Wildfire-Driven Forest Conversion in Western North American Landscapes.

Changing disturbance regimes and climate can overcome forest ecosystem resilience. Following high-severity fire, forest recovery may be compromised by lack of tree seed sources, warmer and drier postfire climate, or short-interval reburning. A potential outcome of the loss of resilience is the conversion of the prefire forest to a different forest type or nonforest vegetation. Conversion implies major, extensive, and enduring changes in dominant species, life forms, or functions, with impacts on ecosystem services. In the present article, we synthesize a growing body of evidence of fire-driven conversion and our understanding of its causes across western North America. We assess our capacity to predict conversion and highlight important uncertainties. Increasing forest vulnerability to changing fire activity and climate compels shifts in management approaches, and we propose key themes for applied research coproduced by scientists and managers to support decision-making in an era when the prefire forest may not return.

Seasonal and drought-related changes in leaf area profiles depend on height and light environment in an Amazon forest.

Seasonal dynamics in the vertical distribution of leaf area index (LAI) may impact the seasonality of forest productivity in Amazonian forests. However, until recently, fine-scale observations critical to revealing ecological mechanisms underlying these changes have been lacking. To investigate fine-scale variation in leaf area with seasonality and drought we conducted monthly ground-based LiDAR surveys over 4 yr at an Amazon forest site. We analysed temporal changes in vertically structured LAI along axes of both canopy height and light environments. Upper canopy LAI increased during the dry season, whereas lower canopy LAI decreased. The low canopy decrease was driven by highly illuminated leaves of smaller trees in gaps. By contrast, understory LAI increased concurrently with the upper canopy. Hence, tree phenological strategies were stratified by height and light environments. Trends were amplified during a 2015-2016 severe El Niño drought. Leaf area low in the canopy exhibited behaviour consistent with water limitation. Leaf loss from short trees in high light during drought may be associated with strategies to tolerate limited access to deep soil water and stressful leaf environments. Vertically and environmentally structured phenological processes suggest a critical role of canopy structural heterogeneity in seasonal changes in Amazon ecosystem function.

Pre-fire drought and competition mediate post-fire conifer mortality in western U.S. National Parks.

Tree mortality is an important outcome of many forest fires. Extensive tree injuries from fire may lead directly to mortality, but environmental and biological stressors may also contribute to tree death. However, there is little evidence showing how the combined effects of two common stressors, drought and competition, influence post-fire mortality. Geographically broad observations of three common western coniferous trees subjected to prescribed fire showed the likelihood of post-fire mortality was related to intermediate-term (10 yr) pre-fire average radial growth, an important component of tree vigor. Path analysis showed that indices of competition and drought stress prior to fire can be described in terms of joint effects on growth, indirectly affecting post-fire mortality. Our results suggest that the conditions that govern the relationship between growth and mortality in unburned stands may also apply to post-fire environments. Thus, biotic and abiotic changes that affect growth negatively (e.g., drought stress) or positively (e.g., growth releases following thinning treatments) prior to fire may influence expressed fire severity, independent of fire intensity (e.g., heat flux, residence time). These relationships suggest that tree mortality may increase under stressful climatic or stand conditions even if fire behavior remains constant.

Direct and indirect climate controls predict heterogeneous early-mid 21st century wildfire burned area across western and boreal North America.

Predicting wildfire under future conditions is complicated by complex interrelated drivers operating across large spatial scales. Annual area burned (AAB) is a useful index of global wildfire activity. Current and antecedent seasonal climatic conditions, and the timing of snowpack melt, have been suggested as important drivers of AAB. As climate warms, seasonal climate and snowpack co-vary in intricate ways, influencing fire at continental and sub-continental scales. We used independent records of seasonal climate and snow cover duration (last date of permanent snowpack, LDPS) and cell-based Structural Equation Models (SEM) to separate direct (climatic) and indirect (snow cover) effects on relative changes in AAB under future climatic scenarios across western and boreal North America. To isolate seasonal climate variables with the greatest effect on AAB, we ran multiple regression models of log-transformed AAB on seasonal climate variables and LDPS. We used the results of multiple regressions to project future AAB using GCM ensemble climate variables and LDPS, and validated model predictions with recent AAB trends. Direct influences of spring and winter temperatures on AAB are larger and more widespread than the indirect effect mediated by changes in LDPS in most areas. Despite significant warming trends and reductions in snow cover duration, projected responses of AAB to early-mid 21st century are heterogeneous across the continent. Changes in AAB range from strongly increasing (one order of magnitude increases in AAB) to moderately decreasing (more than halving of baseline AAB). Annual wildfire area burned in coming decades is likely to be highly geographically heterogeneous, reflecting interacting regional and seasonal climate drivers of fire occurrence and spread.

