Influence of fire history on small mammal distributions: insights from a 100‐year post‐fire chronosequence

Aim Fire affects the structure and dynamics of ecosystems world‐wide, over long time periods (decades and centuries) and at large spatial scales (landscapes and regions). A pressing challenge for ecologists is to develop models that explain and predict faunal responses to fire at broad temporal and spatial scales. We used a 105‐year post‐fire chronosequence to investigate small mammal responses to fire across an extensive area of ‘tree mallee’ (i.e. vegetation characterized by small multi‐stemmed eucalypts). Location The Murray Mallee region (104,000 km²) of semi‐arid Australia. Methods First, we surveyed small mammals at 260 sites and explored the fire responses of four species using nonlinear regression models. Second, we assessed the predictive accuracy of models using cross‐validation and by testing with independent data. Third, we examined our results in relation to an influential model of animal succession, the habitat accommodation model. Results Two of four study species showed a clear response to fire history. The distribution of the Mallee Ningaui Ningaui yvonneae, a carnivorous marsupial, was strongly associated with mature vegetation characterized by its cover of hummock grass. The occurrence of breeding females was predicted to increase up to 40–105 years post‐fire, highlighting the extensive time periods over which small mammal populations may be affected by fire. Evaluation of models for N. yvonneae demonstrated that accurate predictions of species occurrence can be made from fire history and vegetation data, across large geographical areas. The introduced House Mouse Mus domesticus was the only species positively associated with recently burnt vegetation. Main conclusions Understanding the impact of fire over long time periods will benefit ecological and conservation management. In this example, tracts of long‐unburnt mallee vegetation were identified as important habitat for a fire‐sensitive native mammal. Small mammal responses to fire can be predicted accurately at broad spatial scales; however, a conceptual model of post‐fire change in community structure developed in temperate Australia is not, on its own, sufficient for small mammals in semi‐arid systems.     

Why do some species have geographically varying responses to fire history?

A capacity to predict the effects of fire on biota is critical for conservation in fire‐prone regions as it assists managers to anticipate the outcomes of different approaches to fire management. The task is complicated because species’ responses to fire can vary geographically. This poses challenges, both for conceptual understanding of post‐fire succession and fire management. We examine two hypotheses for why species may display geographically varying responses to fire. 1) Species’ post‐fire responses are driven by vegetation structure, but vegetation – fire relationships vary spatially (the ‘dynamic vegetation’ hypothesis). 2) Regional variation in ecological conditions leads species to select different post‐fire ages as habitat (the ‘dynamic habitat’ hypothesis). Our case study uses data on lizards at 280 sites in a ～ 100 000 km² region of south‐eastern Australia. We compared the predictive capacity of models based on 1) habitat associations, with models based on 2) fire history and vegetation type, and 3) fire history alone, for four species of lizards. Habitat association models generally out‐performed fire history models in terms of predictive capacity. For two species, habitat association models provided good discrimination capacity even though the species showed geographically varying post‐fire responses. Our results support the dynamic vegetation hypothesis, that spatial variation in relationships between fire and vegetation structure results in regional variation in fauna–fire relationships. These observations explain how the widely recognised ‘habitat accommodation’ model of animal succession can be conceptually accurate yet predictively weak.     

A conceptual model of plant community changes following cessation of cultivation in semi‐arid grassland

