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Live fuel moisture content (LFMC) plays a critical role in wildfire dynamics, but little is known about responses of LFMC to multivariate climate change, e.g., warming temperature, CO2 fertilization, and altered precipitation patterns, leading to a limited prediction ability of future wildfire risks. Here, we use a hydrodynamic demographic vegetation model to estimate LFMC dynamics of chaparral shrubs, a dominant vegetation type in fire-prone southern California. We parameterize the model based on observed shrub allometry and hydraulic traits and evaluate the model's accuracy through comparisons between observed and simulated LFMC of three plant functional types (PFTs) under current climate conditions. Moreover, we estimate the number of days per year of LFMC below 79 % (which is a critical threshold for wildfire danger rating of southern California chaparral shrubs) from 1960 to 2099 for each PFT and compare the number of days below the threshold for medium and high greenhouse gas emission scenarios (RCP4.5 and 8.5). We find that climate change could lead to more days per year (5.2 %–14.8 % increase) with LFMC below 79 % between the historical (1960–1999) and future (2080–2099) periods, implying an increase in wildfire danger for chaparral shrubs in southern California. Under the high greenhouse gas emission scenario during the dry season, we find that the future LFMC reductions mainly result from a warming temperature, which leads to 9.1 %–18.6 % reduction in LFMC. Lower precipitation in the spring leads to a 6.3 %–8.1 % reduction in LFMC. The combined impacts of warming and precipitation change on fire season length are equal to the additive impacts of warming and precipitation change individually. Our results show that the CO2 fertilization will mitigate fire risk by causing a 3.5 %–4.8 % increase in LFMC. Our results suggest that multivariate climate change could cause a significant net reduction in LFMC and thus exacerbate future wildfire danger in chaparral shrub systems.

© 2020 The Authors., Fire is a powerful ecological and evolutionary force that regulates organismal traits, population sizes, species interactions, community composition, carbon and nutrient cycling and ecosystem function. It also presents a rapidly growing societal challenge, due to both increasingly destructive wildfires and fire exclusion in fire‐dependent ecosystems. As an ecological process, fire integrates complex feedbacks among biological, social and geophysical processes, requiring coordination across several fields and scales of study. Here, we describe the diversity of ways in which fire operates as a fundamental ecological and evolutionary process on Earth. We explore research priorities in six categories of fire ecology: (a) characteristics of fire regimes, (b) changing fire regimes, (c) fire effects on above‐ground ecology, (d) fire effects on below‐ground ecology, (e) fire behaviour and (f) fire ecology modelling. We identify three emergent themes: the need to study fire across temporal scales, to assess the mechanisms underlying a variety of ecological feedbacks involving fire and to improve representation of fire in a range of modelling contexts. Synthesis: As fire regimes and our relationships with fire continue to change, prioritizing these research areas will facilitate understanding of the ecological causes and consequences of future fires and rethinking fire management alternatives., Support was provided by NSF‐DEB‐1743681 to K.K.M. and A.J.T. We thank Shalin Hai‐Jew for helpful discussion of the survey and qualitative methods.

Author(s): Batllori, E; De Caceres, M; Brotons, L; Ackerly, DD; Moritz, MA; Lloret, F | Abstract: Understanding ecosystem responses to compound disturbance regimes and the influence of specific sequences of events in determining ecosystem shifts remains a challenge. We use a modelling framework for Mediterranean-type ecosystems to assess the effects of fire–drought interactions on long-term vegetation dynamics and to identify disturbance-driven changes in trait composition (tree seeder vs. tree resprouter prevalence) and ecosystem state (forest vs. non-forest). Changes in tree seeder and the tree resprouter dominance show nonlinear, threshold-type trends over gradients of increasing compound disturbance frequency. Vegetation composition thresholds mostly occur in a narrow range of the compound fire–drought disturbance space. Additionally, trait compositional switches and the likelihood of sudden changes in ecosystem state are promoted by fire-drought interactions. Distinct sequences of disturbance events cause vegetation transitions, disrupting ecosystem resilience, even under moderate recurrence of individual disturbances. An extreme drought year followed by one or two large fire events promotes shifts from resprouter- to seeder dominance. Contrastingly, a large crown fire followed by an extreme drought promotes changes from seeder to resprouter dominance. This disturbance sequence is also a mechanism strong enough to trigger sudden shifts in ecosystem state (from forest to non-forest). Synthesis. Thresholds of change in vegetation composition occur over a narrow range of the modelled gradients of compound fire-drought recurrence, and the loss of ecosystem resilience is contingent on particular sequences of disturbance events. Overall, our findings highlight that disturbance interactions define the relative location of tipping points in ecosystem state, and that effects and feedbacks of compound disturbance regimes increase the long-term likelihood of sudden ecosystem shifts and, therefore, uncertainty in predicting vegetation state.

