Individual canopy tree species maps for the National Ecological Observatory Network.

The ecology of forest ecosystems depends on the composition of trees. Capturing fine-grained information on individual trees at broad scales provides a unique perspective on forest ecosystems, forest restoration, and responses to disturbance. Individual tree data at wide extents promises to increase the scale of forest analysis, biogeographic research, and ecosystem monitoring without losing details on individual species composition and abundance. Computer vision using deep neural networks can convert raw sensor data into predictions of individual canopy tree species through labeled data collected by field researchers. Using over 40,000 individual tree stems as training data, we create landscape-level species predictions for over 100 million individual trees across 24 sites in the National Ecological Observatory Network (NEON). Using hierarchical multi-temporal models fine-tuned for each geographic area, we produce open-source data available as 1 km2 shapefiles with individual tree species prediction, as well as crown location, crown area, and height of 81 canopy tree species. Site-specific models had an average performance of 79% accuracy covering an average of 6 species per site, ranging from 3 to 15 species per site. All predictions are openly archived and have been uploaded to Google Earth Engine to benefit the ecology community and overlay with other remote sensing assets. We outline the potential utility and limitations of these data in ecology and computer vision research, as well as strategies for improving predictions using targeted data sampling.

North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

The effects of ENSO and the North American monsoon on mast seeding in two Rocky Mountain conifer species.

We aimed to disentangle the patterns of synchronous and variable cone production (i.e. masting) and its relationship to climate in two conifer species native to dry forests of western North America. We used cone abscission scars to reconstruct ca 15 years of recent cone production in Pinus edulis and Pinus ponderosa, and used redundancy analysis to relate time series of annual cone production to climate indices describing the North American monsoon and the El Niño Southern Oscillation (ENSO). We show that the sensitivity to climate and resulting synchrony in cone production varies substantially between species. Cone production among populations of P. edulis was much more spatially synchronous and more closely related to large-scale modes of climate variability than among populations of P. ponderosa. Large-scale synchrony in P. edulis cone production was associated with the North American monsoon and we identified a dipole pattern of regional cone production associated with ENSO phase. In P. ponderosa, these climate indices were not strongly associated with cone production, resulting in asynchronous masting patterns among populations. This study helps frame our understanding of mast seeding as a life-history strategy and has implications for our ability to forecast mast years in these species. This article is part of the theme issue 'The ecology and evolution of synchronized seed production in plants'.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Population collapse and retreat to fire refugia of the Tasmanian endemic conifer Athrotaxis selaginoides following the transition from Aboriginal to European fire management.

Untangling the nuanced relationships between landscape, fire disturbance, human agency, and climate is key to understanding rapid population declines of fire-sensitive plant species. Using multiple lines of evidence across temporal and spatial scales (vegetation survey, stand structure analysis, dendrochronology, and fire history reconstruction), we document landscape-scale population collapse of the long-lived, endemic Tasmanian conifer Athrotaxis selaginoides in remote montane catchments in southern Tasmania. We contextualized the findings of this field-based study with a Tasmanian-wide geospatial analysis of fire-killed and unburned populations of the species. Population declines followed European colonization commencing in 1802 ad that disrupted Aboriginal landscape burning. Prior to European colonization, fire events were infrequent but frequency sharply increased afterwards. Dendrochronological analysis revealed that reconstructed fire years were associated with abnormally warm/dry conditions, with below-average streamflow, and were strongly teleconnected to the Southern Annular Mode. The multiple fires that followed European colonization caused near total mortality of A. selaginoides and resulted in pronounced floristic, structural vegetation, and fuel load changes. Burned stands have very few regenerating A. selaginoides juveniles yet tree-establishment reconstruction of fire-killed adults exhibited persistent recruitment in the period prior to European colonization. Collectively, our findings indicate that this fire-sensitive Gondwanan conifer was able to persist with burning by Aboriginal Tasmanians, despite episodic widespread forest fires. By contrast, European burning led to the restriction of A. selaginoides to prime topographic fire refugia. Increasingly, frequent fires caused by regional dry and warming trends and increased ignitions by humans and lightning are breaching fire refugia; hence, the survival Tasmanian Gondwanan species demands sustained and targeted fire management.

