Relationships of subalpine forest fires in the Colorado Front Range with interannual and multidecadal‐scale climatic variation

Aim An understanding of past relationships between fire occurrence and climate variability will help to elucidate the implications of climate‐change scenarios for future patterns of wildfire. In the present study we investigate the relationships between subalpine‐zone fire occurrence and climate variability and broad‐scale climate patterns in the Pacific and Atlantic Oceans at both interannual and multidecadal time‐scales. Location The study area is the subalpine zone of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa), and lodgepole pine (Pinus contorta) in the southern sector of the Rocky Mountain National Park, which straddles the continental divide of the northern Colorado Front Range. Methods We compared years of widespread fire from AD 1650 to 1978 for the subalpine zone of southern Rocky Mountain National Park, with climate variables such as measures of drought, and indices such as the El Niño–Southern Oscillation (ENSO), the Pacific Decadal Oscillation (PDO), and the Atlantic Multidecadal Oscillation (AMO). Results Years of extensive subalpine‐zone fires are significantly related to climate variability, phases of ENSO, the PDO, and the AMO, as well as to phase combinations of ENSO, the PDO, and the AMO at both interannual and centennial time‐scales. Main conclusions Years of extensive fires are related to extreme drought conditions and are significantly related to the La Niña phase of ENSO, the negative (cool) phase of the PDO, and the positive (warm) phase of the AMO. The co‐occurrence of the phase combination of La Niña‐negative PDO‐positive AMO is more important to fire occurrence than the individual influences of the climate patterns. Low‐frequency trends in the occurrence of this combination of climate‐pattern phases, resulting from trends in the AMO, are the primary climate pattern associated with periods of high fire occurrence (1700–89 and 1851–1919) and a fire‐free period (1790–1850). The apparent controlling influence of the AMO on drought and years of large fires in the subalpine forests of the Colorado Front Range probably applies to an extensive area of western North America.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

Fire regimes and forest changes in mid and upper montane forests of the southern Cascades, Lassen Volcanic National Park, California, U.S.A.

Aim Spatial and temporal variation in fire regime parameters and forest structure were assessed. Location A 2630‐ha area of mid‐ and upper montane forest in Lassen Volcanic National Park (LVNP). Methods Two hypotheses were tested concerned with fire‐vegetation relationships in southern Cascades forests: (1) fire regime parameters (return interval, season of burn, fire size, rotation period) vary by forest dominant, elevation and slope aspect; and (2) fire exclusion since 1905 has caused forest structural and compositional changes in both mid‐ and upper montane forests. The implications of the study for national park management are also discussed. Results Fire regime parameters varied by forest compositional group and elevation in LVNP. Median composite and point fire return intervals were shorter in low elevation Jeffrey pine (Pinus jeffreyi) (JP) (4–6 years, 16 years) and Jeffrey pine–white fir (Abies concolor) (JP‐WF) (5–10 years, 22 years) and longer in high elevation red fir (Abies magnifica)— western white pine (Pinus monticola) (RF‐WWP) forests (9–27 years, 70 years). Median fire return intervals were also shorter on east‐facing (6–9 years, 16.3 years) and longer on south‐ (11 years, 32.5 years) and west‐facing slopes (22–28 years, 54‐years) in all forests and in each forest composition group. Spatial patterns in fire rotation length were the same as those for fire return intervals. More growing season fires also occurred in JP (33.1%) and JP‐WF (17.5%) than in RF‐WWP (1.1%) forests. A dramatic decline in fire frequency occurred in all forests after 1905. Conclusions Changes in forest structure and composition occurred in both mid‐ and upper montane forests due to twentieth‐century fire exclusion. Forest density increased in JP and JP‐WF forests and white fir increased in JP‐WF forests and is now replacing Jeffrey pine. Forest density only increased in some RF‐WWP stands, but not others. Resource managers restoring fire to these now denser forests need to burn larger areas if fire is going to play its pre‐settlement role in montane forest dynamics.     

