The effects of topographic variation and the fire regime on coarse woody debris: insights from a large wildfire

Coarse woody debris (CWD) is a common structural component of terrestrial ecosystems, and provides important habitat for biota. Fires modify the distribution of CWD, both spatially and temporally. Changes in fire regimes, such as those arising from prescribed burning and changing climatic conditions, make it critical to understand the response of this resource to fire. We created a conceptual model of the effects of fire on logs and dead trees in topographically diverse forests in which trees often survive severe fire. We then surveyed paired sites, in a damp gully and adjacent drier slope, ~3.5. years after a large wildfire in south-eastern Australia. Sites were stratified by fire severity (unburnt, understorey burnt and severely burnt), and fire history (burnt ≤3. years or ≥20. years prior to the wildfire). Both components of the fire regime influenced CWD availability in gullies. Severe wildfire and fire history ≤3. years reduced the volume of small logs (10-30. cm diameter) in gullies, while severe wildfire increased the number of large dead trees in gullies. CWD on slopes was not affected by fire severity or history at ~3.5. years post-fire. Log volumes on slopes may recover more quickly after wildfire through rapid collapse of branches and trees. Gullies generally supported more logs than slopes, but longer inter-fire intervals in gullies may allow fuel loads to accumulate and lead to comparatively larger fire impacts. Given that fire severity and fire interval are predicted to change in many fire-prone ecosystems in coming decades, this study highlights the importance of understanding the interacting effects of multiple components of the fire regime with landscape structure. In particular, variation in fire interval and fire severity in relation to topographic position will influence the pattern of accumulation of coarse woody debris across the landscape, and therefore the structure and quality of habitats for biota.

Effects of the fire regime on mammal occurrence after wildfire: site effects vs landscape context in fire-prone forests

Wildfires have major impacts on ecosystems globally. Fire regimes (including fire frequency, intensity, season and type of fire) influence the status of species by altering habitat suitability at the site scale, and by creating heterogeneity at the landscape scale. The relative effects of site and landscape-scale fire attributes on animal species are rarely examined together. Such knowledge is important, given that fire regimes are sensitive to changing land management practices; and that fires are predicted to become larger and more frequent in some regions as a result of climate change. Here, we tested the relative influence of elements of the fire regime (fire severity, fire history) at the site-scale, and the landscape context (extent of surrounding unburnt forest, fire heterogeneity) on the occurrence of native terrestrial mammals after severe wildfire in south-eastern Australia. We conducted surveys by using automatically triggered, infrared cameras at 80 sites in fire-prone eucalypt forests, 2-3. years post-wildfire. Thirteen native mammal species were recorded, eight of which were detected with sufficient frequency for analysis. Most species were widespread (35-90% of sites) and recorded in all fire severity classes. Fire effects at the site-level were more influential than landscape context effects arising from heterogeneity in the fire regime (e.g. extent of surrounding unburnt forest). Fire severity was the most influential of the fire-regime elements investigated, but it affected different species in different ways. This study highlights three main points relevant to conservation of terrestrial mammals after wildfire. First, spatial variation in fire severity associated with wildfire (ranging from unburned to severely burned stands) is an important contributor to the post-fire status of species. Second, post-fire environmental conditions are significant: here, rapid regeneration of vegetation following drought-breaking rains greatly influenced the suitability of post-fire habitats. Third, it is valuable to consider the effects of the fire regime at multiple scales, including both the site (forest stand) and its landscape context. Insights from short-term surveys, such as this, will be enhanced by complementary longitudinal studies, especially where they encompass environmental variation through the post-fire succession.

Fire and its interaction with ecological processes in box-ironbark forests

Box-Ironbark forests extend across a swathe of northern Victoria on the inland side of the Great Dividing Range. Although extensively cleared and modified, they support a distinctive suite of plants and animals. Historical fire regimes in this ecosystem are largely unknown, as are the effects of fire on most of the biota. However, knowledge of the ecological attributes of plant species has been used to determine minimum and maximum tolerable fire intervals for this ecosystem to guide current fire management. Here, we consider the potential effects of planned fire in the context of major ecological drivers of the current box-ironbark forests: namely, the climate and physical environment; historical land clearing and fragmentation; and extractive land uses. We outline an experimental management and research project based on application of planned burns in different seasons (autumn, spring) and at different levels of burn cover (patchy, extensive). A range of ecological attributes will be monitored before and after burns to provide better understanding of the landscape-scale effects of fire in box-ironbark forests. Such integration of management and research is essential to address the many knowledge gaps in fire ecology, particularly in the context of massively increased levels of planned burning currently being implemented in Victoria.

