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Jon Keeley

Publications and source records attributed to Jon Keeley.

At least 19 recordsLinked to original sources
USGS2025

Causal analysis of fire regime drivers in California

Background Understanding the relative contribution of climate and human factors to wildfires is critical for managing risk across California’s diverse ecosystems, in the United States (US). Aims We propose a model that distinguishes between proximate and ultimate drivers of fire regimes and apply it to a century of fire and climate data to assess regional variation in causal mechanisms. Methods We analyzed fire statistics (1910–2021) alongside climate and weather data, stratifying the state by 10 ecoregions. Key results Northern forests had the strongest correlation with the proximate factor fuel aridity, ultimately due to climate. Fire rotation intervals exceeded 100 years, implicating woody fuel accumulation as an additional factor. Lightning ignitions occurred in decadal bursts, with dense strike events potentially overwhelming fire-fighting resources. Lower elevation/latitude foothill ecoregions experienced highest fire activity following wet winters and springs, implicating control by herbaceous fuel loads and a negative effect of global warming on future fires. Human ignitions dominate in these ecoregions, and population growth contributes to expansion of powerlines, a major ignition source. Conclusions While climate change may increase fire activity in forested ecoregions, its role is less pronounced in non-forested ecoregions, where human ignition sources are the dominant factor. Implications Different areas within ecoregions may require different management actions that reflect the specific proximate and ultimate factors at play.

California↗
USGS2025

Sequoia and Sequoiadendron: Two paleoendemic megatrees with markedly different adaptive responses to recent high-severity fires

Premise Coast redwood ( Sequoia sempervirens ) and giant sequoia ( Sequoiadendron giganteum ) are two iconic paleoendemic species with limited distributions, well known for their spectacular size. Recently, they have been exposed to high-severity crown fires, with starkly contrasting responses. Methods We used all available published literature and field observations to understand the responses to fire in an evolutionary context. Results Coast redwoods, found in California's coastal rainforests, were highly resilient to high-severity fires, with most trees surviving due to their ability to resprout from the base and trunk, though seedling regeneration was largely lacking. In contrast, giant sequoias, native to the Sierra Nevada, do not resprout, leading to significant tree mortality after very high-severity fires; they released seeds only in patches where some trees survived moderately high-severity fires. Conclusions These high-severity fires were novel events for giant sequoias, but not for coast redwoods. Fire suppression has disrupted the natural fire regime in the giant sequoia ecosystem by preventing frequent lightning-caused surface fires, resulting in high-severity fires that killed a substantial number of these giants. In coast redwood forests, infrequent but high-severity crown fires were the norm before burning by Native Americans. Frequent, low-severity burning by Native Americans over the past few hundred years was localized and 20 th -century fire suppression has returned the natural fire regime to these forests. The recent crown fires do not represent a threat to redwood conservation; however, other management goals may require emulating Native American burning practices and in some cases may be best termed cultural restoration.

American Journal of Botany↗
USGS2025

The role of fire on Earth

Fire is a defining feature of our biosphere, having appeared when the first plants colonized the land, and it continues to occur across the planet at different frequencies and intensities. Fire has been and remains as an evolutionary force in many plant and animal lineages and contributes to explaining the variability of our biodiversity. Fire has also shaped the structure of many ecosystems and the distribution of biomes, and it is an important contributor to the global biogeochemical cycles. In addition, fire has been a key factor in human evolution, and, in turn, humans have modified fire regimes with important consequences for the biosphere. Consequently, fire is an intrinsic factor on our planet. Our challenge now is to understand and predict the role of fire in a densely populated, highly technological world that imposes significant changes on the Earth.