The Origin and Dynamics of Subalpine White Spruce and Balsam Fir Stands in Boreal Eastern North America

Associations among the few tree species in the North American boreal landscape are the result of complex interactions between climate, biota, and historical disturbances during the Holocene. The closed-crown boreal forest of eastern North America is subdivided into two ecological regions having distinct tree species associations; the balsam fir zone and the black spruce zone, south and north of 49°N, respectively. Subalpine old-growth stands dominated by trees species typical of the balsam fir forest flora (either balsam fir or white spruce) are found on high plateaus, some of which are isolated within the black spruce zone. Here we identified the ecological processes responsible for the distinct forest associations in the subalpine belt across the eastern boreal landscape. Extensive radiocarbon dating, species composition, and size structure analyses indicated contrasted origin and dynamics of the subalpine forests between the two ecological regions. In the black spruce zone, the subalpine belt is a mosaic of post-fire white spruce or balsam fir stands coexisting at similar elevation on the high plateaus. With increasing time without wildfire, the subalpine forests become structurally similar to the balsam fir forest of the fir zone. These results concur with the hypothesis that the subalpine forests of this area are protected remnants of an historical northern expansion of the fir zone. Its replacement by the fire-prone black spruce forest flora was caused by recurrent fires. In the subalpine belt of the fir zone, no fire was recorded for several millennia. Harsh climate at high altitude is the primary factor explaining white spruce dominance over balsam fir forming a distinct subalpine white spruce belt above the balsam fir dominated forest.     

Gradient Analysis of Fire Regimes in Montane Forests of the Southern Cascade Range,Thousand Lakes Wilderness,California, USA

Species distribution and abundance patterns in the southern Cascades are influenced by both environmental gradients and fire regimes. Little is known about fire regimes and variation in fire regimes may not be independent of environmental gradients or vegetation patterns. In this study, we analyze variation in fire regime parameters (i.e., return interval, season, size, severity, and rotation period) with respect to forest composition, elevation, and potential soil moisture in a 2042 ha area of montane forest in the southern Cascades in the Thousand Lakes Wilderness (TLW). Fire regime parameters varied with forest composition, elevation, and potential soil moisture. Median composite and point fire return intervals were shorter (4-9 yr, 14-24 yr) in low elevation and more xeric white fir (Abies concolor)-sugar pine (Pinus lambertiana) and white fir-Jeffrey pine (P. jeffreyi) and longest (20-37 yr, 20-47 yr) in mesic high elevation lodgepole pine (Pinus contorta) and red fir (Abies magnifica)-mountain hemlock (Tsuga mertensiana) forests. Values for mid-elevation red fir-white fir forests were intermediate. The pattern for fire rotation lengths across gradients was the same as for fire return intervals. The percentage of fires that occurred during the growing season was inversely related to elevation and potential soil moisture. Mean fire sizes were larger in lodgepole pine forests (405 ha) than in other forest groups (103-151 ha). In contrast to other parameters, fire severity did not vary across environmental and compositional gradients and >50% of all forests burned at high severity with most of the remainder burning at moderate severity. Since 1905, fire regimes have become similar at all gradient positions because of a policy of suppressing fire and fire regime modification will lead to shifts in landscape scale vegetation patterns.     

Effects of fire and spruce beetle outbreak legacies on the disturbance regime of a subalpine forest in Colorado

Aim There is increasing research attention being given to the role of interactions among natural disturbances in ecosystem processes. We studied the interactions between fire and spruce beetle (Dendroctonus rufipennis Kirkby) disturbances in a Colorado subalpine forest. The central questions of this research were: (1) How does fire history influence stand susceptibility to beetle outbreak? And conversely, (2) How does prior occurrence of a beetle outbreak influence stand susceptibility to subsequent fire? Methods We reconstructed the spatial disturbance history in a c. 4600 ha area by first identifying distinct patches in the landscape on aerial photographs. Then, in the field we determined the disturbance history of each patch by dating stand origin, fire scars, dates of mortality of dead trees, and releases on remnant trees. A geographical information system (GIS) was used to overlay disturbance by fire and spruce beetle. Results and main conclusions The majority of stands in the study area arose following large, infrequent, severe fires occurring in c. 1700, 1796 and 1880. The study area was also affected by a severe spruce beetle outbreak in the 1940s and a subsequent low‐severity fire. Stands that originated following stand‐replacing fire in the late nineteenth century were less affected by the beetle outbreak than older stands. Following the beetle outbreak, stands less affected by the outbreak were more affected by low‐severity fire than stands more severely affected by the outbreak. The reduced susceptibility to low‐severity fire possibly resulted from increased moisture on the forest floor following beetle outbreak. The landscape mosaic of this subalpine forest was strongly influenced by the interactions between fire and insect disturbances.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains

