Fire is a powerful ecological and evolutionary force that regulates organismal traits, population sizes, species interactions, community composition, carbon and nutrient cycling and ecosystem function. It also presents a rapidly growing societal challenge, due to both increasingly destructive wildfires and fire exclusion in fire‐dependent ecosystems. As an ecological process, fire integrates complex feedbacks among biological, social and geophysical processes, requiring coordination across several fields and scales of study. Here, we describe the diversity of ways in which fire operates as a fundamental ecological and evolutionary process on Earth. We explore research priorities in six categories of fire ecology: (a) characteristics of fire regimes, (b) changing fire regimes, (c) fire effects on above‐ground ecology, (d) fire effects on below‐ground ecology, (e) fire behaviour and (f) fire ecology modelling. We identify three emergent themes: the need to study fire across temporal scales, to assess the mechanisms underlying a variety of ecological feedbacks involving fire and to improve representation of fire in a range of modelling contexts. Synthesis: As fire regimes and our relationships with fire continue to change, prioritizing these research areas will facilitate understanding of the ecological causes and consequences of future fires and rethinking fire management alternatives.

The original concept of succession emphasized directional changes in species composition occurring over time on “blank slates” (new substrates or those that followed catastrophic disturbances). In this paper, we explore relationships between different effects of disturbances on residents and the initiation of directional species replacement on sites in a landscape. We present a conceptual model involving recurrent natural disturbances of sites in a landscape containing two species, one of which arrives at sites early and the other arrives late following disturbance. We predict effects of recurrent catastrophic disturbances (no survivors) and non‐catastrophic disturbances (some survivors) on the initiation of directional species replacement, assuming temporal gradients in environmental conditions on sites after disturbance. We predict that directional species replacement will be initiated by catastrophic disturbances, and by non‐catastrophic disturbances when the early species does not survive, but reinvades. Most natural disturbances tend to leave survivors and do not remove early species preferentially. As a result, many post‐disturbance changes are possible on sites; directional species replacement is only one change, and often may not be the most common. One important consequence of the low likelihood that disturbances will initiate directional species replacements should be the precarious existence of early species dependent on disturbances that open sites for colonization. We examined constraints on disturbances, sites, and life histories that might increase colonization by early species. We explored how refuges and life history characteristics might reduce the likelihood of replacement of early species by late species, and we consider how spatial and temporal variation in disturbance effects might influence persistence of early species. In this way, we propose relationships between effects of non‐catastrophic disturbance and the dynamic relationships between species that may be applicable across different ecological systems.Corresponding Editor: S. T. Jackson

Summary: Pyrogenic savannas with a tree–grassland ‘matrix’ experience frequent fires (i.e. every 1–3 yr). Aboveground responses to frequent fires have been well studied, but responses of fungal litter decomposers, which directly affect fuels, remain poorly known. We hypothesized that each fire reorganizes belowground communities and slows litter decomposition, thereby influencing savanna fuel dynamics. In a pine savanna, we established patches near and away from pines that were either burned or unburned in that year. Within patches, we assessed fungal communities and microbial decomposition of newly deposited litter. Soil variables and plant communities were also assessed as proximate drivers of fungal communities. Fungal communities, but not soil variables or vegetation, differed substantially between burned and unburned patches. Saprotrophic fungi dominated in unburned patches but decreased in richness and relative abundance after fire. Differences in fungal communities with fire were greater in litter than in soils, but unaffected by pine proximity. Litter decomposed more slowly in burned than in unburned patches. Fires drive shifts between fire‐adapted and sensitive fungal taxa in pine savannas. Slower fuel decomposition in accordance with saprotroph declines should enhance fuel accumulation and could impact future fire characteristics. Thus, fire reorganization of fungal communities may enhance persistence of these fire‐adapted ecosystems.

