Evolutionary fire ecology: An historical account and future directions.

The idea that fire acts as an evolutionary force contributing to shaping species traits started a century ago, but had not been widely recognized until very recently. Among the first to realize this force were Edward B. Poulton, R. Dale Guthrie, and Edwin V. Komarek in animals and Willis L. Jepson, Walter W. Hough, Tom M. Harris, Philip V. Wells, and Robert W. Mutch in plants. They were all ahead of their time in their evolutionary thinking. Since then, evolutionary fire ecology has percolated very slowly into the mainstream ecology and evolutionary biology; in fact, this topic is still seldom mentioned in textbooks of ecology or evolution. Currently, there is plenty of evidence suggesting that we cannot understand the biodiversity of our planet without considering the key evolutionary role of fire. But there is still research to be done in order to fully understand fire's contribution to species evolution and to predicting species responses to rapid global changes.

Fire-driven vegetation type conversion in Southern California.

One consequence of global change causing widespread concern is the possibility of ecosystem conversions from one type to another. A classic example of this is vegetation type conversion (VTC) from native woody shrublands to invasive annual grasslands in the biodiversity hotspot of Southern California. Although the significance of this problem is well recognized, understanding where, how much, and why this change is occurring remains elusive owing to differences in results from studies conducted using different methods, spatial extents, and scales. Disagreement has arisen particularly over the relative importance of short-interval fires in driving these changes. Chronosequence approaches that use space for time to estimate changes have produced different results than studies of changes at a site over time. Here we calculated the percentage woody and herbaceous cover across Southern California using air photos from ~1950 to 2019. We assessed the extent of woody cover change and the relative importance of fire history, topography, soil moisture, and distance to human infrastructure in explaining change across a hierarchy of spatial extents and regions. We found substantial net decline in woody cover and expansion of herbaceous vegetation across all regions, but the most dramatic changes occurred in the northern interior and southern coastal areas. Variables related to frequent, short-interval fire were consistently top ranked as the explanation for shrub to grassland type conversion, but low soil moisture and topographic complexity were also strong correlates. Despite the consistent importance of fire, there was substantial geographical variation in the relative importance of drivers, and these differences resulted in different mapped predictions of VTC. This geographical variation is important to recognize for management decision-making and, in addition to differences in methodological design, may also partly explain differences in previous study results. The overwhelming importance of short-interval fire has management implications. It suggests that actions should be directed away from imposing fires to preventing fires. Prevention can be controlled through management actions that limit ignitions, fire spread, and the damage sustained in areas that do burn. This study also demonstrates significant potential for changing fire regimes to drive large-scale, abrupt ecological change.

Effects of postfire climate and seed availability on postfire conifer regeneration.

Large, severe fires are becoming more frequent in many forest types across the western United States and have resulted in tree mortality across tens of thousands of hectares. Conifer regeneration in these areas is limited because seeds must travel long distances to reach the interior of large burned patches and establishment is jeopardized by increasingly hot and dry conditions. To better inform postfire management in low elevation forests of California, USA, we collected 5-yr postfire recovery data from 1,234 study plots in 19 wildfires that burned from 2004-2012 and 18 yrs of seed production data from 216 seed fall traps (1999-2017). We used these data in conjunction with spatially extensive climate, topography, forest composition, and burn severity surfaces to construct taxon-specific, spatially explicit models of conifer regeneration that incorporate climate conditions and seed availability during postfire recovery windows. We found that after accounting for other predictors both postfire and historical precipitation were strong predictors of regeneration, suggesting that both direct effects of postfire moisture conditions and biological inertia from historical climate may play a role in regeneration. Alternatively, postfire regeneration may simply be driven by postfire climate and apparent relationships with historical climate could be spurious. The estimated sensitivity of regeneration to postfire seed availability was strongest in firs and all conifers combined and weaker in pines. Seed production exhibited high temporal variability with seed production varying by over two orders of magnitude among years. Our models indicate that during droughts postfire conifer regeneration declines most substantially in low-to-moderate elevation forests. These findings enhance our mechanistic understanding of forecasted and historically documented shifts in the distribution of trees.

