Shifting fire regimes cause continent-wide transformation of threatened species habitat.

Human actions are causing widespread increases in fire size, frequency, and severity in diverse ecosystems globally. This alteration of fire regimes is considered a threat to numerous animal species, but empirical evidence of how fire regimes are shifting within both threatened species' ranges and protected areas is scarce, particularly at large spatial and temporal scales. We used a big data approach to quantify multidecadal changes in fire regimes in southern Australia from 1980 to 2021, spanning 415 reserves (21.5 million ha) and 129 threatened species' ranges including birds, mammals, reptiles, invertebrates, and frogs. Most reserves and threatened species' ranges within the region have experienced declines in unburnt vegetation (≥30 y without fire), increases in recently burnt vegetation (≤5 y since fire), and increases in fire frequency. The mean percentage of unburnt vegetation within reserves declined from 61 to 36% (1980 to 2021), whereas the mean percentage of recently burnt vegetation increased from 20 to 35%, and mean fire frequency increased by 32%, with the latter two trends primarily driven by the record-breaking 2019 to 2020 fire season. The strongest changes occurred for high-elevation threatened species, and reserves of high elevation, high productivity, and strong rainfall decline, particularly in the southeast of the continent. Our results provide evidence for the widely held but poorly tested assumption that threatened species are experiencing widespread declines in unburnt habitat and increases in fire frequency. This underscores the imperative for developing management strategies that conserve fire-threatened species in an increasingly fiery future.

Experimentally testing animal responses to prescribed fire size and severity.

Deserts are often highly biodiverse and provide important habitats for many threatened species. Fire is a dominant disturbance in deserts, and prescribed burning is increasingly being used by conservation managers and Indigenous peoples to mitigate the damaging effects of climate change, invasive plants, and land-use change. The size, severity, and patchiness of fires can affect how animals respond to fire. However, there are almost no studies examining such burn characteristics in desert environments, which precludes the use of such information in conservation planning. Using a before-after control-impact approach with 20 sampling sites, we studied the outcomes of 10 prescribed burns of varying size (5-267 ha), severity, and patchiness to identify which variables best predicted changes in small mammal and reptile species richness and abundance. Three of the 13 species showed a clear response to fire. Captures increased for 2 species (1 mammal, 1 reptile) and decreased for 1 species (a reptile) as the proportional area burned around traps increased. Two other mammal species showed weaker positive responses to fire. Total burn size and burn patchiness were not influential predictors for any species. Changes in capture rates occurred only at sites with the largest and most severe burns. No fire-related changes in capture rates were observed where fires were small and very patchy. Our results suggest that there may be thresholds of fire size or fire severity that trigger responses to fire, which has consequences for management programs underpinned by the patch mosaic burning paradigm. The prescribed burns we studied, which are typical in scale and intensity across many desert regions, facilitated the presence of some taxa and are unlikely to have widespread or persistent negative impacts on small mammal or reptile communities in this ecosystem provided that long unburned habitat harboring threatened species is protected.

Prueba experimental de la respuesta animal al tamaño y gravedad de los incendios controlados Resumen Los desiertos suelen contar con mucha biodiversidad y proporcionar hábitats importantes para una variedad de especies amenazadas. El fuego es una perturbación que domina en los desiertos, y los incendios controlados cada vez se usan más por los gestores de la conservación y los pueblos indígenas para mitigar los efectos dañinos del cambio climático, las plantas invasoras y el cambio de uso de suelo. El tamaño, gravedad y fragmentación de los incendios pueden afectar cómo los animales responden al fuego. Sin embargo, casi no existen estudios que analicen dichas características de la quema en los ambientes desérticos, lo que excluye a dicha información de la planeación de la conservación. Usamos una estrategia de antes­después del control­impacto en 20 sitios de muestreo para estudiar los resultados de diez incendios controlados de diferentes tamaños (5­267 ha), gravedad y fragmentación para identificar cuáles variables pronostican mejor los cambios en la riqueza de especies y abundancia de mamíferos pequeños y reptiles. Tres de las 13 especies mostraron una respuesta clara al incendio. Las capturas incrementaron en dos especies (una de mamífero y una de reptil) y disminuyeron en una especie (un reptil) conforme incrementó el área proporcional incendiada alrededor de las trampas. Otras dos especies de mamíferos mostraron respuestas positivas más débiles ante el fuego. El tamaño total y la fragmentación del incendio no fueron influyentes sobre los pronosticadores de cualquier especie. Los cambios en las tasas de captura ocurrieron solamente en los sitios con los incendios más graves y grandes. No observamos cambios relacionados al incendio en las tasas de captura en donde los incendios fueron pequeños y muy fragmentados. Nuestros resultados sugieren que podría haber umbrales del tamaño o gravedad del incendio que provocan las respuestas al fuego, lo que tiene consecuencias para los programas de manejo sustentados en el paradigma del mosaico de fragmentos del incendio. Los incendios controlados que estudiamos, que son típicos en escala e intensidad en muchas regiones desérticas, facilitaron la presencia de algunos taxones y no tuvieron probabilidad de tener un impacto negativo extenso o persistente sobre las comunidades de mamíferos pequeños y reptiles en este ecosistema, siempre y cuando se proteja el hábitat que lleva mucho tiempo sin incendios y en donde viven las especies amenazadas.

