Contrasting responses of native and introduced mammal communities to fire mosaics in a modified landscape.

Planned fire is increasingly recognized as an important tool in conservation, but other factors such as land-use change may hinder the ability of land managers to use fire for the benefit of biodiversity. The mosaic of past fires in native vegetation may interact with the mosaic of other land-cover types in human-modified landscapes, yet the effects of these interactions on mammal communities are unknown. We investigated the responses of ground-dwelling mammal community composition and species richness to interactions between land cover and post-fire vegetation growth-stage mosaics in southern Australia. This fire-prone, human-modified landscape features a fine-scale fire mosaic in native vegetation patches surrounded by pasture, horticulture, and peri-urban environments. We measured the composition of land-cover types and fire mosaics (landscape structure) at multiple scales of up to 1257 ha surrounding 129 study sites, and considered native and introduced species together and separately. Land-cover composition was the primary driver of community composition: native species favored areas with a greater proportion of native heathy woodland, whereas introduced species were associated with landscapes comprising more cleared land. The fire mosaic also influenced community composition and species richness: greater growth-stage diversity was associated with native habitat-specialist communities and fewer introduced species. In areas with more cleared land, native species richness increased when there was a greater proportion of mid-successional vegetation, demonstrating that the effect of fire mosaics on mammal diversity depended on land-cover composition. The positive relationship between introduced species richness and cleared land extent was also stronger in recently burned sites than in other growth stages, suggesting that introduced species are well suited to more modified areas of the landscape. Land managers need to consider the underlying land-cover composition and the potential interactions it may have with fire mosaics and species composition. In this landscape a greater diversity of growth stages may disadvantage introduced species yet an increase in mid-successional vegetation in more modified areas would be likely to benefit native mammal communities. Our study highlights that fire management may need to be tailored depending on the context of land use and the species of interest.

How fire interacts with habitat loss and fragmentation.

Biodiversity faces many threats and these can interact to produce outcomes that may not be predicted by considering their effects in isolation. Habitat loss and fragmentation (hereafter 'fragmentation') and altered fire regimes are important threats to biodiversity, but their interactions have not been systematically evaluated across the globe. In this comprehensive synthesis, including 162 papers which provided 274 cases, we offer a framework for understanding how fire interacts with fragmentation. Fire and fragmentation interact in three main ways: (i) fire influences fragmentation (59% of 274 cases), where fire either destroys and fragments habitat or creates and connects habitat; (ii) fragmentation influences fire (25% of cases) where, after habitat is reduced in area and fragmented, fire in the landscape is subsequently altered because people suppress or ignite fires, or there is increased edge flammability or increased obstruction to fire spread; and (iii) where the two do not influence each other, but fire interacts with fragmentation to affect responses like species richness, abundance and extinction risk (16% of cases). Where fire and fragmentation do influence each other, feedback loops are possible that can lead to ecosystem conversion (e.g. forest to grassland). This is a well-documented threat in the tropics but with potential also to be important elsewhere. Fire interacts with fragmentation through scale-specific mechanisms: fire creates edges and drives edge effects; fire alters patch quality; and fire alters landscape-scale connectivity. We found only 12 cases in which studies reported the four essential strata for testing a full interaction, which were fragmented and unfragmented landscapes that both span contrasting fire histories, such as recently burnt and long unburnt vegetation. Simulation and empirical studies show that fire and fragmentation can interact synergistically, multiplicatively, antagonistically or additively. These cases highlight a key reason why understanding interactions is so important: when fire and fragmentation act together they can cause local extinctions, even when their separate effects are neutral. Whether fire-fragmentation interactions benefit or disadvantage species is often determined by the species' preferred successional stage. Adding fire to landscapes generally benefits early-successional plant and animal species, whereas it is detrimental to late-successional species. However, when fire interacts with fragmentation, the direction of effect of fire on a species could be reversed from the effect expected by successional preferences. Adding fire to fragmented landscapes can be detrimental for species that would normally co-exist with fire, because species may no longer be able to disperse to their preferred successional stage. Further, animals may be attracted to particular successional stages leading to unexpected responses to fragmentation, such as higher abundance in more isolated unburnt patches. Growing human populations and increasing resource consumption suggest that fragmentation trends will worsen over coming years. Combined with increasing alteration of fire regimes due to climate change and human-caused ignitions, interactions of fire with fragmentation are likely to become more common. Our new framework paves the way for developing a better understanding of how fire interacts with fragmentation, and for conserving biodiversity in the face of these emerging challenges.

Integrating functional connectivity and fire management for better conservation outcomes.

