Post‐fire development of faunal habitat depends on plant regeneration traits

The concept that vegetation structure (and faunal habitat) develops predictably with time since fire has been central to understanding the relationship between fire and fauna. However, because plants regenerate after fire in different ways (e.g. resprouting from above‐ground stems vs. underground lignotubers), use of simple categories based on time since fire might not adequately represent post‐fire habitat development in all ecosystems. We tested the hypothesis that the post‐fire development of faunal habitat structure differs between ecosystems, depending on fire regeneration traits of the dominant canopy trees. We measured 12 habitat components at sites in foothill forests (n = 38), heathy woodlands (n = 38) and mallee woodlands (n = 98) in Victoria, Australia, and used generalised additive models to predict changes in each variable with time since fire. A greater percentage of faunal habitat variables responded significantly to time since fire in mallee woodlands, where fires typically are stand‐replacing, than in foothill forests and heathy woodlands, where canopy tree stems generally persist through fire. In the ecosystem with the highest proportion of epicormic resprouters (foothill forests), only ground cover and understorey vegetation responded significantly to time since fire, compared with all but one variable in the ecosystem dominated by basal resprouters (mallee woodlands). These differences between ecosystems in the post‐fire development of key habitat components suggest there may also be fundamental differences in the role of fire in shaping the distribution of fauna. If so, this challenges the way in which many fire‐prone ecosystems currently are categorised and managed, especially the level of dependence on time since fire and other temporal surrogates such as age‐classes and successional states. Where time since fire is a poor surrogate for habitat structural development, additional complexity (e.g. fire severity, topography and prior land‐use history) could better capture processes that determine faunal occurrence in fire‐prone ecosystems.     

Effects of time since fire on birds in a plant diversity hotspot

Global changes are influencing fire regimes in many parts of the world. In the Fynbos plant diversity hotspot (Cape Floristic Region, South Africa), fire frequency has increased in protected areas where the mean fire interval went from 12–19 to 6–9 years between 1970 and 2000. Fire is one of the main drivers of plant diversity in the Cape Floristic Region. Too frequent fires threaten the persistence of slow-maturing plant species, and such insights have led to the adoption of fire management principles based on plant responses. The effects of fire on Fynbos fauna are much more poorly understood, and have not generally been considered in depth in Fynbos conservation policies, planning or management. We assessed the response of bird communities to long-term fire-induced vegetation changes using space-for-time substitution. We studied bird communities, vegetation structure and plant functional composition in 84 Fynbos plots burnt between two and 18 years before. Ten of the 14 bird species analysed showed a significant change in their abundance with time since fire. We observed a significant species turnover along the post-fire succession due to changes both in vegetation structure and plant functional composition, with a characteristic shift from non-Fynbos specialists and granivorous species to Fynbos specialists and nectarivorous species.If current trends of increasing fire frequency continue, Fynbos endemic birds such as nectarivores may become vulnerable. Conservation management should thus aim more carefully to maintain mosaics of Fynbos patches of different ages. Future research needs to estimate the proportion of vegetation of different ages and patch sizes needed to support dependent fauna, particularly endemics.     

Rainforest expansion reduces understorey plant diversity and density in open forest of eastern Australia

The expansion of rainforest pioneer trees into long‐unburnt open forests has become increasingly widespread across high rainfall regions of Australia. Increasing tree cover can limit resource availability for understorey plant communities and reduce understorey diversity. However, it remains unclear if sclerophyll and rainforest trees differ in their competitive exclusion of understory plant communities, which contain most of the floristic diversity of open forests. Here, we examine dry open forest across contrasting fire histories (burnt and unburnt) and levels of rainforest invasion (sclerophyll or rainforest midstorey) to hindcast changes in understorey plant density, richness and composition. The influence of these treatments and other site variables (midstorey structure, midstorey composition and soil parameters) on understorey plant communities were all examined. This study is the first to demonstrate significantly greater losses of understorey species richness, particularly of dry open‐forest specialists, under an invading rainforest midstorey compared to a typical sclerophyll midstorey. Rainforest pioneers displaced over half of the understorey plant species, and reduced ground cover and density of dry forest specialists by ~90%. Significant understorey declines also occurred with increased sclerophyll midstorey cover following fire exclusion, although losses were typically less than half that of rainforest‐invaded sites over the same period. Understorey declines were closely related to leaf area index and basal area of rainforest and wattle trees, suggesting competitive exclusion through shading and potentially belowground competition for water. Around 20% of displaced species lacked any capacity for population recovery, while transient seed banks or distance‐limited dispersal may hinder recovery for a further 68%. We conclude that rainforest invasion leads to significant declines in understorey plant diversity and cover in open forests. To avoid elimination of local native plant populations in open forests, fires should occur with sufficient frequency to prevent overstorey cover from reaching a level where shade‐intolerant species fail to thrive.     