BioScience↗
USGS2024

Ceanothus: Taxonomic patterns in life history responses to fire

Premise: Ceanothus (Rhamnaceae) is a large genus of shrubs that dominate California chaparral and are resilient to fires. Persistence is ensured by resprouting and/or seedling recruitment from dormant seed banks. Some species do both and others, the obligate seeders, are entirely dependent on seedling recruitment. The distribution of these two modes within the genus is poorly documented. Methods: We used all available publications that document species responses to fire and filled most gaps in the literature based on extensive field studies of more than 60 recent wildfires in California. Results: The genus is divided into two subgenera, Ceanothus and Cerastes . Ceanothus is widely considered to comprise mostly resprouting species and Cerastes to consist of only obligate seeders. The subgenus Ceanothus includes resprouting species throughout their range from the eastern United States and Midwest to western United States. Within the California Floristic Province (CFP), a few species are unique in producing massive lignotubers that develop from repeated fires; however, within the CFP, the majority of species in this subgenus do not resprout and are obligate seeders. Two have disjunct subspecies that are facultative seeders or obligate seeders. Conclusions: Previously, speciation in this genus was contended to have occurred in the late Miocene within the CFP. The syndrome of obligate seeding is most strongly represented in this region, and we hypothesize that evolution of this syndrome was a response to increased predictability of fire driven by the Mediterranean climate and the long interval between fires.

California↗
USGS2024

Climate and weather drivers in southern California Santa Ana Wind and non-Santa Wind fires

Background. Autumn and winter Santa Ana Winds (SAW) are responsible for the largest and most destructive wildfires in southern California. Aims. 1) To contrast fires ignited on SAW days vs non-SAW days, 2) evaluate the predictive ability of the Canadian Fire Weather Index (CFWI) for these two fire types, and 3) determine climate and weather factors responsible for the largest wildfires. Methods. Cal Fire FRAP fire data were coupled with hourly climate data from 4 stations, and with regional indices of SAW wind speed, and with seasonal drought from the Palmer Drought Severity Index. Key results. Fires on non-SAW days were more numerous and burned more area, and were significant May to October. CFWI indices were tied to fire occurrence and size for both non-SAW and SAW days, and in the days following ignition. Multiple regression models for months with greatest area burned explained up to a quarter of variation in area burned. Conclusions. The drivers of fire size differ between non-SAW and SAW fires. The best predictor of fire size for non-SAW fires was drought during the prior five years, followed by current-year vapor pressure deficit. For SAW fires, wind speed followed by drought were most important.

California↗
USGS2023

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 E dward B . Poulton, R. D ale Guthrie, and E dwin V . Komarek in animals and W illis L . Jepson, W alter W . Hough, T om M . Harris, P hilip V . Wells, and R obert 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.

BioScience↗
USGS2023

A decade-long study of repeated prescription burning in California native grassland restoration

Native bunchgrass communities dominated by Stipa pulchra are widely distributed in California but share dominance with non-native annual grasses. Restoration of these grasslands focuses on altering the balance of native to non-native grasses to favor the former. This study investigated the impact of burning on vegetation recovery. In the first postfire year burning showed a 70% reduction in cover of non-native annual grasses ( Bromus diandrus exhibited the greatest reduction) and minimal impact on S. pulchra recovery. In the following 3 years, S. pulchra recovered to levels comparable to controls, whereas the annual grasses remained below control levels until the fifth year. Also, in response to reduced annual grass cover on burned sites several species of non-native Erodium increased from 10 to 30% relative cover, however, the low growth form of these forbs presented a less competitive threat to bunchgrasses than the non-native annual grasses, and by the third postfire year returned to near control levels. The rare native geophyte Brodiaea kinkiensis was present throughout these grasslands and was not inhibited by burning treatments. To document the reliability of these patterns a second prescription burn was conducted on these sites 5 years after the first burn and vegetation recovery followed for the subsequent 4 years. Patterns observed after the first burn were duplicated following the second burn. The cover of S . pulchra varied in response to precipitation, with the 95% credible intervals of precipitation parameters overlapping zero, however, the cover of non-native grasses varied greatly with precipitation and had similar trajectories in unburned and burned plots.