Question: Can vegetation changes that occur following cessation of cultivation for cereal crop production in semi‐arid native grasslands be described using a conceptual model that explains plant community dynamics following disturbance? Location: Eighteen native grasslands with varying time‐since‐last cultivation across northern Victoria, Australia. Methods: We examined recovery of native grasslands after cessation of cultivation along a space for‐ time chronosequence. By documenting floristic composition and soil properties of grasslands with known cultivation histories, we established a conceptual model of the vegetation states that occur following cessation of cultivation and inferred transition pathways for community recovery. Results: Succession from an exotic‐dominated grassland to native grassland followed a linear trajectory. These changes represent an increase in richness and cover of native forbs, a decrease in cover of exotic annual species and little change in native perennial graminoids and exotic perennial forbs. Using a state‐and‐transition model, two distinct vegetation states were evident: (1) an unstable, recently cultivated state, dominated by exotic annuals, and (2) a more diverse, stable state. The last‐mentioned state can be divided into two further states based on species composition: (1) a never‐cultivated state dominated by native perennial shrubs and grasses, and (2) a long‐uncultivated state dominated by a small number of native perennial and native and exotic annual species that is best described as a subset of the never‐cultivated state. Transitions between these states are hypothesized to be dependent upon landscape context, seed availability and soil recovery. Conclusions: Legacies of past land use on soils and vegetation of semi‐arid grasslands are not as persistent as in other Australian communities. Recovery appears to follow a linear, directional model of post‐disturbance regeneration which may be advanced by overcoming dispersal barriers hypothesised to restrict recovery.     

Early post‐fire succession in northwestern Patagonia grasslands

Abstract. Large‐scale disturbances, notably fire and grazing, structure grass and shrubland dynamics in semi‐arid environments. We studied early post‐fire succession in two burned grasslands, one unburned grassland, and one shrubland near the burned area. We observed three processes: (1) establishment of a ‘phantom’ community comprised of fugitive species. Although transient, these species increase diversity and recharge the seed bank before the next disturbance; (2) regeneration of the original community by persistence of resprouter species and by auto‐replacement; (3) early stages of invasion by seedlings of the shrub Fabiana imbricata, which germinate next to shrubland and create new F. imbricata patches. Weed invasion was principally due to the ruderal exotic species Verbascum thapsus from the nearby road verge and by rapid increase of Rumex acetosella cover, another exotic species present before the fire. Although post‐fire climatic conditions are particularly important in semi‐arid environments, succession depends greatly on the regeneration strategies and dispersal abilities of the species present in the burned area. The phantom community occurs only at the first stage of succession when there is little competition for resources. We could call this process ‘the race for occupation of the area’. The second stage, when competition for resources becomes progressively more important, could be called ‘the effort to maintain space’.     

Effects of long‐term fire exclusion and frequent fire on plant community composition: A case study from semi‐arid shrublands

Time since last fire and fire frequency are strong determinants of plant community composition in fire‐prone landscapes. Our study aimed to establish the influence of time since last fire and fire frequency on plant community composition and diversity of a south‐west Australian semi‐arid shrubland. We employed a space‐for‐time approach using four fire age classes: ‘young’, 8–15 years since last fire; ‘medium’, 16–34; ‘old’, 35–50; and ‘very old’, 51–100; and three fire frequency classes: burnt once, twice and three times within the last 50 years. Species diversity was compared using one‐way ANOVA and species composition using PERMANOVA. Soil and climatic variables were included as covariables to partition underlying environmental drivers. We found that time since last fire influenced species richness, diversity and composition. Specifically, we recorded a late successional transition from woody seeders to long‐lived, arid‐zone, resprouting shrub species. Fire frequency did not influence species richness and diversity but did influence species composition via a reduction in cover of longer‐lived resprouter species – presumably because of a reduced ability to replenish epicormic buds and/or sufficient starch stores. The distinct floristic composition of old and very old habitat, and the vulnerability of these areas to wildfires, indicate that these areas are ecologically important and management should seek to preserve them.     

A conceptual model of vegetation dynamics for the unique obligate‐seeder eucalypt woodlands of south‐western Australia

