How does prescribed fire shape bird and plant communities in a temperate dry forest ecosystem?

To mitigate the impact of severe wildfire on human society and the environment, prescribed fire is widely used in forest ecosystems to reduce fuel loads and limit fire spread. To avoid detrimental effects on conservation values, it is imperative to understand how prescribed fire affects taxa having a range of different adaptations to disturbance. Such studies will have greatest benefit if they extend beyond short-term impacts of burning. We used a field study to examine the effects of prescribed fire on birds and plants across a 36-yr post-fire chronosequence in a temperate dry forest ecosystem in southeastern Australia, and by making comparison with long-unburned reference sites (79 yr since wildfire). We modeled changes in the relative abundance of 22 bird species and the cover of 39 plant species, and examined how individual species, functional groups, species richness and community composition differed between sites with different fire history. For most individual bird and plant species modeled, relative abundance or cover at sites subject to prescribed fire did not change significantly with time since fire or differ from that of long-unburned vegetation. When bird species were pooled into functional groups, time since prescribed fire had strong effects on birds that forage in the lower-midstorey, facultative-resprouting shrubs and obligate-seeding shrubs. Species richness for both taxa did not differ between sites subject to prescribed fire and those in long-unburned vegetation. Bird communities varied significantly between the youngest (0-3 yr) and oldest (79 yr) post-fire age classes, driven by species associated with understorey vegetation. Plant community composition showed little evidence of a post-fire successional trajectory. The prevalence of bird species with broad habitat and dietary niches and plant regeneration through resprouting, make bird and plant communities in these forests relatively resilient to small and patchy prescribed fires they have experienced to date. Application of prescribed fire will be most compatible with maintaining biodiversity by taking a landscape approach that (1) plans for a geographic spread of stands with a range of between-prescribed-fire intervals to ensure provision of suitable habitat for all taxa, and (2) avoids burning in moist gullies to maintain their value as fire refuges.

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.

Using decision science to evaluate global biodiversity indices.

Global biodiversity indices are used to measure environmental change and progress toward conservation goals, yet few indices have been evaluated comprehensively for their capacity to detect trends of interest, such as declines in threatened species or ecosystem function. Using a structured approach based on decision science, we qualitatively evaluated 9 indices commonly used to track biodiversity at global and regional scales against 5 criteria relating to objectives, design, behavior, incorporation of uncertainty, and constraints (e.g., costs and data availability). Evaluation was based on reference literature for indices available at the time of assessment. We identified 4 key gaps in indices assessed: pathways to achieving goals (means objectives) were not always clear or relevant to desired outcomes (fundamental objectives); index testing and understanding of expected behavior was often lacking; uncertainty was seldom acknowledged or accounted for; and costs of implementation were seldom considered. These gaps may render indices inadequate in certain decision-making contexts and are problematic for indices linked with biodiversity targets and sustainability goals. Ensuring that index objectives are clear and their design is underpinned by a model of relevant processes are crucial in addressing the gaps identified by our assessment. Uptake and productive use of indices will be improved if index performance is tested rigorously and assumptions and uncertainties are clearly communicated to end users. This will increase index accuracy and value in tracking biodiversity change and supporting national and global policy decisions, such as the post-2020 global biodiversity framework of the Convention on Biological Diversity.

