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Increasing fire severity and warmer, drier postfire conditions are making forests in the western United States (West) vulnerable to ecological transformation. Yet, the relative importance of and interactions between these drivers of forest change remain unresolved, particularly over upcoming decades. Here, we assess how the interactive impacts of changing climate and wildfire activity influenced conifer regeneration after 334 wildfires, using a dataset of postfire conifer regeneration from 10,230 field plots. Our findings highlight declining regeneration capacity across the West over the past four decades for the eight dominant conifer species studied. Postfire regeneration is sensitive to high-severity fire, which limits seed availability, and postfire climate, which influences seedling establishment. In the near-term, projected differences in recruitment probability between low- and high-severity fire scenarios were larger than projected climate change impacts for most species, suggesting that reductions in fire severity, and resultant impacts on seed availability, could partially offset expected climate-driven declines in postfire regeneration. Across 40 to 42% of the study area, we project postfire conifer regeneration to be likely following low-severity but not high-severity fire under future climate scenarios (2031 to 2050). However, increasingly warm, dry climate conditions are projected to eventually outweigh the influence of fire severity and seed availability. The percent of the study area considered unlikely to experience conifer regeneration, regardless of fire severity, increased from 5% in 1981 to 2000 to 26 to 31% by mid-century, highlighting a limited time window over which management actions that reduce fire severity may effectively support postfire conifer regeneration.

Changing wildfire regimes are causing rapid shifts in forests worldwide. In particular, forested landscapes that burn repeatedly in relatively quick succession may be at risk of conversion when pre‐fire vegetation cannot recover between fires. Fire refugia (areas that burn less frequently or severely than the surrounding landscape) support post‐fire ecosystem recovery and the persistence of vulnerable species in fire‐prone landscapes. Observed and projected fire‐induced forest losses highlight the need to understand where and why forests persist in refugia through multiple fires. This research need is particularly acute in the Klamath‐Siskiyou ecoregion of southwest Oregon and northwest California, USA, where expected increases in fire activity and climate warming may result in the loss of up to one‐third of the region's conifer forests, which are the most diverse in western North America. Here, we leverage recent advances in fire progression mapping and weather interpolation, in conjunction with a novel application of satellite smoke imagery, to model the key controls on fire refugia occurrence and persistence through one, two, and three fire events over a 32‐year period. Hotter‐than‐average fire weather was associated with lower refugia probability and higher fire severity. Refugia that persisted through three fire events appeared to be partially entrained by landscape features that offered protection from fire, suggesting that topographic variability may be an important stabilizing factor as forests pass through successive fire filters. In addition, smoke density strongly influenced fire effects, with fire refugia more likely to occur when smoke was moderate or dense in the morning, a relationship attributable to reduced incoming solar radiation resulting from smoke shading. Results from this study could inform management strategies designed to protect fire‐resistant portions of biologically and topographically diverse landscapes., Fire refugia support the survival of mature conifer forest in repeatedly burned portions of the Klamath‐Siskiyou ecoregion, a biologically diverse, fire‐prone landscape in the northwest USA. Our study demonstrates that the probability of fire refugia persistence through repeat burning increases with cooler‐than‐average temperatures and moderate to dense morning smoke, likely the result of temperature inversions. Persistent fire refugia appear to be at least partially entrained by landscape features that offer protection from fire, suggesting topographic variability is an important stabilizing factor for the distribution of mature conifer forest as fire activity in the region continues to increase.