Forest recovery following synchronous outbreaks of spruce and western balsam bark beetle is slowed by ungulate browsing.

Understanding how severe disturbances and their interactions affect forests is key to projecting ecological change under a warming climate. Substantial increases in some biotic disturbances, such as bark beetle outbreaks, in temperate forest ecosystemsmay compromise recovery to a forest vegetation type (i.e., physiognomic recovery or resilience), especially if subsequent biotic disturbances (e.g., herbivory) alter recovery mechanisms. From 2005 to 2017, severe outbreaks (>90% mortality) of spruce bark beetles (SB, Dendroctonus rufipennis) affected Engelmann spruce (Picea engelmannii) across 325,000 ha of spruce and subalpine fir (Abies lasiocarpa) forest in the southern Rocky Mountains, USA. Concurrently, an outbreak of western balsam bark beetle (WBBB, Dryocoetes confuses) infested subalpine fir across at least 47,000 of these hectares. We explored the capacity of 105 stands affected by one or two bark beetle outbreaks and browsing of juvenile trees by ungulates to return to a forest vegetation type in the context of pre-outbreak forest conditions and topography. Nine initial forest trajectories (i.e., at least several decades) were identified from four pre-outbreak forest types affected by three biotic disturbances that occurred at different spatial scales and severities. Most stands (86%) contained surviving nonhost adult trees in the main canopy (fir and aspen [Populus tremuloides]) and many surviving juveniles of all species, implying that they are currently on a trajectory for physiognomic recovery. Stands composed exclusively of large-diameter spruce were affected by a severe SB outbreak and were most vulnerable to a transition to a low-density forest, below regional stocking levels (<370 trees/ha). Greater pre-outbreak stand structural complexity and species diversity were key traits of stands with a higher potential for physiognomic recovery. However, all multispecies stands shifted in relative composition of the main canopy to nonhost species, suggesting low potential for compositional recovery over the next several decades. Most post-outbreak stands (86%) exceeded regional stocking levels with trees taller than the browse zone (<2 m). As such, ungulate browsing on over half of all juveniles will primarily affect the rate of infilling of the forest canopy and preferential browsing of more palatable species will influence the composition of the future forest canopy.

Still standing: Recent patterns of post-fire conifer refugia in ponderosa pine-dominated forests of the Colorado Front Range.

Forested fire refugia (trees that survive fires) are important disturbance legacies that provide seed sources for post-fire regeneration. Conifer regeneration has been limited following some recent western fires, particularly in ponderosa pine (Pinus ponderosa) forests. However, the extent, characteristics, and predictability of ponderosa pine fire refugia are largely unknown. Within 23 fires in ponderosa pine-dominated forests of the Colorado Front Range (1996-2013), we evaluated the spatial characteristics and predictability of refugia: first using Monitoring Trends in Burn Severity (MTBS) burn severity metrics, then using landscape variables (topography, weather, anthropogenic factors, and pre-fire forest cover). Using 1-m resolution aerial imagery, we created a binary variable of post-fire conifer presence ('Conifer Refugia') and absence ('Conifer Absence') within 30-m grid cells. We found that maximum patch size of Conifer Absence was positively correlated with fire size, and 38% of the burned area was ≥ 50m from a conifer seed source, revealing a management challenge as fire sizes increase with warming further limiting conifer recovery. In predicting Conifer Refugia with two MTBS-produced databases, thematic burn severity classes (TBSC) and continuous Relative differenced Normalized Burn Ratio (RdNBR) values, Conifer Absence was high in previously forested areas of Low and Moderate burn severity classes in TBSC. RdNBR more accurately identified post-fire conifer survivorship. In predicting Conifer Refugia with landscape variables, Conifer Refugia were less likely during burn days with high maximum temperatures: while Conifer Refugia were more likely on moister soils and closer to higher order streams, homes, and roads; and on less rugged, valley topography. Importantly, pre-fire forest canopy cover was not strongly associated with Conifer Refugia. This study further informs forest management by mapping post-fire patches lacking conifer seed sources, validating the use of RdNBR for fire refugia, and detecting abiotic and topographic variables that may promote conifer refugia.

