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Fire adaptations

Fire adaptations encompass the suite of evolutionary traits in organisms, particularly plants and animals, that have developed to enhance survival, reproduction, and persistence in fire-prone ecosystems, where recurrent fires act as a major selective force shaping biodiversity and community structure.^[1] These adaptations, which trace back at least 420 million years to the Silurian Period, include morphological features like thick insulating bark in trees such as pines (Pinus spp.) that protect vascular tissues from lethal heat, physiological mechanisms like serotiny in cones that release seeds only after fire exposure, and behavioral responses in animals such as burrowing to escape flames or sensitivity to smoke cues for post-fire foraging.^[2] In plants, resprouting from basal lignotubers or root systems—evident in eucalypts and many grassland species—represents one of the most widespread fire-adaptive strategies, allowing rapid regeneration after aboveground tissues are consumed.^[3] Fire adaptations have profoundly influenced the evolution of terrestrial life, with phylogenetic evidence showing traits like serotiny originating around 100 million years ago in conifers and resprouting emerging independently across angiosperms under diverse disturbance regimes including fire.^[1] In animals, examples include post-fire melanism in insects for camouflage amid charred landscapes and infrared-sensing organs in pyrophilous beetles (Melanophila spp.) that detect distant fires to locate fresh breeding sites.^[1] These traits not only promote individual fitness but also drive ecosystem dynamics, such as maintaining high biodiversity in savannas and Mediterranean shrublands through fire-stimulated germination and nutrient release, while preventing shifts to alternative states like forest encroachment in grasslands.^[2] The study of fire adaptations, rooted in early 20th-century observations but accelerated by modern phylogenetic and paleoecological tools, underscores fire's role as a keystone process integrating biotic and abiotic factors across scales from individual organisms to global biomes.^[3] Contemporary research highlights how alterations in fire regimes—due to climate change, suppression, or land-use shifts—can disrupt these adaptations, leading to biodiversity loss or novel ecosystem configurations, emphasizing the need for informed management to sustain fire-dependent species and habitats.^[3]

Plant Adaptations to Fire

Resistance Strategies

Fire resistance in plants refers to structural and physiological adaptations that protect vital tissues, such as meristems and vascular cambium, from lethal temperatures exceeding 60°C during combustion, thereby enabling survival of the individual plant through an active fire event.^[4]^[5]^[6] A primary resistance strategy is the development of thick bark, which serves as a thermal insulator by limiting heat transfer to underlying tissues due to its low thermal conductivity, typically around 0.06 W/(m·K), and high charring capacity that forms a protective barrier.^[7]^[8] In species like the giant sequoia (Sequoiadendron giganteum), bark thickness can reach up to 75 cm in mature trees, correlating strongly with enhanced fire tolerance in frequent-fire ecosystems, as thicker bark extends the time required for cambial temperatures to reach lethal levels during surface fires.^[9]^[10] Another key feature is self-pruning of lower branches, which reduces the continuity of vertical fuels and prevents the escalation of surface fires into damaging crown fires by eliminating ladder fuels.^[11] In Australian eucalypt forests, many Eucalyptus species exhibit this trait, where shading and natural abscission cause lower branches to die and shed, maintaining a elevated canopy base height that lowers overall fire intensity and spread potential.^[12]^[13] Bark shedding mechanisms in certain species further contribute to resistance by minimizing opportunities for post-fire pathogen invasion through the renewal of the outer protective layer, which can induce defensive resin production and reduce susceptibility to bark beetles and fungi.^[14] For instance, smooth-barked eucalypts periodically exfoliate dead outer layers, creating a fresh barrier that limits entry points for opportunistic pathogens immediately following low-intensity burns.^[15] A notable case study is the jack pine (Pinus banksiana) in North American boreal forests, where mature trees demonstrate resistance to low-intensity surface fires through medium-thick bark that insulates the cambium, allowing survival while succumbing to higher-severity crown fires.^[16] This adaptation aligns with the species' prevalence in fire-prone landscapes, where frequent low-severity fires shape community structure without widespread mortality.^[16]