Disturbances alter biodiversity via their specific characteristics, including severity and extent in the landscape, which act at different temporal and spatial scales. Biodiversity response to disturbance also depends on the community characteristics and habitat requirements of species. Untangling the mechanistic interplay of these factors has guided disturbance ecology for decades, generating mixed scientific evidence of biodiversity responses to disturbance. Understanding the impact of natural disturbances on biodiversity is increasingly important due to human-induced changes in natural disturbance regimes. In many areas, major natural forest disturbances, such as wildfires, windstorms, and insect outbreaks, are becoming more frequent, intense, severe, and widespread due to climate change and land-use change. Conversely, the suppression of natural disturbances threatens disturbance-dependent biota. Using a meta-analytic approach, we analysed a global data set (with most sampling concentrated in temperate and boreal secondary forests) of species assemblages of 26 taxonomic groups, including plants, animals, and fungi collected from forests affected by wildfires, windstorms, and insect outbreaks. The overall effect of natural disturbances on α-diversity did not differ significantly from zero, but some taxonomic groups responded positively to disturbance, while others tended to respond negatively. Disturbance was beneficial for taxonomic groups preferring conditions associated with open canopies (e.g. hymenopterans and hoverflies), whereas ground-dwelling groups and/or groups typically associated with shady conditions (e.g. epigeic lichens and mycorrhizal fungi) were more likely to be negatively impacted by disturbance. Across all taxonomic groups, the highest α-diversity in disturbed forest patches occurred under moderate disturbance severity, i.e. with approximately 55% of trees killed by disturbance. We further extended our meta-analysis by applying a unified diversity concept based on Hill numbers to estimate α-diversity changes in different taxonomic groups across a gradient of disturbance severity measured at the stand scale and incorporating other disturbance features. We found that disturbance severity negatively affected diversity for Hill number q = 0 but not for q = 1 and q = 2, indicating that diversity–disturbance relationships are shaped by species relative abundances. Our synthesis of α-diversity was extended by a synthesis of disturbance-induced change in species assemblages, and revealed that disturbance changes the β-diversity of multiple taxonomic groups, including some groups that were not affected at the α-diversity level (birds and woody plants). Finally, we used mixed rarefaction/extrapolation to estimate biodiversity change as a function of the proportion of forests that were disturbed, i.e. the disturbance extent measured at the landscape scale. The comparison of intact and naturally disturbed forests revealed that both types of forests provide habitat for unique species assemblages, whereas species diversity in the mixture of disturbed and undisturbed forests peaked at intermediate values of disturbance extent in the simulated landscape. Hence, the relationship between α-diversity and disturbance severity in disturbed forest stands was strikingly similar to the relationship between species richness and disturbance extent in a landscape consisting of both disturbed and undisturbed forest habitats. This result suggests that both moderate disturbance severity and moderate disturbance extent support the highest levels of biodiversity in contemporary forest landscapes., British Ecological Society LRB20/1002, Junta de Andalucia, European Commission B-FQM-366-UGR20, Centro ANID Basal FB210015, Direccion de Investigacion Universidad de La Frontera DIUFRO DI20-0066, Federal Ministry of Education & Research (BMBF), Max Planck Society, INCT Madeiras da Amazonia, Federal Ministry of Education & Research (BMBF) 01LB1001A 01LK1602A, Brazilian Ministry of Science, Technology and Innovation (MCTI/FINEP) 01.11.01248.00, Russian Science Foundation (RSF) 21-14-00227, Ministry of Education, Youth & Sports - Czech Republic Czech Republic Government LTC 20058, Czech Academy of Sciences RVO67985939, UK Research & Innovation (UKRI), Natural Environment Research Council (NERC) GR3/11743, Greek project POL-AEGIS, Program THALES MIS 376737, Projekt DEAL TH 2218/5-1

