How predictable are reptile responses to wildfire?

Natural disturbances are key processes in the vast majority of ecosystems and a range of ecological theories have been developed in an attempt to predict biotic responses to them. However, empirical support for these theories has been inconsistent and considerable additional work remains to be done to better understand the response of biodiversity to natural disturbance. We tested predictions from the intermediate disturbance hypothesis and the habitat accommodation model of succession for reptile responses to fire history and a single major fire event. We focused our work on a broad range of vegetation types spanning sedgeland to temperate rainforest located within a national park in south‐eastern Australia. We found no significant relationships between reptile species richness and the number of fires over the past 35 years, the time since the last fire, or the severity of a major fire in 2003. Thus, we found no strong evidence to support the intermediate disturbance hypothesis. A correspondence analysis of reptile assemblages revealed a gradient in species responses to fire history. However, we found limited evidence for an ordered succession of reptiles. Nor could the responses of individual species be readily predicted from life history attributes. Thus, our findings were generally not consistent with predictions from the habitat accommodation model of succession. A possible explanation for the absence of a predictable sequence of recovery following disturbance might be the rapidity of post‐fire recovery of many components of native vegetation cover that were found to be important for reptiles (e.g. the extent of grass cover). This would have limited the time for early successional conditions to prevail and limited opportunities for species associated with such conditions. We found that most reptile species responses were much more strongly linked to vegetation type than fire variables, emphasizing a need to understand relationships with vegetation before being able to understand possible fire effects (if and where they exist). We found the disturbance concepts we examined were limited in their ability to accurately predict reptile responses to past fire history or the impacts of a single major fire in 2003. Practical management might be best guided not by disturbance theory, but by carefully setting objectives to meet conservation goals for particular individual species of reptiles.     

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.     

Predicting the century‐long post‐fire responses of reptiles

Aim We examined the century‐long post‐fire responses of reptiles to (1) determine the time‐scales over which fauna – fire relationships occur, (2) assess the capacity of a conceptual model to predict faunal response to fire, and (3) investigate the degree to which models of fauna – fire relationships can predict species occurrence and are transferable across space. Location A 104,000 km² area in the semi‐arid Murray Mallee region of south‐eastern Australia. Methods We surveyed reptiles at 280 sites across a century‐long post‐fire chronosequence. We developed generalized additive mixed models (GAMMs) of the relationship between time since fire and the occurrence of 17 species in two subregions, and compared modelled responses with predictions derived from the conceptual model. The predictive capacity of GAMMs was then assessed (1) within the subregion the model was developed and (2) when transferred into a novel subregion. Results Eleven species displayed a significant relationship with time‐since‐fire, with changes in species probability of occurrence continuing up to 100 years post‐fire. Predictions of the timing of species post‐fire peak in occurrence were accurate for 9 of 13 species models for which a significant fire response was detected, but little success was achieved in predicting the shape of a species' response. GAMMs predicted species occurrence more accurately when applied within the subregion in which they were developed than when transferred into a novel subregion, primarily due to some species responding to fire more strongly in one part of their geographic range. Main conclusions Fire influences the occurrence of reptiles in semi‐arid ecosystems over century‐long time frames. Habitat‐use conceptual models have value in predicting the peak occurrence of species following fire, particularly for species with distributions strongly shaped by fire. Species relationships with fire can differ across their geographic range, probably associated with variation in climatic influences on post‐fire succession and the consequent provision of habitat resources.     

Post‐fire regeneration in alpine heathland: Does fire severity matter?

Fire severity is thought to be an important determinant of landscape patterns of post‐fire regeneration, yet there have been few studies of the effects of variation in fire severity at landscape scales on floristic diversity and composition, and none within alpine vegetation. Understanding how fire severity affects alpine vegetation is important because fire is relatively infrequent in alpine environments. Globally, alpine ecosystems are at risk from climate change, which, in addition to warming, is likely to increase the severity and frequency of fire in south‐eastern Australia. Here we examine the effects of variation in fire severity on plant diversity and vegetation composition, 5 years after the widespread fires of 2003. We used floristic data from two wide‐spread vegetation types on the Bogong High Plains: open heathland and closed heathland. Three alternative models were tested relating variation in plant community attributes (e.g. diversity, ground cover of dominant species, amount of bare ground) to variation in fire severity. The models were (i) ‘linear’, attributes vary linearly with fire severity; (ii) ‘intermediate disturbance’, attributes are highest at intermediate fire severity and lowest at both low‐ and high‐severity; and (iii) ‘null’, attributes are unaffected by fire severity. In both heathlands, there were few differences in floristic diversity, cover of dominant species and community composition, across the strong fire severity gradient. The null model was most supported in the vast majority of cases, with only limited support for either the linear and intermediate disturbance models. Our data indicate that in both heathlands, vegetation attributes in burnt vegetation were converging towards that of the unburnt state. We conclude that fire severity had little impact on post‐fire regeneration, and that both closed and open alpine heathlands are resilient to variation in fire severity during landscape scale fires.     