Recovery Mechanisms

Recovery mechanisms in fire-adapted plants enable the regeneration of damaged individuals through vegetative resprouting from protected structures, allowing persistence in recurrent fire regimes without relying on seed recruitment. These strategies involve dormant buds and stored reserves that activate post-fire, facilitating rapid canopy recovery and maintenance of adult structures. Unlike resistance traits that minimize damage during combustion, recovery focuses on post-fire regrowth from surviving tissues. Epicormic buds, dormant shoots embedded beneath the bark along stems and branches, serve as a critical recovery mechanism by enabling top-down resprouting after fire kills aboveground foliage. In Mediterranean ecosystems, species like cork oak (Quercus suber) exhibit robust epicormic resprouting, where these buds, protected by thick cork layers, activate following crown scorch to restore the arborescent form and accelerate structural recovery compared to seeding-dependent species. Activation is triggered by fire-induced cues such as increased light penetration, elevated temperatures, and hormonal shifts from the loss of apical dominance, though smoke volatiles play a minor role in some contexts by enhancing bud break in certain resprouters.^[17]^[18]^[19] Lignotubers, woody swellings at the plant base that accumulate carbohydrates and adventitious buds, support basal resprouting by providing energy reserves for rapid post-fire regrowth in fire-prone shrublands. In Australian species such as Banksia attenuata and Banksia oblongifolia, lignotubers store substantial starch in associated roots, fueling shoot production after intense fires and enabling survival in regions with frequent disturbances. Energy allocation models indicate that lignotuber development diverts resources from reproduction to maintenance of these reserves, with frequent fires risking depletion if inter-fire intervals are too short, leading to reduced resprouting vigor over time.^[20]^[21]^[22] Clonal spread through root suckers or rhizomes allows vegetative propagation from underground networks, preserving genetic continuity and enabling colony-level survival even if aboveground stems are destroyed. In trembling aspen (Populus tremuloides), extensive root systems produce suckers post-fire, forming large clones that regenerate across disturbed landscapes via these belowground connections, which remain insulated from lethal heat. This mechanism supports rapid stand replacement in boreal and temperate forests, where clones can span hectares and persist for millennia despite individual tree mortality.^[23] Post-fire nutrient remobilization from charred but viable tissues further bolsters regrowth by reallocating nonstructural carbohydrates (NSCs) and minerals to emerging buds and shoots. In species like ponderosa pine (Pinus ponderosa), surviving inner bark and roots mobilize stored NSCs to fuel recovery, with depletion levels correlating to fire severity—trees with moderate scorch recover NSCs within 14 months, while severe injury leads to exhaustion and mortality. This process prioritizes energy redirection from photosynthesis-limited tissues to support initial sprout establishment.^[24] Resprouting success in fire-adapted species typically ranges from 70-90%, far exceeding the <20% rates in non-adapted plants, underscoring the efficacy of these mechanisms in high-frequency fire environments. For instance, lignotuberous shrubs in restored Banksia woodlands achieve 54-98% initial resprouting within months post-fire, influenced by plant size and soil moisture, while non-adapted taxa often fail due to insufficient reserves. These high rates contribute to ecosystem resilience by minimizing post-fire gaps in vegetation cover.^[25]^[26]

Recruitment Strategies

One prominent recruitment strategy in fire-adapted plants is serotiny, where seeds are retained in woody cones or fruits sealed by resins that melt during fire, enabling synchronized release into a post-fire environment favorable for establishment. In lodgepole pine (Pinus contorta), serotinous cones typically remain closed for years or decades until exposed to temperatures around 60-70°C, which causes them to open and disperse seeds directly onto mineral soil enriched by ash nutrients. Studies have shown that seeds from these cones exhibit high viability, with germination rates approaching 80% when cones are heated optimally, such as at 69°C for 8 hours, demonstrating the strategy's effectiveness in rapid recolonization.^[27]^[28] Fire also directly stimulates seed germination through physical and chemical cues that break dormancy in soil-stored seed banks. Heat from fire scarifies hard seed coats, allowing water imbibition, while smoke contains bioactive compounds like karrikins—specifically, butenolides such as 3-methyl-2H-furo[2,3-c]pyran-2-one—that act as signaling molecules to trigger germination pathways involving gibberellic acid synthesis. Laboratory experiments using smoke water extracts have demonstrated this effect across diverse species; for instance, application of dilute smoke solutions induced germination rates up to 100% in dormant seeds of California chaparral plants like Emmenanthe penduliflora, compared to 0% in controls, highlighting karrikins' role in synchronizing recruitment pulses.^[29]^[30] In addition to seed germination, fire cues promote flowering in certain geophytes and annuals, facilitating immediate reproductive output and seed production in the open post-fire landscape. Heat or smoke exposure induces synchronized blooming, as seen in the fire-lily (Cyrtanthus ventricosus), a South African geophyte where smoke water treatment significantly increases flowering probability in bulbs, leading to prolific seed set within the first growing season after fire. Similarly, in Australian geophytes like Watsonia borbonica, smoke enhances inflorescence development, ensuring reproductive timing aligns with reduced competition and increased resource availability.^[31]^[32] Fire enhances seed dispersal mechanisms, further aiding recruitment by distributing propagules across burned areas. In California chaparral ecosystems, explosive fruit dehiscence in species like Acmispon glaber (previously Lotus scoparius) propels seeds up to several meters at maturity, with post-fire conditions—such as drier fuels and reduced canopy—amplifying dispersal distance and effectiveness. Wind-dispersed seeds, often lightweight and sometimes adhering to ash particles, benefit from fire-cleared skies and turbulent airflow; for example, in Salvia mellifera, ash-coated samaras travel farther post-fire, promoting wider colonization of suitable microsites.^[33]^[34] Demographic models illustrate how these fire-triggered strategies generate recruitment pulses that boost population density. In serotinous conifer systems, matrix population models predict that short fire-return intervals (6-31 years) can elevate post-fire seedling densities from 1,000 to over 100,000 individuals per hectare, driven by massive seed release and high establishment rates in the initial years. These pulses not only restore but often exceed pre-fire densities, underscoring fire's role as a demographic amplifier for resilient species.^[35]

