Reduced fire severity offers near-term buffer to climate-driven declines in conifer resilience across the western United States.

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.

Large-scale disturbance legacies and the climate sensitivity of primary Picea abies forests.

Determining the drivers of shifting forest disturbance rates remains a pressing global change issue. Large-scale forest dynamics are commonly assumed to be climate driven, but appropriately scaled disturbance histories are rarely available to assess how disturbance legacies alter subsequent disturbance rates and the climate sensitivity of disturbance. We compiled multiple tree ring-based disturbance histories from primary Picea abies forest fragments distributed throughout five European landscapes spanning the Bohemian Forest and the Carpathian Mountains. The regional chronology includes 11,595 tree cores, with ring dates spanning the years 1750-2000, collected from 560 inventory plots in 37 stands distributed across a 1,000 km geographic gradient, amounting to the largest disturbance chronology yet constructed in Europe. Decadal disturbance rates varied significantly through time and declined after 1920, resulting in widespread increases in canopy tree age. Approximately 75% of current canopy area recruited prior to 1900. Long-term disturbance patterns were compared to an historical drought reconstruction, and further linked to spatial variation in stand structure and contemporary disturbance patterns derived from LANDSAT imagery. Historically, decadal Palmer drought severity index minima corresponded to higher rates of canopy removal. The severity of contemporary disturbances increased with each stand's estimated time since last major disturbance, increased with mean diameter, and declined with increasing within-stand structural variability. Reconstructed spatial patterns suggest that high small-scale structural variability has historically acted to reduce large-scale susceptibility and climate sensitivity of disturbance. Reduced disturbance rates since 1920, a potential legacy of high 19th century disturbance rates, have contributed to a recent region-wide increase in disturbance susceptibility. Increasingly common high-severity disturbances throughout primary Picea forests of Central Europe should be reinterpreted in light of both legacy effects (resulting in increased susceptibility) and climate change (resulting in increased exposure to extreme events).

Pre-outbreak forest conditions mediate the effects of spruce beetle outbreaks on fuels in subalpine forests of Colorado.

Over the past 30 years, forest disturbances have increased in size, intensity, and frequency globally, and are predicted to continue increasing due to climate change, potentially relaxing the constraints of vegetation properties on disturbance regimes. However, the consequences of the potentially declining importance of vegetation in determining future disturbance regimes are not well understood. Historically, bark beetles preferentially attack older trees and stands in later stages of development. However, as climate warming intensifies outbreaks by promoting growth of beetle populations and compromising tree defenses, smaller diameter trees and stands in early stages of development now are being affected by outbreaks. To date, no study has considered how stand age and other pre-outbreak forest conditions mediate the effects of outbreaks on surface and aerial fuel arrangements. We collected fuels data across a chronosequence of post-outbreak sites affected by spruce beetle (SB) between the 1940s and the 2010s, stratified by young (<130 yr) and old (>130 yr) post-fire stands. Canopy and surface fuel loads were calculated for each tree and stand, and available crown fuel load, crown bulk density, and canopy bulk densities were estimated. Canopy bulk density and density of live canopy individuals were reduced in all stands affected by SB, though foliage loss was proportionally greater in old stands as compared to young stands. Fine surface fuel loads in young stands were three times greater shortly (<30 yr) following outbreak as compared to young stands not affected by outbreak, after which the abundance of fine surface fuels decreased to below endemic (i.e., non-outbreak) levels. In both young and old stands, the net effect of SB outbreaks during the 20th and 21st centuries reduced total canopy fuels and increased stand-scale spatial heterogeneity of canopy fuels following outbreak. Importantly, the decrease in canopy fuels following outbreaks was greater in young post-fire stands than in older stands, suggesting that SB outbreaks may more substantially reduce risk of active crown fire when they affect stands in earlier stages of development. The current study shows that the effects of SB outbreaks on forest structure and on fuel profiles are strongly contingent on pre-outbreak conditions as determined by pre-outbreak disturbance history.

A walk on the wild side: Disturbance dynamics and the conservation and management of European mountain forest ecosystems.