Limitations to recovery following wildfire in dry forests of southern Colorado and northern New Mexico, USA.

Climate warming is contributing to increases in wildfire activity throughout the western United States, leading to potentially long-lasting shifts in vegetation. The response of forest ecosystems to wildfire is thus a crucial indicator of future vegetation trajectories, and these responses are contingent upon factors such as seed availability, interannual climate variability, average climate, and other components of the physical environment. To better understand variation in resilience to wildfire across vulnerable dry forests, we surveyed conifer seedling densities in 15 recent (1988-2010) wildfires and characterized temporal variation in seed cone production and seedling establishment. We then predicted postfire seedling densities at a 30-m resolution within each fire perimeter using downscaled climate data, monthly water balance models, and maps of surviving forest cover. Widespread ponderosa pine (Pinus ponderosa) seed cone production occurred at least twice following each fire surveyed, and pulses of conifer seedling establishment coincided with years of above-average moisture availability. Ponderosa pine and Douglas-fir (Pseudotsuga menziesii) seedling densities were higher on more mesic sites and adjacent to surviving trees, though there were also important interspecific differences, likely attributable to drought and shade tolerance. We estimated that postfire seedling densities in 42% (for ponderosa pine) and 69% (for Douglas-fir) of the total burned area were below the lowest reported historical tree densities in these forests. Spatial models demonstrated that an absence of mature conifers (particularly in the interior of large, high-severity patches) limited seedling densities in many areas, but 30-yr average actual evapotranspiration and climatic water deficit limited densities on marginal sites. A better understanding of the limitations to postfire forest recovery will refine models of vegetation dynamics and will help to improve strategies of adaptation to a warming climate and shifting fire activity.

Wildfires and climate change push low-elevation forests across a critical climate threshold for tree regeneration.

Climate change is increasing fire activity in the western United States, which has the potential to accelerate climate-induced shifts in vegetation communities. Wildfire can catalyze vegetation change by killing adult trees that could otherwise persist in climate conditions no longer suitable for seedling establishment and survival. Recently documented declines in postfire conifer recruitment in the western United States may be an example of this phenomenon. However, the role of annual climate variation and its interaction with long-term climate trends in driving these changes is poorly resolved. Here we examine the relationship between annual climate and postfire tree regeneration of two dominant, low-elevation conifers (ponderosa pine and Douglas-fir) using annually resolved establishment dates from 2,935 destructively sampled trees from 33 wildfires across four regions in the western United States. We show that regeneration had a nonlinear response to annual climate conditions, with distinct thresholds for recruitment based on vapor pressure deficit, soil moisture, and maximum surface temperature. At dry sites across our study region, seasonal to annual climate conditions over the past 20 years have crossed these thresholds, such that conditions have become increasingly unsuitable for regeneration. High fire severity and low seed availability further reduced the probability of postfire regeneration. Together, our results demonstrate that climate change combined with high severity fire is leading to increasingly fewer opportunities for seedlings to establish after wildfires and may lead to ecosystem transitions in low-elevation ponderosa pine and Douglas-fir forests across the western United States.

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.

Correction: Landscape drivers of recent fire activity (2001-2017) in south-central Chile.

[This corrects the article DOI: 10.1371/journal.pone.0201195.].

Landscape drivers of recent fire activity (2001-2017) in south-central Chile.