Meta-analysis reveals impacts of disturbance on reptile and amphibian body condition.

Ecosystem disturbance is increasing in extent, severity and frequency across the globe. To date, research has largely focussed on the impacts of disturbance on animal population size, extinction risk and species richness. However, individual responses, such as changes in body condition, can act as more sensitive metrics and may provide early warning signs of reduced fitness and population declines. We conducted the first global systematic review and meta-analysis investigating the impacts of ecosystem disturbance on reptile and amphibian body condition. We collated 384 effect sizes representing 137 species from 133 studies. We tested how disturbance type, species traits, biome and taxon moderate the impacts of disturbance on body condition. We found an overall negative effect of disturbance on herpetofauna body condition (Hedges' g = -0.37, 95% CI: -0.57, -0.18). Disturbance type was an influential predictor of body condition response and all disturbance types had a negative mean effect. Drought, invasive species and agriculture had the largest effects. The impact of disturbance varied in strength and direction across biomes, with the largest negative effects found within Mediterranean and temperate biomes. In contrast, taxon, body size, habitat specialisation and conservation status were not influential predictors of disturbance effects. Our findings reveal the widespread effects of disturbance on herpetofauna body condition and highlight the potential role of individual-level response metrics in enhancing wildlife monitoring. The use of individual response metrics alongside population and community metrics would deepen our understanding of disturbance impacts by revealing both early impacts and chronic effects within affected populations. This could enable early and more informed conservation management.

Identifying historical and future global change drivers that place species recovery at risk.

Ecosystem management in the face of global change requires understanding how co-occurring threats affect species and communities. Such an understanding allows for effective management strategies to be identified and implemented. An important component of this is differentiating between factors that are within (e.g. invasive predators) or outside (e.g. drought, large wildfires) of a local manager's control. In the global biodiversity hotspot of south-western Australia, small- and medium-sized mammal species are severely affected by anthropogenic threats and environmental disturbances, including invasive predators, fire, and declining rainfall. However, the relative importance of different drivers has not been quantified. We used data from a long-term monitoring program to fit Bayesian state-space models that estimated spatial and temporal changes in the relative abundance of four threatened mammal species: the woylie (Bettongia penicillata), chuditch (Dasyurus geoffroii), koomal (Trichosurus vulpecula) and quenda (Isoodon fusciventor). We then use Bayesian structural equation modelling to identify the direct and indirect drivers of population changes, and scenario analysis to forecast population responses to future environmental change. We found that habitat loss or conversion and reduced primary productivity (caused by rainfall declines) had greater effects on species' spatial and temporal population change than the range of fire and invasive predator (the red fox Vulpes vulpes) management actions observed in the study area. Scenario analysis revealed that a greater extent of severe fire and further rainfall declines predicted under climate change, operating in concert are likely to further reduce the abundance of these species, but may be mitigated partially by invasive predator control. Considering both historical and future drivers of population change is necessary to identify the factors that risk species recovery. Given that both anthropogenic pressures and environmental disturbances can undermine conservation efforts, managers must consider how the relative benefit of conservation actions will be shaped by ongoing global change.