A landscape-scale, applied fire management experiment promotes recovery of a population of the threatened Gouldian Finch, Erythrura gouldiae, in Australia's tropical savannas

Fire is an integral part of savanna ecology and changes in fire patterns are linked to biodiversity loss in savannas worldwide. In Australia, changed fire regimes are implicated in the contemporary declines of small mammals, riparian species, obligate-seeding plants and grass seed-eating birds. Translating this knowledge into management to recover threatened species has proved elusive. We report here on a landscape-scale experiment carried out by the Australian Wildlife Conservancy (AWC) on Mornington Wildlife Sanctuary in northwest Australia. The experiment was designed to understand the response of a key savanna bird guild to fire, and to use that information to manage fire with the aim of recovering a threatened species population. We compared condition indices among three seed-eating bird species-one endangered (Gouldian finch) and two non-threatened (long-tailed finch and double-barred finch)-from two large areas (> 2,830 km2) with initial contrasting fire regimes ('extreme': frequent, extensive, intense fire; versus 'benign': less frequent, smaller, lower intensity fires). Populations of all three species living with the extreme fire regime had condition indices that differed from their counterparts living with the benign fire regime, including higher haematocrit levels in some seasons (suggesting higher levels of activity required to find food), different seasonal haematocrit profiles, higher fat scores in the early wet season (suggesting greater food uncertainty), and then lower muscle scores later in the wet season (suggesting prolonged food deprivation). Gouldian finches also showed seasonally increasing stress hormone concentrations with the extreme fire regime. Cumulatively, these patterns indicated greater nutritional stress over many months for seed-eating birds exposed to extreme fire regimes. We tested these relationships by monitoring finch condition over the following years, as AWC implemented fire management to produce the 'benign' fire regime throughout the property. The condition indices of finch populations originally living with the extreme fire regime shifted to resemble those of their counterparts living with the benign fire regime. This research supports the hypothesis that fire regimes affect food resources for savanna seed-eating birds, with this impact mediated through a range of grass species utilised by the birds over different seasons, and that fire management can effectively moderate that impact. This work provides a rare example of applied research supporting the recovery of a population of a threatened species.

Dominant drivers of seedling establishment in a fire-dependent obligate seeder: climate or fire regimes?

Climate change is causing fire regime shifts in ecosystems worldwide. Plant species with regeneration strategies strongly linked to a fire regime, such as obligate seeders, may be particularly threatened by these changes. It is unclear whether changes in fire regimes or the direct effects of climate change will be the dominant threats to obligate seeders in future. We investigated the relative importance of fire-related variables (fire return interval andfire severity) and environmental factors (climate and topography) on seedling establishment in the world's tallest angiosperm, an obligate seeder, Eucalyptus regnans. Throughout its range, this species dominates the wet montane forests of south-eastern Australia and plays a keystone role in forest structure. Following major wildfires, we investigated seedling establishment in E. regnanswithin 1 year of fire as this is a critical stage in the regeneration niche of obligate seeders. Seedling presence and abundance were strongly related to the occurrence of fire but not to variation in fire severity (moderate vs. high severity). Seedling abundance increased with increasing fire return interval (range 26-300 years). First-year seedling establishment was also strongly associated with low temperatures and with high elevations, high precipitation and persistent soil water availability. Our results show that both climate and fire regimes are strong drivers of E. regnans seedling establishment. The predicted warming and drying of the climate might reduce the regeneration potential for some obligate seeders in future and these threats are likely to be compounded by changes in fire regimes, particularly increases in fire frequency.

