Warming drives a front of white spruce establishment near western treeline, Alaska.

Regional warming has led to increased productivity near the boreal forest margin in Alaska. To date, the effects of warming on seedling recruitment have received little attention, in spite of forecasted forest expansion. Here, we used stand structure and environmental data from 95 white spruce (Picea glauca) plots sampled across a longitudinal gradient in southwest Alaska to explore factors influencing spruce establishment and recruitment near western treeline. We used total counts of live seedlings, saplings, and trees, representing five life stages, to evaluate whether geospatial, climate, and measured plot covariates predicted abundance, using current abundance distributions as a surrogate for climate conditions in the past. We used generalized linear models to test the null hypothesis that conditions favorable for recruitment were similar along the environmental gradient represented by longitude, by exploring relationships between per-plot counts of each life stage and the covariates hypothesized to affect abundance. We also examined the relationship between growing degree days (GDD) and seedling establishment over a period of three decades using tree-ring chronologies obtained from cores taken at a subset of our sites (n = 30). Our results indicated that seedling, sapling, and tree abundance were positively correlated with temperature across the study area. The response to longitude was mixed, with earlier life stages (seedlings, saplings) most abundant at the western end of the gradient, and later life stages (trees) most abundant to the east. The differential relationship between longitude and life-stage abundance suggests a moving front of white spruce establishment through time, driven by changes in environmental conditions near the species' western range limit. Likewise, we found a positive relationship between periods of seedling establishment and GDD, suggesting that longer summers and/or greater heat accumulation might enhance establishment, consistent with the positive relationship we found between life-stage abundance and temperature.

Tree-ring isotopes reveal drought sensitivity in trees killed by spruce beetle outbreaks in south-central Alaska.

Increasing temperatures have resulted in reduced growth and increased tree mortality across large areas of western North American forests. We use tree-ring isotope chronologies (δ13 C and δ18 O) from live and dead trees from four locations in south-central Alaska, USA, to test whether white spruce trees killed by recent spruce beetle (Dendroctonus rufipennis Kirby) outbreaks showed evidence of drought stress prior to death. Trees that were killed were more sensitive to spring/summer temperature and/or precipitation than trees that survived. At two of our sites, we found greater correlations between the δ13 C and δ18 O chronologies and spring/summer temperatures in dead trees than in live trees, suggesting that trees that are more sensitive to temperature-induced drought stress are more likely to be killed. At one site, the difference between δ13 C in live and dead trees was related to winter/spring precipitation, with dead trees showing stronger correlations between δ13 C and precipitation, again suggesting increased water stress in dead trees. At all sites where δ18 O was measured, δ18 O chronologies showed the greatest difference in climate response between live and dead groups, with δ18 O in live trees correlating more strongly with late winter precipitation than dead trees. Our results indicate that sites where trees are already sensitive to warm or dry early growing-season conditions experienced the most beetle-kill, which has important implications for forecasting future mortality events in Alaska.

Historical, observed, and modeled wildfire severity in montane forests of the Colorado Front Range.

Large recent fires in the western U.S. have contributed to a perception that fire exclusion has caused an unprecedented occurrence of uncharacteristically severe fires, particularly in lower elevation dry pine forests. In the absence of long-term fire severity records, it is unknown how short-term trends compare to fire severity prior to 20th century fire exclusion. This study compares historical (i.e. pre-1920) fire severity with observed modern fire severity and modeled potential fire behavior across 564,413 ha of montane forests of the Colorado Front Range. We used forest structure and tree-ring fire history to characterize fire severity at 232 sites and then modeled historical fire-severity across the entire study area using biophysical variables. Eighteen (7.8%) sites were characterized by low-severity fires and 214 (92.2%) by mixed-severity fires (i.e. including moderate- or high-severity fires). Difference in area of historical versus observed low-severity fire within nine recent (post-1999) large fire perimeters was greatest in lower montane forests. Only 16% of the study area recorded a shift from historical low severity to a higher potential for crown fire today. An historical fire regime of more frequent and low-severity fires at low elevations (<2260 m) supports a convergence of management goals of ecological restoration and fire hazard mitigation in those habitats. In contrast, at higher elevations mixed-severity fires were predominant historically and continue to be so today. Thinning treatments at higher elevations of the montane zone will not return the fire regime to an historic low-severity regime, and are of questionable effectiveness in preventing severe wildfires. Based on present-day fuels, predicted fire behavior under extreme fire weather continues to indicate a mixed-severity fire regime throughout most of the montane forest zone. Recent large wildfires in the Front Range are not fundamentally different from similar events that occurred historically under extreme weather conditions.