Extreme climatic events are key factors in initiating gradual or sudden changes in forest ecosystems through the promotion of severe, tree-killing disturbances such as fire, blowdown, and widespread insect outbreaks. In contrast to these climatically-incited disturbances, little is known about the more direct effect of drought on tree mortality, especially in high-elevation forests. Therefore projections of drought-induced mortality under future climatic conditions remain uncertain. For a subalpine forest landscape in the Rocky Mountains of northern Colorado (USA), we quantified lag effects of drought on mortality of Engelmann spruce Picea engelmannii , subalpine fir Abies lasiocarpa , and lodgepole pine Pinus contorta . For the period 1910–2004, we related death dates of 164 crossdated dead trees to early-season and late-season droughts. Following early-season droughts, spruce mortality increased over five years and fir mortality increased sharply over 11 years. Following late-season droughts, spruce showed a small increase in mortality within one year, whereas fir showed a consistent period of increased mortality over two years. Pine mortality was not affected by drought. Low pre-drought radial growth rates predisposed spruce and fir to drought-related mortality. Spruce and fir trees that died during a recent drought (2000–2004) had significantly lower pre-drought growth rates than live neighbour trees. Overall, we found large interspecific differences in drought-related mortality with fir showing the strongest effect followed by spruce and pine. This direct influence of climatic variability on differential tree mortality has the potential for driving large-scale changes in subalpine forests of the Rocky Mountains.     

Selective burning of forest vegetation in Canton Ticino (southern Switzerland)

Abstract

Detailed knowledge of factors controlling fire regime is a prerequisite for efficient fire management. We analyzed the fire selectivity of given forest vegetation classes both in terms of fire frequency and fire size for the present fire regime (1982–2005) in Canton Ticino (southern Switzerland). To this end, we investigated the dataset in four categories (all fires, anthropogenic winter fires, anthropogenic summer fires, and natural summer fires) and performed 1000 random Monte Carlo simulations on frequency and size. Anthropogenic winter and summer fires have a similar selectivity, occurring mostly at low elevations in chestnut stands, broadleaved forests, and in the first 50 m from the forest edge. In winter half of the fires in chestnut stands are significantly larger than 1.0 ha and the average burnt area in some coniferous forests tends to be high. Lightning fires seem to occur more frequently in spruce stands and less often in the summer‐humid chestnut and beech stands and the 50–100 m buffer area. In beech forests, in mixed forests, and in the spruce stands affected by natural fire in summer, the fires tend to be small in size. The selectivity observed, especially the selectivity of anthropogenic fires in terms of fire frequency, seems to be also related to geographical parameters such as altitude and aspect, and to anthropogenic characteristics such as closeness to roads or buildings.     

Influence of forest structure on the abundance of snowshoe hares in western Wyoming

Snowshoe hares (Lepus americanus) are a primary prey species for Canada lynx (Lynx canadensis) in western North America. Lynx management plans require knowledge of potential prey distribution and abundance in the western United States. Whether even-aged regenerating forests or multi-storied forests contain more snowshoe hares is currently unknown. During 2006–2008, we estimated snowshoe hare density in 3 classes of 30–70-year-old lodgepole pine (Pinus contorta) and 4 classes of late seral multi-storied forest with a spruce (Picea engelmannii)-fir (Abies lasiocarpa) component in the Bridger-Teton National Forest, Wyoming. We recorded physiographic variables and forest structure characteristics to understand how these factors influence abundance of snowshoe hares. In many instances, snowshoe hares were more abundant in late seral multi-storied forests than regenerating even-aged forests. Forest attributes predicting hare abundance were often more prevalent in multi-storied forests. Late seral multi-storied forests with a spruce–fir component and dense horizontal cover, as well as 30–70-year-old lodgepole pine with high stem density, were disproportionately influential in explaining snowshoe hare densities in western Wyoming. In order to promote improved habitat conditions for snowshoe hares in this region, management agencies should consider shifting their focus towards maintaining, enhancing, and promoting multi-storied forests with dense horizontal cover, as well as developing 30–70-year-old lodgepole pine stands with high stem density that structurally mimic multi-storied forests. © 2012 The Wildlife Society.     

What explains landscape patterns of tree mortality caused by bark beetle outbreaks in Greater Yellowstone?