Mountain forests are among the most important ecosystems in Europe as they support numerous ecological, hydrological, climatic, social, and economic functions. They are unique relatively natural ecosystems consisting of long-lived species in an otherwise densely populated human landscape. Despite this, centuries of intensive forest management in many of these forests have eclipsed evidence of natural processes, especially the role of disturbances in long-term forest dynamics. Recent trends of land abandonment and establishment of protected forests have coincided with a growing interest in managing forests in more natural states. At the same time, the importance of past disturbances highlighted in an emerging body of literature, and recent increasing disturbances due to climate change are challenging long-held views of dynamics in these ecosystems. Here, we synthesize aspects of this Special Issue on the ecology of mountain forest ecosystems in Europe in the context of broader discussions in the field, to present a new perspective on these ecosystems and their natural disturbance regimes. Most mountain forests in Europe, for which long-term data are available, show a strong and long-term effect of not only human land use but also of natural disturbances that vary by orders of magnitude in size and frequency. Although these disturbances may kill many trees, the forests themselves have not been threatened. The relative importance of natural disturbances, land use, and climate change for ecosystem dynamics varies across space and time. Across the continent, changing climate and land use are altering forest cover, forest structure, tree demography, and natural disturbances, including fires, insect outbreaks, avalanches, and wind disturbances. Projected continued increases in forest area and biomass along with continued warming are likely to further promote forest disturbances. Episodic disturbances may foster ecosystem adaptation to the effects of ongoing and future climatic change. Increasing disturbances, along with trends of less intense land use, will promote further increases in coarse woody debris, with cascading positive effects on biodiversity, edaphic conditions, biogeochemical cycles, and increased heterogeneity across a range of spatial scales. Together, this may translate to disturbance-mediated resilience of forest landscapes and increased biodiversity, as long as climate and disturbance regimes remain within the tolerance of relevant species. Understanding ecological variability, even imperfectly, is integral to anticipating vulnerabilities and promoting ecological resilience, especially under growing uncertainty. Allowing some forests to be shaped by natural processes may be congruent with multiple goals of forest management, even in densely settled and developed countries.

Relative importance of climate and mountain pine beetle outbreaks on the occurrence of large wildfires in the western USA.

Extensive outbreaks of bark beetles have killed trees across millions of hectares of forests and woodlands in western North America. These outbreaks have led to spirited scientific, public, and policy debates about consequential increases in fire risk, especially in the wildland-urban interface (WUI), where homes and communities are at particular risk from wildfires. At the same time, large wildfires have become more frequent across this region. Widespread expectations that outbreaks increase extent, severity, and/or frequency of wildfires are based partly on visible and dramatic changes in foliar moisture content and other fuel properties following outbreaks, as well as associated modeling projections. A competing explanation is that increasing wildfires are driven primarily by climatic extremes, which are becoming more common with climate change. However, the relative importance of bark beetle outbreaks vs. climate on fire occurrence has not been empirically examined across very large areas and remains poorly understood. The most extensive outbreaks of tree-killing insects across the western United States have been of mountain pine beetle (MPB; Dendroctonus ponderosae), which have killed trees over >650,000 km2 , mostly in forests dominated by lodgepole pine (Pinus contorta). We show that outbreaks of MPB in lodgepole pine forests of the western United States have been less important than climatic variability for the occurrence of large fires over the past 29 years. In lodgepole pine forests in general, as well as those in the WUI, occurrence of large fires was determined primarily by current and antecedent high temperatures and low precipitation but was unaffected by preceding outbreaks. Trends of increasing co-occurrence of wildfires and outbreaks are due to a common climatic driver rather than interactions between these disturbances. Reducing wildfire risk hinges on addressing the underlying climatic drivers rather than treating beetle-affected forests.

Large carbon release legacy from bark beetle outbreaks across Western United States.

Warmer conditions over the past two decades have contributed to rapid expansion of bark beetle outbreaks killing millions of trees over a large fraction of western United States (US) forests. These outbreaks reduce plant productivity by killing trees and transfer carbon from live to dead pools where carbon is slowly emitted to the atmosphere via heterotrophic respiration which subsequently feeds back to climate change. Recent studies have begun to examine the local impacts of bark beetle outbreaks in individual stands, but the full regional carbon consequences remain undocumented for the western US. In this study, we quantify the regional carbon impacts of the bark beetle outbreaks taking place in western US forests. The work relies on a combination of postdisturbance forest regrowth trajectories derived from forest inventory data and a process-based carbon cycle model tracking decomposition, as well as aerial detection survey (ADS) data documenting the regional extent and severity of recent outbreaks. We find that biomass killed by bark beetle attacks across beetle-affected areas in western US forests from 2000 to 2009 ranges from 5 to 15 Tg C yr(-1) and caused a reduction of net ecosystem productivity (NEP) of about 6.1-9.3 Tg C y(-1) by 2009. Uncertainties result largely from a lack of detailed surveys of the extent and severity of outbreaks, calling out a need for improved characterization across western US forests. The carbon flux legacy of 2000-2009 outbreaks will continue decades into the future (e.g., 2040-2060) as committed emissions from heterotrophic respiration of beetle-killed biomass are balanced by forest regrowth and accumulation.