Animal mortality during fire.

Earth's rapidly warming climate is propelling us towards an increasingly fire-prone future. Currently, knowledge of the extent and characteristics of animal mortality rates during fire remains rudimentary, hindering our ability to predict how animal populations may be impacted in the future. To address this knowledge gap, we conducted a global systematic review of the direct effects of fire on animal mortality rates, based on studies that unequivocally determined the fate of animals during fire. From 31 studies spanning 1984-2020, we extracted data on the direct impacts of fire on the mortality of 31 species from 23 families. From these studies, there were 43 instances where direct effects were measured by reporting animal survival from pre- to post-fire. Most studies were conducted in North America (52%) and Oceania (42%), focused largely on mammals (53%) and reptiles (30%), and reported mostly on animal survival in planned (82%) and/or low severity (70%) fires. We found no studies from Asia, Europe or South America. Although there were insufficient data to conduct a formal meta-analysis, we tested the effect of fire type, fire severity, fire regime, animal body mass, ecological attributes and class on survival. Only fire severity affected animal mortality, with a higher proportion of animals being killed by high than low severity fires. Recent catastrophic fires across the globe have drawn attention to the plight of animals exposed to wildfire. Yet, our systematic review suggests that a relatively low proportion of animals (mean predicted mortality [95% CI] = 3% [1%-9%]) are killed during fire. However, our review also underscores how little we currently know about the direct effects of fire on animal mortality, and highlights the critical need to understand the effects of high severity fire on animal populations.

Predators, prey or temperature? Mechanisms driving niche use of a foundation plant species by specialist lizards.

Foundation species interact strongly with other species to profoundly influence communities, such as by providing food, refuge from predators or beneficial microclimates. We tested relative support for these mechanisms using spinifex grass (Triodia spp.), which is a foundation species of arid Australia that provides habitat for diverse lizard communities. We first compared the attributes of live and dead spinifex, bare ground and a structurally similar plant (Lomandra effusa), and then tested the relative strength of association of two spinifex specialist lizard species (Ctenophorus spinodomus and Ctenotus atlas) with spinifex using a mesocosm experiment. Temperatures were coolest within spinifex compared to bare ground and Lomandra. Invertebrate abundance and the threat of predation were indistinguishable between treatments, suggesting temperature attenuation may be a more important driver. Overall, the dragon C. spinodomus preferred live over dead spinifex, while the skink C. atlas preferred dead spinifex, particularly at warmer air temperatures. However, both species displayed individual variability in their use of available microhabitats, with some individuals rarely using spinifex. Our results provide an example of temperature attenuation by a foundation species driving niche use by ectothermic animals.

Predator responses to fire: A global systematic review and meta-analysis.

Knowledge of how disturbances such as fire shape habitat structure and composition, and affect animal interactions, is fundamental to ecology and ecosystem management. Predators also exert strong effects on ecological communities, through top-down regulation of prey and competitors, which can result in trophic cascades. Despite their ubiquity, ecological importance and potential to interact with fire, our general understanding of how predators respond to fire remains poor, hampering ecosystem management. To address this important knowledge gap, we conducted a systematic review and meta-analysis of the effects of fire on terrestrial, vertebrate predators world-wide. We found 160 studies spanning 1978-2018. There were 36 studies with sufficient information for meta-analysis, from which we extracted 96 effect sizes (Hedges' g) for 67 predator species relating to changes in abundance indices, occupancy or resource selection in burned and unburned areas, or before and after fire. Studies spanned geographic locations, taxonomic families and study designs, but most were located in North America and Oceania (59% and 24%, respectively), and largely focussed on felids (24%) and canids (25%). Half (50%) of the studies reported responses to wildfire, and nearly one third concerned prescribed (management) fires. There were no clear, general responses of predators to fire, nor relationships with geographic area, biome or life-history traits (e.g. body mass, hunting strategy and diet). Responses varied considerably between species. Analysis of species for which at least three effect sizes had been reported in the literature revealed that red foxes Vulpes vulpes mostly responded positively to fire (e.g. higher abundance in burned compared to unburned areas) and eastern racers Coluber constrictor negatively, with variances overlapping zero only slightly for both species. Our systematic review and meta-analysis revealed strong variation in predator responses to fire, and major geographic and taxonomic knowledge gaps. Varied responses of predator species to fire likely depend on ecosystem context. Consistent reporting of ongoing monitoring and management experiments is required to improve understanding of the mechanisms driving predator responses to fire, and any broader effects (e.g. trophic interactions). The divergent responses of species in our study suggest that adaptive, context-specific management of predator-fire relationships is required.