Uso de las Ciencias de la Decisión para Evaluar los Índices Globales de Biodiversidad Resumen Los índices globales de biodiversidad se usan para medir el cambio ambiental y el avance hacia los objetivos de conservación, aunque pocos han sido evaluados completamente en cuanto a su capacidad para detectar las tendencias de interés como las declinaciones de especies amenazadas o la función del ecosistema. Evaluamos cualitativamente nueve índices de uso común para dar seguimiento a la biodiversidad a escala global y regional contra cinco criterios relacionados con los objetivos, diseño, comportamiento, incorporación de la incertidumbre y restricciones (p. ej.: costos y disponibilidad de datos) mediante una estrategia estructurada basada en las ciencias de la decisión. La evaluación se basó en la literatura de referencia para los índices disponibles al momento del análisis. Identificamos cuatro vacíos importantes en los índices estudiados: las vías para lograr los objetivos (objetivos medios) no fueron siempre claras o relevantes para los resultados deseados (objetivos fundamentales); el análisis del índice y el entendimiento del comportamiento esperado casi siempre fueron escasos; pocas veces se consideró o explicó la incertidumbre; y casi nunca se consideraron los costos de la implementación. Estos vacíos pueden hacer que los índices sean inadecuados en ciertos contextos de toma de decisiones y son problemáticos para los índices vinculados a los objetivos de biodiversidad y las metas de sustentabilidad. Es de suma importancia asegurarse que los objetivos del índice sean claros y que su diseño esté respaldado por un modelo de procesos relevantes para tratar con los vacíos identificados en nuestro estudio. La aceptación y el uso productivo de los índices mejorarán si el desempeño del índice es evaluado rigurosamente y las suposiciones e incertidumbres se les comunican claramente a los usuarios finales. Lo anterior aumentará la precisión y valor del índice en el seguimiento de los cambios de la biodiversidad y en el apoyo a las decisiones políticas nacionales y mundiales, como el marco de trabajo para la biodiversidad post-2020 establecido por la Convención sobre la Diversidad Biológica.

Fire and biodiversity in the Anthropocene.

Fire has been a source of global biodiversity for millions of years. However, interactions with anthropogenic drivers such as climate change, land use, and invasive species are changing the nature of fire activity and its impacts. We review how such changes are threatening species with extinction and transforming terrestrial ecosystems. Conservation of Earth's biological diversity will be achieved only by recognizing and responding to the critical role of fire. In the Anthropocene, this requires that conservation planning explicitly includes the combined effects of human activities and fire regimes. Improved forecasts for biodiversity must also integrate the connections among people, fire, and ecosystems. Such integration provides an opportunity for new actions that could revolutionize how society sustains biodiversity in a time of changing fire activity.

Animal movements in fire-prone landscapes.

Movement is a trait of fundamental importance in ecosystems subject to frequent disturbances, such as fire-prone ecosystems. Despite this, the role of movement in facilitating responses to fire has received little attention. Herein, we consider how animal movement interacts with fire history to shape species distributions. We consider how fire affects movement between habitat patches of differing fire histories that occur across a range of spatial and temporal scales, from daily foraging bouts to infrequent dispersal events, and annual migrations. We review animal movements in response to the immediate and abrupt impacts of fire, and the longer-term successional changes that fires set in train. We discuss how the novel threats of altered fire regimes, landscape fragmentation, and invasive species result in suboptimal movements that drive populations downwards. We then outline the types of data needed to study animal movements in relation to fire and novel threats, to hasten the integration of movement ecology and fire ecology. We conclude by outlining a research agenda for the integration of movement ecology and fire ecology by identifying key research questions that emerge from our synthesis of animal movements in fire-prone ecosystems.

Do multiple fires interact to affect vegetation structure in temperate eucalypt forests?