Fire seasonality, an important characteristic of fire regimes, commonly is delineated using seasons based on single weather variables (rainfall or temperature). We used nonparametric cluster analyses of a 17-year (1993–2009) data set of weather variables that influence likelihoods and spread of fires (relative humidity, air temperature, solar radiation, wind speed, soil moisture) to explore seasonality of fire in pine savanna-grassland landscapes at the Avon Park Air Force Range in southern Florida. A four-variable, three-season model explained more variation within fire weather variables than models with more seasons. The three-season model also delineated intra-annual timing of fire more accurately than a conventional rainfall-based two-season model. Two seasons coincided roughly with dry and wet seasons based on rainfall. The third season, which we labeled the fire season, occurred between dry and wet seasons and was characterized by fire-promoting conditions present annually: drought, intense solar radiation, low humidity, and warm air temperatures. Fine fuels consisting of variable combinations of pyrogenic pine needles, abundant C4 grasses, and flammable shrubs, coupled with low soil moisture, and lightning ignitions early in the fire season facilitate natural landscape-scale wildfires that burn uplands and across wetlands. We related our three season model to fires with different ignition sources (lightning, military missions, and prescribed fires) over a 13-year period with fire records (1997–2009). Largest wildfires originate from lightning and military ignitions that occur within the early fire season substantially prior to the peak of lightning strikes in the wet season. Prescribed ignitions, in contrast, largely occur outside the fire season. Our delineation of a pronounced fire season provides insight into the extent to which different human-derived fire regimes mimic lightning fire regimes. Delineation of a fire season associated with timing of natural lightning ignitions should be useful as a basis for ecological fire management of humid savanna-grassland landscapes worldwide.

Natural fires ignited by lightning strikes following droughts frequently are posited as the ecological mechanism maintaining discontinuous tree cover and grass‐dominated ground layers in savannas. Such fires, however, may not reliably maintain humid savannas. We propose that savanna trees producing pyrogenic shed leaves might engineer fire characteristics, affecting ground‐layer plants in ways that maintain humid savannas. We explored our hypothesis in a high‐rainfall, frequently burned pine savanna in which the dominant tree, longleaf pine (Pinus palustris), produces resinous needles that become highly flammable when shed and dried. We postulated that pyrogenic needles should have much greater influence on fire characteristics at ground level, and hence post‐fire responses of dominant shrubs and grasses, than other abundant fine fuels (shed oak leaves and grass culms). We further reasoned that these effects should increase with amounts of needles. We managed site conditions that affect fuels (time since fire, dominant vegetation), manipulated amounts of needles in ground‐layer plots, prescribed burned the plots, and measured fire characteristics at ground level. We also measured characteristics of ground‐layer oaks and grasses before, then 2 and 8 months after fires. We tested our hypotheses regarding effects of pyrogenic pine fuels on fire characteristics and vegetation regrowth and explored direct and indirect effects of fuels on fire characteristics and vegetation using a structural equation model. Pine needles influenced fire characteristics, elevating maximum temperature increases, durations of heating above 60°C, and fine fuel consumption considerably above measurements when fuels only included other savanna plants. Presence of pine needles depressed post‐fire numbers of oak stems and grass culms, especially in the interior of grass genets, as well as post‐fire flowering of grasses. The structural equation model indicated strong direct and indirect pathways from pine needles to post‐fire responses of oaks and grasses. The experimental field tests of hypotheses, bolstered by structural equation modeling, indicate pyrogenic fine fuels modify characteristics of prescribed fires at ground level, negatively affecting dominant ground‐layer oaks and grasses. Frequent fires fueled by pyrogenic needles should maintain humid savannas and generate spatial pyrodiversity that affects composition and dynamics of pine savanna ground‐layer vegetation.

Research Highlights: Spatial patterns of fire spread and severity influence survival of juvenile pines in longleaf pine savannas. Small areas that do not burn during frequent fires facilitate formation of patches of even-aged longleaf pine juveniles. These regeneration patches are especially associated with inner portions of openings (gaps) and where canopy trees have died in recent decades. Patterns of prescribed fire can thus have an important influence on stand dynamics of the dominant tree in pine savannas. Background and Objectives: Savannas are characterized by bottlenecks to tree regeneration. In pine savannas, longleaf pine is noted for recruitment in discrete clusters located within gaps away from canopy trees. Various mechanisms promoting this pattern have been hypothesized: light limitations, soil moisture, soil nutrients, pine needle mulching, competition with canopy tree roots, and fire severity associated with pine needle litter. We tested the hypothesis that regeneration patches are associated with areas that remain unburned during some prescribed fires, as mediated by gaps in the canopy, especially inner portions of gaps, and areas re-opened by death of canopy trees. Materials and Methods: We mapped areas that were unburned during prescribed fires applied at 1–2 year intervals from 2005–2018 in an old-growth pine savanna in Georgia, USA. We compared the maps to locations of longleaf pine juveniles (<1.5 m height) measured in 2018 and canopy cover and canopy tree deaths using a long-term (40 year) tree census. Results: Logistic regression analysis showed juveniles to be associated with unburned areas, gaps, inner gaps, and areas where canopy trees died. Conclusions: Patterns of fire spread and severity limit survival of longleaf pine juveniles to patches away from canopy trees, especially where canopy trees have died in recent decades. These processes contribute to a buffering mechanism that maintains the savanna structure and prevents transition to closed canopy forest or open grassland communities.