RESEARCH ◥ extinction by the International Union for the REVIEW Conservation of Nature (IUCN) (24)and found that for at least 4403 (15%), modification of FIRE ECOLOGY fire regimes is a recorded threat. Changes in fire activity threaten a range of taxonomic groups Fire and biodiversity in the Anthropocene that have been assessed comprehensively or through sampling representative species or Luke T. Kelly1*, Katherine M. Giljohann2, Andrea Duane3, Núria Aquilué3,4 , Sally Archibald5,6, multiple regions, from birds, dragonflies, and Enric Batllori7,8 , Andrew F. Bennett9, Stephen T. Buckland10, Quim Canelles3, Michael F. Clarke9, mammals to gymnosperms, legumes, and Marie-Josée Fortin11, Virgilio Hermoso3, Sergi Herrando12, Robert E. Keane13, Frank K. Lake14, monocots(Fig.1A).Somegroups, such as Michael A. McCarthy2, Alejandra Morán-Ordez3, Catherine L. Parr5,15,16, Juli G. Pausas17, gymnosperms, are at greater risk of fire-driven Trent D. Penman1, Adrián Regos18,19, Libby Rumpff2, Julianna L. Santos1, Annabel L. Smith20,21 , extinction: Changed fire activity is a threat Alexandra D. Syphard22,23,24 , Morgan W. Tingley25, Lluís Brotons3,7,26 to 28% of these taxa classified as critically endangered, endangered, or vulnerable (Fig. 1A). Fire has been a source of global biodiversity for millions of years. However, interactions with Changes in fire activity threaten biodiversity anthropogenic drivers such as climate change, land use, and invasive species are changing the nature of in habitat types worldwide (Fig. 1B). Propor- fire activity and its impacts. We review how such changes are threatening species with extinction and tionally, the threat from changed fire activity transforming terrestrial ecosystems. Conservation of Earth’s biological diversity will be achieved only to species at risk of extinction is greater for by recognizing and responding to the critical role of fire. In the Anthropocene, this requires that savannas (28%), closely followed by grasslands conservation planning explicitly includes the combined effects of human activities and fire regimes. (26%), rocky areas (26%), shrublands (26%), Improved forecasts for biodiversity must also integrate the connections among people, fire, and and forests (19%) (Fig. 1B). ecosystems. Such integration provides an opportunity for new actions that could revolutionize how Across nine taxonomic groups that have society sustains biodiversity in a time of changing fire activity. been assessed systematically (Fig. 1A), we foundthatatleast1071 speciesarecatego- rizedasthreatenedbyanincreaseinfire ire has shaped the diversity of life on by synthesizing how changes in fire activity frequency or intensity and 55 species by Earth for millions of years (1). Variation threaten species with extinction across the exclusion of fire. This delineation, however, in fire regimes enables many plants to globe. Next, we examine how multiple human oversimplifies the nature of threats; for example, F complete their life cycles (2), creates drivers are causing these changes in fire acti- it masks the relationship in some ecosystems habitats for a range of animals (3), and vity and biodiversity. We then highlight forward- maintains a diversity of ecosystems (4). Al- looking methods for predicting changes in though people have used fire to modify envi- ecosystems and forecasting the positive and 1School of Ecosystem and Forest Sciences, University of ronments for millennia (5–7), the cumulative negative effects of fire on biodiversity. Finally, Melbourne, Parkville, Victoria 3010, Australia. 2School of BioSciences, University of Melbourne, Parkville, Victoria 3010, effects of human activities are now changing we foreshadow emerging actions and strategies Australia. 3InForest JRU (CTFC-CREAF), 25280 Solsona, Lleida, patterns of fire at a global scale—to the det- that could revolutionize how society manages Spain. 4Centre d'Étude de la Forêt, Université du Québec à riment of human society, biodiversity, and biodiversity in ecosystems that experience fire. Montréal, Montreal, Quebec H3C 3P8, Canada. 5Centre for ecosystems. Our review concludes that conservation of African Ecology, School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Johannesburg, South Many recent fires have burned ecosystems Earth’s biodiversity is unlikely to be achieved Africa. 6Natural Resources and the Environment, CSIR, Pretoria, wherefirehas historically been rareor absent, without incorporating the critical role of fire South Africa. 7CREAF, Edifici C. Autonomous, University of 8 from the tropical forests of Southeast Asia (8) in national biodiversity strategies and action Barcelona, 08193 Bellaterra, Barcelona, Spain. Department of Evolutionary Biology, Ecology, and Environmental Sciences, and South America (9) to the tundra of the plans and in the implementation of interna- University of Barcelona, 08028 Barcelona, Spain. 