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.

Circadian rhythms enable efficient resource selection in a human-modified landscape.

Animals access resources such as food and shelter, and acquiring these resources has varying risks and benefits, depending on the suitability of the landscape. Some animals change their patterns of resource selection in space and time to optimize the trade-off between risks and benefits. We examine the circadian variation in resource selection of swamp wallabies (Wallabia bicolor) within a human-modified landscape, an environment of varying suitability. We used GPS data from 48 swamp wallabies to compare the use of landscape features such as woodland and scrub, housing estates, farmland, coastal areas, wetlands, waterbodies, and roads to their availability using generalized linear mixed models. We investigated which features were selected by wallabies and determined whether the distance to different landscape features changed, depending on the time of the day. During the day, wallabies were more likely to be found within or near natural landscape features such as woodlands and scrub, wetlands, and coastal vegetation, while avoiding landscape features that may be perceived as more risky (roads, housing, waterbodies, and farmland), but those features were selected more at night. Finally, we mapped our results to predict habitat suitability for swamp wallabies in human-modified landscapes. We showed that wallabies living in a human-modified landscape selected different landscape features during day or night. Changing circadian patterns of resource selection might enhance the persistence of species in landscapes where resources are fragmented and disturbed.

Biotelemetry marches on: A cost-effective GPS device for monitoring terrestrial wildlife.

The availability of low-cost wildlife trackers increases the capacity to collect valuable ecological data when research budgets are limited. We converted a commercially available global positioning system (GPS) product into a low-cost tracking device that sends data via the mobile phone network, and assessed its performance under varying conditions. We established a stationary test, deploying devices along a continuum from open urban areas to topographically and structurally complex forested sites. We tested three features of the device: (a) the GPS, by measuring fix success rate, fix precision and horizontal dilution of precision (HDOP), (b) remote download capacity via the mobile phone network and (c) battery drain. Measures of GPS performance demonstrated high fix success rates and precision. HDOP values were influenced by habitat type and topographical position, but generally remained very low, giving an acceptable degree of error for most applications in wildlife research. Devices experienced delayed data transmission at sites with less phone reception, and faster battery drain at sites with denser vegetation. We recorded device malfunctions in 8.2% of the 110 sampling locations, but these were not associated with habitat type or topography. Our device was effective under a wide range of conditions, and the development process we used provides guidance to other researchers aiming to develop cost-effective wildlife trackers. Reducing the financial and labour costs of acquiring high-quality movement data will improve the capacity to increase sample size in animal movement studies.

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.

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.

Human-modified habitats facilitate forest-dwelling populations of an invasive predator, Vulpes vulpes.

Invasive and over-abundant predators pose a major threat to biodiversity and often benefit from human activities. Effective management requires understanding predator use of human-modified habitats (including resource subsidies and disturbed environments), and individual variation within populations. We investigated selection for human-modified habitats by invasive red foxes, Vulpes vulpes, within two predominantly forested Australian landscapes. We predicted that foxes would select for human-modified habitats in their range locations and fine-scale movements, but that selection would vary between individuals. We GPS-tracked 19 foxes for 17-166 days; ranges covered 33 to >2500 ha. Approximately half the foxes selected for human-modified habitats at the range scale, with some 'commuting' more than five kilometres to farmland or townships at night. Two foxes used burnt forest intensively after a prescribed fire. In their fine-scale nocturnal movements, most foxes selected for human-modified habitats such as reservoirs, forest edges and roads, but there was considerable individual variation. Native fauna in fragmented and disturbed habitats are likely to be exposed to high rates of fox predation, and anthropogenic food resources may subsidise fox populations within the forest interior. Coordinating fox control across land-tenures, targeting specific landscape features, and limiting fox access to anthropogenic resources will be important for biodiversity conservation.

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.

Ecological specialisation in habitat selection within a macropodid herbivore guild.

Specialist species show stronger resource selection, narrower niches and lower niche overlap than generalist species. We examined ecological specialisation with respect to habitat selection in a macropodid community comprising the western grey kangaroo Macropus fuliginosus, red-necked wallaby M. rufogriseus and swamp wallaby Wallabia bicolor in the Grampians National Park, Victoria, Australia. We used radio tracking to quantify habitat selection. We predicted that because the fitness benefits of generalisation and specialisation differ, there would be a mix of generalised and specialised species in our community. As all three macropodid species show marked sexual dimorphism, we also expected that they would show sex-based specialisation. Finally, because many large herbivores select different habitats for foraging and resting, we predicted that our species would specialise on a subset of their overall selected habitat based on activity period (diurnal or nocturnal). All three species specialised on the available resources to some degree. Western grey kangaroos were specialists, at least during the active period. Niche data for the two wallaby species were harder to interpret so we could not determine their degree of specialisation. Within species, we found no evidence of sex-based specialisation. However, we found clear evidence of specialisation by activity period in western grey kangaroos and red-necked wallabies, but not in swamp wallabies. The strength of behavioural decisions made during the active period in influencing specialisation points to the likelihood that bottom-up processes regulate this community.

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.