Wildfires in the western United States (US) are increasingly expensive, destructive, and deadly. Reducing wildfire losses is particularly challenging when fires frequently start on one land tenure and damage natural or developed assets on other ownerships. Managing wildfire risk in multijurisdictional landscapes has recently become a centerpiece of wildfire strategic planning, legislation, and risk research. However, important empirical knowledge gaps remain regarding cross-boundary fire activity in the western US. Here, we use lands administered by the US Forest Service as a study system to assess the causes, ignition locations, structure loss, and social and biophysical factors associated with cross-boundary fire activity over the past three decades. Results show that cross-boundary fires were primarily caused by humans on private lands. Cross-boundary ignitions, area burned, and structure losses were concentrated in California. Public lands managed by the US Forest Service were not the primary source of fires that destroyed the most structures. Cross-boundary fire activity peaked in moderately populated landscapes with dense road and jurisdictional boundary networks. Fire transmission is increasing, and evidence suggests it will continue to do so in the future. Effective cross-boundary fire risk management will require cross-scale risk co-governance. Focusing on minimizing damages to high-value assets may be more effective than excluding fire from multijurisdictional landscapes.

Fire regimes in many dry forests of western North America are substantially different from historical conditions, and there is concern about the ability of these forests to recover following severe wildfire. Fire refugia, unburned or low-severity burned patches where trees survived fire, may serve as essential propagule sources that enable forest regeneration. To quantify the influence of fire refugia spatial pattern and other biophysical factors on the process of post-fire tree regeneration; in particular examining both the proximity and density of surrounding refugia to characterize the landscape of refugial seed sources. We surveyed regeneration at 135 sites in stand-replacement patches across a gradient of fire refugia density in eastern Oregon, USA. We characterized the influence of refugial seed source pattern and other environmental factors on the abundance of regenerating seedlings, and examined the relationship between post-fire climate and the temporal pattern of ponderosa pine seedling establishment. Tree seedlings were present in 83% of plots 12–17 years post-fire, and densities varied substantially (0–67800 stems ha−1, median = 1100). Variation in seedling abundance was driven by the spatial patterns of refugial seed sources. Despite widespread post-fire shrub cover, high-severity burned forests have not undergone a persistent type conversion to shrublands. Ponderosa pine seedling establishment peaked 5–11 years after fire and was not closely associated with post-fire climate. Fire refugia and the seed sources they contain fostered tree regeneration in severely burned patches. Management practices that reduce refugia within post-fire landscapes may negatively influence essential forest recovery processes.

Wildfires in forest ecosystems produce landscape mosaics that include relatively unaffected areas, termed fire refugia. These patches of persistent forest cover can support fire-sensitive species and the biotic legacies important for post-fire forest recovery, yet little is known about their abundance and distribution within fire perimeters. Readily accessible 30-m resolution satellite imagery and derived burn severity products are commonly employed to characterize post-fire landscapes, however, coarse image resolution, generalized burn severity thresholds, and other limitations can constrain accurate representation of fire refugia. This study quantifies the abundance and pattern of fire refugia within 10 fires occurring in ponderosa pine and dry mixed-conifer forests between 2000 and 2003. We developed high-resolution maps of post-fire landscapes using semi-automated, object-based classification of 1-m aerial imagery, conducted imagery- and field-based accuracy assessments, and contrasted these with Landsat-derived burn severity metrics. Fire refugia area within burn perimeters ranged from 20% to 57%. Refugia proportion generally decreased with increasing Landsat-derived burn severity, but still accounted for 3&ndash, 12% of areas classified as high severity. Patch size ranged from 1-m2 isolated trees to nearly 8000 ha, and median patch size was 0.01 ha&mdash, substantially smaller than a 30-m Landsat pixel. Patch size was negatively related to burn severity, distance to fire refugia from open areas was positively related to burn severity. Finally, optimized thresholds of 30-m post-fire normalized burn ratio (NBR) and relative differenced normalized burn ratio (RdNBR) delineated fire refugia with an accuracy of 77% when validated against the 1-m resolution maps. Estimations of fire refugia abundance based on Landsat-derived burn severity metrics are unlikely to detect small, isolated fire refugia patches. Finer-resolution maps can improve understanding of the distribution of forest legacies and inform post-fire management activities including reforestation and treatments.