Interest in translational ecology (TE) – a research approach that yields useful scientific outcomes through ongoing collaboration between scientists and stakeholders – is growing among both of these groups. Translational ecology brings together participants from different cultures and with different professional incentives. We address ways to cultivate a culture of TE, such as investing time in understanding one another's decision context and incentives, and outline common entry points to translational research, such as working through boundary organizations, building place-based research programs, and being open to opportunities as they arise. We also highlight common institutional constraints on scientists and practitioners, and ways in which collaborative research can overcome these limitations, emphasizing considerations for navigating TE within current institutional frameworks, but also pointing out ways in which institutions are evolving to facilitate translational research approaches.

Author(s): Newman, Erica A; Wilber, Mark Q; Kopper, Karen E; Moritz, Max A; Falk, Donald A; McKenzie, Don; Harte, John | Abstract: Macroecological studies have established widespread patterns of species diversity and abundance in ecosystems but have generally restricted their scope to relatively steady‐state systems. As a result, how macroecological metrics are expected to scale in ecosystems that experience natural disturbance regimes is unknown. We examine macroecological patterns in a fire‐dependent forest of Bishop pine (Pinus muricata ). We target two different‐aged stands in a stand‐replacing fire regime: a mature stand with a diverse understory and with no history of major disturbance for at least 40 yr, and one disturbed by a stand‐replacing fire 17 yr prior to measurement. We compare properties of these stands with macroecological predictions from the Maximum Entropy Theory of Ecology (METE ), an information entropy‐based theory that has proven highly successful in predicting macroecological metrics in multiple ecosystems and taxa. Ecological patterns in the mature stand more closely match METE predictions than do data from the more recently disturbed, mid‐seral stage stand. This suggests METE 's predictions are more robust in late‐successional, slowly changing, or steady‐state systems than those in rapid flux with respect to species composition, abundances, and organisms’ sizes. Our findings highlight the need for a macroecological theory that incorporates natural disturbance, perturbations, and ecological dynamics into its predictive capabilities, because most natural systems are not in a steady state.

Ecologists who specialize in translational ecology (TE) seek to link ecological knowledge to decision making by integrating ecological science with the full complement of social dimensions that underlie today's complex environmental issues. TE is motivated by a search for outcomes that directly serve the needs of natural resource managers and decision makers. This objective distinguishes it from both basic and applied ecological research and, as a practice, it deliberately extends research beyond theory or opportunistic applications. TE is uniquely positioned to address complex issues through interdisciplinary team approaches and integrated scientist–practitioner partnerships. The creativity and context-specific knowledge of resource managers, practitioners, and decision makers inform and enrich the scientific process and help shape use-driven, actionable science. Moreover, addressing research questions that arise from on-the-ground management issues – as opposed to the top-down or expert-oriented perspectives of traditional science – can foster the high levels of trust and commitment that are critical for long-term, sustained engagement between partners.