Invasive predators and global biodiversity loss.

Invasive species threaten biodiversity globally, and invasive mammalian predators are particularly damaging, having contributed to considerable species decline and extinction. We provide a global metaanalysis of these impacts and reveal their full extent. Invasive predators are implicated in 87 bird, 45 mammal, and 10 reptile species extinctions-58% of these groups' contemporary extinctions worldwide. These figures are likely underestimated because 23 critically endangered species that we assessed are classed as "possibly extinct." Invasive mammalian predators endanger a further 596 species at risk of extinction, with cats, rodents, dogs, and pigs threatening the most species overall. Species most at risk from predators have high evolutionary distinctiveness and inhabit insular environments. Invasive mammalian predators are therefore important drivers of irreversible loss of phylogenetic diversity worldwide. That most impacted species are insular indicates that management of invasive predators on islands should be a global conservation priority. Understanding and mitigating the impact of invasive mammalian predators is essential for reducing the rate of global biodiversity loss.

Coupling movement and landscape ecology for animal conservation in production landscapes.

Habitat conversion in production landscapes is among the greatest threats to biodiversity, not least because it can disrupt animal movement. Using the movement ecology framework, we review animal movement in production landscapes, including areas managed for agriculture and forestry. We consider internal and external drivers of altered animal movement and how this affects navigation and motion capacities and population dynamics. Conventional management approaches in fragmented landscapes focus on promoting connectivity using structural changes in the landscape. However, a movement ecology perspective emphasizes that manipulating the internal motivations or navigation capacity of animals represents untapped opportunities to improve movement and the effectiveness of structural connectivity investments. Integrating movement and landscape ecology opens new opportunities for conservation management in production landscapes.

BioTIME: A database of biodiversity time series for the Anthropocene.

MOTIVATION: The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. MAIN TYPES OF VARIABLES INCLUDED: The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. SPATIAL LOCATION AND GRAIN: BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). TIME PERIOD AND GRAIN: BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. MAJOR TAXA AND LEVEL OF MEASUREMENT: BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. SOFTWARE FORMAT: .csv and .SQL.

How do invasive predators and their native prey respond to prescribed fire?

Fire shapes animal communities by altering resource availability and species interactions, including between predators and prey. In Australia, there is particular concern that two highly damaging invasive predators, the feral cat (Felis catus) and European red fox (Vulpes vulpes), increase their activity in recently burnt areas and exert greater predation pressure on the native prey due to their increased exposure. We tested how prescribed fire occurrence and extent, along with fire history, vegetation, topography, and distance to anthropogenic features (towns and farms), affected the activity (detection frequency) of cats, foxes, and the native mammal community in south-eastern Australia. We used camera traps to quantify mammal activity before and after a prescribed burn and statistically tested how the fire interacted with these habitat variables to affect mammal activity. We found little evidence that the prescribed fire influenced the activity of cats and foxes and no evidence of an effect on kangaroo or small mammal (<800 g) activity. Medium-sized mammals (800-2000 g) were negatively associated with prescribed fire extent, suggesting that prescribed fire has a negative impact on these species in the short term. The lack of a clear activity increase from cats and foxes is likely a positive outcome from a fire management perspective. However, we highlight that their response is likely dependent upon factors like fire size, severity, and prey availability. Future experiments should incorporate GPS-trackers to record fine-scale movements of cats and foxes in temperate ecosystems immediately before and after prescribed fire to best inform management within protected areas.

A global synthesis and assessment of free-ranging domestic cat diet.