In recent decades large fires have affected communities throughout central and southern Chile with great social and ecological consequences. Despite this high fire activity, the controls and drivers and the spatiotemporal pattern of fires are not well understood. To identify the large-scale trends and drivers of recent fire activity across six regions in south-central Chile (~32-40° S Latitude) we evaluated MODIS satellite-derived fire detections and compared this data with Chilean Forest Service records for the period 2001-2017. MODIS burned area estimates provide a spatially and temporally comprehensive record of fire activity across an important bioclimatic transition zone between dry Mediterranean shrublands/sclerophyllous forests and wetter deciduous-broadleaf evergreen forests. Results suggest fire activity was highly variable in any given year, with no statistically significant trend in the number of fires or mean annual area burned. Evaluation of the variables associated with spatiotemporal patterns of fire for the 2001-2017 period indicate vegetation type, biophysical conditions (e.g., elevation, slope), mean annual and seasonal climatic conditions (e.g., precipitation) and mean population density have the greatest influence on the probability of fire occurrence and burned area for any given year. Both the number of fires and annual area burned were greatest in warmer, biomass-rich lowland Bío-Bío and Araucanía regions. Resource selection analyses indicate fire 'preferentially' occurs in exotic plantation forests, mixed native-exotic forests, native sclerophyll forests, pasture lands and matorral, vegetation types that all provide abundant, flammable and connected biomass for burning. Structurally and compositionally homogenous exotic plantation forests may promote fire spread greater than native deciduous-Nothofagaceae forests which were once widespread in the southern parts of the study area. In the future, the coincidence of warmer and drier conditions in landscapes dominated by flammable and fuel-rich forest plantations and mixed native-exotic and sclerophyll forests are likely to further promote large fires in south-central Chile.

Moisture availability limits subalpine tree establishment.

In the absence of broad-scale disturbance, many temperate coniferous forests experience successful seedling establishment only when abundant seed production coincides with favorable climate. Identifying the frequency of past establishment events and the climate conditions favorable for seedling establishment is essential to understanding how climate warming could affect the frequency of future tree establishment events and therefore future forest composition or even persistence of a forest cover. In the southern Rocky Mountains, USA, research on the sensitivity of establishment of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa)-two widely distributed, co-occurring conifers in North America-to climate variability has focused on the alpine treeline ecotone, leaving uncertainty about the sensitivity of these species across much of their elevation distribution. We compared annual germination dates for >450 Engelmann spruce and >500 subalpine fir seedlings collected across a complex topographic-moisture gradient to climate variability in the Colorado Front Range. We found that Engelmann spruce and subalpine fir established episodically with strong synchrony in establishment events across the study area. Broad-scale establishment events occurred in years of high soil moisture availability, which were characterized by above-average snowpack and/or cool and wet summer climatic conditions. In the recent half of the study period (1975-2010), a decrease in the number of fir and spruce establishment events across their distribution coincided with declining snowpack and a multi-decadal trend of rising summer temperature and increasing moisture deficits. Counter to expected and observed increases in tree establishment with climate warming in maritime subalpine forests, our results show that recruitment declines will likely occur across the core of moisture-limited subalpine tree ranges as warming drives increased moisture deficits.

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.

Evidence for declining forest resilience to wildfires under climate change.

Forest resilience to climate change is a global concern given the potential effects of increased disturbance activity, warming temperatures and increased moisture stress on plants. We used a multi-regional dataset of 1485 sites across 52 wildfires from the US Rocky Mountains to ask if and how changing climate over the last several decades impacted post-fire tree regeneration, a key indicator of forest resilience. Results highlight significant decreases in tree regeneration in the 21st century. Annual moisture deficits were significantly greater from 2000 to 2015 as compared to 1985-1999, suggesting increasingly unfavourable post-fire growing conditions, corresponding to significantly lower seedling densities and increased regeneration failure. Dry forests that already occur at the edge of their climatic tolerance are most prone to conversion to non-forests after wildfires. Major climate-induced reduction in forest density and extent has important consequences for a myriad of ecosystem services now and in the future.

Seed origin and warming constrain lodgepole pine recruitment, slowing the pace of population range shifts.

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here, we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had more than three-fold greater recruitment to their third year than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating halved recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering more than doubled recruitment, alleviating some of the negative effects of heating. Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid postfire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species' range could enhance recruitment and facilitate upslope range shifts with climate change.

Southern Annular Mode drives multicentury wildfire activity in southern South America.