Globally, the mean abundance of terrestrial animals has fallen by 50% since 1970, and populations face ongoing threats associated with habitat loss, fragmentation, climate change, and disturbance. Climate change can influence the quality of remaining habitat directly and indirectly by precipitating increases in the extent, frequency, and severity of natural disturbances, such as fire. Species face the combined threats of habitat clearance, changing climates, and altered disturbance regimes, each of which may interact and have cascading impacts on animal populations. Typically, conservation agencies are limited in their capacity to mitigate rates of habitat clearance, habitat fragmentation, or climate change, yet fire management is increasingly used worldwide to reduce wildfire risk and achieve conservation outcomes. A popular approach to ecological fire management involves the creation of fire mosaics to promote animal diversity. However, this strategy has 2 fundamental limitations: the effect of fire on animal movement within or among habitat patches is not considered and the implications of the current fire regime for long-term population persistence are overlooked. Spatial and temporal patterns in fire history can influence animal movement, which is essential to the survival of individual animals, maintenance of genetic diversity, and persistence of populations, species, and ecosystems. We argue that there is rich potential for fire managers to manipulate animal movement patterns; enhance functional connectivity, gene flow, and genetic diversity; and increase the capacity of populations to persist under shifting environmental conditions. Recent methodological advances, such as spatiotemporal connectivity modeling, spatially explicit individual-based simulation, and fire-regime modeling can be integrated to achieve better outcomes for biodiversity in human-modified, fire-prone landscapes. Article impact statement: Land managers may conserve populations by using fire to sustain or enhance functional connectivity.

Integración de la Conectividad Funcional y el Manejo de Incendios para Mejores Resultados de Conservación Resumen La abundancia media de animales terrestres ha disminuido en un 50% desde 1970 en todo el mundo y las poblaciones de animales enfrentan amenazas continuas asociadas con la pérdida del hábitat, la fragmentación, el cambio climático y la perturbación. El cambio climático puede influir directa e indirectamente sobre la calidad de los hábitats permanecientes al precipitar incrementos en la extensión, frecuencia y severidad de la perturbación natural, como por ejemplo los incendios. Las especies enfrentan las amenazas combinadas de la modificación de hábitats, climas cambiantes y regímenes alterados de perturbación, cada uno de los cuales puede interactuar con los otros y tener impactos en cascada sobre las poblaciones animales. Habitualmente, las agencias de conservación tienen capacidades limitadas para mitigar las tasas de alteración de hábitats, fragmentación del hábitat o cambio climático. A pesar de esto, el manejo de incendios cada vez se usa más a nivel mundial para reducir el riesgo de incendios forestales y para producir resultados de conservación. Una estrategia popular para el manejo de incendios ecológicos involucra la creación de mosaicos de incendios para promover la diversidad de animales. Sin embargo, esta estrategia tiene dos limitaciones fundamentales: no considera el efecto del fuego sobre el movimiento de los animales dentro o entre los fragmentos de hábitat y pasa por alto las implicaciones del régimen actual de incendios para la persistencia a largo plazo de la población. Los patrones espaciales y temporales en la historia de los incendios pueden influir sobre el movimiento animal, lo cual es esencial para la supervivencia de los animales individuales, el mantenimiento de la diversidad genética y la persistencia de las poblaciones, especies y ecosistemas. Argumentamos que existe un amplio potencial para que los manejadores de incendios manipulen los patrones de movimiento de los animales; mejoren la conectividad funcional, el flujo génico y la diversidad genética; e incrementen la capacidad de las poblaciones de persistir bajo condiciones ambientales cambiantes. Los avances metodológicos recientes, como el modelado de la conectividad espaciotemporal, la simulación espacial basada en individuos y el modelado de regímenes de incendios, pueden integrarse para obtener mejores resultados para la biodiversidad en paisajes propensos a incendios y modificados por el humano.

Survey design for precise fire management conservation targets.

Common goals of ecological fire management are to sustain biodiversity and minimize extinction risk. A novel approach to achieving these goals determines the relative proportions of vegetation growth stages (equivalent to successional stages, which are categorical representations of time since fire) that maximize a biodiversity index. The method combines data describing species abundances in each growth stage with numerical optimization to define an optimal growth-stage structure that provides a conservation-based operational target for managers. However, conservation targets derived from growth-stage optimization are likely to depend critically on choices regarding input data. There is growing interest in the use of growth-stage optimization as a basis for fire management, thus understanding of how input data influence the outputs is crucial. Simulated data sets provide a flexible platform for systematically varying aspects of survey design and species inclusions. We used artificial data with known properties, and a case-study data set from southeastern Australia, to examine the influence of (1) survey design (total number of sites and their distribution among growth stages) and (2) species inclusions (total number of species and their level of specialization) on the precision of conservation targets. Based on our findings, we recommend that survey designs for precise estimates would ideally involve at least 80 sites, and include at least 80 species. Greater numbers of sites and species will yield increasingly reliable results, but fewer might be sufficient in some circumstances. An even distribution of sites among growth stages was less important than the total number of sites, and omission of species is unlikely to have a major influence on results as long as several species specialize on each growth stage. We highlight the importance of examining the responses of individual species to growth stage before feeding survey data into the growth-stage optimization black box, and advocate use of a resampling procedure to determine the precision of results. Collectively, our findings form a reproducible guide to designing ecological surveys that yield precise conservation targets through growth-stage optimization, and ultimately help sustain biodiversity in fire-prone systems.