California↗
USGS2022

Autumn precipitation: The competition with Santa Ana winds in determining fire outcomes in southern California

Background: California’s South Coast has experienced peak burned area in autumn. Following typically dry, warm summers, precipitation events and Santa Ana winds (SAWs) each occur with increasing frequency from autumn to winter and may affect fire outcomes. Aims: We investigate historical records to understand how these counteracting influences have affected fires. Methods: We defined autumn precipitation onset as the first 3 days when precipitation ≥8.5 mm, and assessed how onset timing and SAWs were associated with frequency of ≥100 ha fires and area burned during 1948–2018. Key results: Timing of autumn precipitation onset had negligible trend but varied considerably from year to year. A total of 90% of area burned in autumn through winter occurred from fires started before onset. Early onset autumns experienced considerably fewer fires and area burned than late onset autumns. SAWs were involved in many of the large fires before onset and nearly all of the lesser number after onset. Conclusions: Risk of large fires is reduced after autumn precipitation onset, but may resurge during SAWs, which provide high risk weather required to generate a large fire. Implications: During autumn before onset, and particularly during late onset autumns, high levels of preparation and vigilance are needed to avoid great fire impacts.

California↗
USGS2022

What makes wildfires destructive in California?

As human impacts from wildfires mount, there is a pressing need to understand why structures are lost in destructive fires. Despite growing research on factors contributing to structure loss, fewer studies have focused on why some fires are destructive and others are not. We characterized overall differences between fires that resulted in structure loss (“destructive fires”) and those that did not (“non-destructive wildfires”) across three California regions. Then, we performed statistical analyses on large fires only (≥100 ha) to distinguish the primary differences between large destructive large fires and large non-destructive fires. Overall, destructive fires were at least an order of magnitude larger than non-destructive fires, with the largest area burned varying by season in different regions. Fire severity was also significantly higher in destructive than non-destructive fires. The statistical analysis showed that, in the San Francisco Bay Area and the northern Sierra Nevada foothills, proximity to the Wildland Urban Interface (WUI) was by far the most important factor differentiating destructive and non-destructive wildfires, followed by different combinations of short-term weather, seasonal climate, topography, and vegetation productivity. In Southern California, wind velocity on the day of the fire ignition was the top factor, which is consistent with previous assumptions that wind-driven fires tend to be most destructive and most of the destruction occurs within the first 24 h. Additionally, Southern California’s high population density increases the odds that a human-caused wildfire may occur during a severe fire-weather event. The geographical differences among regions and the variation of factors explaining the differences between large destructive and large non-destructive fires reflects the complexity inherent in decision-making for reducing wildfire risk. Land use planning to reduce future exposure of housing development to fire and increased focus on wildfire ignition prevention emerge as two approaches with substantial potential.

California↗
USGS2022

The effects of prolonged drought on vegetation dieback and megafires in southern California chaparral

Drought contributed to extensive dieback of southern California chaparral, and normalized difference vegetation index before drought and near the end of the drought was used to estimate this dieback, after accounting for other disturbances recorded in aerial photographs. Within the perimeters of two megafires that occurred after the drought, the 2017 Thomas Fire and the 2018 Woolsey Fire, there had been extensive areas of dieback. Comparing dieback with Monitoring Trends in Burn Severity measures of fire severity, there was a highly significant negative relationship between drought-caused shrub dieback and fire-caused dieback as measured by fire severity. We interpret this as further support for our remote sensing methodology for prefire dieback. Models of fire behavior suggest that one means by which dieback contributes to fire size is through increasing the density and distance of spot fires, particularly under extreme wind conditions. Lower elevation chaparral associations appear to be most vulnerable and are closer to urban environments, which should be a concern to fire managers in regions subjected to extended droughts.

California↗
USGS2022

Vegetation type conversion in the US Southwest: Frontline observations and management responses

Forest and nonforest ecosystems of the western United States are experiencing major transformations in response to land-use change, climate warming, and their interactive effects with wildland fire. Some ecosystems are transitioning to persistent alternative types, hereafter called “vegetation type conversion” (VTC). VTC is one of the most pressing management issues in the southwestern US, yet current strategies to intervene and address change often use trial-and-error approaches devised after the fact. To better understand how to manage VTC, we gathered managers, scientists, and practitioners from across the southwestern US to collect their experiences with VTC challenges, management responses, and outcomes.