Free-ranging cats (Felis catus) are globally distributed invasive carnivores that markedly impact biodiversity. Here, to evaluate the potential threat of cats, we develop a comprehensive global assessment of species consumed by cats. We identify 2,084 species eaten by cats, of which 347 (16.65%) are of conservation concern. Islands contain threefold more species of conservation concern eaten by cats than continents do. Birds, reptiles, and mammals constitute ~90% of species consumed, with insects and amphibians being less frequent. Approximately 9% of known birds, 6% of known mammals, and 4% of known reptile species are identified in cat diets. 97% of species consumed are <5 kg in adult body mass, though much larger species are also eaten. The species accumulation curves are not asymptotic, indicating that our estimates are conservative. Our results demonstrate that cats are extreme generalist predators, which is critical for understanding their impact on ecological systems and developing management solutions.

Distinctive diets of eutherian predators in Australia.

Introduction of the domestic cat and red fox has devastated Australian native fauna. We synthesized Australian diet analyses to identify traits of prey species in cat, fox and dingo diets, which prey were more frequent or distinctive to the diet of each predator, and quantified dietary overlap. Nearly half (45%) of all Australian terrestrial mammal, bird and reptile species occurred in the diets of one or more predators. Cat and dingo diets overlapped least (0.64 ± 0.27, n = 24 location/time points) and cat diet changed little over 55 years of study. Cats were more likely to have eaten birds, reptiles and small mammals than foxes or dingoes. Dingo diet remained constant over 53 years and constituted the largest mammal, bird and reptile prey species, including more macropods/potoroids, wombats, monotremes and bandicoots/bilbies than cats or foxes. Fox diet had greater overlap with both cats (0.79 ± 0.20, n = 37) and dingoes (0.73 ± 0.21, n = 42), fewer distinctive items (plant material, possums/gliders) and significant spatial and temporal heterogeneity over 69 years, suggesting the opportunity for prey switching (especially of mammal prey) to mitigate competition. Our study reinforced concerns about mesopredator impacts upon scarce/threatened species and the need to control foxes and cats for fauna conservation. However, extensive dietary overlap and opportunism, as well as low incidence of mesopredators in dingo diets, precluded resolution of the debate about possible dingo suppression of foxes and cats.

Fire as a driver and mediator of predator-prey interactions.

Both fire and predators have strong influences on the population dynamics and behaviour of animals, and the effects of predators may either be strengthened or weakened by fire. However, knowledge of how fire drives or mediates predator-prey interactions is fragmented and has not been synthesised. Here, we review and synthesise knowledge of how fire influences predator and prey behaviour and interactions. We develop a conceptual model based on predator-prey theory and empirical examples to address four key questions: (i) how and why do predators respond to fire; (ii) how and why does prey vulnerability change post-fire; (iii) what mechanisms do prey use to reduce predation risk post-fire; and (iv) what are the outcomes of predator-fire interactions for prey populations? We then discuss these findings in the context of wildlife conservation and ecosystem management before outlining priorities for future research. Fire-induced changes in vegetation structure, resource availability, and animal behaviour influence predator-prey encounter rates, the amount of time prey are vulnerable during an encounter, and the conditional probability of prey death given an encounter. How a predator responds to fire depends on fire characteristics (e.g. season, severity), their hunting behaviour (ambush or pursuit predator), movement behaviour, territoriality, and intra-guild dynamics. Prey species that rely on habitat structure for avoiding predation often experience increased predation rates and lower survival in recently burnt areas. By contrast, some prey species benefit from the opening up of habitat after fire because it makes it easier to detect predators and to modify their behaviour appropriately. Reduced prey body condition after fire can increase predation risk either through impaired ability to escape predators, or increased need to forage in risky areas due to being energetically stressed. To reduce risk of predation in the post-fire environment, prey may change their habitat use, increase sheltering behaviour, change their movement behaviour, or use camouflage through cryptic colouring and background matching. Field experiments and population viability modelling show instances where fire either amplifies or does not amplify the impacts of predators on prey populations, and vice versa. In some instances, intense and sustained post-fire predation may lead to local extinctions of prey populations. Human disruption of fire regimes is impacting faunal communities, with consequences for predator and prey behaviour and population dynamics. Key areas for future research include: capturing data continuously before, during and after fires; teasing out the relative importance of changes in visibility and shelter availability in different contexts; documenting changes in acoustic and olfactory cues for both predators and prey; addressing taxonomic and geographic biases in the literature; and predicting and testing how changes in fire-regime characteristics reshape predator-prey interactions. Understanding and managing the consequences for predator-prey communities will be critical for effective ecosystem management and species conservation in this era of global change.