Disturbance and productivity interactions mediate stability of forest composition and structure.

Fire is returning to many conifer-dominated forests where species composition and structure have been altered by fire exclusion. Ecological effects of these fires are influenced strongly by the degree of forest change during the fire-free period. Response of fire-adapted species assemblages to extended fire-free intervals is highly variable, even in communities with similar historical fire regimes. This variability in plant community response to fire exclusion is not well understood; however, ecological mechanisms such as individual species' adaptations to disturbance or competition and underlying site characteristics that facilitate or impede establishment and growth have been proposed as potential drivers of assemblage response. We used spatially explicit dendrochronological reconstruction of tree population dynamics and fire regimes to examine the influence of historical disturbance frequency (a proxy for adaptation to disturbance or competition), and potential site productivity (a proxy for underlying site characteristics) on the stability of forest composition and structure along a continuous ecological gradient of pine, dry mixed-conifer, mesic mixed-conifer, and spruce-fir forests following fire exclusion. While average structural density increased in all forests, species composition was relatively stable in the lowest productivity pine-dominated and highest productivity spruce-fir-dominated sites immediately following fire exclusion and for the next 100 years, suggesting site productivity as a primary control on species composition and structure in forests with very different historical fire regimes. Species composition was least stable on intermediate productivity sites dominated by mixed-conifer forests, shifting from primarily fire-adapted species to competition-adapted, fire-sensitive species within 20 years of fire exclusion. Rapid changes to species composition and stand densities have been interpreted by some as evidence of high-severity fire. We demonstrate that the very different ecological process of fire exclusion can produce similar changes by shifting selective pressures from disturbance-mediated to productivity-mediated controls. Restoring disturbance-adapted species composition and structure to intermediate productivity forests may help to buffer them against projected increasing temperatures, lengthening fire seasons, and more frequent and prolonged moisture stress. Fewer management options are available to promote adaptation in forest assemblages historically constrained by underlying site productivity.

Average Stand Age from Forest Inventory Plots Does Not Describe Historical Fire Regimes in Ponderosa Pine and Mixed-Conifer Forests of Western North America.

Quantifying historical fire regimes provides important information for managing contemporary forests. Historical fire frequency and severity can be estimated using several methods; each method has strengths and weaknesses and presents challenges for interpretation and verification. Recent efforts to quantify the timing of historical high-severity fire events in forests of western North America have assumed that the "stand age" variable from the US Forest Service Forest Inventory and Analysis (FIA) program reflects the timing of historical high-severity (i.e. stand-replacing) fire in ponderosa pine and mixed-conifer forests. To test this assumption, we re-analyze the dataset used in a previous analysis, and compare information from fire history records with information from co-located FIA plots. We demonstrate that 1) the FIA stand age variable does not reflect the large range of individual tree ages in the FIA plots: older trees comprised more than 10% of pre-stand age basal area in 58% of plots analyzed and more than 30% of pre-stand age basal area in 32% of plots, and 2) recruitment events are not necessarily related to high-severity fire occurrence. Because the FIA stand age variable is estimated from a sample of tree ages within the tree size class containing a plurality of canopy trees in the plot, it does not necessarily include the oldest trees, especially in uneven-aged stands. Thus, the FIA stand age variable does not indicate whether the trees in the predominant size class established in response to severe fire, or established during the absence of fire. FIA stand age was not designed to measure the time since a stand-replacing disturbance. Quantification of historical "mixed-severity" fire regimes must be explicit about the spatial scale of high-severity fire effects, which is not possible using FIA stand age data.

Multiscale perspectives of fire, climate and humans in western North America and the Jemez Mountains, USA.

Interannual climate variations have been important drivers of wildfire occurrence in ponderosa pine forests across western North America for at least 400 years, but at finer scales of mountain ranges and landscapes human land uses sometimes over-rode climate influences. We reconstruct and analyse effects of high human population densities in forests of the Jemez Mountains, New Mexico from ca 1300 CE to Present. Prior to the 1680 Pueblo Revolt, human land uses reduced the occurrence of widespread fires while simultaneously adding more ignitions resulting in many small-extent fires. During the 18th and 19th centuries, wet/dry oscillations and their effects on fuels dynamics controlled widespread fire occurrence. In the late 19th century, intensive livestock grazing disrupted fuels continuity and fire spread and then active fire suppression maintained the absence of widespread surface fires during most of the 20th century. The abundance and continuity of fuels is the most important controlling variable in fire regimes of these semi-arid forests. Reduction of widespread fires owing to reduction of fuel continuity emerges as a hallmark of extensive human impacts on past forests and fire regimes.This article is part of the themed issue 'The interaction of fire and mankind'.