Subspecies differentiation in an enigmatic chaparral shrub species.

PREMISE: Delimiting biodiversity units is difficult in organisms in which differentiation is obscured by hybridization, plasticity, and other factors that blur phenotypic boundaries. Such work is more complicated when the focal units are subspecies, the definition of which has not been broadly explored in the era of modern genetic methods. Eastwood manzanita (Arctostaphylos glandulosa Eastw.) is a widely distributed and morphologically complex chaparral shrub species with much subspecific variation, which has proven challenging to categorize. Currently 10 subspecies are recognized, however, many of them are not geographically segregated, and morphological intermediates are common. Subspecies delimitation is of particular importance in this species because two of the subspecies are rare. The goal of this study was to apply an evolutionary definition of "subspecies" to characterize structure within Eastwood manzanita. METHODS: We used publicly available geospatial environmental data and reduced-representation genome sequencing to characterize environmental and genetic differentiation among subspecies. In addition, we tested whether subspecies could be differentiated by environmentally associated genetic variation. RESULTS: Our analyses do not show genetic differentiation among subspecies of Eastwood manzanita, with the exception of one of the two rare subspecies. In addition, our environmental analyses did not show ecological differentiation, though limitations of the analysis prevent strong conclusions. CONCLUSIONS: Genetic structure within Eastwood manzanita does not correspond to current subspecies circumscriptions, but rather reflects geographic distribution. Our study suggests that subspecies concepts need to be reconsidered in long-lived plant species, especially in the age of next-generation sequencing.

Human presence diminishes the importance of climate in driving fire activity across the United States.

Growing human and ecological costs due to increasing wildfire are an urgent concern in policy and management, particularly given projections of worsening fire conditions under climate change. Thus, understanding the relationship between climatic variation and fire activity is a critically important scientific question. Different factors limit fire behavior in different places and times, but most fire-climate analyses are conducted across broad spatial extents that mask geographical variation. This could result in overly broad or inappropriate management and policy decisions that neglect to account for regionally specific or other important factors driving fire activity. We developed statistical models relating seasonal temperature and precipitation variables to historical annual fire activity for 37 different regions across the continental United States and asked whether and how fire-climate relationships vary geographically, and why climate is more important in some regions than in others. Climatic variation played a significant role in explaining annual fire activity in some regions, but the relative importance of seasonal temperature or precipitation, in addition to the overall importance of climate, varied substantially depending on geographical context. Human presence was the primary reason that climate explained less fire activity in some regions than in others. That is, where human presence was more prominent, climate was less important. This means that humans may not only influence fire regimes but their presence can actually override, or swamp out, the effect of climate. Thus, geographical context as well as human influence should be considered alongside climate in national wildfire policy and management.

Towards understanding resprouting at the global scale.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Evolutionary ecology of resprouting and seeding in fire-prone ecosystems.

There are two broad mechanisms by which plant populations persist under recurrent disturbances: resprouting from surviving tissues, and seedling recruitment. Species can have one of these mechanisms or both. However, a coherent framework explaining the differential evolutionary pressures driving these regeneration mechanisms is lacking. We propose a bottom-up approach in addressing this question that considers the relative survivorship of adults and juveniles in an evolutionary context, based on two assumptions. First, resprouting and seeding can be interpreted by analogy with annual versus perennial life histories; that is, if we consider disturbance cycles to be analogous to annual cycles, then resprouting species are analogous to the perennial life history with iteroparous reproduction, and obligate seeding species that survive disturbances solely through seed banks are analogous to the annual life history with semelparous reproduction. Secondly, changes in the selective regimes differentially modify the survival rates of adults and juveniles and thus the relative costs and benefits of resprouting versus seeding. Our approach provides a framework for understanding temporal and spatial variation in resprouting and seeding under crown-fire regimes. It accounts for patterns of coexistence and environmental changes that contribute to the evolution of seeding from resprouting ancestors.