Human disturbance causes widespread disruption of animal movement.

Disturbance and habitat modification by humans can alter animal movement, leading to negative impacts on fitness, survival and population viability. However, the ubiquity and nature of these impacts across diverse taxa has not been quantified. We compiled 208 studies on 167 species from terrestrial and aquatic ecosystems across the globe to assess how human disturbance influences animal movement. We show that disturbance by humans has widespread impacts on the movements of birds, mammals, reptiles, amphibians, fish and arthropods. More than two-thirds of 719 cases represented a change in movement of 20% or more, with increases in movement averaging 70% and decreases -37%. Disturbance from human activities, such as recreation and hunting, had stronger impacts on animal movement than habitat modification, such as logging and agriculture. Our results point to a global restructuring of animal movement and emphasize the need to reduce the negative impacts of humans on animal movement.

Artificial refuges for wildlife conservation: what is the state of the science?

Artificial refuges are human-made structures that aim to create safe places for animals to breed, hibernate, or take shelter in lieu of natural refuges. Artificial refuges are used across the globe to mitigate the impacts of a variety of threats on wildlife, such as habitat loss and degradation. However, there is little understanding of the science underpinning artificial refuges, and what comprises best practice for artificial refuge design and implementation for wildlife conservation. We address this gap by undertaking a systematic review of the current state of artificial refuge research for the conservation of wildlife. We identified 224 studies of artificial refuges being implemented in the field to conserve wildlife species. The current literature on artificial refuges is dominated by studies of arboreal species, primarily birds and bats. Threatening processes addressed by artificial refuges were biological resource use (26%), invasive or problematic species (20%), and agriculture (15%), yet few studies examined artificial refuges specifically for threatened (Vulnerable, Endangered, or Critically Endangered) species (7%). Studies often reported the characteristics of artificial refuges (i.e. refuge size, construction materials; 87%) and surrounding vegetation (35%), but fewer studies measured the thermal properties of artificial refuges (18%), predator activity (17%), or food availability (3%). Almost all studies measured occupancy of the artificial refuges by target species (98%), and over half measured breeding activity (54%), whereas fewer included more detailed measures of fitness, such as breeding productivity (34%) or animal body condition (4%). Evaluating the benefits and impacts of artificial refuges requires sound experimental design, but only 39% of studies compared artificial refuges to experimental controls, and only 10% of studies used a before-after-control-impact (BACI) design. As a consequence, few studies of artificial refuges can determine their overall effect on individuals or populations. We outline a series of key steps in the design, implementation, and monitoring of artificial refuges that are required to avoid perverse outcomes and maximise the chances of achieving conservation objectives. This review highlights a clear need for increased rigour in studies of artificial refuges if they are to play an important role in wildlife conservation.

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.

A guide to ecosystem models and their environmental applications.

Applied ecology has traditionally approached management problems through a simplified, single-species lens. Repeated failures of single-species management have led us to a new paradigm - managing at the ecosystem level. Ecosystem management involves a complex array of interacting organisms, processes and scientific disciplines. Accounting for interactions, feedback loops and dependencies between ecosystem components is therefore fundamental to understanding and managing ecosystems. We provide an overview of the main types of ecosystem models and their uses, and discuss challenges related to modelling complex ecological systems. Existing modelling approaches typically attempt to do one or more of the following: describe and disentangle ecosystem components and interactions; make predictions about future ecosystem states; and inform decision making by comparing alternative strategies and identifying important uncertainties. Modelling ecosystems is challenging, particularly when balancing the desire to represent many components of an ecosystem with the limitations of available data and the modelling objective. Explicitly considering different forms of uncertainty is therefore a primary concern. We provide some recommended strategies (such as ensemble ecosystem models and multi-model approaches) to aid the explicit consideration of uncertainty while also meeting the challenges of modelling ecosystems.

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.