Life history influences how fire affects genetic diversity in two lizard species

'Fire mosaics' are often maintained in landscapes to promote successional diversity in vegetation with little understanding of how this will affect ecological processes in animal populations such as dispersal, social organization and re-establishment. To investigate these processes, we conducted a replicated, spatiotemporal landscape genetics study of two Australian woodland lizard species [Amphibolurus norrisi (Agamidae) and Ctenotus atlas (Scincidae)]. Agamids have a more complex social and territory structure than skinks, so fire might have a greater impact on their population structure and thus genetic diversity. Genetic diversity increased with time since fire in C. atlas and decreased with time since fire in A. norrisi. For C. atlas, this might reflect its increasing population size after fire, but we could not detect increased gene flow that would reduce the loss of genetic diversity through genetic drift. Using landscape resistance analyses, we found no evidence that postfire habitat succession or topography affected gene flow in either species and we were unable to distinguish between survival and immigration as modes of postfire re-establishment. In A. norrisi, we detected female-biased dispersal, likely reflecting its territorial social structure and polygynous mating system. The increased genetic diversity in A. norrisi in recently burnt habitat might reflect a temporary disruption of its territoriality and increased male dispersal, a hypothesis that was supported with a simulation experiment. Our results suggest that the effects of disturbance on genetic diversity will be stronger for species with territorial social organization.

Fire severity and landscape context effects on arboreal marsupials

Although fire is a major form of natural disturbance worldwide, both fire-derived landscape context effects and the impacts of fire severity are poorly known for many species. To address this knowledge gap, we quantified the response of Australian arboreal marsupials to: (1) the spatial effects of fire, (2) fire severity, and (3) fire impacts on the availability of critical nesting resources - hollow-bearing trees.We identified substantial differences among species in response to fire severity and landscape-scale fire. The Sugar Glider (Petaurus breviceps) and the endangered Leadbeater's Possum (Gymnobelideus leadbeateri) were extremely rare on burned sites irrespective of fire severity. In addition, these two species declined with the amount of burned forest in the surrounding landscape even when their habitat remained unburnt. The Mountain Brushtail Possum (Trichosurus cunninghami) and the Greater Glider (Petauroides volans) both occurred on burned and unburned sites. The Greater Glider responded negatively to fire severity at the site level and also negatively to the amount of forest burned in the surrounding landscape. The abundance of the Mountain Brushtail Possum was lowest on sites subject to moderate severity fire.On unburned sites, the presence and abundance of virtually all species was characterised by a common positive response to the availability of nesting resources in hollow-bearing trees.Our findings underscore the importance of management practices to better protect species that decline after fire. These include conserving areas of unburned forest, particularly those with hollow-bearing trees which are critical nest sites for arboreal marsupials. These recommendations are currently the opposite of existing management practices.

Can the intermediate disturbance hypothesis and information on species traits predict anuran responses to fire?

Fire is a common form of recurrent disturbance in many ecosystems, but ecological theory has a poor record of predicting animal responses to fire, at both species and assemblage levels. As a consequence, there is limited information to guide fire regime management for biodiversity conservation. We investigated a key research gap in the fire ecology literature; that is, the response of an anuran assemblage to variation in the fire return interval. We tested two hypotheses using a spatially-explicit fire database collected over a 40 year period: 1) species richness would peak at intermediate levels of disturbance. 2) Species with traits which enabled them to escape fire - burrowing or canopy dwelling - would be better able to survive fires, resulting in higher levels of occurrence in frequently burned sites. We found no evidence for either a reduction in species richness at locations with short fire return intervals, or a peak in species richness at intermediate levels of disturbance. Although we found some support for individual species responses to fire return intervals, these were inconsistent with the interpretation of burrowing or climbing being functional traits for fire-avoidance. Instead burrowing and climbing species may be more likely to be disadvantaged by frequent fire than surface dwelling frogs. More generally, our results show that many species in our study system have persisted despite a range of fire frequencies, and therefore that active management of fire regimes for anuran persistence may be unnecessary. The responses of anurans to fire in this location are unlikely to be predictable using simple life-history traits. Future work should focus on understanding the mechanistic underpinnings of fire responses, by integrating information on animal behavior and species' ecological requirements.

Post-fire succession affects abundance and survival but not detectability in a knob-tailed gecko

Altered fire regimes threaten the persistence of many animal species globally, thus understanding how fire affects demographic processes is critical for conservation. Using 2. years of mark-recapture data from the Australian gecko Nephrurus stellatus, we investigated the effect of fire on (i) detectability to reliably measure post-fire changes in abundance, and (ii) survival and reproductive rates to investigate the mechanisms of successional change. Data were collected from two conservation reserves each with three different fire categories based on time since the last fire "Early", "medium" and "late" sites had 2-3, 7-9 and 42-48. years since fire, respectively. A robust design modelling framework was used to estimate the effect of fire category on abundance, survival and capture probability while also examining the influence of temperature and behaviour on detectability. Geckos showed trap-shy behaviour and detectability increased significantly with increasing temperature but was not affected by time since fire. Accounting for detectability, geckos were more abundant in the medium than the early sites, and were rare in the late sites. Although trends in survival are more difficult to address with short-term data, our results showed lower monthly survival rates, but higher fecundity in the early than the medium sites. These results were possibly related to successional changes in predation, the thermal environment, and food availability. We demonstrated how mark-recapture analysis can show the causes of animal fire responses while realistically accounting for detectability. Such information is necessary to provide a predictive framework to guide fire management for biodiversity. .