Plants in Fire Regimes

Adaptation Matching to Fire Patterns

Fire regimes describe the recurring patterns of wildfires in an ecosystem, primarily characterized by their interval (the time between consecutive fires, often termed fire return interval or FRI), severity (the intensity and effects on vegetation, such as low-severity surface fires versus high-severity crown fires), and size (the spatial extent of individual fires), along with attributes like seasonality and predictability.^[36] These regimes vary markedly across biomes due to climatic, fuel, and ignition differences; for example, savannas typically experience frequent, low-intensity surface fires at annual or subannual intervals driven by grassy fuels and lightning, whereas boreal forests are shaped by infrequent, high-intensity crown fires occurring every 30–100 years or longer, fueled by accumulated woody biomass in cold, moist environments.^[36]^[37] Plant adaptations align with these regime characteristics to optimize survival and reproduction, but mismatches arise when fire patterns deviate from historical norms. In the fynbos shrublands of South Africa, serotinous species such as Proteaceae (e.g., Leucadendron and Protea spp.) are well-suited to short fire return intervals of 10–30 years, where heat from crown fires triggers the release of canopy-stored seeds into nutrient-rich, ash-bed environments for rapid establishment.^[38]^[39] However, when fires recur too frequently—such as intervals under 10 years—individuals fail to replenish seed banks before reburning, resulting in depleted recruitment, reduced population viability, and local extinctions of these non-resprouting species.^[40]^[39] Resource allocation trade-offs further illustrate adaptation matching, as plants balance investments in fire resistance (e.g., resprouting from lignotubers or epicormic buds for immediate post-fire recovery) against recruitment via seeds (e.g., building persistent banks for episodic germination).^[41] In regimes with short return intervals (e.g., 4–8 years), resprouting strategies predominate because they enable quick regrowth without relying on seedling establishment in competitive, frequently disturbed conditions, whereas longer intervals (e.g., >20 years) favor seeding to accumulate viable propagules over time.^[41] Conceptual models, such as the fire interval-adaptation curve, depict this dynamic: species fitness peaks at regime-specific frequencies, with resprouters declining at very short intervals due to energy costs of repeated recovery and seeders faltering at long intervals from competition with established vegetation.^[42]^[41] Case studies from Amazonian forests underscore how infrequent fire regimes select for recovery-oriented traits over resistance. In these moist tropical systems, where historical FRI exceeds centuries and fires are rare, canopy trees like those in Lecythidaceae and Myrtaceae exhibit limited fire resistance (e.g., thin bark vulnerable to lethal heating) but strong recovery potential through basal resprouting and vegetative cloning, allowing persistence after isolated events without the metabolic burden of chronic defenses.^[43]^[44] This contrasts with fire-prone biomes, where frequent burning would favor bark-thickening or serotiny instead. Post-2020 research reveals how climate change exacerbates adaptation mismatches by intensifying fire regimes beyond historical variability. In southwestern Australian shrublands, warming prolongs juvenile periods in serotinous species (e.g., Banksia spp.), heightening immaturity risk when shortened FRI depletes canopy seed stores before maturity, potentially shifting communities toward less diverse states.^[45] Similarly, in southeastern Australian eucalypt forests, altered seasonality and increased frequency threaten obligate-seeder Eucalyptus regnans by preventing seed production cycles, elevating collapse risks under novel drought-fire interactions.^[45] These shifts drive evolutionary pressures for regime-specific traits, though rapid alterations often outpace genetic adaptation, amplifying extinction vulnerabilities.^[46]

Ecological and Biodiversity Impacts

Fire-adapted plants play a crucial role in maintaining mosaic landscapes in fire-prone ecosystems by creating heterogeneous patches through varied post-fire recovery patterns, which in turn promote beta-diversity via patch dynamics.^[47] Diverse fire regimes, including differences in frequency and season, enhance environmental heterogeneity, leading to increased species turnover and beta-diversity across landscapes, as observed in South African Drakensberg grasslands where intermediate fire intervals maximized forb richness and patch variation.^[48] This patchiness prevents dominance by a single vegetation type and supports overall ecosystem resilience to disturbances.^[48] In biodiversity hotspots such as Mediterranean shrublands, recruitment strategies of fire-adapted plants, including serotiny and resprouting, drive post-fire species turnover and elevate community diversity.^[49] For instance, in chaparral ecosystems of California, plant diversity exhibits a hump-shaped relationship with fire severity, peaking at moderate levels where diverse recruitment enhances Simpson's diversity index, reflecting greater evenness among regenerating species.^[49] Similarly, in Spanish Garraf shrublands following the 1994 wildfire, Simpson's index tracked community composition recovery, initially increasing due to establishment of fire-responsive taxa before stabilizing, though drought can delay this turnover.^[50] Fire adaptations contribute to key ecosystem services in these landscapes, including accelerated nutrient cycling from the combustion of killed biomass, which rapidly mineralizes organic matter and releases nutrients like nitrogen and phosphorus into the soil for plant uptake.^[51] Prescribed fires, in particular, enhance this process by reducing excess fuels and promoting nutrient availability in grasslands and forests.^[52] Additionally, resprouting species bolster carbon storage by rapidly reallocating belowground reserves to aboveground regrowth, maintaining ecosystem carbon stocks even after canopy fires, as seen in eucalypt-dominated forests where resprouters resist losses compared to non-resprouters.^[53] Altered fire regimes, often intensified by climate change, can have negative impacts by favoring invasive non-adapted species that outcompete natives, disrupting biodiversity in fire-prone areas.^[54] In the US West, post-wildfire invasions by annual grasses like cheatgrass (Bromus tectorum) in sagebrush steppes exemplify this, as these invaders establish rapidly in disturbed patches, altering fuel continuity and perpetuating more frequent fires that hinder native recovery.^[55] Such invasions have converted native shrublands to grass-dominated systems across multiple ecoregions, reducing overall plant diversity.^[56] Interactions between fire-adapted plants and soil microbes further influence ecosystem recovery, with fire-tolerant mycorrhizae playing a vital role in facilitating plant establishment post-fire.^[57] Ectomycorrhizal fungal spore banks, for example, survive severe fires and dominate seedling colonization, as demonstrated in California pine forests after the 2013 Rim Fire, where spore bank richness decreased from 31.8 to 20.0 taxa but the community composition remained largely intact (Pearson r=0.88), aiding rapid regeneration without external inoculum.^[57] These microbes enhance nutrient uptake for host plants, supporting biodiversity restoration in burned soils. Long-term, such plant-microbe dynamics contribute to stable habitats for animals by preserving vegetation mosaics essential for foraging and shelter.

