for conservation management of threatened species. Small savanna mammals in northern Australia are currently undergoing widespread declines and regional extinctions partly attributable to fire regimes. This study investigates mammal trophic and ecosystem responses to fire in order to identify possible mechanisms underlying these declines. Mammal trophic responses to fire were investigated by surveying mammal abundance, mammal diet, vegetation structure and non‐mammal fauna dynamics in savannas six times at eight sites over a period of 3 years. Known site‐specific fire history was used to test for trophic responses to post‐fire interval and fire frequency. Mammal and non‐mammal fauna showed only minor responses of post‐fire interval and no effect of fire frequency. Lack of fauna responses differed from large post‐fire vegetation responses. Dietary analysis showed that two mammal species, Dasyurus hallucatus and Isoodon auratus, increased their intake of large prey groups in recently burnt, compared to longer unburnt vegetation. This suggests a fire‐related change in trophic interactions among predators and their prey, after removal of ground‐layer vegetation. No evidence was found for other changes in food resource uptake by mammals after fire. These data provide support for a fire‐related top‐down ecosystem response among savanna mammals, rather than a bottom‐up resource limitation response. Future studies need to investigate fire responses among other predators, including introduced cats and dingoes, to determine their roles in fire‐related mammal declines in savannas of northern Australia.     

Contrasting responses of small mammals to fire and topographic refugia

Unburnt patches within burnt landscapes are expected to provide an important resource for fauna, potentially acting as a refuge from direct effects of fire and allowing animals to persist in burnt landscapes. Nevertheless, there is little information about the way refugia are used by fauna and how populations may be affected by them. Planned burns are often patchy, with unburnt areas generally associated with gully systems providing a good opportunity to study faunal use of refugia. We used a before–after control‐impact design associated with a planned burn in south eastern Australia to investigate how two small mammal species, the bush rat Rattus fuscipes and agile antechinus Antechinus agilis, used unburnt gully systems within a larger burnt area. We tested three alternative hypotheses relating to post‐fire abundance: (i) active refugia – abundance would increase in unburnt patches because of a post‐fire shift of individuals from burnt to unburnt areas; (ii) passive refugia – abundance in unburnt patches would remain similar to pre‐fire levels; and (iii) limited or no refugia – abundance would reduce in unburnt patches related to the change induced by fire in the wider landscape. We found the two species responded differently to the presence of unburnt refugia in the landscape. Relative to controls, fire had little effect on bush rat abundance in gullies, supporting hypothesis 2. In contrast, agile antechinus abundance increased in gullies immediately post‐fire consistent with a shift of individuals from burnt parts of the landscape, supporting hypothesis 1. Differences in site fidelity, habitat use and intraspecific competition between these species are suggested as likely factors influencing responses to refugia. The way unburnt patches function as faunal refugia and the subsequent influence they have on post‐fire population dynamics, will to some extent depend on the life history attributes of individual species.     

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.     

Immediate and short‐term responses of bird and mammal assemblages to a subalpine wildfire in the Snowy Mountains,Australia

Abstract Over 35 days in January–February 2003, wildfires burnt across much of the subalpine/alpine landscape of south‐eastern Australia, including about 70% of the land above 1500 m in the Snowy Mountains. At the time of the fire, studies of two subalpine faunal assemblages were being undertaken. The opportunity to resurvey the assemblages was taken in order to examine the immediate impact of fire in an environment where it is uncommon but predicted to occur increasingly with global warming. A study area in the Whites River Valley, where the number of bird species was counted monthly from 1996 to 2003, and weekly in late winter–spring from 2000 to 2003, was burnt in one fire. As well as the expected decrease in the number of individual birds, the fire resulted in an immediate decrease in the number of bird species, unlike in previously studied montane forest, with only the regularly wintering species plus the olive whistler and the ground‐feeding flame robin remaining. During the post‐winter avian immigration, few regular spring migrants appeared on burnt sites despite their nearby presence on the unburnt side of the valley. Five of six small mammal trapping grids were burnt. As with fires at lower altitudes, there was an immediate reduction in mammal numbers on burnt grids following the fire, but in addition, one species, Mastacomys fuscus, declined further in the ensuing 2 months both on burnt and unburnt sites. Numbers of Antechinus swainsonii and Rattus fuscipes stabilized until autumn/winter when there was a further decline due to the unavailability of subnivean space to allow winter foraging, allied with a concentration of fox predation on areas still carrying populations of small mammals.     