Fire-driven alien invasion in a fire-adapted ecosystem.

Disturbance plays a key role in many alien plant invasions. However, often the main driver of invasion is not disturbance per se but alterations in the disturbance regime. In some fire-adapted shrublands, the community is highly resilient to infrequent, high-intensity fires, but changes in the fire regime that result in shorter fire intervals may make these communities more susceptible to alien plant invasions. This study examines several wildfire events that resulted in short fire intervals in California chaparral shrublands. In one study, we compared postfire recovery patterns in sites with different prefire stand ages (3 and 24 years), and in another study we compared sites that had burned once in four years with sites that had burned twice in this period. The population size of the dominant native shrub Adenostoma fasciculatum was drastically reduced following fire in the 3-year sites relative to the 24-year sites. The 3-year sites had much greater alien plant cover and significantly lower plant diversity than the 24-year sites. In a separate study, repeat fires four years apart on the same sites showed that annual species increased significantly after the second fire, and alien annuals far outnumbered native annuals. Aliens included both annual grasses and annual forbs and were negatively correlated with woody plant cover. Native woody species regenerated well after the first fire but declined after the second fire, and one obligate seeding shrub was extirpated from two sites by the repeat fires. It is concluded that some fire-adapted shrublands are vulnerable to changes in fire regime, and this can lead to a loss of native diversity and put the community on a trajectory towards type conversion from a woody to an herbaceous system. Such changes result in alterations in the proportion of natives to non-natives, changes in functional types from deeply rooted shrubs to shallow rooted grasses and forbs, increased fire frequency due to the increase in fine fuels, and changes in carbon storage.

The impact of antecedent fire area on burned area in southern California coastal ecosystems.

Frequent wildfire disasters in southern California highlight the need for risk reduction strategies for the region, of which fuel reduction via prescribed burning is one option. However, there is no consensus about the effectiveness of prescribed fire in reducing the area of wildfire. Here, we use 29 years of historical fire mapping to quantify the relationship between annual wildfire area and antecedent fire area in predominantly shrub and grassland fuels in seven southern California counties, controlling for annual variation in weather patterns. This method has been used elsewhere to measure leverage: the reduction in wildfire area resulting from one unit of prescribed fire treatment. We found little evidence for a leverage effect (leverage = zero). Specifically our results showed no evidence that wildfire area was negatively influenced by previous fires, and only weak relationships with weather variables rainfall and Santa Ana wind occurrences, which were variables included to control for inter-annual variation. We conclude that this is because only 2% of the vegetation burns each year and so wildfires rarely encounter burned patches and chaparral shrublands can carry a fire within 1 or 2 years after previous fire. Prescribed burning is unlikely to have much influence on fire regimes in this area, though targeted treatment at the urban interface may be effective at providing defensible space for protecting assets. These results fit an emerging global model of fire leverage which position California at the bottom end of a continuum, with tropical savannas at the top (leverage = 1: direct replacement of wildfire by prescribed fire) and Australian eucalypt forests in the middle (leverage ~ 0.25).

Ignitions explain more than temperature or precipitation in driving Santa Ana wind fires.

Autumn and winter Santa Ana wind (SAW)-driven wildfires play a substantial role in area burned and societal losses in southern California. Temperature during the event and antecedent precipitation in the week or month prior play a minor role in determining area burned. Burning is dependent on wind intensity and number of human-ignited fires. Over 75% of all SAW events generate no fires; rather, fires during a SAW event are dependent on a fire being ignited. Models explained 40 to 50% of area burned, with number of ignitions being the strongest variable. One hundred percent of SAW fires were human caused, and in the past decade, powerline failures have been the dominant cause. Future fire losses can be reduced by greater emphasis on maintenance of utility lines and attention to planning urban growth in ways that reduce the potential for powerline ignitions.

Large, high-intensity fire events in southern California shrublands: debunking the fine-grain age patch model.