9Department Arctic Circle (10). Large, severe fires have also tional agreements and initiatives such as the of Ecology, Environment and Evolution, Centre for Future 10 been observed in areas with a long history of UN Convention on Biological Diversity. Landscapes, La Trobe University, Bundoora, Australia. Centre for Research into Ecological and Environmental Modelling, recurrent fire, and this is consistent with University of St Andrews, St Andrews, Fife KY16 9LZ, UK. predictions of increased wildfire activity in Extinction risk in a fiery world 11Department of Ecology & Evolutionary Biology, University of 12 the boreal forests of Canada and Russia (11, 12) A central concept in fire science is the fire re- Toronto, Toronto, Ontario M5S 3B2, Canada. Catalan Ornithological Institute, Natural History Museum of Barcelona, and the mixed forests and shrublands of gime, which describes the type, frequency, in- 08019 Barcelona, Catalonia, Spain. 13U.S. Department of Australia, southern Europe, and the western tensity, seasonality, and spatial dimensions of Agriculture Forest Service Rocky Mountain Research Station, United States (13–15). Conversely, fire-dependent recurrent fire (19). Many species are adapted to Missoula Fire Sciences Laboratory, Missoula, MT 59808, USA. 14U.S. Department of Agriculture Forest Service Pacific grassland and savanna ecosystems in coun- a particular fire regime, so substantial changes Southwest Research Station, Albany, CA 94710, USA. tries such as Brazil, Tanzania, and the United to these fire characteristics can harm popula- 15Department of Earth, Ocean & Ecological Sciences, University States have had fire activity reduced and even tions (20) and shift ecosystems (21). For example, of Liverpool, Liverpool, UK. 16Department of Zoology & Entomology, University of Pretoria, Pretoria, South Africa. excluded (16–18). These emerging changes pose plants that require fire to release seeds can be 17 Centro de Investigaciones sobre Desertificacin (CIDE-CSIC), a global challenge for understanding how to threatened by fire intervals shorter than the 46113 Montcada, Valencia, Spain. 18Departamento de Zooloxía, sustain biodiversity in a new era of fire. This time needed for them to mature and reestab- Xenética e Antropoloxía Fisica, Universidade de Santiago de 19 requires improved knowledge of the interactions lish a seed bank or by fire intervals longer than Compostela, Santiago de Compostela, Spain. CIBIO/InBIO, Research Center in Biodiversity and Genetic Resources, among fire, biodiversity, and human drivers and seed and plant life spans (22). For animals, ECOCHANGE Group, Vairão, Portugal. 20School of Agriculture new insights into conservation actions that will changes in the frequency and intensity of fire and Food Science, University of Queensland, Gatton 4343, 21 be effective in this changing environment. can reduce the availability of key resources Australia. Zoology, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. 22Vertus Wildfire, San Francisco, CA In this review, we explore the causes and for foraging and shelter, limit the capacity to 94108, USA. 23San Diego State University, San Diego, CA 92182, consequences of fire-induced changes to bio- recolonize regenerating habitats, and, in the case USA. 24Conservation Biology Institute, Corvallis, OR 97333, USA. 25 diversity in the Anthropocene, the current era of severe fires, directly increase mortality (23). Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095, USA. 26Spanish Research Council (CSIC), characterized by the prominent role of human We reviewed the 29,304 terrestrial and fresh- 08193 Bellaterra, Barcelona, Spain. activity in shaping global ecosystems. We start water species categorized as threatened with *Corresponding author. Email: [email protected] Kelly et al., Science 370, eabb0355 (2020) 20 November 2020 1of 10 RESEARCH | REVIEW Vulnerable Endangered Critically endangered toward a hotter climate and a markedly different biosphere (27). Fire is both a con- A Gymnosperms n = 395 27.8% sequence of and a contributor to this acceler- ation (28) but it is not acting alone: Interactions Legumes n = 1032 19.4% between fire and anthropogenic drivers such as Birds n = 1484 18.6% global climate change, land use, and invasive Monocots n = 2533 18.2% species are reshaping ecosystems worldwide. Recent work describing global fire regimes has Amphibians n = 2200 17.9% shown that patterns of fire are closely linked Dragonfies and damselfies n = 372 17.2% to climate, vegetation, and human activity (7, 17, 29, 30). Here, we synthesize linked Mammals