The response of cerambycid beetles (Coleoptera: Cerambycidae) to long‐term fire frequency regimes in subtropical eucalypt forest

Fire has a varied influence on plant and animal species through direct (e.g. fire‐induced mortality) and indirect (e.g. modification of habitat) effects. Our understanding of the influence of fire regime on invertebrates and their response to fire‐induced modifications to habitat is poor. We aimed to determine the response of a beetle family (Coleoptera: Cerambycidae) to varying fire treatments and hypothesised that the abundance of cerambycid beetles is influenced by fire frequency due to modifications in habitat associated with the fire treatments. Arthropods were sampled across 3 months in annually and triennially burnt areas (treatments starting in 1952 and 1973 respectively), an area unburnt since 1946, and a former unburnt treatment, burnt by wildfire in 2006. Eleven different cerambycid taxa were collected using flight intercept panel traps, dominated by three species (Ipomoria tillides, Adrium sp. and Bethelium signiferum) which made up 99% of individuals collected. Over the sampling period the long unburnt treatment had significantly lower species richness than the triennial and wildfire treatments. Cerambycid abundance was significantly higher in the triennially burnt treatment than in all other fire treatments. Ipomoria tillides was more abundant in both frequently burnt treatments, Adrium sp. was more common in triennially burnt areas, whereas B. signiferum, was more common in the wildfire affected treatment. Some, but not all, cerambycid beetles were more common in areas with a more open understorey (i.e. resulting from frequent burning), and lower tree basal area, as this likely influences their ability to fly easily between food sources. Cerambycid abundance was positively related to the volume of coarse woody debris and healthy tree crowns. Cerambycid beetles were clearly influenced by historic fire regime, suggesting that changes in fire regime can potentially have a profound influence on arthropod assemblages, and subsequent influences on ecosystem processes, which are currently poorly understood.     

Resprouting of saplings following a tropical rainforest fire in north‐east Queensland,Australia

Abstract In 2002, fire burnt areas of Mesophyll‐ and Notophyll Vine Forest in the Smithfield Conservation Park near Cairns, Australia. We assessed the ability of rainforest plant species to persist through fire via resprouting. Natural rates of mortality and resprouting in unburnt areas were assessed for all saplings (stems < 2 m) via 13, 2 × 50 m belt transects, and compared to estimates of mortality and resprouting in 26 transects in burnt areas. We also tested the resprouting ability per‐individual stem of each species against all other stems with which it co‐occurred. Totals of 1242 stems (138 species) were sampled in burnt transects and 503 stems (95 species) in unburnt transects (total number of unique species = 169). There was no difference in the number of stems existing prior to the fire in burnt and unburnt areas when expressed on a per‐sample area basis. Resprouting from basal shoots and root suckers was significantly greater in burnt than in unburnt areas, but rates of stem sprouting were not different. In burnt areas 72 species were tested for resprouting ability and most (65/72) resprouted at similar rates. All species analysed contained individuals that resprouted. The resprouting response of five species was significantly lower, and in two species was significantly higher. For these species especially, fire may act as a mechanism altering relative abundances. The fire coincided with an extreme El Niño event. Current predictions indicate El Niño conditions may become increasingly common, suggesting fire events within rainforest could become more frequent. Resprouting as a general phenomenon of rainforest species, and differential resprouting ability between species should therefore be an important consideration in assessing the potential path of vegetation change in rainforests after fire.     

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.     