Resin duct size and density as ecophysiological traits in fire scars of Pseudotsuga menziesii and Larix occidentalis.

BACKGROUND AND AIMS: Resin ducts (RDs) are features present in most conifer species as defence structures against pests and pathogens; however, little is known about RD expression in trees following fire injury. This study investigates changes in RD size and density in fire scars of Douglas fir (Pseudotsuga menziesii) and western larch (Larix occidentalis) as a means to evaluate the ecophysiological significance of traumatic resinosis for tree defence and survival. METHODS: Transverse and tangential microsections were prepared for light microscopy and image analysis in order to analyse axial and radial RDs, respectively. Epithelial cells of RDs and fusiform rays associated with radial RDs were also examined. RDs were compared between normal xylem and wound xylem at four different section heights along the fire-injured stem. KEY RESULTS: Following fire injury, P. menziesii axial RDs narrowed by 38-43 % in the first year after injury, and the magnitude of this change increased with stem height. Larix occidentalis axial RDs widened by 46-50 % in the second year after injury. Radial RDs were of equivalent size in P. menziesii, but widened by 162-214 % in L. occidentalis. Fusiform rays were larger following fire injury, by 4-14 % in P. menziesii and by 23-38 % in L. occidentalis. Furthermore, axial RD density increased in both species due to the formation of tangential rows of traumatic RDs, especially in the first and second years after injury. However, radial RD density did not change significantly. CONCLUSIONS: These results highlight traumatic resinosis as a species-specific response. Pseudotsuga menziesii produce RDs of equivalent or reduced size, whereas L. occidentalis produce wider RDs in both the axial and radial duct system, thereby increasing resin biosynthesis and accumulation within the whole tree. Larix occidentalis thus appears to allocate more energy to defence than P. menziesii.

Changes in tracheid and ray traits in fire scars of North American conifers and their ecophysiological implications.

BACKGROUND AND AIMS: Fire scars have been widely used as proxies for the reconstruction of fire history; however, little is known about the impact of fire injury on wood anatomy. This study investigates changes in tracheid and ray traits in fire scars of Douglas fir (Pseudotsuga menziesii), western larch (Larix occidentalis) and ponderosa pine (Pinus ponderosa), and discusses their ecophysiological implications for tree recovery from fire. METHODS: Transverse and tangential microsections were prepared for light microscopy and image analysis. Measurements of tracheids and rays were made in the three spatial dimensions: axially (at different section heights), radially (in different rings) and tangentially (with increasing distance from the wound margin). KEY RESULTS: Changes were strongest in the first year after fire injury, with a decrease in tracheid size (by 25-30 %) and an increase in tracheid density (by 21-53 %) for the three species. In addition, an increase in ray size (by 5-27 %) and an increase in ray density (by 19-36 %) were found in P. menziesii and L. occidentalis. Changes were comparable along the fire-injured stem and were often most marked close to the fire scar. CONCLUSIONS: The differentiation after fire injury of narrower and more numerous tracheids expresses a trade-off between hydraulic safety and hydraulic efficiency, while that of larger and more numerous rays serves compartmentalization and wound closure, mechanical strength and defence responses. Pinus ponderosa does not generally produce more ray tissue after fire injury and thus appears to be more adapted to fire.

Testing a pyroclimatic hypothesis on the Mexico-United States border.

The "pyroclimatic hypothesis" proposed by F. Biondi and colleagues provides a basis for testable expectations about climatic and other controls of fire regimes. This hypothesis asserts an a priori relationship between the occurrence of widespread fire and values of a relevant climatic index. Such a hypothesis provides the basis for predicting spatial and temporal patterns of fire occurrence based on climatic control. Forests near the Mexico-United States border offer a place to test the relative influence of climatic and other controls in mountain ranges that are ecologically similar and subject to broadly similar top-down climatic influence, but with differing cultural influences. We tested the pyroclimatic hypothesis by comparing fire history information from the Mesa de las Guacamayas, a mountain range in northwestern Chihuahua, with previously published fire data from the Chiricahua Mountains, in southeastern Arizona, approximately 150 km away. We developed a priori hypothetical models of fire occurrence and compared their performance to empirical climate-based models. Fires were frequent at all Mesa de las Guacamayas study sites through the mid-20th century and continued uninterrupted to the present at one site, in contrast to nearly complete fire exclusion after 1892 at sites in the Chiricahua Mountains. The empirical regression models explained a higher proportion of the variability in fire regime associated with climate than did the a priori models. Actual climate-fire relationships diverged in each country after 1892. The a priori models predicted continuing fires at the same rate per century as prior to 1892; fires did in fact continue in Mexico, albeit with some alteration of fire regimes, but ceased in the United States, most likely due to changes in land use. The cross-border comparison confirms that a frequent-fire regime could cease without a climatic cause, supporting previous arguments that bottom-up factors such as livestock grazing can rapidly and drastically alter surface fire regimes. Understanding the historical patterns of climate controls on fire could inform the use of historical data as ecological reference conditions and for future sustainability.