Aim Bark beetle outbreaks have recently affected extensive areas of western North American forests, and factors explaining landscape patterns of tree mortality are poorly understood. The objective of this study was to determine the relative importance of stand structure, topography, soil characteristics, landscape context (the characteristics of the landscape surrounding the focal stand) and beetle pressure (the abundance of local beetle population eruptions around the focal stand a few years before the outbreak) to explain landscape patterns of tree mortality during outbreaks of three species: the mountain pine beetle, which attacks lodgepole pine and whitebark pine; the spruce beetle, which feeds on Engelmann spruce; and the Douglas‐fir beetle, which attacks Douglas‐fir. A second objective was to identify common variables that explain tree mortality among beetle–tree host pairings during outbreaks. Location Greater Yellowstone ecosystem, Wyoming, USA. Methods We used field surveys to quantify stand structure, soil characteristics and topography at the plot level in susceptible stands of each forest type showing different severities of infestation (0–98% mortality; n= 129 plots). We then used forest cover and beetle infestation maps derived from remote sensing to develop landscape context and beetle pressure metrics at different spatial scales. Plot‐level and landscape‐level variables were used to explain outbreak severity. Results Engelmann spruce and Douglas‐fir mortality were best predicted using landscape‐level variables alone. Lodgepole pine mortality was best predicted by both landscape‐level and plot‐level variables. Whitebark pine mortality was best – although poorly – predicted by plot‐level variables. Models including landscape context and beetle pressure were much better at predicting outbreak severity than models that only included plot‐level measures, except for whitebark pine. Main conclusions Landscape‐level variables, particularly beetle pressure, were the most consistent predictors of subsequent outbreak severity within susceptible stands of all four host species. These results may help forest managers identify vulnerable locations during ongoing outbreaks.     

Resilience and regime change in a southern Rocky Mountain ecosystem during the past 17?000 years

Paleoecological records indicate that subalpine forests in western North America have been resilient in response to multiple influences, including severe droughts, insect outbreaks, and widely varying fire regimes, over many millennia. One hypothesis for explaining this ecosystem resilience centers on the disruption of forest dynamics by frequent disturbance and climatic variability, and the resulting development of non-steady-state regimes dominated by early-successional conifers with broad climatic tolerances, such as lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.). To evaluate this hypothesis, we independently reconstructed the vegetation, fire, and effective-moisture histories of a small, forested watershed at 2890 m elevation in southeastern Wyoming, USA, using sedimentary pollen and charcoal counts in conjunction with sedimentary lake-level indicators. The data indicate that prominent vegetation shifts (from sagebrush steppe to spruce–fir parkland at ca. 10.7 ka and spruce–fir parkland to pine-dominated forest at ca. 8.5 ka [ka stands for thousands of years before the present, defined as AD 1950]) coincided with changes in effective moisture. However, after lodgepole pine forests established at ca. 8.5 ka, similar hydroclimatic changes did not produce detectable vegetation responses. Fire history data show that other aspects of the ecosystem were responsive to changes in effective moisture at centennial timescales with prolonged fire-free episodes coinciding with periods of low effective moisture ca. 7.2–5.6 and 3.7–1.6 ka. Throughout our record, the ratio of ecosystem perturbation time (i.e., fire frequency and changes in effective moisture) to recovery time (assuming 200–600-year successional processes) falls within estimates of the ratio for non-steady state ecosystems. Frequent perturbations, therefore, may have prevented this ecosystem from reaching compositional equilibrium with the varied climatic conditions over the past 8.5 ka. Equilibrium states could have included more abundant spruce (Picea spp.) and fir (Abies spp.) than presently observed based on brief increases in pollen abundances of these taxa during prolonged dry, fire-free intervals. Our results show that, although current climate changes favor widespread disturbance in Rocky Mountain forests, the composition of these ecosystems could be highly resilient and recover through successional dynamics over the next few decades to centuries.     

Macrocharcoal-Based Chronosequences Reveal Shifting Dominance of Conifer Boreal Forests Under Changing Fire Regime

Balsam fir (Abies balsamea) and black spruce (Picea mariana) forests are the main conifer forest types in the North American boreal zone. The coexistence of the two species as well as their respective canopy dominance in distinct stands raises questions about the long-term evolution from one forest type to the other in relation to environmental factors including climate and stand disturbance. We tested the hypothesis that repetitive fire events promote the succession of balsam fir forest to black spruce forest and vice versa. Postfire chronosequences of one black spruce (BSP) and one balsam fir (BFI) sites were reconstructed based on the botanical composition and ¹⁴C-dated soil macrocharcoals. The results support the hypothesis of a successional dynamics. The BSP site has been affected by fires for the last 7600 years, whereas the BFI site, after having been impacted by several fires during the first half of the Holocene, evolved in a fire-free environment for the last 4400 years. Periods of fire activity facilitated the dominance of black spruce forests. The cessation of fires around 4400 cal. years BP on BFI site marks the beginning of the transition from black spruce to balsam fir stands. This succession is a long process, due to the ability of black spruce to regenerate by layering in the absence of fire. The resulting balsam fir stands are ancient and precarious ecosystems, since fire generally leads to the return of black spruce. The increase in balsam fir to the detriment of black spruce in boreal forests is a response to a decrease in fire frequency.     