Fine‐scale refuges can buffer demographic and genetic processes against short‐term climatic variation and disturbance: a 22‐year case study of an arboreal marsupial

Ecological disturbance and climate are key drivers of temporal dynamics in the demography and genetic diversity of natural populations. Microscale refuges are known to buffer species’ persistence against environmental change, but the effects of such refuges on demographic and genetic patterns in response to short‐term environmental variation are poorly understood. We quantified demographic and genetic responses of mountain brushtail possums (Trichosurus cunninghami) to rainfall variability (1992–2013) and to a major wildfire. We hypothesized that there would be underlying differences in demographic and genetic processes between an unburnt mesic refuge and a topographically exposed zone that was burnt in 2009. Fire caused a 2‐year decrease in survival in the burnt zone, but the population grew after the fire due to immigration, leading to increased expected heterozygosity. We documented a fire‐related behavioural shift, where the rate of movement by individuals in the unburnt refuge to the burnt zone decreased after fire. Irrespective of the fire, there were long‐term differences in demographic and genetic parameters between the mesic/unburnt refuge and the nonmesic/burnt zone. Survival was high and unaffected by rainfall in the refuge, but lower and rainfall‐dependent in the nonmesic zone. Net movement of individuals was directional, from the mesic refuge to the nonmesic zone, suggesting fine‐scale source–sink dynamics. There were higher expected heterozygosity (H_E) and temporal genetic stability in the refuge, but lower H_E and marked temporal genetic structure in the exposed habitat, consistent with reduced generational overlap caused by elevated mortality and immigration. Thus, fine‐scale refuges can mediate the short‐term demographic and genetic effects of climate and ecological disturbance.     

Post‐fire seed predation: Does distance to unburnt vegetation matter?

Human‐induced changes to fire regimes result in smaller, more patchy fires in many peri‐urban areas, with a concomitant increase in potential edge effects. In sclerophyll vegetation, many structurally dominant serotinous plants rely on the immediate post‐fire environment for recruitment. However, there is little information about how fire attributes affect seed predation or recruitment for these species. We examined the influence of distance to unburnt vegetation on post‐dispersal seed predation for five serotinous species from sclerophyll vegetation in the Sydney region, south‐eastern Australia; Banksia serrata L.f., Banksia spinulosa Sm. var. spinulosa, Hakea gibbosa (Sm.) Cav., Hakea teretifolia (Salisb.) Britten (all Proteaceae) and Allocasuarina distyla (Vent.) L. Johnson (Casuarinaceae). We used cafeteria trials and differential exclusion of vertebrates and invertebrates to test whether rates of seed removal for these five species differed among (i) unburnt, (ii) burnt‐edge (approx. 10 m from unburnt vegetation) and (iii) burnt‐interior (approx. 100 m from unburnt vegetation) locations. When all animals had access to seeds, seeds were removed at lower rates from burnt‐interior areas than from other locations. Vertebrates (small mammals) showed this pattern markedly the first time the experiment was run, but in a repeat trial this effect disappeared. Rate of seed removal by invertebrates differed among plant species but we did not detect any such differences for removal by vertebrates. Overall rates of seed removal also differed significantly between the two fires studied. Our results indicate that small mammal seed predation can be substantial for large‐seeded serotinous shrubs, and that differences in the perimeter: area ratio, severity or size of a fire are likely to affect seed predation.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

Can stable carbon isotopes (δ13C) in soil carbon be used to describe the dynamics of Eucalyptus savanna–rainforest boundaries in the Australian monsoon tropics?