Altered fire regimes can drive major and enduring compositional shifts or losses of forest ecosystems. In western North America, ponderosa pine and dry mixed‐conifer forest types appear increasingly vulnerable to uncharacteristically extensive, high‐severity wildfire. However, unburned or only lightly impacted forest stands that persist within burn mosaics—termed fire refugia—may serve as tree seed sources and promote landscape recovery. We sampled tree regeneration along gradients of fire refugia proximity and density at 686 sites within the perimeters of 12 large wildfires that occurred between 2000 and 2005 in the interior western United States. We used generalized linear mixed‐effects models to elucidate statistical relationships between tree regeneration and refugia pattern, including a new metric that incorporates patch proximity and proportional abundance. These relationships were then used to develop a spatially explicit landscape simulation model. We found that regeneration by ponderosa pine and obligate‐seeding mixed‐conifer tree species assemblages was strongly and positively predicted by refugia proximity and density. Simulation models revealed that for any given proportion of the landscape occupied by refugia, small patches produced greater landscape recovery than large patches. These results highlight the disproportionate importance of small, isolated islands of surviving trees, which may not be detectable with coarse‐scale satellite imagery. Findings also illustrate the interplay between patch‐scale resistance and landscape‐scale resilience: Disturbance‐resistant settings (fire refugia) can entrain resilience (forest regeneration) across the burn matrix. Implications and applications for land managers and conservation practitioners include strategies for the promotion and maintenance of fire refugia as components of resilient forest landscapes.

Local and regional species extirpations may become more common as changing climate and disturbance regimes accelerate species’ in situ range contractions. Identifying locations that function as both climate and disturbance refugia is critical for biodiversity conservation. Here, we investigate the persistence of a disjunct, fire-sensitive conifer population, yellow-cedar (Callitropsis nootkatensis), in the historically frequent-fire landscape of the Blue Mountains in eastern Oregon, USA. We used tree rings to reconstruct multi-century fire histories, which were then used to compare historical mean fire return intervals (MFRIs) inside of the cedar grove to the surrounding dry forest matrix, as well as to examine relationships between historical fire occurrence and reconstructed climate. We also examined trends in post-fire yellow-cedar mortality and regeneration between 2006, when the grove burned in a wildfire, through 2017. Results indicated that fire was less frequent in the cedar grove (MFRI = 35.8 years) than in the surrounding dry mixed-conifer forest (MFRI = 14.4 years). Historical fire occurrence was associated with hotter and drier conditions. Following the 2006 fire, cedar mortality was high (>90%), but by 2017 post-fire regeneration was abundant (median = 8125 seedlings ha-1). The eastern Oregon cedar grove appears to occupy a fire refugium historically decoupled from the frequent-fire regime of the broader landscape. The topographic position of the grove suggests it is also a climate refugium. Despite high mortality following fire in 2006, yellow-cedar is regenerating in the grove. Managing for climate and disturbance refugia has been identified as an important option for conservation in the context of rapid global change, and this study provides evidence that locations where climate and disturbance refugia overlap may be disproportionately important for the maintenance of regional biodiversity. However, active management may be required for populations to persist in some of these locations as anthropogenic change is capable of overwhelming the underlying buffering capacity of refugia.

Fire refugia are landscape elements that remain unburned or minimally affected by fire, thereby supporting postfire ecosystem function, biodiversity, and resilience to disturbances. Although fire refugia have been studied across continents, scales, and affected taxa, they have not been characterized systematically over space and time, which is crucial for understanding their role in facilitating resilience in the context of global change. We identify four dichotomies that delineate an overarching conceptual framework of fire refugia: unburned versus lower severity, species-specific versus landscape-process characteristics, predictable versus stochastic, and ephemeral versus persistent. We outline the principal concepts underlying the ecological function of fire refugia and describe both the role of fire refugia and uncertainties regarding their persistence under global change. An improved understanding of fire refugia is crucial to conservation given the role that humans play in shaping disturbance regimes across landscapes.