We evaluate the fine-grain age patch model of fire regimes in southern California shrublands. Proponents contend that the historical condition was characterized by frequent small to moderate size, slow-moving smoldering fires, and that this regime has been disrupted by fire suppression activities that have caused unnatural fuel accumulation and anomalously large and catastrophic wildfires. A review of more than 100 19th-century newspaper reports reveals that large, high-intensity wildfires predate modern fire suppression policy, and extensive newspaper coverage plus first-hand accounts support the conclusion that the 1889 Santiago Canyon Fire was the largest fire in California history. Proponents of the fine-grain age patch model contend that even the very earliest 20th-century fires were the result of fire suppression disrupting natural fuel structure. We tested that hypothesis and found that, within the fire perimeters of two of the largest early fire events in 1919 and 1932, prior fire suppression activities were insufficient to have altered the natural fuel structure. Over the last 130 years there has been no significant change in the incidence of large fires greater than 10,000 ha, consistent with the conclusion that fire suppression activities are not the cause of these fire events. Eight megafires (> or = 50,000 ha) are recorded for the region, and half have occurred in the last five years. These burned through a mosaic of age classes, which raises doubts that accumulation of old age classes explains these events. Extreme drought is a plausible explanation for this recent rash of such events, and it is hypothesized that these are due to droughts that led to increased dead fine fuels that promoted the incidence of firebrands and spot fires. A major shortcoming of the fine-grain age patch model is that it requires age-dependent flammability of shrubland fuels, but seral stage chaparral is dominated by short-lived species that create a dense surface layer of fine fuels. Results from the Behave Plus fire model with a custom fuel module for young chaparral shows that there is sufficient dead fuel to spread fire even under relatively little winds. Empirical studies of fuel ages burned in recent fires illustrate that young fuels often comprise a major portion of burned vegetation, and there is no difference between evergreen chaparral and semi-deciduous sage scrub. It has also been argued that the present-day fire size distribution in northern Baja California is a model of the historical patterns that were present on southern California landscapes. Applying this model with historical fire frequencies shows that the Baja model is inadequate to maintain these fire-prone ecosystems and further demonstrates that fire managers in southern California are not likely to learn much from studying modern Baja California fire regimes. Further supporting this conclusion are theoretical cellular automata models of fire spread, which show that, even in systems with age dependent flammability, landscapes evolve toward a complex age mosaic with a plausible age structure only when there is a severe stopping rule that constrains fire size, and only if ignitions are saturating.

Fire treatment effects on vegetation structure, fuels, and potential fire severity in western U.S. forests.

Forest structure and species composition in many western U.S. coniferous forests have been altered through fire exclusion, past and ongoing harvesting practices, and livestock grazing over the 20th century. The effects of these activities have been most pronounced in seasonally dry, low and mid-elevation coniferous forests that once experienced frequent, low to moderate intensity, fire regimes. In this paper, we report the effects of Fire and Fire Surrogate (FFS) forest stand treatments on fuel load profiles, potential fire behavior, and fire severity under three weather scenarios from six western U.S. FFS sites. This replicated, multisite experiment provides a framework for drawing broad generalizations about the effectiveness of prescribed fire and mechanical treatments on surface fuel loads, forest structure, and potential fire severity. Mechanical treatments without fire resulted in combined 1-, 10-, and 100-hour surface fuel loads that were significantly greater than controls at three of five FFS sites. Canopy cover was significantly lower than controls at three of five FFS sites with mechanical-only treatments and at all five FFS sites with the mechanical plus burning treatment; fire-only treatments reduced canopy cover at only one site. For the combined treatment of mechanical plus fire, all five FFS sites with this treatment had a substantially lower likelihood of passive crown fire as indicated by the very high torching indices. FFS sites that experienced significant increases in 1-, 10-, and 100-hour combined surface fuel loads utilized harvest systems that left all activity fuels within experimental units. When mechanical treatments were followed by prescribed burning or pile burning, they were the most effective treatment for reducing crown fire potential and predicted tree mortality because of low surface fuel loads and increased vertical and horizontal canopy separation. Results indicate that mechanical plus fire, fire-only, and mechanical-only treatments using whole-tree harvest systems were all effective at reducing potential fire severity under severe fire weather conditions. Retaining the largest trees within stands also increased fire resistance.