Habitat Conditions Associated With Lynx Hunting Behavior During Winter in Northern Washington

Abstract Effectively managing habitat for threatened populations of Canada lynx (Lynx canadensis) requires knowledge of habitat conditions that provide for the ecological needs of lynx. We snow-tracked lynx to identify habitat conditions associated with hunting behavior and predation during winters of 2002–2003 and 2003–2004 in the northern Cascade Range in Washington state, USA. We recorded number and success of predation attempts, prey species killed, and trail sinuosity on 149 km of lynx trails. Lynx killed snowshoe hares (Lepus americanus), red squirrels (Tamiasciurus hudsonicus), and cricetids more than expected in Englemann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests, where snowshoe hare densities were highest. Lynx killed prey less than expected in Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) forests and forest openings. We used the sinuosity of lynx trails as an index of quality of habitat hunted. Lynx trails that included predation attempts were more sinuous than trail segments without predation attempts. Lynx trails had greater sinuosity in forest stands with high hare densities dominated by Engelmann spruce and subalpine fir than in stands with low hare densities dominated by Douglas-fir and ponderosa pine or in forest openings. We encourage forest managers to maintain or create sufficient understory cover to support high densities of snowshoe hares as foraging habitat for lynx.     

Fire, fuels and restoration of ponderosa pine–Douglas fir forests in the Rocky Mountains, USA

Aim Forest restoration in ponderosa pine and mixed ponderosa pine–Douglas fir forests in the US Rocky Mountains has been highly influenced by a historical model of frequent, low‐severity surface fires developed for the ponderosa pine forests of the Southwestern USA. A restoration model, based on this low‐severity fire model, focuses on thinning and prescribed burning to restore historical forest structure. However, in the US Rocky Mountains, research on fire history and forest structure, and early historical reports, suggest the low‐severity model may only apply in limited geographical areas. The aim of this article is to elaborate a new variable‐severity fire model and evaluate the applicability of this model, along with the low‐severity model, for the ponderosa pine–Douglas fir forests of the Rocky Mountains. Location Rocky Mountains, USA. Methods The geographical applicability of the two fire models is evaluated using historical records, fire histories and forest age‐structure analyses. Results Historical sources and tree‐ring reconstructions document that, near or before ad 1900, the low‐severity model may apply in dry, low‐elevation settings, but that fires naturally varied in severity in most of these forests. Low‐severity fires were common, but high‐severity fires also burned thousands of hectares. Tree regeneration increased after these high‐severity fires, and often attained densities much greater than those reconstructed for Southwestern ponderosa pine forests. Main conclusions Exclusion of fire has not clearly and uniformly increased fuels or shifted the fire type from low‐ to high‐severity fires. However, logging and livestock grazing have increased tree densities and risk of high‐severity fires in some areas. Restoration is likely to be most effective which seeks to (1) restore variability of fire, (2) reverse changes brought about by livestock grazing and logging, and (3) modify these land uses so that degradation is not repeated.     

Climatic influences on fire regimes in the northern Sierra Nevada mountains, Lake Tahoe Basin, Nevada, USA