Evolution of Fire Traits in Plants

Fossil and Historical Evidence

The earliest evidence of fire in Earth's history appears in the fossil record during the Late Devonian period, approximately 370 million years ago, marked by the presence of fossil charcoal (fusain) in sedimentary deposits such as those from the Catskill Formation in Pennsylvania. These charcoalified remains, primarily from early vascular plants like Rhacophyton, indicate that wildfires were occurring in seasonally dry environments, with anatomical preservation suggesting combustion under low-oxygen conditions typical of the time. Concurrently, the development of periderm—a bark-like tissue in Devonian plants such as Archaeopteris—provided insulation against desiccation and likely offered early protection during surface fires, representing a foundational adaptation in terrestrial vegetation.^[58]^[59] During the Cretaceous period, around 100 million years ago, the expansion of angiosperms coincided with increasing fire activity, as evidenced by fossil charcoal and resin deposits that point to more frequent wildfires in a warming, oxygen-rich atmosphere. While serotiny— the retention of seeds in fire-resistant cones until triggered by heat—was well-established in gymnosperms, these adaptations, including thick bark and serotinous cones in co-occurring conifers, facilitated post-fire recruitment in polar and mid-latitude ecosystems during the global hothouse climate.^[60]^[61]^[62] Following the Cretaceous-Paleogene extinction, fire-adapted traits diversified rapidly in Gondwanan floras, particularly in Australia, where Eocene sediments (around 50 million years ago) preserve evidence of resprouting mechanisms in early eucalypt relatives (Myrtaceae) and other scleromorphic plants, reflecting adaptation to recurrent fires in increasingly seasonal, aridifying environments as Gondwana fragmented, enabling the persistence of fire-prone vegetation across southern continents.^[63]^[64] Historical records from the 1800s onward document how European colonization and subsequent fire suppression policies altered plant trait frequencies in fire-dependent ecosystems, such as Mediterranean woodlands and North American prairies, favoring shade-tolerant, non-sprouting species over fire-adapted resprouters and serotinous plants. This anthropogenic shift reduced fire return intervals from decades to centuries, leading to adaptation lags where populations of fire-dependent taxa declined due to insufficient selective pressure, as seen in pollen and charcoal profiles from European lake sediments showing decreased abundance of pyrophytic pollen types since the Industrial Revolution.^[65]^[66] Charcoal accumulation rates in lake sediments serve as reliable proxies for reconstructing ancient fire regimes, with Miocene records (23–5 million years ago) from sites in the Black Sea region (eastern Eurasia) and the western United States indicating 1–5 fires per century in grassland-forest mosaics, based on peaks in charcoal influx corresponding to intervals of 20–100 years. These data, derived from quantitative analyses of macroscopic charcoal particles, highlight how elevated fire frequencies during the Miocene drove the expansion of C4 grasslands and associated adaptations like enhanced resprouting, providing a timeline for the co-evolution of vegetation and fire. Recent phylogenetic studies (as of 2023) confirm multiple independent origins of key fire traits like serotiny and resprouting since the Paleogene.^[67]^[68]^[1]

Genetic and Selective Processes

Fire acts as a potent abiotic stressor in fire-prone ecosystems, imposing strong natural selection pressures that favor traits enhancing post-fire survival and reproduction in plants. Recurrent fires drive the evolution of protective structures and regenerative mechanisms, with empirical evidence showing that populations exposed to frequent, high-intensity fires exhibit higher frequencies of adaptive alleles compared to those in low-fire environments. Heritability estimates for key traits like bark thickness, which insulates vascular tissues from lethal heat, often exceed 0.5 in field studies of fire-adapted species, underscoring the genetic potential for rapid evolutionary responses to changing fire regimes.^[1] At the molecular level, fire adaptations are underpinned by specific genes, such as the KAI2 gene encoding karrikin receptors that perceive smoke-derived butenolides to promote germination in fire-following species. Activation of KAI2 signaling reduces seed dormancy and enhances seedling establishment in post-fire environments, a mechanism conserved across diverse taxa from Arabidopsis to monocots and eudicots, reflecting an ancient evolutionary origin in seed plants predating frequent fire exposure. This conservation suggests that co-option of existing developmental pathways enabled the fine-tuning of fire-specific responses.^[69]^[70] Polyploidy and interspecific hybridization further accelerate trait evolution by generating novel genetic combinations, particularly in fire-prone lineages like the Proteaceae family. In genera such as Adenanthos and Banksia, hybridization events have introduced allelic diversity, facilitating the spread of fire-resilient traits like serotiny and resprouting amid fluctuating fire intervals. These processes provide raw material for selection, enabling quicker adaptation than mutation alone in dynamic ecosystems.^[71] Convergent evolution illustrates how analogous fire pressures yield similar traits across distantly related lineages; for instance, serotiny—the retention of seeds in fire-released structures—has independently arisen in Pinus (conifers) and Banksia (Proteaceae) through parallel mutations affecting tissue lignification and heat sensitivity. Recent genomic analyses, including 2020s GWAS on Eucalyptus species, have pinpointed QTLs for resprouting capacity, accounting for 3.5–20% of trait variation and revealing pleiotropic loci that integrate growth, damage resistance, and recovery under fire stress.^[72]^[73]