Drought induces spruce beetle (Dendroctonus rufipennis) outbreaks across northwestern Colorado.

This study examines influences of climate variability on spruce beetle (Dendroctonus rufipennis) outbreak across northwestern Colorado during the period 1650 2011 CE. Periods of broad-scale outbreak reconstructed using documentary records and tree rings were dated to 1843-1860, 1882-1889, 1931-1957, and 2004-2010. Periods of outbreak were compared with seasonal temperature, precipitation, vapor pressure deficit (VPD), the Palmer Drought Severity Index (PDSI), and indices of ocean-atmosphere oscillation that include the El Niño Southern Oscillation (ENSO), Pacific Decadal Oscillation (PDO), and Atlantic Multidecadal Oscillation (AMO). Classification trees showed that outbreaks can be predicted most successfully from above average annual AMO values and above average summer VPD values, indicators of drought across Colorado. Notably, we find that spruce beetle outbreaks appear to be predicted best by interannual to multidecadal variability in drought, not by temperature alone. This finding may imply that spruce beetle outbreaks are triggered by decreases in host tree defenses, which are hypothesized to occur with drought stress. Given the persistence of the AMO, the shift to a positive AMO phase in the late 1990s is likely to promote continued spruce beetle disturbance.

Patterns and drivers of recent disturbances across the temperate forest biome.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001-2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes.

Effect of prior disturbances on the extent and severity of wildfire in Colorado subalpine forests.

Disturbances are important in creating spatial heterogeneity of vegetation patterns that in turn may affect the spread and severity of subsequent disturbances. Between 1997 and 2002 extensive areas of subalpine forests in northwestern Colorado were affected by a blowdown of trees, bark beetle outbreaks, and salvage logging. Some of these stands were also affected by severe fires in the late 19th century. During a severe drought in 2002, fires affected extensive areas of these subalpine forests. We evaluated and modeled the extent and severity of the 2002 fires in relation to these disturbances that occurred over the five years prior to the fires and in relation to late 19th century stand-replacing fires. Occurrence of disturbances prior to 2002 was reconstructed using a combination of tree-ring methods, aerial photograph interpretation, field surveys, and geographic information systems (GIS). The extent and severity of the 2002 fires were based on the normalized difference burn ratio (NDBR) derived from satellite imagery. GIS and classification trees were used to analyze the effects of prefire conditions on the 2002 fires. Previous disturbance history had a significant influence on the severity of the 2002 fires. Stands that were severely blown down (> 66% trees down) in 1997 burned more severely than other stands, and young (approximately 120 year old) postfire stands burned less severely than older stands. In contrast, prefire disturbances were poor predictors of fire extent, except that young (approximately 120 years old) postfire stands were less extensively burned than older stands. Salvage logging and bark beetle outbreaks that followed the 1997 blowdown (within the blowdown as well as in adjacent forest that was not blown down) did not appear to affect fire extent or severity. Conclusions regarding the influence of the beetle outbreaks on fire extent and severity are limited, however, by spatial and temporal limitations associated with aerial detection surveys of beetle activity. Thus, fire extent in these forests is largely independent of prefire disturbance history and vegetation conditions. In contrast, fire severity, even during extreme fire weather and in conjunction with a multiyear drought, is influenced by prefire stand conditions, including the history of previous disturbances.

Multidecadal climate variability and climate interactions affect subalpine fire occurrence, western Colorado (USA).

