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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Area burned in the western United States is unaffected by recent mountain pine beetle outbreaks.

In the western United States, mountain pine beetles (MPBs) have killed pine trees across 71,000 km(2) of forest since the mid-1990s, leading to widespread concern that abundant dead fuels may increase area burned and exacerbate fire behavior. Although stand-level fire behavior models suggest that bark beetle-induced tree mortality increases flammability of stands by changing canopy and forest floor fuels, the actual effect of an MPB outbreak on subsequent wildfire activity remains widely debated. To address this knowledge gap, we superimposed areas burned on areas infested by MPBs for the three peak years of wildfire activity since 2002 across the western United States. Here, we show that the observed effect of MPB infestation on the area burned in years of extreme fire appears negligible at broad spatial extents. Contrary to the expectation of increased wildfire activity in recently infested red-stage stands, we found no difference between observed area and expected area burned in red-stage or subsequent gray-stage stands during three peak years of wildfire activity, which account for 46% of area burned during the 2002-2013 period. Although MPB infestation and fire activity both independently increased in conjunction with recent warming, our results demonstrate that the annual area burned in the western United States has not increased in direct response to bark beetle activity. Therefore, policy discussions should focus on societal adaptation to the effects of recent increases in wildfire activity related to increased drought severity.

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 unaffected by spruce beetle outbreak in spruce-fir forests in southwestern Colorado.

Recent large and severe outbreaks of native bark beetles have raised concern among the general public and land managers about potential for amplified fire activity in western North America. To date, the majority of studies examining bark beetle outbreaks and subsequent fire severity in the U.S. Rocky Mountains have focused on outbreaks of mountain pine beetle (MPB; Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests, but few studies, particularly field studies, have addressed the effects of the severity of spruce beetle (Dendroctonus rufipennis Kirby) infestation on subsequent fire severity in subalpine Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests. In Colorado, the annual area infested by spruce beetle outbreaks is rapidly rising, while MPB outbreaks are subsiding; therefore understanding this relationship is of growing importance. We collected extensive field data in subalpine forests in the eastern San Juan Mountains, southwestern Colorado, USA, to investigate whether a gray-stage (< 5 yr from outbreak to time of fire) spruce beetle infestation affected fire severity. Contrary to the expectation that bark beetle infestation alters subsequent fire severity, correlation and multivariate generalized linear regression analysis revealed no influence of pre-fire spruce beetle severity on nearly all field or remotely sensed measurements of fire severity. Findings were consistent across moderate and extreme burning conditions. In comparison to severity of the pre-fire beetle outbreak, we found that topography, pre-outbreak basal area, and weather conditions exerted a stronger effect on fire severity. Our finding that beetle infestation did not alter fire severity is consistent with previous retrospective studies examining fire activity following other bark beetle outbreaks and reiterates the overriding influence of climate that creates conditions conducive to large, high-severity fires in the subalpine zone of Colorado. Both bark beetle outbreaks and wildfires have increased autonomously due to recent climate variability, but this study does not support the expectation that post-beetle outbreak forests will alter fire severity, a result that has important implications for management and policy decisions.

Biogeochemistry of beetle-killed forests: explaining a weak nitrate response.

A current pine beetle infestation has caused extensive mortality of lodgepole pine (Pinus contorta) in forests of Colorado and Wyoming; it is part of an unprecedented multispecies beetle outbreak extending from Mexico to Canada. In United States and European watersheds, where atmospheric deposition of inorganic N is moderate to low (<10 kg⋅ha⋅y), disturbance of forests by timber harvest or violent storms causes an increase in stream nitrate concentration that typically is close to 400% of predisturbance concentrations. In contrast, no significant increase in streamwater nitrate concentrations has occurred following extensive tree mortality caused by the mountain pine beetle in Colorado. A model of nitrate release from Colorado watersheds calibrated with field data indicates that stimulation of nitrate uptake by vegetation components unaffected by beetles accounts for significant nitrate retention in beetle-infested watersheds. The combination of low atmospheric N deposition (<10 kg⋅ha⋅y), tree mortality spread over multiple years, and high compensatory capacity associated with undisturbed residual vegetation and soils explains the ability of these beetle-infested watersheds to retain nitrate despite catastrophic mortality of the dominant canopy tree species.

Effects of high-severity fire drove the population collapse of the subalpine Tasmanian endemic conifer Athrotaxis cupressoides.