The dynamic regeneration niche of a forest following a rare disturbance event

Aim: Knowledge of how climate and fire regimes affect regeneration in foundation species is critical to the conservation of entire ecosystems. Different stages of regeneration often require different ecological conditions, but dynamic constraints on regeneration are poorly known for species that regenerate only after infrequent wildfires. Focussing on a long-lived, foundation tree species (Eucalyptus regnans), we tested the hypothesis that the relative importance of fire regime variables (fire severity and time since previous fire) and environmental gradients on post-fire regeneration would shift as seedlings developed. Location: South-eastern Australia. Methods: Following a large (> 59,000 ha) summer wildfire in 2009, we sampled 131 sites (61 burnt) annually for four years (2009-2012), representing the range of environmental conditions in which E. regnans occurs. We analysed the effect of fire severity, time since fire and environmental variables on early regeneration processes critical for post-fire species distributions: seedling establishment, seedling density and growth through different height stages (10 cm, 25 cm, 50 cm and 200 cm). Results: The regeneration niche of E. regnans was defined by different factors at different stages of development. Initially, seedlings established prolifically on burnt sites, regardless of severity. Three years into the regeneration process, high-severity fire became the dominant driver of seedling persistence and growth over 25 cm. Growth over 50 cm was dependent on environmental conditions relating to elevation and precipitation. Main conclusions: Our results describe how fire occurrence, fire severity and environmental gradients affected seedling establishment, persistence and growth. The dynamic constraints on regeneration likely reflect temporal changes in the biotic and abiotic environment and variation in resource requirements during the early post-fire years. Our findings will enable more accurate forecasts of species distributions to assist forest conservation in the face of global changes in climate and fire regimes.

Conserving long unburnt vegetation is important for bird species, guilds and diversity

Landscape-level wildfires have a major role in structuring faunal assemblages, particularly in fire-prone landscapes. These effects are mediated by changes to vegetation structure and composition that directly influence the availability of shelter, feeding and breeding resources. We investigated the response of a semi-arid shrubland bird community in Western Australia to the prevailing fire regime by examining the abundance, diversity and guild structure in relation to time since fire. We also examined vegetation structural attributes in relation to time since fire. We surveyed 32 sites ranging in age from 12 to 84 years since last fire. A total of 845 birds from 40 species were recorded. Vegetation structure varied with fire history with old and very old sites characterised by less bare ground, more leaf litter cover and greater canopy cover. Bird community composition varied with time since fire, driven by increased bird species richness and abundance of insectivores, granivores/frugivores, golden whistlers, grey shrike-thrush and red-capped robins with time since fire. Frequent, intense landscape-scale fires transform the landscape into homogeneous young shrublands, which may render vegetation unsuitable for several species and guilds.

Reptile responses to fire and the risk of post-disturbance sampling bias

Altered fire regimes are a driver of biodiversity decline. To plan effective management, we need to know how species are influenced by fire and to develop theory describing fire responses. Animal responses to fire are usually measured using methods that rely on animal activity, but animal activity may vary with time since fire, potentially biasing results. Using a novel approach for detecting bias in the pit-fall trap method, we found that leaf-litter dependent reptiles were more active up to 6 weeks after fire, giving a misleading impression of abundance. This effect was not discovered when modelling detectability with zero-inflated binomial models. Two species without detection bias showed early-successional responses to time since fire, consistent with a habitat-accommodation succession model. However, a habitat specialist did not have the predicted low abundance after fire due to increased post-fire movement and non-linear recovery of a key habitat component. Interactions between fire and other processes therefore must be better understood to predict reptile responses to changing fire-regimes. We conclude that there is substantial bias when trapping reptiles after fire, with species that are otherwise hard to detect appearing to be abundant. Studies that use a survey method based on animal activity such as bird calls or animal movements, likely face a similar risk of bias when comparing recently-disturbed with control sites.