Models of vegetation dynamics framed as testable hypotheses provide powerful tools for predicting vegetation change in response to contemporary disturbances or climate change. Synthesizing existing information and applying new data, we develop a conceptual model of vegetation states and transitions for the previously overlooked woodlands dominated by obligate‐seeder eucalypts of dry to semi‐arid south‐western Australia. These comprise the largest extant temperate woodland globally, are uniquely dominated by a high diversity of obligate‐seeder eucalypts (55 taxa), but are under threat from wildfire. Our conceptual model incorporates four critical ecological processes that also distinguish obligate‐seeder woodlands from temperate woodlands dominated by resprouting eucalypts: (i) a lack of well‐protected epicormic buds results in major disturbances (prominently fire) being stand‐replacing. The pre‐disturbance tree cohort is killed, followed by dense post‐disturbance recruitment from seed shed from a serotinous seed bank; (ii) competition between saplings leads to self‐thinning over a multi‐century timeframe, with surviving individuals having great longevity (regularly >400 years); (iii) multiple processes limit recruitment in the absence of stand‐replacement disturbances, leading to frequent single‐cohort stands. However, unlike the few other obligate‐seeder eucalypt communities, trickle recruitment in very long‐unburnt stands can facilitate indefinite community persistence in the absence of stand‐replacement disturbances; and (iv) discontinuous fuels, relatively low understorey flammability (low grass and often high chenopod cover) and topographic barriers to fire (salt lakes) allow mature woodlands to persist for centuries without burning. Notably though, evidence suggests that flammability peaks at intermediate times since fire, establishing a ‘flammability bottleneck’ (or landscape fire trap) through which regenerating woodlands must pass. Our model provides a framework to support management to conserve obligate‐seeder eucalypt woodlands. Research into reasons for exceptional tree heights relative to ecosystem productivity, the evolution of diverse and dominant obligate‐seeder eucalypts, the paucity of grass, and the recent spatial distribution of fires, will further inform conservation management.     

Edge effects in fire‐prone landscapes: Ecological importance and implications for fauna

Edges are ecologically important environmental features and have been well researched in agricultural and urban landscapes. However, little work has been conducted in flammable ecosystems where spatially and temporally dynamic fire edges are expected to influence important processes such as recolonization of burnt areas and landscape connectivity. We review the literature on fire, fauna, and edge effects to summarize current knowledge of faunal responses to fire edges and identify knowledge gaps. We then develop a conceptual model to predict faunal responses to fire edges and present an agenda for future research. Faunal abundance at fire edges changes over time, but patterns depend on species traits and resource availability. Responses are also influenced by edge architecture (e.g., size and shape), site and landscape context, and spatial scale. However, data are limited and the influence of fire edges on both local abundance and regional distributions of fauna is largely unknown. In our conceptual model, biophysical properties interact with the fire regime (e.g., patchiness, frequency) to influence edge architecture. Edge architecture and species traits influence edge permeability, which is linked to important processes such as movement, resource selection, and species interactions. Predicting the effect of fire edges on fauna is challenging, but important for biodiversity conservation in flammable landscapes. Our conceptual model combines several drivers of faunal fire responses (biophysical properties, regime attributes, species traits) and will therefore lead to improved predictions. Future research is needed to understand fire as an agent of edge creation; the spatio‐temporal flux of fire edges across landscapes; and the effect of fire edges on faunal movement, resource selection, and biotic interactions. To aid the incorporation of new data into our predictive framework, our model has been designed as a Bayesian Network, a statistical tool capable of analyzing complex environmental relationships, dealing with data gaps, and generating testable hypotheses.     

Time‐since‐fire and climate interact to affect the structural recovery of an Australian semi‐arid plant community

How does time‐since‐fire influence the structural recovery of semi‐arid, eucalypt‐dominated Murray‐Mallee shrublands after fire, and is recovery affected by spatial variation in climate? We assessed the structure and dynamics of a hummock grass, Triodia scariosa N.T. Burb, and mallee eucalypts – two key structural components of mallee shrublands – using a >100 year time‐since‐fire chronosequence. The relative influence of climatic variables, both individually and combined with time‐since‐fire, was modelled to account for spatial variation in the recovery of vegetation structural components. Time‐since‐fire was the primary determinant of the structural recovery of T. scariosa and eucalypts. However, climate, notably mean annual rainfall and rainfall variability, also influenced the recovery of the eucalypt overstorey, T. scariosa cover and mean hummock height. We observed that (i) the mean number of live eucalypt stems per individual decreased while mean individual basal area increased, (ii) cover of T. scariosa peaked at ~30 years post‐fire and gradually decreased thereafter, and (iii) the ‘hummock’ form of T. scariosa occurred throughout the chronosequence, whereas the ‘ring’ form tended not to occur until ~30 years post‐fire. Time‐since‐fire was the key determinant of the structural recovery of eucalypt‐dominated mallee shrublands, but there is geographical variation in recovery related to rainfall and its variability. Fire regimes are likely to have different effects across the geographic range of mallee shrublands.     