The occurrence of multiple disturbances can jointly affect the recovery capacity of ecosystems, potentially leading to changes in vegetation dynamics or loss of resilience. The effects of interacting disturbances on ecosystems are, however, not well understood. We use a model system based on Mediterranean‐type ecosystems (MTE s) to examine how the interplay between vegetation regeneration traits and compound, stochastic disturbances modulate ecosystem dynamics. We developed a state‐and‐transition simulation model including two tree species with contrasting regeneration strategies (seeder vs. resprouter) and a shrubland formation. We aim to assess potential compositional switches under contrasted scenarios of compound fire–drought regimes, and to characterize the cumulative effects of fire–drought (synergism vs. antagonism) relative to the effects of individual disturbance regimes. Our simulation results indicate that interaction between moderate fire and sporadic drought recurrence—as opposed to chronic dryness—can act as a strong mechanism generating highly heterogeneous landscapes in which different regeneration types coexist, as observed in MTE s. Resprouters dominated under individual, moderate disturbance regimes of fire or drought, whereas the interaction of the two disturbances promoted the long‐term coexistence of both tree regeneration strategies. However, shrubland expansion and persistence at the expanse of forests was favored by increases in drought recurrence and associated fire–drought interactions, highlighting the potential for important vegetation changes in MTE s under climate change. Overall, the cumulative effects of fire and drought can lead to distinct landscape configurations under moderate disturbance regimes that are otherwise only attained under high frequency of individual disturbances. At the ecosystem level, however, we suggest that disturbance‐induced vegetation dynamics can modify vegetation sensitivity and resilience to further disturbances precluding the prevalence of synergistic effects of the two disturbances. Enric Batllori thanks the support of a Marie Curie International Incoming Fellowship (PIFF‐GA‐2013‐625547) and Miquel De Cáceres the support of a Spanish Ramon y Cajal Fellowship. Lluís Brotons and Francisco Lloret thank the support of the research group 2014 SGR 00453 and the projects FORESTCAST, NEWFORESTS, INFORMED, SECADIN, and BIOCLIM.

Aim Pyrogeographical theory suggests that fire is controlled by spatial gradients in resources to burn (fuel amount) and climatic conditions promoting combustion (fuel moisture). Examining trade-offs among these environmental constraints is critical to understanding future fire activity. We evaluate constraints on fire frequency in modern fire records over the entire Mediterranean biome and identify potential shifts in fire activity under an ensemble of global climate projections. Location The biome encompassing the Mediterranean-type ecosystems (MTEs). Methods We evaluate potential changes in fire over the 21st century in MTEs based on a standardized global framework. Future fire predictions are generated from statistical fire−climate models driven by ensembles of climate projections under the IPCC A2 emissions scenario depicting warmer–drier and warmer–wetter syndromes. We test the hypothesis that MTEs lie in the transition zone discriminating fuel moisture versus fuel amount as the dominant constraint on fire activity. Results Fire increases reported in MTEs in recent decades may not continue throughout the century. MTEs occupy a sensitive portion of global fire−climate relationships, especially for precipitation-related variables, leading to highly divergent fire predictions under drier versus wetter syndromes. Warmer–drier conditions could result in decreased fire activity over more than half the Mediterranean biome by 2070–2099, and the opposite is predicted under a warmer–wetter future. MTEs encompass, however, a climate space broad and complex enough to include spatially varied fire responses and potential conversions to non-MTE biomes. Main conclusions Our results strongly support the existence of both fuel amount and fuel moisture constraints on fire activity and show their geographically variable influence throughout MTEs. Climatic controls on fire occurrence in MTEs lie close to ‘tipping points’, where relatively small changes in future climates could translate into drastic and divergent shifts in fire activity over the Mediterranean biome, mediated by productivity alterations.

Seedling establishment is a critical step that may ultimately govern tree species’ distribution shifts under environmental change. Annual variation in the location of seed rain and microclimates results in transient “windows of opportunity” for tree seedling establishment across the landscape. These establishment windows vary at fine spatiotemporal scales that are not considered in most assessments of climate change impacts on tree species range dynamics and habitat displacement. We integrate field seedling establishment trials conducted in the southern Sierra Nevada and western Tehachapi Mountains of southern California with spatially downscaled grids of modeled water‐year climatic water deficit (CWDwy) and mean August maximum daily temperature (Tmax) to map historical and projected future microclimates suitable for establishment windows of opportunity for Quercus douglasii, a dominant tree species of warm, dry foothill woodlands, and Q. kelloggii, a dominant of cooler, more mesic montane woodlands and forests. Based on quasi‐binomial regression models, Q. douglasii seedling establishment is significantly associated with modeled CWDwy and to a lesser degree with modeled Tmax. Q. kelloggii seedling establishment is most strongly associated with Tmax and best predicted by a two‐factor model including CWDwy and Tmax. Establishment niche models are applied to explore recruitment window dynamics in the western Tehachapi Mountains, where these species are currently widespread canopy dominants. Establishment windows are projected to decrease by 50–95%, shrinking locally to higher elevations and north‐facing slopes by the end of this century depending on the species and climate scenario. These decreases in establishment windows suggest the potential for longer‐term regional population declines of the species. While many additional processes regulate seedling establishment and growth, this study highlights the need to account for topoclimatic controls and interannual climatic variation when assessing how seedling establishment and colonization processes could be affected by climate change.