The National Fire and Fire Surrogate study: effects of fuel reduction methods on forest vegetation structure and fuels.

Changes in vegetation and fuels were evaluated from measurements taken before and after fuel reduction treatments (prescribed fire, mechanical treatments, and the combination of the two) at 12 Fire and Fire Surrogate (FFS) sites located in forests with a surface fire regime across the conterminous United States. To test the relative effectiveness of fuel reduction treatments and their effect on ecological parameters we used an information-theoretic approach on a suite of 12 variables representing the overstory (basal area and live tree, sapling, and snag density), the understory (seedling density, shrub cover, and native and alien herbaceous species richness), and the most relevant fuel parameters for wildfire damage (height to live crown, total fuel bed mass, forest floor mass, and woody fuel mass). In the short term (one year after treatment), mechanical treatments were more effective at reducing overstory tree density and basal area and at increasing quadratic mean tree diameter. Prescribed fire treatments were more effective at creating snags, killing seedlings, elevating height to live crown, and reducing surface woody fuels. Overall, the response to fuel reduction treatments of the ecological variables presented in this paper was generally maximized by the combined mechanical plus burning treatment. If the management goal is to quickly produce stands with fewer and larger diameter trees, less surface fuel mass, and greater herbaceous species richness, the combined treatment gave the most desirable results. However, because mechanical plus burning treatments also favored alien species invasion at some sites, monitoring and control need to be part of the prescription when using this treatment.

Fire severity and ecosytem responses following crown fires in California shrublands.

Chaparral shrublands burn in large high-intensity crown fires. Managers interested in how these wildfires affect ecosystem processes generally rely on surrogate measures of fire intensity known as fire severity metrics. In shrublands burned in the autumn of 2003, a study of 250 sites investigated factors determining fire severity and ecosystem responses. Using structural equation modeling we show that stand age, prefire shrub density, and the shortest interval of the prior fire history had significant direct effects on fire severity, explaining > 50% of the variation in severity. Fire severity per se is of interest to resource managers primarily because it is presumed to be an indicator of important ecosystem processes such as vegetative regeneration, community recovery, and erosion. Fire severity contributed relatively little to explaining patterns of regeneration after fire. Two generalizations can be drawn: fire severity effects are mostly shortlived, i.e., by the second year they are greatly diminished, and fire severity may have opposite effects on different functional types. Species richness exhibited a negative relationship to fire severity in the first year, but fire severity impacts were substantially less in the second postfire year and varied by functional type. Much of this relationship was due to alien plants that are sensitive to high fire severity; at all scales from 1 to 1000 m2, the percentage of alien species in the postfire flora declined with increased fire severity. Other aspects of disturbance history are also important determinants of alien cover and richness as both increased with the number of times the site had burned and decreased with time since last fire. A substantial number of studies have shown that remote-sensing indices are correlated with field measurements of fire severity. Across our sites, absolute differenced normalized burn ratio (dNBR) was strongly correlated with field measures of fire severity and with fire history at a site but relative dNBR was not. Despite being correlated with fire severity, absolute dNBR showed little or no relationship with important ecosystem responses to wildfire such as shrub resprouting or total vegetative regeneration. These findings point to a critical need for further research on interpreting remote sensing indices as applied to postfire management of these shrublands.

Fire and Plant Diversification in Mediterranean-Climate Regions.