A field trial to test effects of watering,seed addition and disturbance on perennial species recruitment in Belah woodland

Failure of perennial species to regenerate is a significant threat to semi‐arid woodlands across south‐eastern Australia. High grazing pressure eliminates the recruitment of many perennial species in semi‐arid woodlands, but little is known about requirements for regeneration under low grazing pressure. We tested the effects of addition of water (irrigation to match the largest rainfall events of the last century), seed, soil disturbance and fire within a grazing exclosure in Belah (Casuarina pauper) woodland in the Murray‐Sunset National Park, Victoria. Recruitment was observed in 13 perennial species and was dominated by chenopods. Addition of water, seed and soil disturbance increased abundance of juvenile perennial species above the low‐level background recruitment that occurred in the prevailing drought conditions. This supports the view that continuous recruitment occurs for many semi‐arid perennials. Low seed availability and an inability to maintain soil moisture conditions matching that of regeneration events are likely factors in the lack of recruitment for tree species and limited response of shrubs in this experiment.     

Landscape‐scale effects of fire on bird assemblages: does pyrodiversity beget biodiversity?

Aim A common strategy for conserving biodiversity in fire‐prone environments is to maintain a diversity of post‐fire age classes at the landscape scale, under the assumption that ‘pyrodiversity begets biodiversity’. Another strategy is to maintain extensive areas of a particular seral state regarded as vital for the persistence of threatened species, under the assumption that this will also cater for the habitat needs of other species. We investigated the likely effects of these strategies on bird assemblages in tree mallee vegetation, characterized by multi‐stemmed Eucalyptus species, where both strategies are currently employed. Location The semi‐arid Murray Mallee region of south‐eastern Australia. Methods We systematically surveyed birds in 26 landscapes (each 4‐km diameter), selected to represent gradients in the diversity of fire age classes and the proportion of older vegetation (> 35 years since fire). Additional variables were measured to represent underlying vegetation‐ or fire‐mediated properties of the landscape, as well as its biogeographic context. We used an information‐theoretic approach to investigate the relationships between these predictor variables and the species richness of birds (total species, threatened species and rare species). Results Species richness of birds was not strongly associated with fire‐mediated heterogeneity. Species richness was associated with increasing amounts of older vegetation in landscapes, but not with the proportion of recently burned vegetation in landscapes. Main conclusions The preference of many mallee birds for older vegetation highlights the risk of a blanket application of the ‘pyrodiversity begets biodiversity’ paradigm. If application of this paradigm involved converting large areas from long unburned to recently burned vegetation to increase fire‐mediated heterogeneity in tree mallee landscapes, our findings suggest that this could threaten birds. This research highlights the value of adopting a landscape‐scale perspective when evaluating the utility of fire‐management strategies intended to benefit biodiversity.     

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.     

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.     

Response of vertebrates to fenceline contrasts in grazing intensity in semi‐arid woodlands of eastern Australia