Humans and their ancestors are unique in being a fire-making species, but ‘natural’ (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from ‘natural’ background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.

The spatial pattern of fire observed across boreal landscapes is the outcome of complex interactions among components of the fire environment. We investigated how the naturally occurring patterns of ignitions, fuels, and weather generate spatial pattern of burn probability (BP) in a large and highly fireprone boreal landscape of western Canada, Wood Buffalo National Park. This was achieved by producing a high-resolution map of BP using a fire simulation model that models the ignition and spread of individual fires for the current state of the study landscape (that is, the ‘control’). Then, to extract the effect of the variability in ignitions, fuels, and weather on spatial BP patterns, we subtracted the control BP map to those produced by ‘‘homogenizing’’ a single environmental factor of interest (that is, the ‘experimental treatments’). This yielded maps of spatial residuals that represent the spatial BP patterns for which the heterogeneity of each factor of interest is responsible. Residuals were analyzed within a structural equation modeling framework. The results showed unequal contributions of fuels (67.4%), weather (29.2%), and ignitions (3.4%) to spatial BP patterning. The large contribution of fuels reflects how substantial heterogeneity of land cover on this landscape strongly affects BP. Although weather has a chiefly temporal control on fire regimes, the variability in fire-conducive weather conditions exerted a surprisingly large influence on spatial BP patterns. The almost negligible effect of spatial ignition patterns was surprising but explainable in the context of this area’s fire regime. Similar contributions of fuels, weather, and ignitions could be expected in other parts of the boreal forest that lack a strong anthropogenic imprint, but are likely to be altered in human-dominated fire regimes.

Fire is an important factor in determining plant community composition and distribution. This study quantifies landscape-scale vegetation change following a large fire at Point Reyes National Seashore, California, USA. Vegetation in the Point Reyes region is characterized by a complex mosaic of grassland, shrub, and forest plant communities, and by high levels of plant diversity. Although large fires are relatively rare on the coast of California north of San Francisco Bay, they are important in determining the distributions of plant communities at the landscape scale. We mapped vegetation communities throughout the study area using aerial imagery and analyzed how vegetation shifted following fire. We found substantial areas had transitioned from coastal scrub to ceanothus scrub (Ceanothus thyrsiflorus Eschsch.) or Bishop pine (Pinus muricata D. Don) forest following fire. Transitions from shrub to tree vegetation following fire have rarely been documented in this region. Logistic regression analysis was used to examine the factors influencing the post-fire distribution of Bishop pine and ceanothus scrub. Proximity to pre-fire Bishop pine stands and pre-fire vegetation type were the most important predictors of post-fire Bishop pine regeneration. Pre-fire vegetation type, burn severity, and topography were the most important predictors of post-fire ceanothus scrub distribution. This study demonstrates the capacity of these ecosystems for substantial change over short time periods in response to fire, and identifies some of the factors driving this change.

We provide an empirical, global test of the varying constraints hypothesis, which predicts systematic heterogeneity in the relative importance of biomass resources to burn and atmospheric conditions suitable to burning (weather/climate) across a spatial gradient of long-term resource availability. Analyses were based on relationships between monthly global wildfire activity, soil moisture, and mid-tropospheric circulation data from 2001 to 2007, synthesized across a gradient of long-term averages in resources (net primary productivity), annual temperature, and terrestrial biome. We demonstrate support for the varying constraints hypothesis, showing that, while key biophysical factors must coincide for wildfires to occur, the relative influence of resources to burn and moisture/weather conditions on fire activity shows predictable spatial patterns. In areas where resources are always available for burning during the fire season, such as subtropical/tropical biomes with mid-high annual long-term net primary productivity, fuel moisture conditions exert their strongest constraint on fire activity. In areas where resources are more limiting or variable, such as deserts, xeric shrublands, or grasslands/savannas, fuel moisture has a diminished constraint on wildfire, and metrics indicating availability of burnable fuels produced during the antecedent wet growing seasons reflect a more pronounced constraint on wildfire. This macro-scaled evidence for spatially varying constraints provides a synthesis with studies performed at local and regional scales, enhances our understanding of fire as a global process, and indicates how sensitivity to future changes in temperature and precipitation may differ across the world.