Arizona, California, Colorado, New Mexico↗
USGS2022

Climate change and plant regeneration from seeds in Mediterranean regions of the Northern Hemisphere

Mediterranean regions are biodiversity hotspots whose landscapes are characterized by evergreen sclerophyllous vegetation, mild-wet winters, and hot-dry summers. In the Northern Hemisphere, they occur in the Mediterranean Basin and California regions. In these areas, whose habitats are heavily shaped by centuries of anthropic activities, the main effects of climate change include a decrease in precipitation and change in its regime, a pronounced warming, an increase in frequency of extreme temperature events, and altered and intensified fire regimes. Climate change poses a double challenge to the regeneration from seeds of Mediterranean plants. Warmer winters mainly affect the seed germination phase by limiting “typical” Mediterranean cold-cued germination in autumn/winter and reducing overwinter dormancy release for those species whose seeds germinate in spring. Harsher summers are detrimental for the establishment phase, compromising seedling survival. However, evidence of phenotypic plasticity in some Mediterranean plants suggests potential adaptation to a changing climate in the short- to medium-term for species of these regions.

Book chapter↗
USGS2022

Divergent successional trajectories of soil seed bank and post-fire vegetation in a semiarid oak forest: Implications for post-fire ecological restoration

Wildfires are a major disturbance in forest ecosystems around the world and may lead to changes in vegetation succession trajectories. This study examined the impact of time since wildfires on the successional gradients of the degraded Zagros semi-arid oak forest in Iran. Here, we investigated the role of soil seed bank in postfire understory vegetation successional trajectories after wildfires and how time-since-fire influenced plant recovery of this disturbed site. Three adjacent high severity burned areas with different fire histories and the same physiographic conditions were considered. In sampling, we surveyed both aboveground understory vegetation and soil seed bank in all the 96 plots taken along the transects of each area. Soil samples were also collected from each plot and physicochemical properties were analysed in the laboratory. Species composition in the seed bank showed divergent successional trajectories compared to the aboveground vegetation after wildfire. The diversity of soil seed banks followed a gradual decrease, while aboveground understory plants revealed an increasing trend of diversity over time. In addition, the physical and chemical composition of soils was significantly altered by fire. This study presents important insights into soil seed bank dynamics compared to the corresponding aboveground vegetation during postfire succession. The observed changes in diversity and vegetation composition after wildfire can give important insights to management strategies involving prescribed fire in the restoration efforts of highly disturbed semiarid oak forest.

Ecological Engineering↗
USGS2022

Mechanisms of forest resilience

Ecosystems are dynamic systems with complex responses to environmental variation. In response to pervasive stressors of changing climate and disturbance regimes, many ecosystems are realigning rapidly across spatial scales, in many cases moving outside of their observed historical range of variation into alternative ecological states. In some cases, these new states are transitory and represent successional stages that may ultimately revert to the pre-disturbance condition; in other cases, alternative states are persistent and potentially self-reinforcing, especially under conditions of altered climate, disturbance regimes, and influences of non-native species. These reorganized states may appear novel, but reorganization is a characteristic ecosystem response to environmental variation that has been expressed and documented throughout the paleoecological record. Resilience, the ability of an ecosystem to recover or adapt following disturbance, is an emergent property that results from the expression of multiple mechanisms operating across levels of organism, population, and community. We outline a unifying framework of ecological resilience based on ecological mechanisms that lead to outcomes of persistence, recovery, and reorganization. Persistence is the ability of individuals to tolerate exposure to environmental stress, disturbance, or competitive interactions. As a direct expression of life history evolution and adaptation to environmental variation and stress, persistence is manifested most directly in survivorship and continued growth and reproduction of established individuals. When persistence has been overcome (e.g., following mortality from stress, disturbance, or both), populations must recover by reproduction. Recovery requires the establishment of new individuals from seed or other propagules following dispersal from the parent plant. When recovery fails to re-establish the pre-disturbance community, the ecosystem will assemble into a new state. Reorganization occurs along a gradient of magnitude, from changes in the relative dominance of species present in a community, to individual species replacements within an essentially intact community, to complete species turnover and shift to dominance by plants of different functional types, e.g. transition from forest to shrub or grass dominance. When this latter outcome is persistent and involves reinforcing mechanisms, the resulting state represents a vegetation type conversion (VTC), which in this framework represents an end member of reorganization processes. We explore reorganization in greater detail as this phase is increasingly observed but the least understood of the resilience responses. This resilience framework provides a direct and actionable basis for ecosystem management in a rapidly changing world, by targeting specific components of ecological response and managing for sustainable change.