Athrotaxis cupressoides is a slow-growing and long-lived conifer that occurs in the subalpine temperate forests of Tasmania, a continental island to the south of Australia. In 1960-1961, human-ignited wildfires occurred during an extremely dry summer that killed many A. cupressoides stands on the high plateau in the center of Tasmania. That fire year, coupled with subsequent regeneration failure, caused a loss of ca. 10% of the geographic extent of this endemic Tasmanian forest type. To provide historical context for these large-scale fire events, we (i) collected dendroecological, floristic, and structural data, (ii) documented the postfire survival and regeneration of A. cupressoides and co-occurring understory species, and (iii) assessed postfire understory plant community composition and flammability. We found that fire frequency did not vary following the arrival of European settlers, and that A. cupressoides populations were able to persist under a regime of low-to-mid severity fires prior to the 1960 fires. Our data indicate that the 1960 fires were (i) of greater severity than previous fires, (ii) herbivory by native marsupials may limit seedling survival in both burned and unburned A. cupressoides stands, and (iii) the loss of A. cupressoides populations is largely irreversible given the relatively high fuel loads of postfire vegetation communities that are dominated by resprouting shrubs. We suggest that the feedback between regeneration failure and increased flammability will be further exacerbated by a warmer and drier climate causing A. cupressoides to contract to the most fire-proof landscape settings.

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.

Spatiotemporal patterns of mountain pine beetle activity in the southern Rocky Mountains.

The current mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak in the southern Rocky Mountains has impacted approximately 750 000 ha of forest. Weather and habitat heterogeneity influence forest insect population dynamics at multiple spatial and temporal scales. Comparison of forest insect population dynamics in two principal host species may elucidate the relative contribution of weather and landscape factors in initiating and driving extensive outbreaks. To investigate potential drivers of the current MPB outbreak, we compared broadscale spatiotemporal patterns of MPB activity in lodgepole pine (Pinus contorta) and ponderosa pine (Pinus ponderosa) from 1996 to 2010 in Colorado and southern Wyoming with regional weather fluctuations, and then tracked the annual meso-scale progression of the epidemic in lodgepole pine with respect to weather, topographic, previous MPB activity, and forest stand attributes. MPB activity in lodgepole pine compared to ponderosa pine showed higher magnitude and extent of spatial synchrony. Warm temperatures and low annual precipitation favorable to beetle populations showed high regional synchrony across areas of both pine species, suggesting that habitat interacts with weather in synchronizing MPB populations. Cluster analysis of time series patterns identified multiple, disjunct locations of incipient MPB activity (epicenters) in lodgepole pine, which overlapped an earlier 1980s MPB outbreak, and suggests a regional trigger (drought) across this homogenous forest type. Negative departures from mean annual precipitation played a key role in subsequent spread of MPB outbreak. Development of the outbreak was also associated with lower elevations, greater dominance by lodgepole pine, stands of larger tree size, and stands with higher percentage canopy cover. After epidemic levels of MPB activity were attained, MPB activity was less strongly associated with stand and weather variables. These results emphasize the importance of considering differences in patterns of MPB dynamics for different host pine species even under similar regional-scale weather variation and the nonstationarity of outbreak dynamics over time.

Fire history and tree recruitment in the Colorado Front Range upper montane zone: implications for forest restoration.

Forests experiencing moderate- or mixed-severity fire regimes are presumed to be widespread across the western United States, but few studies have characterized these complex disturbance regimes and their effects on contemporary forest structure. Restoration of pre-fire-suppression open-forest structure to reduce the risk of uncharacteristic stand-replacing fires is a guiding principle in forest management policy, but identifying which forests are clear candidates for restoration remains a challenge. We conducted dendroecological reconstructions of fire history and stand structure at 40 sites in the upper montane zone of the Colorado Front Range (2400-2800 m), sampled in proportion to the distribution of forest types in that zone (50% dominated by ponderosa pine, 28% by lodgepole pine, 12% by aspen, 10% by Douglas-fir). We characterized past fire severity based on remnant criteria at each site in order to assess the effect of fire history on tree establishment patterns, and we also evaluated the influence of fire suppression and climate. We found that 62% of the sites experienced predominantly moderate-severity fire, 38% burned at high severity, and no sites burned exclusively at low severity. The proportion of total tree and sapling establishment was significantly different among equal time periods based on a chi-square test, with highest tree and sapling establishment during the pre-fire-suppression period (1835-1919). Superposed epoch analysis revealed that fires burned during years of extreme drought (95% CI). The major pulse of tree establishment in the upper montane zone occurred during a multidecadal period of extreme drought conditions in the Colorado Front Range (1850-1889), during which 53% of the fires from the 1750-1989 period burned. In the upper montane zone of the Colorado Front Range, historical evidence suggests that these forests are resilient to prolonged periods of severe drought and associated severe fires.