Aim Substantial overlap in the climate characteristics of the United States and China results in similar land-cover types and weather conditions, especially in the eastern half of the two countries. These parallels suggest similarities in fire regimes as well, yet relatively little is known about the historical role of fire in Chinese ecosystems. Consequently, we aimed to infer fire regime characteristics for China based on our understanding of climate–fire relationships in the United States. Location The conterminous United States and the People's Republic of China. Methods We used generalized additive models to quantify the relationship between reference fire regime classes adopted by the LANDFIRE initiative in the United States, and a global climate data set. With the models, we determined which climate variables best described the distribution of fire regimes in the United States then used these models to predict the spatial distribution of fire regimes in China. The fitted models were validated quantitatively using receiver operating characteristic area under the curve (AUC). We validated the predicted fire regimes in China by comparison with palaeoecological fire data and satellite-derived estimates of current fire activity. Results Quantitative validation using the AUC indicated good discrimination of the distribution of fire regimes by models for the United States. Overall, fire regimes with more frequent return intervals were more likely in the east than in the west. The resolution of available historical and prehistorical fire data for China, including sediment cores, allowed only coarse, qualitative validation, but provided supporting evidence that fire has long been a part of ecosystem function in eastern China. MODIS satellite data illustrated that fire frequency within the last decade supported the classification of much of western China as relatively fire-free; however, much of south-eastern China experiences more fire activity than predicted with our models, probably as a function of the extensive use of fire by people. Conclusions While acknowledging there are many cultural, environmental and historical differences between the United States and China, our fire regime models based on climate data demonstrate potential historical fire regimes for China, and propose that large areas of China share historical fire–vegetation–climate complexes with the United States.

Despite its widespread occurrence globally, wildfire preferentially occupies an environmental middle ground and is significantly less prevalent in biomes characterized by environmental extremes (e.g., tundra, rain forests, and deserts). We evaluated the biophysical “environmental space” of wildfire from regional to subcontinental extents, with methods widely used for modeling habitat distributions. This approach is particularly suitable for the biogeographic study of wildfire, because it simultaneously considers patterns in multiple factors controlling wildfire suitability over large areas. We used the Maxent and boosted regression tree algorithms to assess wildfire–environment relationships for three levels of complexity (in terms of inclusion of variables) at three spatial scales: the conterminous United States, the state of California, and five wildfire-prone ecoregions of California. The resulting models were projected geographically to obtain spatial predictions of wildfire suitability and were also ...

Due to a unique combination of environmental conditions, the chaparral shrublands of southern California are prone to large, intense wildland fires. There is ongoing work in the fire research community to establish whether fuel accumulation or weather conditions are the determining factor in the prevalence of large chaparral fires. This study introduces a framework for contributing a modeling perspective to understanding these alternative hypotheses. As models formalize our understanding of the physical process of fire spread, the sensitivity of the models to the meteorological and fuel inputs should be indicators of their relative importance. A global sensitivity analysis (GSA) was conducted on HFire, a spatially explicit raster model developed for modeling fire spread in chaparral fuels, based on the Rothermel spread equations. The GSA provided a quantitative measure of the importance of each of the model inputs on the predicted fire size. The results indicate that, under extreme weather conditions, wind speed was over three times more influential on predicted fire sizes than any other single model input. This finding supports the idea that fires burning under Santa Ana conditions are primarily driven by high wind speeds. Future research will involve extending the GSA methodology to quantify the relative importance of these inputs in terms of the long-term fire regime in chaparral ecosystems.