Aim The goal of this study was to understand better the role of interannual and interdecadal climatic variation on local pre‐EuroAmerican settlement fire regimes in fire‐prone Jeffrey pine (Pinus jeffreyi Grev. & Balf.) dominated forests in the northern Sierra Nevada Mountains. Location Our study was conducted in a 6000‐ha area of contiguous mixed Jeffrey pine‐white fir (Abies concolor Gordon & Glend.) forest on the western slope of the Carson Range on the eastern shore of Lake Tahoe, Nevada. Methods Pre‐EuroAmerican settlement fire regimes (i.e. frequency, return interval, extent, season) were reconstructed in eight contiguous watersheds for a 200‐year period (1650–1850) from fire scars preserved in the annual growth rings of nineteenth century cut stumps and recently dead pre‐settlement Jeffrey pine trees. Superposed epoch analysis (SEA) and correlation analysis were used to examine relationships between tree ring‐based reconstructions of the Palmer Drought Severity Index (PDSI), Southern Oscillation Index (SOI), Pacific Decadal Oscillation (PDO) and pre‐EuroAmerican fire regimes in order to assess the influence of drought and equatorial and north Pacific teleconnections on fire occurrence and fire extent. Results For the entire period of record (1650–1850), wet conditions were characteristic of years without fires. In contrast, fire years were associated with drought. Drought intensity also influenced fire extent and the most widespread fires occurred in the driest years. Years with widespread fires were also preceded by wet conditions 3 years before the fire. Widespread fires were also associated with phase changes of the PDO, with the most widespread burns occurring when the phase changed from warm (positive) to cold (negative) conditions. Annual SOI and fire frequency or extent were not associated in our study. At decadal time scales, burning was more widespread during decades that were dryer and characterized by La Niña and negative PDO conditions. Interannual and interdecadal fire–climate relationships were not stable over time. From 1700 to 1775 there was no interannual relationship between drought, PDO, and fire frequency or extent. However, from 1775 to 1850, widespread fires were associated with dry years preceded by wet years. This period also had the strongest association between fire extent and the PDO. In contrast, fire–climate associations at interdecadal time scales were stronger in the earlier period than in the later period. The change from strong interdecadal to strong interannual climate influence was associated with a breakdown in decadal scale constructive relationships between PDO and SOI. Main conclusions Climate strongly influenced pre‐settlement pine forest fire regimes in northern Sierra Nevada. Both interannual and interdecadal climatic variation regulated conditions conducive to fire activity, and longer term changes in fire frequency and extent correspond with climate‐mediated changes observed in both the northern and southern hemispheres. The sensitivity of fire regimes to shifts in modes of climatic variability suggests that climate was a key regulator of pine forest ecosystem structure and dynamics before EuroAmerican settlement. An understanding of pre‐EuroAmerican fire–climate relationships may provide useful insights into how fire activity in contemporary forests may respond to future climatic variation.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

Origin of the lichen–spruce woodland in the closed-crown forest zone of eastern Canada

Aim We investigate the timing and factors responsible for the transformation of closed‐crown forests into lichen–spruce woodlands. Location The study area extends between 70° and 72° W in the closed‐crown forest zone from its southern limit near 47°30′ N to its northern limit at the contact with the lichen–spruce woodland zone around 52°10′ N. A total of 24 lichen–spruce woodlands were selected. Methods Radiocarbon dating of charcoals at mineral soil contact and within the organic horizons allowed the principal factors causing the degradation of the closed‐crown forest to be identified, i.e. light fires, successive fires and the occurrence of a spruce budworm epidemic followed by a fire. Results Charcoals dated in the organic horizon were less than 200 years old, suggesting a recent transformation of the closed‐crown forest following surface fires. Before their transformation into lichen–spruce woodlands, stands were occupied by old, dense forests that originated from fires dating back to 1000 yr bp . The radiocarbon dating of charcoals in the organic horizon indicated that several stands burned twice in less than 50 years, while others burned shortly after a spruce budworm epidemic. Light fires are frequent within the lichen–spruce woodlands according to multiple charcoal layers found within the organic matter horizon. Main conclusions While closed‐crown forests are predicted to expand under climate warming, compound disturbances diminish the natural regeneration of the closed‐crown forests in the south and favour the expansion of lichen–spruce woodlands. As black spruce germinates on mineral soils, surface fires accentuate the expansion of the lichen–spruce woodlands southward. Under global warming, warmer springs will lead to earlier low‐intensity fires that do not remove as much organic matter, and hence prevent conditions suitable for black spruce regeneration. Also, spruce budworm reduces seed production for a certain time. The occurrence of fire during this period is critical for regeneration of black spruce.     

Lodgepole pine forest age class dynamics and susceptibility to mountain pine beetle attack

Because mountain pine beetle attack mature pine stands, an understanding of forest age class dynamics is important to managing forests within the distribution of the beetle. The assumed theoretical negative exponential forest age distribution provides an estimate when ecosystem dynamics are in equilibrium. This study investigates the dynamics of forest age distribution for non-equilibrium ecosystem dynamics, which result primarily from large and irregular stand-replacement fire disturbances that alter the forest age distribution. A model experiment using the SEM-LAND model on a 1 million ha lodgepole pine forest landscape was conducted to estimate how the proportion of susceptible area could be influenced by different fire regimes. The results of the simulation suggest that the temporal dynamics of the area susceptible to mountain pine beetle attack are complex and depend on the fire history of the study area, if the area is experiencing large and irregular stand-replacement fires. The age range of the lodgepole pine forest stands susceptible to mountain pine beetle attack might significantly affect the estimate of the area susceptible to attack.