Detecting impacts of non‐native species on associated invertebrate assemblages depends on microhabitat

Invasive plants that displace native floral communities can cause changes to associated invertebrate species assemblages. Using a mini‐review of the literature and our own data we add to the still considerable debate about the most effective methods for testing community‐level impacts by invasive species. In endangered saltmarshes of southeast Australia, the non‐native rush Juncus acutus L. is displacing its native congener J. kraussii Hochst., with concurrent changes to floral and faunal assemblages. In two coastal saltmarshes, we tested the hypothesis that the ability to detect differences in the invertebrate assemblage associated with these congeneric rushes depends on the microhabitat of the plant sampled. We used three sampling methods, each targeting specific microhabitats: sweep netting of the plant stems, vacuum sampling of the plant tussock, and vacuum sampling of the ground directly below the plants. Over 3800 individuals and 92 morphospecies were collected across four main taxa: gastropods, crustaceans, hexapods and arachnids. Detection of differences in invertebrate density, richness and composition associated with native compared with non‐native rushes was dependent on the microhabitat sampled and these differences were spatially variable. For example, at one saltmarsh the stems and tussock of J. acutus had a lower density and richness of total invertebrates and hexapods than those of the native J. kraussii. In contrast, crustaceans on the ground were in greater abundance below J. acutus than J. kraussii. This study demonstrates that on occasions where overall differences in the assemblage are not detected between species, differences may become apparent when targeting different microhabitats of the plant. In addition, separately targeting multiple microhabitats likely leads to a greater probability of detecting impacts of invasion. Comparing the invertebrate assemblage without differentiating between or sampling an array of microhabitats can fail to determine the impact of invasive species. These results highlight that a combination of methods targeting different microhabitats is important for detecting differences within the invertebrate community, even for phylogenetically related species.     

Germination responses of native and invasive Cerrado grasses to simulated fire temperatures

Background: Fire is an important ecological factor in the Cerrado (Brazilian savanna). However, comparative studies on the effect of high temperatures experienced during fires on seed germination of native and invasive grass species are few.

Aims: To assess germination responses to simulated fire temperatures by seeds of invasive and native Cerrado grasses.

Methods: Heat-shock treatments (50 °C, 70 °C, 90 °C, 110 °C, 130 °C or 150 °C) were applied to seeds of 10 species of native and invasive grasses. For each temperature, the seeds were heated in a dry-air flow for 2 or 5 min. This combination of temperatures and exposure times simulated the soil conditions during typical Cerrado fires.

Results: Temperature treatment was significantly related to germination, and the effect varied according to species. Heat shock did not increase germination in either the native or the invasive species. Exposure time was important for only two species, and four species showed a significant increase in mean germination time.

Conclusions: Species showed different tolerances to high temperatures. It was not possible to differentiate the native and invasive grasses only by their tolerance to high temperatures, suggesting that fire alone may not be an efficient management tool to control the invasive species studied here.     

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.     

Grass bud responses to fire in a semiarid savanna system

Increasingly, land managers have attempted to use extreme prescribed fire as a method to address woody plant encroachment in savanna ecosystems. The effect that these fires have on herbaceous vegetation is poorly understood. We experimentally examined immediate (<24 hr) bud response of two dominant graminoids, a C₃ caespitose grass, Nassella leucotricha, and a C₄ stoloniferous grass, Hilaria belangeri, following fires of varying energy (J/m²) in a semiarid savanna in the Edwards Plateau ecoregion of Texas. Treatments included high‐ and low‐energy fires determined by contrasting fuel loading and a no burn (control) treatment. Belowground axillary buds were counted and their activities classified to determine immediate effects of fire energy on bud activity, dormancy, and mortality. High‐energy burns resulted in immediate mortality of N. leucotricha and H. belangeri buds (p < .05). Active buds decreased following high‐energy and low‐energy burns for both species (p < .05). In contrast, bud activity, dormancy, and mortality remained constant in the control. In the high‐energy treatment, 100% (n = 24) of N. leucotricha individuals resprouted while only 25% (n = 24) of H. belangeri individuals resprouted (p < .0001) 3 weeks following treatment application. Bud depths differed between species and may account for this divergence, with average bud depths for N. leucotricha 1.3 cm deeper than H. belangeri (p < .0001). Synthesis and applications: Our results suggest that fire energy directly affects bud activity and mortality through soil heating for these two species. It is imperative to understand how fire energy impacts the bud banks of grasses to better predict grass response to increased use of extreme prescribed fire in land management.     