Opposing Responses of Bird Functional Diversity to Vegetation Structural Diversity in Wet and Dry Forest.

Disturbance regimes are changing worldwide, and the consequences for ecosystem function and resilience are largely unknown. Functional diversity (FD) provides a surrogate measure of ecosystem function by capturing the range, abundance and distribution of trait values in a community. Enhanced understanding of the responses of FD to measures of vegetation structure at landscape scales is needed to guide conservation management. To address this knowledge gap, we used a whole-of-landscape sampling approach to examine relationships between bird FD, vegetation diversity and time since fire. We surveyed birds and measured vegetation at 36 landscape sampling units in dry and wet forest in southeast Australia during 2010 and 2011. Four uncorrelated indices of bird FD (richness, evenness, divergence and dispersion) were derived from six bird traits, and we investigated responses of these indices and species richness to both vertical and horizontal vegetation diversity using linear mixed models. We also considered the extent to which the mean and diversity of time since fire were related to vegetation diversity. Results showed opposing responses of FD to vegetation diversity in dry and wet forest. In dry forest, where fire is frequent, species richness and two FD indices (richness and dispersion) were positively related to vertical vegetation diversity, consistent with theory relating to environmental variation and coexistence. However, in wet forest subject to infrequent fire, the same three response variables were negatively associated with vertical diversity. We suggest that competitive dominance by species results in lower FD as vegetation diversity increases in wet forest. The responses of functional evenness were opposite to those of species richness, functional richness and dispersion in both forest types, highlighting the value of examining multiple FD metrics at management-relevant scales. The mean and diversity of time since fire were uncorrelated with vegetation diversity in wet forest, but positively correlated with vegetation diversity in dry forest. We therefore suggest that protection of older vegetation is important, but controlled application of low-severity fire in dry forest may sustain ecosystem function by enhancing different elements of FD.

Bird functional diversity decreases with time since disturbance: Does patchy prescribed fire enhance ecosystem function?.

Animal species diversity is often associated with time since disturbance, but the effects of disturbances such as fire on functional diversity are unknown. Functional diversity measures the range, abundance, and distribution of trait values in a community, and links changes in species composition with the consequences for ecosystem function. Improved understanding of the relationship between time since fire (TSF) and functional diversity is critical given that the frequency of both prescribed fire and wildfire is expected to increase. To address this knowledge gap, we examined responses of avian functional diversity to TSF and two direct measures of environmental heterogeneity, plant diversity, and structural heterogeneity. We surveyed birds across a 70-year chronosequence spanning four vegetation types in southeast Australia. Six bird functional traits were used to derive four functional diversity indices (richness, evenness, divergence, and dispersion) and the effects of TSF, plant diversity and structural heterogeneity on species richness and the functional diversity indices were examined using mixed models. We used a regression tree method to identify traits associated with species more common in young vegetation. Functional richness and dispersion were negatively associated with TSF in all vegetation types, suggesting that recent prescribed fire generates heterogeneous vegetation and provides greater opportunities for resource partitioning. Species richness was not significantly associated with TSF, and is probably an unreliable surrogate for functional diversity in fire-prone systems. A positive, relationship between functional evenness and structural heterogeneity was comnon to all vegetation types, suggesting that fine-scale (tens of meters) structural variation can enhance ecosystem function. Species more common in young vegetation were primarily linked by their specialist diets, indicating that ecosystem services such as seed dispersal and insect control are enhanced in more recently burnt vegetation. We suggest that patchy prescribed fire sustains functional diversity, and that controlled use of patchy fire to break up large expanses of mature vegetation will enhance ecosystem function.

Predicting faunal fire responses in heterogeneous landscapes: the role of habitat structure.

Predicting the effects of fire on biota is important for biodiversity conservation in fire-prone landscapes. Time since fire is often used to predict the occurrence of fauna, yet for many species, it is a surrogate variable and it is temporal change in resource availability to which animals actually respond. Therefore prediction of fire-fauna relationships will be uncertain if time since fire is not strongly related to resources. In this study, we used a space-for-time substitution across a large diverse landscape to investigate interrelationships between the occurrence of ground-dwelling mammals, time since fire, and structural resources. We predicted that much variation in habitat structure would remain unexplained by time since fire and that habitat structure would predict species' occurrence better than time since fire. In line with predictions, we found that time since fire was moderately correlated with habitat structure yet was a poor surrogate for mammal occurrence. Variables representing habitat structure were better predictors of occurrence than time since fire for all species considered. Our results suggest that time since fire is unlikely to be a useful surrogate for ground-dwelling mammals in heterogeneous landscapes. Faunal conservation in fire-prone landscapes will benefit from a combined understanding of fauna-resource relationships and the ways in which fire (including planned fires and wildfires) alters the spatial and temporal distribution of faunal resources.