Fire can be a dominant process in the ecology of forest vegetation and can also affect forest disease dynamics. Little is known about the relationship between fire and an emerging disease epidemic called Sudden Oak Death, which is caused by a new pathogen, Phytophthora ramorum. This disease has spread across a large, fire-prone portion of California, killing great numbers of oaks and tanoaks and infecting most associated woody plants. Suitable hosts cover a much broader geographic range, raising concern over where the disease may spread. To understand the strength and potential sensitivities of a fire-disease relationship, we examined geographic patterns of confirmed P. ramorum infections in relation to past fire history. We found these infections to be extremely rare within the perimeter of any area burned since 1950. This finding is not caused by spatial bias in sampling for the disease, and is robust to variation in host abundance scenarios and to aggregation of closely spaced sampling locations. We therefore investigated known fire-related factors that could result in significantly lower incidence of the disease in relatively recently burned landscapes. Chemical trends in post-fire environments can influence the success of pathogens like P. ramorum, either by increasing plant nutrient stress or by reducing the occurrence of chemicals antagonistic to Phytophthoras. Succession in the absence of fire leads to greater abundance of host species, which will provide increased habitat for P. ramorum; this will also increase intraspecific competition where these trees are abundant, and other density-dependent effects (e.g. shading) can reduce resource allocation to defenses. Despite these findings about a fire-disease relationship, a much deeper understanding is necessary before fire can be actively used as a tool in slowing the epidemic.

The Klamath-Siskiyou region of northwestern California and southwestern Oregon supports glob- ally outstanding temperate biodiversity. Fire has been important in the evolutionary history that shaped this diversity, but recent human influences have altered the fire environment. We tested for modern human impacts on the fire regime by analyzing temporal patterns in fire extent and spatial patterns of fire severity in relation to vegetation structure, past fire occurrence, roads, and timber management in a 98,814-ha area burned in 1987. Fire severity was mapped by the U.S. Department of Agriculture Forest Service as low, moderate, and high based on levels of canopy scorch and consumption. We found (1) a trend of increasing fire size in recent decades; (2) that overall fire-severity proportions were 59% low, 29% moderate, and 12% high, which is compa- rable to both contemporary and historic fires in the region; (3) that multiaged, closed forests, the predominant vegetation, burned with much lower severity than did open forest and shrubby nonforest vegetation; (4) that considerably less high-severity fire occurred where fire had previously be absent since 1920 in closed forests compared to where the forests had burned since 1920 (7% vs. 16%); (5) that nonforest vegetation burned with greater severity where there was a history of fire since 1920 and in roaded areas; and (6) that tree plantations experienced twice as much severe fire as multi-aged forests. We concluded that fuel buildup in the absence of fire did not cause increased fire severity as hypothesized. Instead, fuel that is receptive to combustion may decrease in the long absence of fire in the closed forests of our study area, which will favor the fire regime that has maintained these forests. However, plantations are now found in one-third of the roaded landscape. Together with warming climate, this may increase the size and severity of future fires, favoring further establishment of structurally and biologically simple plantations.

This year's catastrophic wildfires in southern California highlight the need for effective planning and management for fire-prone landscapes. Fire frequency analysis of several hundred wildfires over a broad expanse of California shrublands reveals that there is generally not, as is commonly assumed, a strong relationship between fuel age and fire probabilities. Instead, the hazard of burning in most locations increases only moderately with time since the last fire, and a marked age effect of fuels is observed only in limited areas. Results indicate a serious need for a re-evaluation of current fire management and policy, which is based largely on eliminating older stands of shrubland vegetation. In many shrubland ecosystems exposed to extreme fire weather, large and intense wildfires may need to be factored in as inevitable events.

Large fires in chaparral-dominated shrublands of southern and central California are widely attributed to decades of fire suppression. Prehistoric shrubland landscapes are hypothesized to have exhibited fine-grained age-patch mosaics in which fire spread was limited by the age and spatial pattern of fuels. In contrast, I hypothesize that fires during extreme weather conditions have been capable of burning through all age classes of the vegetation mosaic. Using the mapped fire history (1911–1995) of Los Padres National Forest, I analyzed burning patterns for hundreds of fires using a geographic information system (GIS). To estimate the degree of age dependency exhibited by the fire regime at different spatial scales, I employed methods of fire frequency analysis (i.e., fitting a generalized Weibull function to fire interval distributions). Statistics were also calculated using a temporal breakpoint of 1950 to assess possible effects of suppression. Results indicated that shrubland fires have frequently bur...