Examining historical and current mixed-severity fire regimes in ponderosa pine and mixed-conifer forests of western North America.

There is widespread concern that fire exclusion has led to an unprecedented threat of uncharacteristically severe fires in ponderosa pine (Pinus ponderosa Dougl. ex. Laws) and mixed-conifer forests of western North America. These extensive montane forests are considered to be adapted to a low/moderate-severity fire regime that maintained stands of relatively old trees. However, there is increasing recognition from landscape-scale assessments that, prior to any significant effects of fire exclusion, fires and forest structure were more variable in these forests. Biota in these forests are also dependent on the resources made available by higher-severity fire. A better understanding of historical fire regimes in the ponderosa pine and mixed-conifer forests of western North America is therefore needed to define reference conditions and help maintain characteristic ecological diversity of these systems. We compiled landscape-scale evidence of historical fire severity patterns in the ponderosa pine and mixed-conifer forests from published literature sources and stand ages available from the Forest Inventory and Analysis program in the USA. The consensus from this evidence is that the traditional reference conditions of low-severity fire regimes are inaccurate for most forests of western North America. Instead, most forests appear to have been characterized by mixed-severity fire that included ecologically significant amounts of weather-driven, high-severity fire. Diverse forests in different stages of succession, with a high proportion in relatively young stages, occurred prior to fire exclusion. Over the past century, successional diversity created by fire decreased. Our findings suggest that ecological management goals that incorporate successional diversity created by fire may support characteristic biodiversity, whereas current attempts to "restore" forests to open, low-severity fire conditions may not align with historical reference conditions in most ponderosa pine and mixed-conifer forests of western North America.

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.

Climate variability and spruce beetle (Dendroctonus rufipennis) outbreaks in south-central and southwest Alaska.

We used tree ring data (AD 1601-2007) to examine the occurrence of and climatic influences on spruce beetle (Dendroctonus rufipennis) outbreaks in south-central and southwest Alaska and found evidence of regional-scale outbreaks dating from the mid-1700s, related to climate variability at multiple temporal scales. Over interannual time scales (approximately 1-3 years), El Niño years, combined with severe late-summer drought, appeared to contribute significantly to spruce beetle outbreaks in the study area. Over multidecadal time scales (up to approximately 40 years), cool-phase Pacific Decadal Oscillation (PDO) conditions tended to precede beetle outbreaks, regardless of the phase of El Niño-Southern Oscillation (ENSO). All sites showed low-severity disturbances attributed to spruce beetle damage, most notably during the 1810s. During other major periods of disturbance (i.e., 1870s, 1910s, 1970s), the effects of spruce beetle outbreaks were of moderate or higher severity. The highly synchronized timing of spruce beetle outbreaks at interannual to multidecadal scales, and particularly the association between cool-phase PDO conditions and beetle disturbance, suggests that climate (i.e., temperature, precipitation) is a primary driver of outbreaks in the study area. Our disturbance chronologies (mid-1700s to present) suggest that recent irruptions (1990s to present) in south-central and southwest Alaska are within the historical geographic range, but that outbreaks since the 1990s show greater spatiotemporal synchrony (i.e., more sites record high-severity infestations) than at any other time in the past approximatly 250 years.