Sampling grain influences trends in vegetation composition and diversity with time since fire in Australian heathland

Scale‐dependency of pattern and process is well‐understood for many ecological communities; however, the influence of spatial scale (sampling grain) in detecting temporal change in communities is less well‐understood. The temperate lowland heathlands of south‐east Australia are one of the most fire‐prone ecosystems on earth. Despite the extensive literature documenting the effect of time since fire on heathlands, we know little about how sampling grain influences trends in vegetation variables over time, and whether these trends are scale‐dependent. Using 3500 ha of heathland in the Gippsland Lakes Coastal Park, south‐east Australia, we investigated how above‐ground species composition and diversity, and trends in these variables with increasing time since fire, were influenced by sampling grain (1 m², 10 m², 100 m², 900 m², 1 ha, 4 ha). Sampling grain influenced patterns detected in vegetation variables and in some instances, significantly affected their relationship with time since fire. Richness decreased with time since fire, with mean richness decreasing at three of the four grains, while total richness decreased at half of the sampled grains. Evenness (J) decreased with increasing time since fire for all grains except 1 m². The decline in diversity (H) with time since fire appeared to be independent of scale, as all grains decreased significantly with increasing time since fire. Community heterogeneity demonstrated a weak response to time since fire across most grains. Changes in composition among young (0–6 years since fire), intermediate (9–19 years) and old (23–27 years) sites were dependent on sampling grain, with all grains exhibiting significant differences in composition, apart from the 1 m² grain and the 100 m² grain (presence/absence data). Overall, species composition, richness, evenness, diversity and community heterogeneity were dependent on the scale at which the vegetation was sampled. In addition, trends in many of these vegetation variables with increasing time since fire were scale‐dependent. This work provides strong evidence that sampling at multiple grains contributes substantially to understanding pattern and process in heathlands.     

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.     

Herbaceous vegetation responses to experimental fire in savannas and forests depend on biome and climate

Fire–vegetation feedbacks potentially maintain global savanna and forest distributions. Accordingly, vegetation in savanna and forest ecosystems should have differential responses to fire, but fire response data for herbaceous vegetation have yet to be synthesized across biomes. Here, we examined herbaceous vegetation responses to experimental fire at 30 sites spanning four continents. Across a variety of metrics, herbaceous vegetation increased in abundance where fire was applied, with larger responses to fire in wetter and in cooler and/or less seasonal systems. Compared to forests, savannas were associated with a 4.8 (±0.4) times larger difference in herbaceous vegetation abundance for burned versus unburned plots. In particular, grass cover decreased with fire exclusion in savannas, largely via decreases in C₄ grass cover, whereas changes in fire frequency had a relatively weak effect on grass cover in forests. These differential responses underscore the importance of fire for maintaining the vegetation structure of savannas and forests.     

Ant community succession within eucalypt plantations on used pasture and implications for taxonomic sufficiency in biomonitoring

Abstract Ground‐active ants were sampled from three habitats: (i) a 10‐year‐old Eucalyptus punctata plantation, (ii) native woodland regrowth, and (iii) the surrounding pasture, at a study site in the Hunter Valley, New South Wales, Australia. A previous study, undertaken 6 years earlier at the same study sites, revealed no difference in species richness or composition between the eucalypt plantation and pasture. The aims of the present study were: (i) to investigate the successional change in ant community structure within the plantations; and (ii) to evaluate what levels of taxonomic identification were sufficient to indicate a change had taken place. Univariate statistics (anova ) were used to compare estimates of assemblage richness between habitats using data classified at five levels of taxonomic resolution: species, morphospecies, easily recognisable taxonomic units, genus and functional group. Multivariate statistics (anosim and non‐metric multidimensional scaling) were used to compare ant assemblages between habitats and between sampling events at a range of taxonomic resolutions from species to functional group. This study found: (i) a significant temporal change in community composition was evident using species, genus and functional group level data, but no change was detected in the pasture or woodland; (ii) mean ant species, morphospecies and easily recognisable taxonomic units richness were significantly greater within the plantations than the pasture; (iii) compositional differences between the plantation and pasture assemblages were evident at all levels of taxonomic resolution; (iv) mean ant species and genus richness were significantly higher in the woodland than in the plantation, and these two habitats were compositionally distinct at all levels of taxonomic resolution. This is the first case study to have documented a successional response from ants to the revegetation of agricultural land with eucalypt plantations. Reasons for the temporal and interhabitat differences in community structure are discussed, as well as the implications for taxonomic sufficiency in monitoring ant community successions.     