Animal Adaptations to Fire

Behavioral Responses

Animals exhibit a range of behavioral responses to fire that enable them to evade immediate threats from heat, flames, and smoke. Flight reflexes are common among mobile species, allowing rapid escape to safer areas. For instance, kangaroos in Australian bushlands detect smoke plumes and flee toward water bodies or previously burned areas, often moving in groups to increase survival odds during intense wildfires.^[74] Similarly, many terrestrial taxa, including mammals and reptiles, display instinctive fleeing behaviors triggered by sensory cues such as heat and smoke, which can initiate movements tens to hundreds of kilometers from the fire front.^[75] These responses are often modulated by prior experience with fire regimes, enhancing escape efficiency in fire-prone habitats.^[76] Burrowing and shelter-seeking behaviors provide refuge for less mobile animals during fire passage. Certain lizards quickly seek cracks in the soil or deep burrows to avoid lethal surface temperatures, a reflex triggered by thermal and olfactory cues from approaching flames.^[77] This behavior minimizes direct mortality, as evidenced by low post-fire lizard carcasses in burned areas, indicating successful evasion through rapid sheltering.^[78] In savanna ecosystems, ungulates like impalas (Aepyceros melampus) and zebras (Equus quagga) shift foraging patterns pre-fire in response to early smoke detection or heterospecific alarm signals, such as bird calls signaling danger, allowing them to relocate to less flammable grasslands ahead of the blaze.^[76] Post-fire, opportunistic scavenging emerges as a key behavioral adaptation to exploit newly available resources. Corvids, particularly common ravens (Corvus corax), increase group foraging activities on carrion from fire-killed animals, with feeding bouts extending longer in burned landscapes due to abundant, unburied carcasses.^[79] This social scavenging enhances detection and defense of food patches, boosting raven survival and reproduction in the immediate aftermath.^[80] Fire events also prompt migratory adjustments in avian species to avoid smoke inhalation and disorientation. Radar data from western North American flyways reveal that songbirds and waterfowl, such as Tule white-fronted geese (Anser albifrons), deviate from standard routes when encountering smoke plumes, resulting in paths extended by over 25% and durations doubled, with some individuals traveling an additional 470 miles to circumvent hazardous areas.^[81]^[82] Parental care behaviors during fires prioritize offspring protection, often at risk to adults. Female marsupials, including red kangaroos (Osphranter rufus), carry dependent joeys in their pouches while fleeing, shielding them from radiant heat and embers as they bound to safety zones like riverbanks.^[83] This instinctive transport sustains joey survival rates, though mothers may succumb if unable to outpace the fire front.^[84]

Physiological and Morphological Traits

Animals in fire-prone environments have evolved physiological and morphological traits that mitigate the effects of high temperatures, smoke, and post-fire conditions. These traits include enhanced thermal tolerance, where species can withstand body temperatures up to approximately 50°C during brief exposure to fire heat, preventing immediate lethality from radiant or convective heat.^[85] For example, small mammals like rodents exhibit morphological adaptations such as thick pelage that provides insulation, reducing the depth of burns from surface fires. Similarly, mammals like armadillos possess bony scales that offer physical protection, with their armored exoskeleton and burrowing behavior aiding survival in grassland fires by providing refuge from heat.^[85]^[86] Burrow-dwelling morphology is a key trait in many rodents, with reinforced tunnel systems that insulate against high surface temperatures during fires. Pocket gophers (Thomomys bottae) in California grasslands construct deep, multi-chambered burrows with compacted soil walls that maintain internal temperatures below lethal thresholds, enabling high survival rates even in intense wildfires.^[85] These structures often include ventilation shafts that facilitate gas exchange in smoky conditions. Sensory adaptations further enhance post-fire survival; some snakes, such as pit vipers (Crotalus spp.), possess infrared-sensitive pit organs that detect thermal signatures of prey from up to 1 meter away, allowing efficient foraging in ash-covered landscapes where visual cues are obscured by smoke or debris.^[87] Invertebrates also show notable fire adaptations. For example, some insects exhibit post-fire melanism for camouflage in charred landscapes, while pyrophilous beetles like Melanophila spp. have infrared-sensing organs to detect distant fires and locate fresh breeding sites.^[1] These traits collectively enable animals to endure and exploit fire-disturbed habitats, though their effectiveness depends on fire intensity and individual condition.

Animals in Fire-Affected Ecosystems

Immediate Fire Impacts

Wildfires impose severe immediate threats to animal individuals and small groups through direct physical harm, primarily via burns, smoke inhalation, and dehydration. Direct mortality rates vary by species and fire intensity, but systematic reviews indicate that overall animal mortality during fires averages around 3% (95% CI: 1-9%), with higher proportions in high-severity events where heat and smoke overwhelm escape capabilities. For small mammals, such as arboreal marsupials in crown fires, predicted mortality from partial and full-thickness burns combined with asphyxiation can reach 62-79%, as modeled in simulations of intense blazes. Ground-dwelling small mammals experience lower direct mortality from burns due to burrowing, but smoke inhalation remains a significant killer, contributing to population declines immediately post-fire. Dehydration exacerbates these effects in arid environments, where heat stress compounds fluid loss during evasion. Habitat destruction from wildfires triggers rapid displacement of animals, forcing them to seek refuge in unburned patches and increasing vulnerability to secondary threats. Understory vegetation, critical for nesting and cover, is often obliterated, leading to nest failure rates as high as 62.8% for scorched cavities in cavity-nesting birds, with broader impacts on understory species where up to 48.6% of potential sites become unusable. This immediate loss compels birds and small mammals to relocate, disrupting foraging and breeding, and in severe cases, resulting in mass exodus from burned areas. Post-fire, predation rates surge as flames and smoke disorient prey, exposing them to opportunistic hunters. In Australian ecosystems, predators like raptors and feral cats flock to burned zones to exploit fleeing or stunned animals, with studies documenting dramatic increases in predator activity targeting disoriented avifauna, including ground birds such as quails that become easy targets amid reduced cover. This spike can decimate local prey populations in the hours and days following the blaze. Surviving animals endure acute physiological stress, evidenced by elevated cortisol levels that signal heightened fight-or-flight responses. In the 2019-2020 Australian bushfires, rescued koalas exhibited significantly higher stress hormones compared to unburned counterparts, with limited data indicating persistent elevation for weeks post-rescue, impairing immune function and recovery. Similar patterns occur in other wildlife, where wildfire exposure triggers cortisol surges that correlate with dehydration and injury. The lethality of immediate fire impacts varies markedly by fire type, with surface fires posing lower risks to ground-dwelling animals than crown fires. Surface fires, which burn low-lying vegetation, allow many burrowers and understory species to shelter underground with minimal heat penetration, resulting in limited direct mortality. In contrast, crown fires generate intense radiant heat, widespread smoke, and embers that penetrate refuges, elevating mortality for the same ground-dwellers through asphyxiation and burns. Behavioral evasions, such as fleeing to water sources, can mitigate some of these effects in less intense fires.