Accepted Ar tic le Averaged 30 year climate change projections mask opportunities for species establishment Josep M. Serra-Diaz 1,2 , Janet Franklin 1 , Lynn C. Sweet 3 , Ian M. McCullough 3 , Alexandra D. Syphard 4 , Helen M. Regan 5 , Lorraine E. Flint 6 , Alan L. Flint 6 , John R. Dingman 7 , Max Moritz 8 , Kelly Redmond 9 , Lee Hannah 10 , Frank W. Davis 3 School of Geographical Sciences and Urban Planning, Arizona State University, Tempe, AZ, USA Harvard Forest, Harvard University, Petersham, MA, USA Bren School of Environmental Science and Management, University of California Santa Barbara, CA, USA Conservation Biology Institute, La Mesa, CA, USA Department of Biology, University of California, Riverside, CA, USA U.S. Geological Survey, California Water Science Center, Sacramento, CA, USA Air Resources Board, California Environmental Protection Agency, Sacramento, CA, USA Department of Environmental Science, Policy and Management, University of California Berkeley, CA, USA Division of atmospheric sciences, Desert Research Institute, Reno, NV, USA Conservation International, Arlington, VA, USA Corresponding author: Josep M. Serra-Diaz, School of Geographical Sciences and Urban Planning, Arizona State University, Tempe, AZ, USA. E-mail: pep.science@gmail.com, jserradi@asu.edu Decision date: 30-Sep-2015 This article has been accepted for publication and undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process, which may lead to differences between this version and the Version of Record. Please cite this article as doi: [10.1111/ecog.02074]. ‘This article is protected by copyright. All rights reserved.’

Conservation in the face of climate change is hindered by a preoccupation with place. ‘Refugial capacity’ [1] adds useful descriptive power to the terminology of place, while ‘stepping stones’ and ‘holdouts’ are terms that usefully describe the dynamics of place [2]. Ultimately, no matter what happens in the lexicon, conservation needs to be implemented with a shift in focus from place to process. It will not be easy.

In this article, we develop a heuristic model of ecosystem-disturbance dynamics that illustrates a range of responses of disturbance impact to gradients of increasing disturbance extent, intensity, or duration. Three general kinds of response are identified and illustrated: (a) threshold response, (b) scale-independent response, and (c) continuous response. Threshold responses are those in which the response curve shows a discontinuity or a sudden change in slope along the axis of increasing disturbance extent, intensity, or duration. The response threshold occurs at a point where the force of the disturbance exceeds the capacity of internal mechanisms to resist disturbance, or where new mechanisms of recovery become involved. Within this conceptual framework, we find that some unusually large or intense disturbances, but not all, produce qualitatively different responses compared with similar disturbances of lesser magnitude. If disturbance impact does not increase with increasing disturbance extent, intensity, or duration, or if the response curve changes monotonically, then large disturbances are not qualitatively different from small ones. For example, jack pine tends to become reestablished after stand-replacing fire in boreal forests, regardless of fire size, because its serotinous cones provide an adequate seed source throughout the burned area. Thus, large fires are not qualitatively different from small fires in terms of jack pine reproduction. However, if disturbance impact does increase abruptly at some point with increasing disturbance extent, intensity, or duration, often because of thresholds in the capacity of internal mechanisms to resist or respond to disturbance impact, then large disturbances are qualitatively different from small ones, at least for some parameters of ecological response. For example, balsam fir and white cedar can recolonize a small burned patch of boreal forest in close proximity to surviving individuals of these species, but they will be eliminated from a large burn because of their susceptibility to fire-caused mortality and their inability to disperse their seeds over long distances. The conceptual framework presented here permits some new insights into the dynamics of natural systems and may provide a useful tool with which managers can assess the potential for catastrophic damages resulting from large, infrequent disturbances.

In our 2011 synthesis (Bowman et al., Journal of Biogeography, 2011, 38, 2223–2236), we argued for a holistic approach to human issues in fire science that we term ‘pyrogeography’. Coughlan & Petty (Journal of Biogeography, 2013, 40, 1010–1012) critiqued our paper on the grounds that our ‘pyric phase’ model was built on outdated views of cultural development, claiming we developed it to be the unifying explanatory framework for all human–fire sciences. Rather, they suggest that ‘historical ecology’ could provide such a framework. We used the ‘pyric transition’ for multiple purposes but did not offer it as an exclusive explanatory framework for pyrogeography. Although ‘historical ecology’ is one of many useful approaches to studying human–fire relationships, scholars should also look to political and evolutionary ecology, ecosystems and complexity theories, as well as empirical generalizations to build an interdisciplinary fire science that incorporates human, ecological and biophysical dimensions of fire regimes.