Abstract Changes in the abundance, species richness and assemblage composition of vertebrates due to grazing by domestic stock were investigated in the semi‐arid woodlands of eastern Australia. Analyses were based on the differences found at 10 fenceline contrast sites. Two species of amphibians, 22 species of reptiles and two species of small mammal were captured in pit traps during the surveys. Kangaroos (red and eastern grey), sheep, goats and 66 species of birds were recorded along line transects. Analyses revealed that abundance of diurnal reptiles and species richness of diurnal reptiles and birds were significantly lower on heavily grazed sites than they were on lightly grazed sites. At a local scale, the gecko, Gehyra variegata, was more abundant where grazing was heavier, while Diplodactylus conspicillatus, Diplodactylus steindachneri and Rhynchoedura ornata responded to variables indirectly related to grazing intensity (kangaroo density, sheep and goat dung mass and sheep density, respectively). Birds more commonly sighted on lightly grazed areas than heavily grazed areas were the apostlebird, brown treecreeper, crested bellbird, grey butcherbird, hooded robin, jacky winter, little woodswallow, Australian magpie‐lark, mulga parrot, splendid wren, white‐browed treecreeper and yellow‐rumped thornbill. Birds more commonly sighted on heavily grazed areas than on lightly grazed areas were the Australian raven and chestnut‐crowned babbler. Most variation in species composition between sites was due to spatial separation and no regional‐level indicator species of grazing were evident. A combination of direct grazing‐related changes (e.g. loss of ground cover) and indirect effects of the pastoral industry (e.g. introduction of artificial sources of water) lead to changes in fauna at different scales of analysis across regions.     

Is the reassembly of an arid spider assemblage following fire deterministic?

Fire is a common disturbance in many ecosystems, including arid Australia. Understanding whether fauna respond in a deterministic manner towards a single end‐point, or to multiple states, is of crucial importance for conservation management. Why different taxa or assemblages display single or multiple end‐points is also important to develop a synthetic theory of succession. To examine the post‐fire changes in assemblages of spiders, we established a chronosequence study in spinifex habitat of central Western Australia. Ground‐active spiders were pitfall‐trapped over nine months in sites representing experimental fires (0 and 0.5 years post‐fire) and wildfires (3, 5, 8 and 20 years post‐fire). There were significant non‐linear changes in species richness, evenness and composition of spiders with increasing post‐fire age. For all three measures, the assemblage appeared highly deterministic, converging towards the long unburnt state. Similarity in richness, evenness and species composition to the 20‐year‐old sites all increased with increasing time since fire (3–8 years). However, experimentally burnt sites did not neatly fit this sequence. We consider two alternative hypotheses to explain this second trajectory: inertia within the system or the rapid migration and recolonization from nearby surrounding unburnt areas. Analyses indicated that half of the 179 species had significant preferences for, or were restricted to, particular post‐fire ages. This suggests that adequate pyrodiversity, both in terms of post‐fire ages and/or scale and intensity of fires, may be important for the conservation of spiders in this habitat. However, owing to the high number of singletons and low indicator values, the significance of this result for conservation management remains equivocal. Despite this, the high degree of determinism provides hope that managers can develop a good predictive understanding of post‐fire successional changes in spider assemblages in arid Australia.     

Climatic thresholds shape northern high‐latitude fire regimes and imply vulnerability to future climate change

Boreal forests and arctic tundra cover 33% of global land area and store an estimated 50% of total soil carbon. Because wildfire is a key driver of terrestrial carbon cycling, increasing fire activity in these ecosystems would likely have global implications. To anticipate potential spatiotemporal variability in fire‐regime shifts, we modeled the spatially explicit 30‐yr probability of fire occurrence as a function of climate and landscape features (i.e. vegetation and topography) across Alaska. Boosted regression tree (BRT) models captured the spatial distribution of fire across boreal forest and tundra ecoregions (AUC from 0.63–0.78 and Pearson correlations between predicted and observed data from 0.54–0.71), highlighting summer temperature and annual moisture availability as the most influential controls of historical fire regimes. Modeled fire–climate relationships revealed distinct thresholds to fire occurrence, with a nonlinear increase in the probability of fire above an average July temperature of 13.4°C and below an annual moisture availability (i.e. P‐PET) of approximately 150 mm. To anticipate potential fire‐regime responses to 21st‐century climate change, we informed our BRTs with Coupled Model Intercomparison Project Phase 5 climate projections under the RCP 6.0 scenario. Based on these projected climatic changes alone (i.e. not accounting for potential changes in vegetation), our results suggest an increasing probability of wildfire in Alaskan boreal forest and tundra ecosystems, but of varying magnitude across space and throughout the 21st century. Regions with historically low flammability, including tundra and the forest–tundra boundary, are particularly vulnerable to climatically induced changes in fire activity, with up to a fourfold increase in the 30‐yr probability of fire occurrence by 2100. Our results underscore the climatic potential for novel fire regimes to develop in these ecosystems, relative to the past 6000–35 000 yr, and spatial variability in the vulnerability of wildfire regimes and associated ecological processes to 21st‐century climate change.     