Long-Term Community Effects

Repeated fires in fire-prone ecosystems can lead to long-term shifts in animal community composition, often favoring species that are tolerant or opportunistic in altered post-fire landscapes. For instance, fire-tolerant taxa, such as certain small mammals and birds adapted to early-successional stages, tend to increase in abundance over time as vegetation recovers, while sensitive species decline. This shift is particularly evident in granivorous animals, which benefit from post-fire seed pulses released from serotinous cones or soil seed banks; studies in pine forests show that granivores like rodents exhibit heightened seed removal rates in moderately burned areas, enhancing their population growth for several years following fire events.^[88] Such changes in species abundance can persist across multiple fire cycles, reducing overall community diversity and promoting dominance by generalist or fire-resilient groups.^[89] Burn patches from recurrent fires contribute to habitat fragmentation, which disrupts animal dispersal and connectivity, often resulting in range contractions for less mobile species. Fire creates a mosaic of burned and unburned areas, isolating habitat remnants and increasing edge effects that alter microclimates and resource availability; connectivity models indicate that this fragmentation can reduce effective dispersal distances by 20-40% in fragmented landscapes, limiting gene flow and exacerbating local extinctions. In particular, species reliant on contiguous habitats, such as forest-dwelling mammals, experience constrained movement across burn boundaries, leading to population isolation over decades. These effects compound with ongoing fire regimes, altering long-term community structure by favoring species with high dispersal capabilities. Trophic cascades in fire-affected ecosystems can arise from herbivore declines due to reduced forage availability post-fire, subsequently impacting predators through diminished prey resources and potential starvation risks. In Yellowstone National Park, the 1988 fires altered vegetation dynamics, contributing to shifts in elk (Cervus elaphus) foraging patterns and abundance; combined with subsequent wolf (Canis lupus) predation after their 1995 reintroduction, this led to a trophic cascade where elk declines reduced browsing pressure on aspen (Populus tremuloides), indirectly benefiting vegetation recovery while straining predator populations during periods of low herbivore density. Such cascades illustrate how fire-induced bottom-up effects on herbivores can propagate upward, destabilizing predator-prey balances over generations.^[90]^[91] Stressed animal populations in repeatedly burned areas are more susceptible to disease outbreaks, exacerbated by changes in disease vectors and hosts following fire.^[92] Interactions between climate change and altered fire regimes intensify these community effects, driving projected declines in animal biodiversity through more frequent and severe burns. Models from the 2020s forecast that warming temperatures will expand fire-prone areas, exacerbating habitat loss and species declines; in Australia, projections indicate substantial biodiversity loss in bush ecosystems by mid-century under moderate emissions scenarios, as intensified fire intervals outpace recovery for fire-sensitive fauna. These climate-fire synergies amplify fragmentation and trophic disruptions, threatening long-term community stability across continents.^[93]^[94]