Forest Ecology and Management↗
USGS2022

Bet-hedging and best-bet strategies shape seed dormancy

Seed dormancy (i.e. delayed germination even when conditions are favourable) is a key plant characteristic that occurs among many species worldwide. But, what selective pressures led to seed dormancy? A recent study provides a major analysis of the factors driving this trait at the global scale (Zhang et al ., 2022 ). Using c. 12 000 species and 10 million records across the globe, they conclude that dormancy is a strategy for plants living under ‘seasonal/unpredictable’ environments; and suggest that bet-hedging could be the major mechanism behind the pattern. To reach their conclusions the authors relate the proportion of species with dormant seeds in a grid-cell global map against climate variables related to annual precipitation, temperature and seasonality. Then they showed that the most significant variables were those related to climate seasonality that they equate with unpredictable climates (although seasonal climates usually are highly predictable in their seasonal cycles).

New Phytologist↗
USGS2022

Evolutionary ecology of fire

Fire has been an ecosystem process since plants colonized land over 400 million years ago. Many diverse traits provide a fitness benefit following fires, and these adaptive traits vary with the fire regime. Some of these traits enhance fire survival, while others promote recruitment in the postfire environment. Demonstrating that these traits are fire adaptations is challenging, since many arose early in the paleontological record, although increasingly better fossil records and phylogenetic analysis make timing of these trait origins to fire more certain. Resprouting from the base of stems is the most widely distributed fire-adaptive trait, and it is likely to have evolved under a diversity of disturbance types. The origins of other traits like serotiny, thick bark, fire-stimulated germination, and postfire flowering are more tightly linked to fire. Fire-adaptive traits occur in many environments: boreal and temperate forests, Mediterranean-type climate (MTC) shrublands, savannas, and other grasslands. MTC ecosystems are distinct in that many taxa in different regions have lost the resprouting ability and depend solely on postfire recruitment for postfire recovery. This obligate seeding mode is perhaps the most vulnerable fire-adaptive syndrome in the face of current global change, particularly in light of increasing anthropogenic fire frequency.

Annual Review of Ecology, Evolution, and Systemati↗
USGS2021

Large California wildfires: 2020 fires in historical context

Background California in the year 2020 experienced a record breaking number of large fires. Here, we place this and other recent years in a historical context by examining records of large fire events in the state back to 1860. Since drought is commonly associated with large fire events, we investigated the relationship of large fire events to droughts over this 160 years period. Results This study shows that extreme fire events such as seen in 2020 are not unknown historically, and what stands out as distinctly new is the increased number of large fires (defined here as > 10,000 ha) in the last couple years, most prominently in 2020. Nevertheless, there have been other periods with even greater numbers of large fires, e.g., 1929 had the second greatest number of large fires. In fact, the 1920’s decade stands out as one with many large fires. Conclusions In the last decade, there have been several years with exceptionally large fires. Earlier records show fires of similar size in the nineteenth and early twentieth century. Lengthy droughts, as measured by the Palmer Drought Severity Index (PDSI), were associated with the peaks in large fires in both the 1920s and the early twenty-first century.

California↗