Land‐use and fire history effects on post‐fire vegetation dynamics in eastern Spain

Question: Is post‐fire, medium‐term vegetation dynamics determined by land‐use or fire history prior to fire? Location: South‐facing slope in the Gallinera valley, Alicante province, eastern Spain. Methods: After mapping the land‐use and fire history of the study site using photo‐interpretation, we sampled vegetation structure on a set of plots representing the most frequent land‐use and fire history combinations on an area burned six years before sampling. We studied the effects of land‐use history, comparing the one‐fire land‐use trajectories. We analysed the effects of fire history; comparing one‐ and two‐fire plots for both previously cropped and uncropped areas. Results: Most variables were not significantly different between the earliest abandoned plots (abandoned at least 38 years before the fire) and the uncropped plots. On the most recently abandoned plots (abandoned between one and four years before the fire), the therophyte richness and the ratio of seeder: resprouter richness were significantly greatest. Different fire recurrences did not determine different post‐fire vegetation on either the uncropped or the early abandoned plots (all dominated by fire‐recruited seeder shrubs). The most recently abandoned plots had a lower resilience to fire. Conclusions: Land‐use history and recent pre‐fire land use, in particular, determined the post‐fire vegetation in the medium term. The vegetation composition converged during secondary succession among land‐use histories. Increasing fire recurrence had a small effect on mature plant communities, due to the combination of life‐history traits determining the response to fire of the dominant species.     

Belowground community responses to fire: meta‐analysis reveals contrasting responses of soil microorganisms and mesofauna

Global fire regimes are shifting due to climate and land use changes. Understanding the responses of belowground communities to fire is key to predicting changes in the ecosystem processes they regulate. We conducted a comprehensive meta‐analysis of 1634 observations from 131 empirical studies to investigate the effect of fire on soil microorganisms and mesofauna. Fire had a strong negative effect on soil biota biomass, abundance, richness, evenness, and diversity. Fire reduced microorganism biomass and abundance by up to 96%. Bacteria were more resistant to fire than fungi. Fire reduced nematode abundance by 88% but had no significant effect on soil arthropods. Fire reduced richness, evenness and diversity of soil microorganisms and mesofauna by up to 99%. We found little evidence of temporal trends towards recovery within 10 years post‐disturbance suggesting little resilience of the soil community to fire. Interactions between biome, fire type, and depth explained few of these negative trends. Future research at the intersection of fire ecology and soil biology should aim to integrate soil community structure with the ecosystem processes they mediate under changing global fire regimes.     

Root responses of Quercus ilex L. seedlings to drought and fire

Abstract

Drought treatments in holm-oak (Quercus ilex) seedlings induce variations in total root length, number of root apices, shoot/root dry weight, and root electrolyte leakage. When drought treatments last for more than 50 days a considerable number of fine lateral roots die, irrespective of branching order or distribution within the root system. Scorching of drought-treated seedlings induces a transient stimulation of root growth. These results indicate that root turnover is deeply affected during treatments, with survival of seedlings being entrusted to the tolerance of a number of roots situated in the deeper region of the root system. Activity of the meristematic tissue present within the apices of these surviving roots supports regeneration of above-ground lost organs during recovery. Knowledge of the mechanisms ensuring the survival of Mediterranean tree seedlings following drought and fire is useful for developing models of vegetation dynamics.     

The responses of small mammals to patches regenerating after fire and rainfall in the Simpson Desert,central Australia

Abstract Patch‐burning is frequently advocated as a management tool to enhance the biodiversity and pasture values of spinifex (Triodia) grasslands. In this study we compare the capture rates of small mammals in habitats regenerating shortly after fire (aged 1–5 years) and in long‐unburnt habitats (aged >25 years). To unravel the effects of temporally and spatially variable rainfall on capture rates, the study was replicated at three locations spaced over 50 km apart that experience different rainfall regimes. Ten species of small mammals were captured over the course of the study, between October 1999 and June 2001. Pseudomys desertor showed a strong preference for long‐unburnt habitats. Notomys alexis, Sminthopsis youngsoni and Sminthopsis hirtipes showed some preferences for regenerating habitats, but these were not consistent throughout the study. Factors indicative of temporal and spatial variation in rainfall, time and site had important effects on capture rates. High rainfalls associated with the La Niña phase of the El Niño/Southern Oscillation in 2000 increased seed production and prompted eruptions of rodent species and the carnivorous Dasycercus cristicauda. The greatest numbers of captures were made at the sites that received the highest rainfalls. We conclude that patch‐burning regimes do not benefit small mammals directly, but are likely to increase the resilience of ‘fire‐sensitive’ species that are dependent on dense spinifex by reducing the extent of wildfires.