Despite decades of broad interest in global patterns of biodiversity, little attention has been given to understanding the remarkable levels of plant diversity present in the world's five Mediterranean-type climate (MTC) regions, all of which are considered to be biodiversity hotspots. Comprising the Mediterranean Basin, California, central Chile, the Cape Region of South Africa, and southwestern Australia, these regions share the unusual climatic regime of mild wet winters and warm dry summers. Despite their small extent, covering only about 2.2% of world land area, these regions are home to approximately one-sixth of the world vascular plant flora. The onset of MTCs in the middle Miocene brought summer drought, a novel climatic condition, but also a regime of recurrent fire. Fire has been a significant agent of selection in assembling the modern floras of four of the five MTC regions, with central Chile an exception following the uplift of the Andes in the middle Miocene. Selection for persistence in a fire-prone environment as a key causal factor for species diversification in MTC regions has been under-appreciated or ignored. Mechanisms for fire-driven speciation are diverse and may include both directional (novel traits) and stabilizing selection (retained traits) for appropriate morphological and life-history traits. Both museum and nursery hypotheses have important relevance in explaining the extant species richness of the MTC floras, with fire as a strong stimulant for diversification in a manner distinct from other temperate floras. Spatial and temporal niche separation across topographic, climatic and edaphic gradients has occurred in all five regions. The Mediterranean Basin, California, and central Chile are seen as nurseries for strong but not spectacular rates of Neogene diversification, while the older landscapes of southwestern Australia and the Cape Region show significant components of both Paleogene and younger Neogene speciation in their diversity. Low rates of extinction suggesting a long association with fire more than high rates of speciation have been key to the extant levels of species richness.

Human influence on California fire regimes.

Periodic wildfire maintains the integrity and species composition of many ecosystems, including the mediterranean-climate shrublands of California. However, human activities alter natural fire regimes, which can lead to cascading ecological effects. Increased human ignitions at the wildland-urban interface (WUI) have recently gained attention, but fire activity and risk are typically estimated using only biophysical variables. Our goal was to determine how humans influence fire in California and to examine whether this influence was linear, by relating contemporary (2000) and historic (1960-2000) fire data to both human and biophysical variables. Data for the human variables included fine-resolution maps of the WUI produced using housing density and land cover data. Interface WUI, where development abuts wildland vegetation, was differentiated from intermix WUI, where development intermingles with wildland vegetation. Additional explanatory variables included distance to WUI, population density, road density, vegetation type, and ecoregion. All data were summarized at the county level and analyzed using bivariate and multiple regression methods. We found highly significant relationships between humans and fire on the contemporary landscape, and our models explained fire frequency (R2 = 0.72) better than area burned (R2 = 0.50). Population density, intermix WUI, and distance to WUI explained the most variability in fire frequency, suggesting that the spatial pattern of development may be an important variable to consider when estimating fire risk. We found nonlinear effects such that fire frequency and area burned were highest at intermediate levels of human activity, but declined beyond certain thresholds. Human activities also explained change in fire frequency and area burned (1960-2000), but our models had greater explanatory power during the years 1960-1980, when there was more dramatic change in fire frequency. Understanding wildfire as a function of the spatial arrangement of ignitions and fuels on the landscape, in addition to nonlinear relationships, will be important to fire managers and conservation planners because fire risk may be related to specific levels of housing density that can be accounted for in land use planning. With more fires occurring in close proximity to human infrastructure, there may also be devastating ecological impacts if development continues to grow farther into wildland vegetation.

Epicormic Resprouting in Fire-Prone Ecosystems.

Many plants resprout from basal buds after disturbance, and this is common in shrublands subjected to high-intensity fires. However, resprouting after fire from epicormic (stem) buds is globally far less common. Unlike basal resprouting, post-fire epicormic resprouting is a key plant adaptation for retention of the arborescent skeleton after fire, allowing rapid recovery of the forest or woodland and leading to greater ecosystem resilience under recurrent high-intensity fires. Here we review the biogeography of epicormic resprouting, the mechanisms of protection, the fire regimes where it occurs, and the evolutionary drivers that shaped this trait. We propose that epicormic resprouting is adaptive in ecosystems with high fire frequency and relatively high productivity, at moderate-high fire intensities.