Assessing biodiversity with species accumulation curves; inventories of small reptiles by pit‐trapping in Western Australia

Abstract We examined 11 non‐linear regression models to determine which of them best fitted curvilinear species accumulation curves based on pit‐trapping data for reptiles in a range of heterogeneous and homogenous sites in mesic, semi‐arid and arid regions of Western Australia. A well‐defined plateau in a species accumulation curve is required for any of the models accurately to estimate species richness. Two different measures of effort (pit‐trapping days and number of individuals caught) were used to determine if the measure of effort influenced the choice of the best model(s). We used species accumulation curves to predict species richness, determined the trapping effort required to catch a nominated percentage (e.g. 95%) of the predicted number of species in an area, and examined the relationship between species accumulation curves with diversity and rarity. Species richness, diversity and the proportion of rare species in a community influenced the shape of species accumulation curves. The Beta‐P model provided the best overall fit (highest r²) for heterogeneous and homogeneous sites. For heterogeneous sites, Hill, Rational, Clench, Exponential and Weibull models were the next best. For homogeneous habitats, Hill, Weibull and Chapman–Richards were the next best models. There was very little difference between Beta‐P and Hill models in fitting the data to accumulation curves, although the Hill model generally over‐estimated species richness. Most models worked equally well for both measures of trapping effort. Because the number of individuals caught was influenced by both pit‐trapping effort and the abundance of individuals, both measures of effort must be considered if species accumulation curves are to be used as a planning tool. Trapping effort to catch a nominated percentage of the total predicted species in homogeneous and heterogeneous habitats varied among sites, but even for only 75% of the predicted number of species it was generally much higher than the typical effort currently being used for terrestrial vertebrate fauna surveys in Australia. It was not possible to provide a general indication of the effort required to predict species richness for a site, or to capture a nominated proportion of species at a site, because species accumulation curves are heavily influenced by the characteristics of particular sites.     

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.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

Western myall groves (Acacia papyrocarpa) determine the fine‐scale distribution of soil Collembola in semi‐arid South Australia

Vegetation can exert a strong influence on the distribution and activity of biotic communities across a broad range of spatial scales, especially in arid and semi‐arid ecosystems. At fine spatial scales, patches created by individual plants can support different faunal and floral communities even at locations distant from the plant. These differences can have profound effects on a range of ecosystem processes, including seed dispersal, nutrient cycling and resource distribution. In semi‐arid Australia, areas surrounding groves of western myall (Acacia papyrocarpa) trees are largely devoid of vegetation, being referred to as ‘halos’. Here, we investigate the soil‐dwelling Collembola in groves of western myall trees, the surrounding halos and nearby chenopod shrubland. We also investigated whether the abundance of Collembola was influenced by soil depth (0–5 cm layer vs. 6–10 cm layer) in groves. We found that collembolan density was approximately nine times lower and taxonomic richness half that in a halo compared with the grove and chenopod vegetation. Furthermore, analyses at finer taxonomic levels indicate that vegetation patches differed in species composition, with some species restricted to or preferring particular patches. In the grove, we found a higher abundance of Collembola in the 0–5 cm soil layer compared with the 6–10 layer. Our results indicate vegetation patches strongly influence collembolan abundance and species composition in bare patches around western myall. As patches created by vegetation are a common feature of semi‐arid and arid regions, we suspect that these effects are widespread although seldom reported. Furthermore, as Collembola are involved in the decomposition process, Acacia papyrocarpa patches will be influencing nutrient cycling through their effects on the soil biota. Our results also emphasize that comprehensive fauna survey and management of woodland ecosystems need to consider fine‐scale processes.