Interactions Between Animals and Fire Regimes

Habitat and Foraging Dynamics

Fire regimes profoundly influence habitat structure and resource availability, creating dynamic opportunities for animal foraging in post-fire environments. Following wildfires, nutrient-rich ash deposits enhance soil fertility by recycling minerals such as phosphorus, potassium, and calcium, leading to temporary surges in plant productivity that support increased food resources for herbivores and their predators. This nutrient pulse often triggers foraging booms, particularly through insect irruptions in the early post-fire period, where herbivorous insects proliferate on succulent regrowth, attracting insectivorous birds and mammals; for instance, bark beetle abundance can drive a five-fold increase in hairy woodpecker densities in burned conifer forests. Such surges in insect populations exemplify how fire-induced resource pulses facilitate opportunistic foraging by species like ground-foraging birds and small mammals.^[95] Animals frequently exhibit shifts in habitat selection toward early-successional burns, which provide enhanced cover and forage amid reduced canopy competition. In fire-prone landscapes, ungulates such as white-tailed deer demonstrate preferential use of recently burned patches for their abundance of palatable grasses and forbs, as evidenced by GPS collar data showing concentrated movements into low-severity burn areas shortly after fire. Mule deer, for example, expand home ranges post-megafire and may favor open, regenerating habitats in some contexts, though immediate responses to severe fires can involve avoidance of heavily burned areas while selecting for older disturbances (6–15 years post-fire) over unburned mature forests to exploit emergent vegetation while balancing predation risks. These behavioral adjustments underscore fire's role in resetting habitat mosaics, enabling species to track optimal conditions for cover and nutrition in heterogeneous post-fire terrains.^[96]^[97] Seasonal alignment with fire cycles further optimizes foraging dynamics for mobile species in fire-dependent ecosystems. Nomadic birds in African savannas often track post-fire green-up patches, where fresh grass flushes attract insect and seed resources during dry seasons, though specific timing varies by species. This opportunistic tracking of fire mosaics—maintained by frequent burns—allows these species to exploit transient productivity pulses, synchronizing migrations or nomadism with the spatiotemporal variability of burned landscapes. Such adaptations highlight how fire regimes structure seasonal resource landscapes, promoting efficient energy acquisition across expansive grassland systems.^[98] Competition dynamics intensify during post-fire resource pulses, where fire-adapted species gain advantages in exploiting ephemeral abundances. In regenerating habitats, generalist small mammals like deer mice experience competitive release from displaced competitors, leading to population expansions and shifts in community composition as they dominate seed and insect resources. Fire-adapted taxa, such as certain rodents and birds, outcompete less resilient species by rapidly colonizing burns, leveraging enhanced mobility and dietary flexibility to capitalize on nutrient flushes before succession closes opportunities. These interactions reinforce the role of fire in maintaining biodiversity through pulsed competition, favoring species attuned to disturbance-driven variability.^[99]^[100] Human-altered fire regimes, particularly exclusion policies, disrupt these dynamics by homogenizing habitats and favoring generalist over specialist species. Fire suppression in ecosystems like Rocky Mountain forests reduces early-successional patches critical for fire-dependent animals, allowing dense overstories to suppress understory growth and diminish resource pulses that specialists rely on for foraging. Consequently, generalists proliferate in the altered, less disturbed landscapes, while specialists—such as certain cavity-nesting birds or burn-preferring herbivores—decline due to lost habitat heterogeneity and competitive disadvantages. This shift exemplifies how anthropogenic interventions can erode the adaptive foraging strategies evolved under natural fire cycles.^[101]^[102]

Population and Evolutionary Responses

Fire regimes profoundly influence animal population dynamics, often synchronizing fluctuations with fire return intervals in fire-prone ecosystems. In arid regions like central Australia, rodent populations exhibit boom-bust cycles driven by post-fire resource pulses and rainfall, where densities can surge within 1–2 years due to enhanced vegetation regrowth and reduced competition, before declining as habitats mature and predators recover. These cycles, observed in species such as the spinifex hopping mouse (Notomys alexis), highlight how frequent fires (every 3–5 years) maintain low baseline densities punctuated by rapid increases, preventing long-term overpopulation while promoting resilience to disturbance.^[103] Mass mortalities during intense wildfires can create genetic bottlenecks, drastically reducing population diversity and increasing vulnerability to future stressors. For instance, koala (Phascolarctos cinereus) populations in southeastern Australia experienced severe declines during the 2019–2020 bushfires, with up to 80% mortality in affected areas leading to localized losses in genetic diversity, including elevated inbreeding coefficients and reduced heterozygosity in surviving groups. Genomic analyses post-fires revealed runs of homozygosity indicating recent inbreeding, underscoring the risk of diminished adaptive potential in fragmented remnants.^[104] Natural selection under recurrent fire exposure favors traits enhancing survival, such as improved escape capabilities in response to disturbance cues. In fire-adapted species like the Australian frilled lizard (Chlamydosaurus kingii), populations in high-fire regimes show evolved behavioral traits for rapid evasion, including faster sprint speeds and heightened sensitivity to smoke or heat, selected over generations to minimize mortality during blazes. Similarly, studies on lizards in fire-maintained savannas demonstrate directional selection for locomotor performance, where individuals with superior escape abilities contribute disproportionately to post-fire reproduction.^[105] Metapopulation dynamics in heterogeneous burn mosaics sustain gene flow among animal subpopulations, buffering against local extinctions. In fire-prone landscapes, unburned patches act as refugia, facilitating dispersal and recolonization that maintain genetic connectivity; for example, small mammals like the agile antechinus (Antechinus agilis) in Australian forests exhibit gene flow across burn patches, with effective migration rates preventing isolation in fragmented habitats. This mosaic structure, shaped by variable fire severity, promotes overall metapopulation persistence despite periodic local crashes.^[106] Recent studies from the 2020s provide evidence of rapid evolutionary responses to environmental stressors, including those compounded by wildfires and urban heat. These shifts illustrate how intensifying interactions between fire and human-modified landscapes can accelerate trait evolution, potentially aiding resilience in altered ecosystems.^[107]

Animal Use of Fire

Observed Behaviors Across Species

One of the most compelling examples of animals intentionally spreading fire for ecological benefits is observed in Australian "firehawk" raptors, including the black kite (Milvus migrans), whistling kite (Haliastur sphenurus), and brown falcon (Falco berigora). These birds have been documented carrying burning sticks, twigs, or embers in their beaks or talons from active wildfires and dropping them into unburned vegetation up to a kilometer away, thereby extending the fire front to flush out hidden prey such as small mammals, reptiles, insects, and birds.^[108] This behavior enables the raptors to hunt more effectively by creating chaos that drives prey into the open, where the birds can swoop down to capture it.^[108] Ethnographic evidence from Australian Aboriginal communities provides the primary documentation of this practice, with at least 12 distinct Indigenous groups in northern Australia reporting firsthand observations of fire-spreading by these raptors, often linking it to cultural narratives such as Dreaming stories.^[108] Non-Indigenous eyewitness accounts, including those from firefighters and researchers, corroborate these reports, describing flocks of firehawks working individually or cooperatively to ignite new patches of dry grass during wildfires.^[108] Although video footage capturing the act has proven elusive despite targeted efforts, the consistency across oral histories and direct sightings collected since the 1960s supports the intentional nature of the behavior.^[109] Such fire-spreading occurs predominantly in opportunistic contexts during Australia's dry season (May to October), when natural wildfires are frequent in arid and savanna landscapes, allowing the raptors to exploit existing blazes without initiating them from scratch.^[108] While primarily documented in these avian species, anecdotal reports suggest similar tactics in other birds, though these lack empirical verification and are not habitual.^[110] In contrast, no confirmed instances of intentional fire use exist for mammals, including debated claims of primates like chimpanzees employing fire to flush prey in African savannas, which remain unverified in wild settings.^[110]