The Southern Annular Mode (SAM) is the main driver of climate variability at mid to high latitudes in the Southern Hemisphere, affecting wildfire activity, which in turn pollutes the air and contributes to human health problems and mortality, and potentially provides strong feedback to the climate system through emissions and land cover changes. Here we report the largest Southern Hemisphere network of annually resolved tree ring fire histories, consisting of 1,767 fire-scarred trees from 97 sites (from 22 °S to 54 °S) in southern South America (SAS), to quantify the coupling of SAM and regional wildfire variability using recently created multicentury proxy indices of SAM for the years 1531-2010 AD. We show that at interannual time scales, as well as at multidecadal time scales across 37-54 °S, latitudinal gradient elevated wildfire activity is synchronous with positive phases of the SAM over the years 1665-1995. Positive phases of the SAM are associated primarily with warm conditions in these biomass-rich forests, in which widespread fire activity depends on fuel desiccation. Climate modeling studies indicate that greenhouse gases will force SAM into its positive phase even if stratospheric ozone returns to normal levels, so that climate conditions conducive to widespread fire activity in SAS will continue throughout the 21st century.

Summer and winter drought drive the initiation and spread of spruce beetle outbreak.

This study used Landsat-based detection of spruce beetle (Dendroctonus rufipennis) outbreak over the years 2000-2014 across the Southern Rocky Mountain Ecoregion to examine the spatiotemporal patterns of outbreak and assess the influence of temperature, drought, forest characteristics, and previous spruce beetle activity on outbreak development. During the 1999-2013 period, time series of spruce beetle activity were highly spatially correlated (r > 0.5) at distances <5 km, but remained weakly correlated (r = 0.08) at distances >400 km. Furthermore, cluster analysis on time series of outbreak activity revealed the outbreak developed at multiple incipient locations and spread to unaffected forest, highlighting the importance of both local-scale dispersal and regional-scale drivers in synchronizing spruce beetle outbreak. Spatial overlay analysis and Random Forest modeling of outbreak development show that outbreaks initiate in areas characterized by summer, winter, and multi-year drought and that outbreak spread is strongly linked to the proximity and extent of nearby outbreak, but remains associated with drought. Notably, we find that spruce beetle outbreak is associated with low peak snow water equivalent, not just summer drought. As such, future alterations to both winter and summer precipitation regimes are likely to drive important changes in subalpine forests.

Mixed-severity fire history at a forest-grassland ecotone in west central British Columbia, Canada.

This study examines spatially variable stand structure and fire-climate relationships at a low elevation forest-grassland ecotone in west central British Columbia, Canada. Fire history reconstructions were based on samples from 92 fire-scarred trees and stand demography from 27 plots collected over an area of about 7 km2 . We documented historical chronologies of widespread fires and localized grassland fires between AD 1600 and 1900. Relationships between fire events, reconstructed values of the Palmer Drought Severity Index, and annual precipitation were examined using superposed epoch and bivariate event analyses. Widespread fires occurred during warm, dry years and were preceded by multiple anomalously dry, warm years. Localized fires that affected only grassland-proximal forests were more frequent than widespread fires. These localized fires showed a lagged, positive relationship with wetter conditions. The landscape pattern of forest structure provided further evidence of complex fire activity with multiple plots shown to have experienced low-, mixed-, and/or high-severity fires over the last four centuries. We concluded that this forest-grassland ecotone was characterized by fires of mixed severity, dominated by frequent, low-severity fires punctuated by widespread fires of moderate to high severity. This landscape-level variability in fire-climate relationships and patterns in forest structure has important implications for fire and grassland management in west central British Columbia and similar environments elsewhere. Forest restoration techniques such as prescribed fire and thinning are oftentimes applied at the forest-grassland ecotone on the basis that historically high frequency, low-severity fires defined the character of past fire activity. This study provides forest managers and policy makers with important information on mixed-severity fire activity at a low elevation forest-grassland ecotone, a crucial prerequisite for the effective management of these complex ecosystems.

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.