Land management explains major trends in forest structure and composition over the last millennium in California's Klamath Mountains.

SignificanceWe provide the first assessment of aboveground live tree biomass in a mixed conifer forest over the late Holocene. The biomass record, coupled with local Native oral history and fire scar records, shows that Native burning practices, along with a natural lightning-based fire regime, promoted long-term stability of the forest structure and composition for at least 1 millennium in a California forest. This record demonstrates that climate alone cannot account for observed forest conditions. Instead, forests were also shaped by a regime of frequent fire, including intentional ignitions by Native people. This work suggests a large-scale intervention could be required to achieve the historical conditions that supported forest resiliency and reflected Indigenous influence.

Persistent yet vulnerable: resurvey of an Abies ecotone reveals few differences but vulnerability to climate change.

Climate change is shifting forest tree species distributions across elevational and latitudinal gradients, and these changes are often pronounced at ecotones where species meet their climatic bounds and are replaced by other species. Using an extensive ecotone composed of lower-montane white fir (Abies concolor var. lowiana) and upper-montane red fir (Abies magnifica var. magnifica) in the central Sierra Nevada range of California, USA, we (1) examined how the demographics of the ecotone have responded to recent climate using a field observational study and a historical dataset, (2) quantified climate drivers across species life stages using contemporary demographic data, and (3) tested the potential impacts of future climate on species-specific seedling survival and growth in a fully factorial growth chamber experiment that varied temperature, growing season length, and water availability. A re-examination of the ecotone midpoint after 35 yr suggested a reduction in A. concolor sapling and tree densities and a rise in A. magnifica proportional dominance between surveys. Seedling abundances across the ecotone indicated that A. magnifica tends to dominate the regeneration layer and currently forms an important component of the seedling community at elevations below those where A. magnifica saplings or trees begin to co-dominate stands. Observational and experimental assessments suggest that temperature and precipitation serve as important drivers, differentiating A. concolor vs. A. magnifica distributions, and are primary stressors at the seedling stage. Seedlings of both species were adversely affected by experimental climate treatments, although A. concolor exhibited greater survival and a more conservative growth strategy under extreme climatic stress than A. magnifica. Projections indicate that historical climate conditions will rise by an amount greater than the ecotone's current elevational extent by the end of the 21st century. Differential drivers of species abundances suggest that the projected climate will expand conditions that promote A. concolor abundance and impede A. magnifica abundance across the ecotone; however, disturbance activity and microclimatic conditions will also influence regeneration and overstory tree dynamics. Our study demonstrates the importance of quantifying species-specific responses to climate and indicates that widespread regeneration failure may be one possible consequence in which species exhibit strong sensitivity to projected climate conditions.

Pyric tree spatial patterning interactions in historical and contemporary mixed conifer forests, California, USA.

Tree spatial patterns in dry coniferous forests of the western United States, and analogous ecosystems globally, were historically aggregated, comprising a mixture of single trees and groups of trees. Modern forests, in contrast, are generally more homogeneous and overstocked than their historical counterparts. As these modern forests lack regular fire, pattern formation and maintenance is generally attributed to fire. Accordingly, fires in modern forests may not yield historically analogous patterns. However, direct observations on how selective tree mortality among pre-existing forest structure shapes tree spatial patterns is limited. In this study, we (a) simulated fires in historical and contemporary counterpart plots in a Sierra Nevadan mixed-conifer forest, (b) estimated tree mortality, and (c) examined tree spatial patterns of live trees before and after fire, and of fire-killed trees. Tree mortality in the historical period was clustered and density-dependent, because trees were aggregated and segregated by tree size before fire. Thus, fires maintained an aggregated distribution of tree groups. Tree mortality in the contemporary period was widespread, except for dispersed large trees, because most trees were a part of large, interconnected tree groups. Thus, postfire tree patterns were more uniform and devoid of moderately sized tree groups. Postfire tree patterns in the historical period, unlike the contemporary period, were within the historical range of variability identified for the western United States. This divergence suggests that decades of forest dynamics without significant disturbances have altered the historical means of pyric pattern formation. Our results suggest that ecological silvicultural treatments, such as forest restoration thinnings, which emulate qualities of historical forests may facilitate the reintroduction of fire as a means to reinforce forest structural heterogeneity.