Lizard responses to wildfire in arid interior Australia: Long‐term experimental data and commonalities with other studies

Life in terrestrial Australian ecosystems has evolved over the past 10 million years to thrive in habitats kept in a dynamic state through fire succession cycles. Previous studies support the notion that wildfires promote species diversity in plant and animal communities by creating a heterogeneous mix of habitats, each habitat more suitable for particular subsets of species. We document population and community responses to fire in a species‐rich lizard assemblage in the Great Victoria Desert of Western Australia. Lizards were censused by pit trapping at a long‐unburned flat spinifex site in the Great Victoria Desert in Austral springs of 1992 and 1995. A controlled burn was undertaken in mid‐October of 1995, and lizards were censused thereafter in late 1995 and early 1996, and then again in the Austral springs of 1998, 2003 and 2008. Forty‐six species of lizards (2872 individuals) were collected and their stomach contents analysed over the course of a 16‐year fire succession cycle at this single study site. Most strikingly, relative abundances of two species of agamids varied inversely, responding oppositely to habitat clearing effects of fire. The military dragon Ctenophorus isolepis reached higher abundances when vegetation was dense, and decreased in abundance in open vegetation following fire. The netted dragon Ctenophorus nuchalis was rare when vegetation coverage was high but increased rapidly after fire. Abundances of five species of Ctenotus skinks, C. ariadnae, C. calurus, C. hanloni, C. pantherinus and C. piankai, tracked those of C. isolepis. Abundance of a termite‐specialized nocturnal gecko, Rhynchoedura ornata, increased in abundance following fire. Lizard diets changed during the course of the fire succession cycle, returning to near pre‐burn conditions after 16 years. In addition to short‐term fire succession cycles that contribute to structuring local communities, changes in long‐term rainfall also impact desert food webs and regional biotas.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

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.     

Contrasting changes in vegetation structure and diversity with time since fire in two Australian Mediterranean‐climate plant communities

Mallee‐heath and mallee communities occur in a mosaic across large areas of south‐western Australia, in topographically subdued and fire‐prone landscapes. Consequently, it could be expected that these communities would have historically experienced similar fire regimes, and would respond similarly to variation in aspects of the fire regime. We studied the response of mallee‐heath and mallee to time since the last fire, measuring species density, species–area relationships, diversity indices and vegetation structure. Floristic responses to time since fire accorded with the initial floristic composition model of plant succession, with declining species density and Shannon diversity with age in mallee‐heath. Mallee‐heath exhibited structural senescence when > approximately 45–55 years since fire, with increasing standing dead vegetation, bare ground and stagnating or declining size in sprouting Eucalyptus spp. Mallee showed no such evidence of senescence, and indeed continued to increase in stature beyond the mean fire interval but without the compositional change required to provide support for the relay floristic model of plant succession. These results indicate that mallee‐heath is a fire maintained community and as such is reliant upon periodic burning to maintain diversity and vigour. Mallee, in contrast, is modified but not maintained by fire (at least over the period of time since fire examined) and hence is less susceptible to fire interval effects. Indeed, structural attributes likely to be significant for fauna habitat and carbon sequestration continue to develop in mallee unburnt for 55 years or more. Different responses to time since fire will create challenges for management, particularly in fragmented landscapes where fire potentially interacts with other threatening processes.     

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.     

Fire and regeneration: the role of seed banks in the dynamics of northern heathlands

Questions: How do species composition and abundance of soil seed bank and standing vegetation vary over the course of a post‐fire succession in northern heathlands? What is the role of seed banks – do they act as a refuge for early successional species or can they simply be seen as a spillover from the extant local vegetation? Location: Coastal Calluna heathlands, Western Norway. Methods: We analysed vegetation and seed bank along a 24‐year post‐fire chronosequence. Patterns in community composition, similarity and abundances were tested using multivariate analyses, Sørensen's index of similarity, vegetation cover (%) and seedling counts. Results: The total diversity of vegetation and seed bank were 60 and 54 vascular plant taxa, respectively, with 39 shared species, resulting in 68% similarity overall. Over 24 years, the heathland community progressed from open newly burned ground via species rich graminoid‐ and herb‐dominated vegetation to mature Calluna heath. Post‐fire succession was not reflected in the seed bank. The 10 most abundant species constituted 98% of the germinated seeds. The most abundant were Calluna vulgaris (49%; 12 018 seeds m^?2) and Erica tetralix (34%; 8 414 seeds m^?2). Calluna showed significantly higher germination the first 2 years following fire. Conclusions: Vegetation species richness, ranging from 23 to 46 species yr^?1, showed a unimodal pattern over the post‐fire succession. In contrast, the seed bank species richness, ranging from 21 to 31 species yr^?1, showed no trend. This suggests that the seed bank act as a refuge; providing a constant source of recruits for species that colonise newly burned areas. The traditional management regime has not depleted or destroyed the seed banks and continued management is needed to ensure sustainability of northern heathlands.