The history of isolated patches of monsoon rainforest within large tracts of Eucalyptus savanna is poorly understood because of the scarcity of reliable palaeoecological records in the Australian monsoon tropics. Elsewhere in the world, the ratio of the stable isotopes ¹³C to ¹²C (δ¹³C) in soil organic matter has shed light on the dynamics of rainforest–savanna boundaries because tropical grasses with the C4 photosynthetic pathway have a distinct δ¹³C signature (–17 to –9‰) compared with that of woody plants with the C3 photosynthetic pathway (–32 to –22‰). In order to determine the magnitude of the variation in δ¹³C, unreplicated soil profiles were sampled beneath different vegetation types on three boundaries between Eucalyptus savanna and rainforest that were both growing on Tertiary age laterite parent material. Replicated (n = 3) soil profiles, which were also derived from Tertiary age laterite, were sampled from beneath: (i) dense stands of African grasses within a frequently burnt Eucalyptus savanna; and within the same long unburnt Eucalyptus savanna, (ii) patches of African and natives grasses and (iii) clumps of Acacia trees. The strongly negative δ¹³C values of soil organic matter derived from the frequently burnt and long unburnt grassy understoreys in the Eucalyptus savannas showed that a considerable amount of the soil carbon was derived from C3 (woody) species despite the presence of a ground layer dominated by C4 grasses. However, a feature of these data was the considerable variability among the three ‘replicate’ profiles. The surface soil samples from beneath three clumps of Acacia trees in the unburnt Eucalyptus savanna had much less variable δ¹³C values and were similar to two of the three monsoon rainforests sampled. The pattern of δ¹³C values from unreplicated soil profiles from different vegetation types across three rainforest boundaries was also very variable and not always obviously related the known disturbance history of the extant vegetation. Given the considerable variability within and between vegetation types with contrasting disturbance histories, it is concluded that the use of carbon stable isotopes to advance understanding of the dynamics of rainforest and Eucalyptus savanna boundaries will require further development, such as determination of the ¹⁴C age and δ¹³C values of different soil carbon fractions.     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.     

Variable rainfall has a greater effect than fire on the demography of the dominant tree in a semi‐arid Eucalyptus savanna

Rainfall, fire and competition are emphasized as determinants of the density and basal area of woody vegetation in savanna. The semi‐arid savannas of Australia have substantial multi‐year rainfall deficits and insufficient grass fuel to carry annual fire in contrast to the mesic savannas in more northern regions. This study investigates the influence of rainfall deficit and excess, fire and woody competition on the population dynamics of a dominant tree in a semi‐arid savanna. All individuals of Eucalyptus melanophloia were mapped and monitored in three, 1‐ha plots over an 8.5 year period encompassing wet and dry periods. The plots were unburnt, burnt once and burnt twice. A competition index incorporating the size and distance of neighbours to target individuals was determined. Supplementary studies examined seedling recruitment and the transition of juvenile trees into the sapling layer. Mortality of burnt seedlings was related to lignotuber area but the majority of seedlings are fire resistant within 12 months of germination. Most of the juveniles (≤1 cm dbh) of E. melanophloia either died in the dry period or persisted as juveniles throughout 8.5 years of monitoring. Mortality of juveniles was positively related to woody competition and was higher in the dry period than the wet period. The transition of juveniles to a larger size class occurred at extremely low rates, and a subsidiary study along a clearing boundary suggests release from woody competition allows transition into the sapling layer. From three fires the highest proportion of saplings (1–10 cm dbh) reduced to juveniles was only 5.6% suggesting rates of ‘top‐kill’ of E. melanophloia as a result of fire are relatively low. Girth growth was enhanced in wet years, particularly for larger trees (>10 cm dbh), but all trees regardless of size or woody competition levels are vulnerable to drought‐induced mortality. Overall the results suggest that variations in rainfall, especially drought‐induced mortality, have a much stronger influence on the tree demographics of E. melanophloia in a semi‐arid savanna of north‐eastern Australia than 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.     

Demographic responses of an arboreal marsupial,the common brushtail possum (<Emphasis Type="Italic">Trichosurus vulpecula</Emphasis>), to a prescribed fire

We investigated demographic responses of the common brushtail possum Trichosurus vulpecula, a medium-sized arboreal marsupial, after a prescribed fuel reduction burn on Magnetic Island, tropical north Queensland, Australia. Possums were live-trapped every month for 14 months before the fire and 11 months after the fire in both the burnt and unburnt areas; measurements of individuals were taken each month and demographic parameters were modelled using capture–mark–recapture data. Significant differences between the burnt and unburnt sites were found following the fire; recruitment was lower in the unburnt area, where population size also declined. In the burnt area, population size and recruitment displayed a tendency to increase after the fire, while capture probability declined, suggesting that an influx of new individuals, attracted to re-sprouting vegetation, had resulted in trap saturation. There was no detectable effect of the fire on survival, and no fire-induced mortalities were observed. We conclude that a low-intensity, prescribed, fuel-reduction burn had no obvious negative consequences for this possum population.