Spatial and temporal corroboration of a fire-scar-based fire history in a frequently burned ponderosa pine forest.

Fire scars are used widely to reconstruct historical fire regime parameters in forests around the world. Because fire scars provide incomplete records of past fire occurrence at discrete points in space, inferences must be made to reconstruct fire frequency and extent across landscapes using spatial networks of fire-scar samples. Assessing the relative accuracy of fire-scar fire history reconstructions has been hampered due to a lack of empirical comparisons with independent fire history data sources. We carried out such a comparison in a 2780-ha ponderosa pine forest on Mica Mountain in southern Arizona (USA) for the time period 1937-2000. Using documentary records of fire perimeter maps and ignition locations, we compared reconstructions of key spatial and temporal fire regime parameters developed from documentary fire maps and independently collected fire-scar data (n = 60 plots). We found that fire-scar data provided spatially representative and complete inventories of all major fire years (> 100 ha) in the study area but failed to detect most small fires. There was a strong linear relationship between the percentage of samples recording fire scars in a given year (i.e., fire-scar synchrony) and total area burned for that year (y = 0.0003x + 0.0087, r2 = 0.96). There was also strong spatial coherence between cumulative fire frequency maps interpolated from fire-scar data and ground-mapped fire perimeters. Widely reported fire frequency summary statistics varied little between fire history data sets: fire-scar natural fire rotations (NFR) differed by < 3 yr from documentary records (29.6 yr); mean fire return intervals (MFI) for large-fire years (i.e., > or = 25% of study area burned) were identical between data sets (25.5 yr); fire-scar MFIs for all fire years differed by 1.2 yr from documentary records. The known seasonal timing of past fires based on documentary records was furthermore reconstructed accurately by observing intra-annual ring position of fire scars and using knowledge of tree-ring growth phenology in the Southwest. Our results demonstrate clearly that representative landscape-scale fire histories can be reconstructed accurately from spatially distributed fire-scar samples.

El Niño-southern oscillation effect on a fire regime in northeastern Mexico has changed over time.

The El Niño Southern Oscillation (ENSO) is a climate-forcing mechanism that has been shown to affect precipitation and the occurrence of wildfires in many parts of the world. In the southern United States and northern Mexico, warm events (El Niño) are associated with moist winter conditions and fewer fires, while cool events (La Niñia) tend to favor dry winters and more fires. We tested this relationship in a region of northeastern Mexico by characterizing the historical fire regime and climatic influences: Fire regimes were reconstructed from fire-scar samples collected from 100 trees in three high-elevation sites on Peña Nevada in southern Nuevo Le6n. The sites were approximately 25 ha each, and the site centers were approximately 1 km apart. The earliest recorded fire occurred in 1521 and the time period we used for analysis was 1645-1929. The sites were characterized by frequent surface fires before the 1920s. In the three sites, mean fire intervals ranged from 8.6 to 9.6 years (all fires) and 11.9 to 18.6 years (fires that scarred > or = 25% of recording trees). The per-tree mean fire return interval was 17 years, and all three sites burned in the same year seven times between 1774 and 1929. After 1929, fires were nearly eliminated in all sites, likely due to human causes. We found a temporal change in the association between ENSO events and fires; before the 1830s La Niña events were significantly associated with fire years, while after the 1830s this association was not significant. In 1998, when the most severe El Niño event of the past century occurred, the three sites experienced severe, stand-replacing fires that killed many trees that had survived multiple surface fires in the past. Prior to the 1830s, fires tended to occur during dry La Niña years, but since then both La Niña and El Niño have been associated with dry years in this region, especially during the last three decades. This result suggests that ENSO effects have changed over time in this location and that phases of ENSO are not consistent indicators of precipitation, fire occurrence, or fire behavior in this area of northeastern Mexico.