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.

Long-term effects of fire severity on oak-conifer dynamics in the southern Cascades.

We studied vegetation composition and structure in a mixed conifer-oak ecosystem across a range of fire severity 10 years following wildfire. Sample plots centered on focal California black oaks (Quercus kelloggii) were established to evaluate oak and neighboring tree and shrub recovery across a gradient of fire severity in the southern Cascade Range, USA. Shrub and oak resprouting was strongest around focal oaks where conifer mortality was greatest. Linear modeling revealed negative relationships between California black oak sprout height or basal area and residual overstory tree survival, primarily white fir (Abies concolor). The two dominant competing species, California black oak and white fir, showed opposite responses to fire severity. Sprouting California black oak and associated shrubs dominated in severely burned areas, while surviving, non-sprouting white fir maintained dominance by its height advantage and shading effects in areas that burned with low fire severity. Our results indicate that high-severity fire promotes persistence and restoration of ecosystems containing resprouting species, such as California black oak, that are increasingly rare due to widespread fire exclusion in landscapes that historically experienced more frequent fire. We present a conceptual model based on our results and supported by a synthesis of postfire resprouting dynamics literature. Our results and conceptual model help illuminate long-term postfire vegetation responses and the potential ability of fire to catalyze formation of alternate vegetation community structures that may not be apparent in studies that evaluate postfire effects at shorter time-since-fire intervals or at coarser scales.

Climatic stress increases forest fire severity across the western United States.

Pervasive warming can lead to chronic stress on forest trees, which may contribute to mortality resulting from fire-caused injuries. Longitudinal analyses of forest plots from across the western US show that high pre-fire climatic water deficit was related to increased post-fire tree mortality probabilities. This relationship between climate and fire was present after accounting for fire defences and injuries, and appeared to influence the effects of crown and stem injuries. Climate and fire interactions did not vary substantially across geographical regions, major genera and tree sizes. Our findings support recent physiological evidence showing that both drought and heating from fire can impair xylem conductivity. Warming trends have been linked to increasing probabilities of severe fire weather and fire spread; our results suggest that warming may also increase forest fire severity (the number of trees killed) independent of fire intensity (the amount of heat released during a fire).

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

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

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

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

Pollen-limited reproduction in blue oak: implications for wind pollination in fragmented populations.

Human activities are fragmenting forests and woodlands worldwide, but the impact of reduced tree population densities on pollen transfer in wind-pollinated trees is poorly understood. In a 4-year study, we evaluated relationships among stand density, pollen availability, and seed production in a thinned and fragmented population of blue oak (Quercus douglasii). Geographic coordinates were established and flowering interval determined for 100 contiguous trees. The number of neighboring trees within 60 m that released pollen during each tree's flowering period was calculated and relationships with acorn production explored using multiple regression. We evaluated the effects of female flower production, average temperature, and relative humidity during the pollination period, and number of pollen-producing neighbors on individual trees' acorn production. All factors except temperature were significant in at least one of the years of our study, but the combination of factors influencing acorn production varied among years. In 1996, a year of large acorn crop size, acorn production was significantly positively associated with number of neighboring pollen producers and density of female flowers. In 1997, 1998, and 1999, many trees produced few or no acorns, and significant associations between number of pollen-producing neighbors and acorn production were only apparent among moderately to highly reproductive trees. Acorn production by these reproductive trees in 1997 was significantly positively associated with number of neighboring pollen producers and significantly negatively associated with average relative humidity during the pollination period. In 1998, no analysis was possible, because too few trees produced a moderate to large acorn crop. Only density of female flowers was significantly associated with acorn production of moderately to highly reproductive trees in 1999. The effect of spatial scale was also investigated by conducting analyses with pollen producers counted in radii ranging from 30 m to 80 m. The association between number of pollen-producing neighbors and acorn production was strongest when neighborhood sizes of 60 m or larger were considered. Our results suggest that fragmentation and thinning of blue oak woodlands may reduce pollen availability and limit reproduction in this wind-pollinated species.