Fire plays an important role in structuring vegetation in fire-prone regions worldwide. Progress has been made towards documenting the effects of individual fire events and fire regimes on vegetation structure; less is known of how different fire history attributes (e.g., time since fire, fire frequency) interact to affect vegetation. Using the temperate eucalypt foothill forests of southeastern Australia as a case study system, we examine two hypotheses about such interactions: (1) post-fire vegetation succession (e.g., time-since-fire effects) is influenced by other fire regime attributes and (2) the severity of the most recent fire overrides the effect of preceding fires on vegetation structure. Empirical data on vegetation structure were collected from 540 sites distributed across central and eastern Victoria, Australia. Linear mixed models were used to examine these hypotheses and determine the relative influence of fire and environmental attributes on vegetation structure. Fire history measures, particularly time since fire, affected several vegetation attributes including ground and canopy strata; others such as low and sub-canopy vegetation were more strongly influenced by environmental characteristics like rainfall. There was little support for the hypothesis that post-fire succession is influenced by fire history attributes other than time since fire; only canopy regeneration was influenced by another variable (fire type, representing severity). Our capacity to detect an overriding effect of the severity of the most recent fire was limited by a consistently weak effect of preceding fires on vegetation structure. Overall, results suggest the primary way that fire affects vegetation structure in foothill forests is via attributes of the most recent fire, both its severity and time since its occurrence; other attributes of fire regimes (e.g., fire interval, frequency) have less influence. The strong effect of environmental drivers, such as rainfall and topography, on many structural features show that foothill forest vegetation is also influenced by factors outside human control. While fire is amenable to human management, results suggest that at broad scales, structural attributes of these forests are relatively resilient to the effects of current fire regimes. Nonetheless, the potential for more frequent severe fires at short intervals, associated with a changing climate and/or fire management, warrant further consideration.

Historical Maps from Modern Images: Using Remote Sensing to Model and Map Century-Long Vegetation Change in a Fire-Prone Region.

Understanding the age structure of vegetation is important for effective land management, especially in fire-prone landscapes where the effects of fire can persist for decades and centuries. In many parts of the world, such information is limited due to an inability to map disturbance histories before the availability of satellite images (~1972). Here, we describe a method for creating a spatial model of the age structure of canopy species that established pre-1972. We built predictive neural network models based on remotely sensed data and ecological field survey data. These models determined the relationship between sites of known fire age and remotely sensed data. The predictive model was applied across a 104,000 km(2) study region in semi-arid Australia to create a spatial model of vegetation age structure, which is primarily the result of stand-replacing fires which occurred before 1972. An assessment of the predictive capacity of the model using independent validation data showed a significant correlation (rs = 0.64) between predicted and known age at test sites. Application of the model provides valuable insights into the distribution of vegetation age-classes and fire history in the study region. This is a relatively straightforward method which uses widely available data sources that can be applied in other regions to predict age-class distribution beyond the limits imposed by satellite imagery.

Optimal fire histories for biodiversity conservation.

Fire is used as a management tool for biodiversity conservation worldwide. A common objective is to avoid population extinctions due to inappropriate fire regimes. However, in many ecosystems, it is unclear what mix of fire histories will achieve this goal. We determined the optimal fire history of a given area for biological conservation with a method that links tools from 3 fields of research: species distribution modeling, composite indices of biodiversity, and decision science. We based our case study on extensive field surveys of birds, reptiles, and mammals in fire-prone semi-arid Australia. First, we developed statistical models of species' responses to fire history. Second, we determined the optimal allocation of successional states in a given area, based on the geometric mean of species relative abundance. Finally, we showed how conservation targets based on this index can be incorporated into a decision-making framework for fire management. Pyrodiversity per se did not necessarily promote vertebrate biodiversity. Maximizing pyrodiversity by having an even allocation of successional states did not maximize the geometric mean abundance of bird species. Older vegetation was disproportionately important for the conservation of birds, reptiles, and small mammals. Because our method defines fire management objectives based on the habitat requirements of multiple species in the community, it could be used widely to maximize biodiversity in fire-prone ecosystems.

Does fire influence the landscape-scale distribution of an invasive mesopredator?