Microclimates have played a critical role in past species range shifts, suggesting that they could be important in biological response to future change. Terms are needed to discuss these future effects. We propose that populations occupying microclimates be referred to as holdouts, stepping stones and microrefugia. A holdout is a population that persists in a microclimate for a limited period of time under deteriorating climatic conditions. Stepping stones successively occupy microclimates in a way that facilitates species' range shifts. Microrefugia refer to populations that persist in microclimates through a period of unfavorable climate. Because climate projections show that return to present climate is highly unlikely, conservation strategies need to be built around holdouts and stepping stones, rather than low-probability microrefugia.

Smoke from forest fires can persist in the environment for weeks and while there is a substantial amount of literature examining the effects of smoke exposure on seed germination, the effects of smoke on leaf function are nearly uninvestigated. The objective of this study was to compare growth and primary and secondary metabolic responses of deciduous angiosperm and evergreen conifer tree species to short smoke exposure. Twenty minutes of smoke exposure resulted in a greater than 50% reduction in photosynthetic capacity in five of the six species we examined. Impairment of photosynthesis in response to smoke was a function of reductions in stomatal conductance and biochemical limitations. In general, deciduous angiosperm species showed a greater sensitivity than evergreen conifers. While there were significant decreases in photosynthesis and stomatal conductance, smoke had no significant effect on growth or secondary defense compound production in any of the tree species examined.

Fire in high-elevation forest ecosystems can have severe impacts on forest structure, function and biodiversity. Using a 105-year data set, we found increasing elevation extent of fires in the Sierra Nevada, and pose five hypotheses to explain this pattern. Beyond the recognized pattern of increasing fire frequency in the Sierra Nevada since the late 20th century, we find that the upper elevation extent of those fires has also been increasing. Factors such as fire season climate and fuel build up are recognized potential drivers of changes in fire regimes. Patterns of warming climate and increasing stand density are consistent with both the direction and magnitude of increasing elevation of wildfire. Reduction in high elevation wildfire suppression and increasing ignition frequencies may also contribute to the observed pattern. Historical biases in fire reporting are recognized, but not likely to explain the observed patterns. The four plausible mechanistic hypotheses (changes in fire management, climate, fuels, ignitions) are not mutually exclusive, and likely have synergistic interactions that may explain the observed changes. Irrespective of mechanism, the observed pattern of increasing occurrence of fire in these subalpine forests may have significant impacts on their resilience to changing climatic conditions.

"Future disruptions to fire activity will threaten ecosystems and human well-being throughout the world, yet there are few fire projections at global scales and almost none from a broad range of global climate models (GCMs). Here we integrate global fire datasets and environmental covariates to build spatial statistical models of fire probability at a 0.58 resolution and examine environmental controls on fire activity. Fire models are driven by climate norms from 16 GCMs (A2 emissions scenario) to assess the magnitude and direction of change over two time periods, 2010-2039 and 2070-2099. From the ensemble results, we identify areas of consensus for increases or decreases in fire activity, as well as areas where GCMs disagree. Although certain biomes are sensitive to constraints on biomass productivity and others to atmospheric conditions promoting combustion, substantial and rapid shifts are projected for future fire activity across vast portions of the globe. In the near term, the most consistent increases in fire activity occur in biomes with already somewhat warm climates; decreases are less pronounced and concentrated primarily in a few tropical and subtropical biomes. However, models do not agree on the direction of nearterm changes across more than 50% of terrestrial lands, highlighting major uncertainties in the next few decades. By the end of the century, the magnitude and the agreement in direction of change are projected to increase substantially. Most far-term model agreement on increasing fire probabilities (;62%) occurs at mid- to high-latitudes, while agreement on decreasing probabilities (;20%) is mainly in the tropics. Although our global models demonstrate that long-term environmental norms are very successful at capturing chronic fire probability patterns, future work is necessary to assess how much more explanatory power would be added through interannual variation in climate variables. This study provides a first examination of global disruptions to fire activity using an empirically based statistical framework and a multi-model ensemble of GCM projections, an important step toward assessing fire-related vulnerabilities to humans and the ecosystems upon which they depend."