Ecological and Evolutionary Roles

Certain raptors, such as the black kite (Milvus migrans), whistling kite (Haliastur sphenurus), and brown falcon (Falco berigora), exhibit fire-foraging behaviors in fire-prone savannas of northern Australia, where they intentionally spread wildfires by carrying burning sticks in their talons or beaks to ignite new areas, often across firebreaks.^[111] This "firehawk" behavior, documented through Indigenous ecological knowledge and direct observations from 2011 to 2017, allows these birds to flush out small mammals, reptiles, and insects for easier capture, enhancing their hunting efficiency during active burns.^[111] Similar pyric-carnivory—carnivory facilitated by fire—occurs in other raptor species, like Swainson's hawk (Buteo swainsoni), which aggregate at prescribed fires in North American grasslands, with abundances increasing up to sevenfold as prey becomes exposed or injured.^[112] Ecologically, these behaviors play a pivotal role in shaping fire regimes and community dynamics within ecosystems. By propagating fires, fire-foraging raptors contribute to pyrodiversity—the spatial and temporal variation in fire patterns—which maintains habitat heterogeneity essential for biodiversity in savannas and grasslands.^[113] This process influences predator-prey interactions by temporarily boosting prey vulnerability, potentially regulating herbivore populations and promoting nutrient cycling through post-fire vegetation regrowth that supports diverse foraging opportunities.^[112] In managed landscapes, such as those in northern Australia, raptor-induced fire spread complicates suppression efforts but also aids in fuel load reduction, indirectly benefiting ecosystem resilience against megafires.^[111] Overall, these interactions position raptors as keystone actors in fire-mediated trophic cascades, where their foraging amplifies fire's role in sustaining open habitats favored by many species.^[113] From an evolutionary perspective, fire-foraging represents a behavioral adaptation honed in pyrogenic environments, where fire acts as a selective force driving trait evolution in animals.^[113] In raptors, sensitivity to fire cues like smoke plumes has likely evolved to exploit ephemeral foraging boons, with fire-spreading behaviors suggesting cognitive advancements in tool use that parallel early human fire manipulation and may trace back to ancestral responses in fire-prone biomes.^[111] Such adaptations contribute to broader evolutionary patterns in the "Pyrocene," an era of intensified human-altered fire regimes, where rapid selection for fire-resilient traits—like enhanced dispersal or physiological tolerance—occurs across taxa, potentially influencing gene flow and speciation in fire-dependent communities.^[113] For instance, the persistence of these behaviors in Australian raptors underscores co-evolutionary dynamics between animals and fire, fostering lineages specialized for recurrent disturbances.^[111]

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Nov 18, 2024 · In forest management settings, disturbance resets forests to earlier successional stages, typically improving forage conditions for mule ...
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Fire mosaics build up in regions with low cost of access to water, high intrinsic food availability, and good access to trade opportunities.
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Effects of Controlled Fire and Livestock Grazing on Bird ...
Aug 7, 2025 · We investigated the effects of prescribed fires and bomas on savanna bird communities in East Africa during the first and second dry seasons of ...
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Competitive release during fire succession influences ecological ...
One possibility is competitive release, which predicts a reduction in vole competition may contribute to niche expansion and population growth in deermice.
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Small mammal responses to fire severity mediated by vegetation ...
May 19, 2022 · In particular, small mammal communities are vulnerable to post‐fire shifts in resource availability and play critical roles in forest ecosystems ...
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[PDF] Cascading Effects of Fire Exclusion in Rocky Mountain Ecosystem
The health of many Rocky Mountain ecosystems is in decline because of the policy of excluding fire in the management of these ecosystems. Fire exclusion has ...
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Is Response to Fire Influenced by Dietary Specialization and ...
Feb 7, 2014 · In a general way, the disturbance theory states that specialist species are negatively affected by disturbance, while generalist species ...
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Genomics identifies koala populations at risk across eastern Australia
Nov 29, 2024 · We identified several regions of the continent where koalas have low genomic diversity and high inbreeding, as measured by runs of homozygosity.
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Understanding the genetic implications of fire regimes for fauna and ...
This review confirms that fire regimes impact the genetic patterns of fauna at an individual, population and meta-population scale.
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Continent‐wide parallel urban evolution of increased heat tolerance ...
Dec 26, 2023 · Our results indicate widespread urban evolution of increased heat tolerance in the adult stage only, suggesting that the UHI may be a stronger selective agent ...Missing: fire interfaces 2020s
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Intentional Fire-Spreading by “Firehawk” Raptors in Northern Australia
Dec 1, 2017 · In Australia, IEK of fire-spreading by the firehawk is often related to Dreaming fire ceremonies such as Lorrkon (Garde et al. 2009) and ...
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Feb 5, 2018 · A recent paper published in Journal of Ethnobiology gathers reports that all three species do spread wildfires for hunting purposes.
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Nov 25, 2021 · ... chimpanzees. Indeed, whoever and whenever and wherever they were ... Using fire also carried the risk of smoke inhalation, especially ...
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