Predation and fire shape the structure and function of ecosystems globally. However, studies exploring interactions between these two processes are rare, especially at large spatial scales. This knowledge gap is significant not only for ecological theory, but also in an applied context, because it limits the ability of landscape managers to predict the outcomes of manipulating fire and predators. We examined the influence of fire on the occurrence of an introduced and widespread mesopredator, the red fox (Vulpes vulpes), in semi-arid Australia. We used two extensive and complimentary datasets collected at two spatial scales. At the landscape-scale, we surveyed red foxes using sand-plots within 28 study landscapes - which incorporated variation in the diversity and proportional extent of fire-age classes - located across a 104 000 km2 study area. At the site-scale, we surveyed red foxes using camera traps at 108 sites stratified along a century-long post-fire chronosequence (0-105 years) within a 6630 km2 study area. Red foxes were widespread both at the landscape and site-scale. Fire did not influence fox distribution at either spatial scale, nor did other environmental variables that we measured. Our results show that red foxes exploit a broad range of environmental conditions within semi-arid Australia. The presence of red foxes throughout much of the landscape is likely to have significant implications for native fauna, particularly in recently burnt habitats where reduced cover may increase prey species' predation risk.

Effects of time since fire on birds: how informative are generalized fire response curves for conservation management?

Fire is both a widespread natural disturbance that affects the distribution of species and a tool that can be used to manage habitats for species. Knowledge of temporal changes in the occurrence of species after fire is essential for conservation management in fire-prone environments. Two key issues are: whether postfire responses of species are idiosyncratic or if multiple species show a limited number of similar responses; and whether such responses to time since fire can predict the occurrence of species across broad spatial scales. We examined the response of bird species to time since fire in semiarid shrubland in southeastern Australia using data from surveys at 499 sites representing a 100-year chronosequence. We used nonlinear regression to model the probability of occurrence of 30 species with time since fire in two vegetation types, and compared species' responses with generalized response shapes from the literature. The occurrence of 16 species was significantly influenced by time since fire: they displayed six main responses consistent with generalized response shapes. Of these 16 species, 15 occurred more frequently in mid- or later-successional vegetation (> 20 years since fire), and only one species occurred more often in early succession (< 5 years since fire). The models had reasonable predictive ability for eight species, some predictive ability for seven species, and were little better than random for one species. Bird species displayed a limited range of responses to time since fire; thus a small set of fire ages should allow the provision of habitat for most species. Postfire successional changes extend for decades and management of the age class distribution of vegetation will need to reflect this timescale. Response curves revealed important seral stages for species and highlighted the importance of mid- to late-successional vegetation (> 20 years). Although time since fire clearly influences the distribution of numerous bird species, predictive models of the spatial distribution of species in fire-prone landscapes need to incorporate other factors in addition to time since fire.

The decoupling of abundance and species richness in lizard communities.

1. Patterns of species richness often correlate strongly with measures of energy. The more individuals hypothesis (MIH) proposes that this relationship is facilitated by greater resources supporting larger populations, which are less likely to become extinct. Hence, the MIH predicts that community abundance and species richness will be positively related. 2. Recently, Buckley & Jetz (2010, Journal of Animal Ecology, 79, 358-365) documented a decoupling of community abundance and species richness in lizard communities in south-west United States, such that richer communities did not contain more individuals. They predicted, as a consequence of the mechanisms driving the decoupling, a more even distribution of species abundances in species-rich communities, evidenced by a positive relationship between species evenness and species richness. 3. We found a similar decoupling of the relationship between abundance and species richness for lizard communities in semi-arid south-eastern Australia. However, we note that a positive relationship between evenness and richness is expected because of the nature of the indices used. We illustrate this mathematically and empirically using data from both sets of lizard communities. When we used a measure of evenness, which is robust to species richness, there was no relationship between evenness and richness in either data set. 4. For lizard communities in both Australia and the United States, species dominance decreased as species richness increased. Further, with the iterative removal of the first, second and third most dominant species from each community, the relationship between abundance and species richness became increasingly more positive. 5. Our data support the contention that species richness in lizard communities is not directly related to the number of individuals an environment can support. We propose an alternative hypothesis regarding how the decoupling of abundance and richness is accommodated; namely, an inverse relationship between species dominance and species richness, possibly because of ecological release.