This study investigates the influence of climatic variability on subalpine forest fire occurrence in western Colorado during the AD 1600-2003 period. Interannual and multidecadal relationships between fire occurrence and the El Niño Southern Oscillation (ENSO), Pacific Decadal Oscillation (PDO), and Atlantic Multidecadal Oscillation (AMO) were examined, in addition to the effects of phase interactions among these oscillations. Fires occurred during short-term periods of significant drought and extreme cool (negative) phases of ENSO and PDO and during positive departures from mean AMO index. At longer time scales, fires exhibited 20-year periods of synchrony with the cool phase of the PDO, and 80-year periods of synchrony with extreme warm (positive) phases of the AMO. Years of combined positive AMO and negative ENSO and PDO phases represent "triple whammies" that significantly increased the occurrence of drought-induced fires. Fires were synchronous with this phase combination over 0-30 year periods and distinctly asynchronous with the opposite phase combination. Overall, because fires are synchronous at supra-annual to multidecadal time scales with warm AMO events, particularly when combined with cool ENSO and PDO phases, this suggests that we may be entering a qualitatively different fire regime in the next few decades due to the recent shift in 1998 to a likely long-term warm AMO phase. Although uncertainty remains regarding the effects of CO2-induced warming at regional scales, given the multidecadal persistence of the AMO there is mounting evidence that the recent shift to the positive phase of the AMO will promote higher fire frequencies in the region.

Fire Severity Controlled Susceptibility to a 1940s Spruce Beetle Outbreak in Colorado, USA.

The frequency, magnitude, and size of forest disturbances are increasing globally. Much recent research has focused on how the occurrence of one disturbance may affect susceptibility to subsequent disturbances. While much has been learned about such linked disturbances, the strength of the interactions is likely to be contingent on the severity of disturbances as well as climatic conditions, both of which can affect disturbance intensity and tree resistance to disturbances. Subalpine forests in western Colorado were affected by extensive and severe wildfires in the late 19th century and an extensive and severe outbreak of spruce beetle (Dendroctonus rufipennis) in the 1940s. Previous research found that most, but not all, of the stands that burned and established following the late 19th century fires were not susceptible to the 1940s outbreak as beetles preferentially attack larger trees and stands in advanced stages of development. However, previous research also left open the possibility that some stands that burned and established following the 19th century fires may have been attacked during the 1940s outbreak. Understanding how strongly stand structure, as shaped by disturbances of varying severity, affected susceptibility to past outbreaks is important to provide a baseline for assessing the degree to which recent climate change may be relaxing the preferences of beetles for larger trees and for stands in latter stages of structural development and thereby changing the nature of linked disturbances. Here, dendroecological methods were used to study disturbance history and tree age of stands in the White River National Forest in Western Colorado that were identified in historical documents or remotely-sensed images as having burned in the 19th century and having been attacked by spruce beetle in the 1940s. Dendroecological reconstructions indicate that in young post-fire stands only old remnant trees that survived the otherwise stand-replacing fires were killed in the 1940s outbreak. No young post-fire trees (< ca. 128 years) were susceptible to the 1940s outbreak, implying that under the relatively cool and wet conditions of the mid-20th century, susceptibility to and spatial patterns of spruce beetle outbreak were most likely controlled by variations in severity of prior disturbance by fire. This study provides a baseline for comparing linked disturbances under the relatively warmer and drier conditions of recent (e.g. post-1990) outbreaks in order to assess how climate mitigates the degree to which pre-disturbance history and structure affect susceptibility to disturbances.

Negative feedbacks on bark beetle outbreaks: widespread and severe spruce beetle infestation restricts subsequent infestation.

Understanding disturbance interactions and their ecological consequences remains a major challenge for research on the response of forests to a changing climate. When, where, and how one disturbance may alter the severity, extent, or occurrence probability of a subsequent disturbance is encapsulated by the concept of linked disturbances. Here, we evaluated 1) how climate and forest habitat variables, including disturbance history, interact to drive 2000s spruce beetle (Dendroctonus rufipennis) infestation of Engelmann spruce (Picea engelmannii) across the Southern Rocky Mountains; and 2) how previous spruce beetle infestation affects subsequent infestation across the Flat Tops Wilderness in northwestern Colorado, which experienced a severe landscape-scale spruce beetle infestation in the 1940s. We hypothesized that drought and warm temperatures would promote infestation, whereas small diameter and non-host trees, which may reflect past disturbance by spruce beetles, would inhibit infestation. Across the Southern Rocky Mountains, we found that climate and forest structure interacted to drive the 2000s infestation. Within the Flat Tops study area we found that stands infested in the 1940s were composed of higher proportions of small diameter and non-host trees ca. 60 years later. In this area, the 2000s infestation was constrained by a paucity of large diameter host trees (> 23 cm at diameter breast height), not climate. This suggests that there has not been sufficient time for trees to grow large enough to become susceptible to infestation. Concordantly, we found no overlap between areas affected by the 1940s infestation and the current infestation. These results show a severe spruce beetle infestation, which results in the depletion of susceptible hosts, can create a landscape template reducing the potential for future infestations.