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Singular and interactive effects of blowdown, salvage logging, and wildfire in sub-boreal pine systems

The role of disturbance in structuring vegetation is widely recognized; however, we are only beginning to understand the effects of multiple interacting disturbances on ecosystem recovery and development. Of particular interest is the impact of post-disturbance management interventions, particularly in light of the global controversy surrounding the effects of salvage logging on forest ecosystem recovery. Studies of salvage logging impacts have focused on the effects of post-disturbance salvage logging within the context of a single natural disturbance event. There have been no formal evaluations of how these effects may differ when followed in short sequence by a second, high severity natural disturbance. To evaluate the impact of this management practice within the context of multiple disturbances, we examined the structural and woody plant community responses of sub-boreal Pinus banksiana systems to a rapid sequence of disturbances. Specifically, we compared responses to Blowdown (B), Fire (F), Blowdown–Fire, and Blowdown–Salvage–Fire (BSF) and compared these to undisturbed control (C) stands. Comparisons between BF and BSF indicated that the primary effect of salvage logging was a decrease in the abundance of structural legacies, such as downed woody debris and snags. Both of these compound disturbance sequences (BF and BSF), resulted in similar woody plant communities, largely dominated by Populus tremuloides ; however, there was greater homogeneity in community composition in salvage logged areas. Areas experiencing solely fire (F stands) were dominated by P. banksiana regeneration, and blowdown areas (B stands) were largely characterized by regeneration from shade tolerant conifer species. Our results suggest that salvage logging impacts on woody plant communities are diminished when followed by a second high severity disturbance; however, impacts on structural legacies persist. Provisions for the retention of snags, downed logs, and surviving trees as part of salvage logging operations will minimize these structural impacts and may allow for greater ecosystem recovery following these disturbance combinations.

Forest Ecology and Management

Forest disturbance interactions and successional pathways in the Southern Rocky Mountains

The pine forests in the southern portion of the Rocky Mountains are a heterogeneous mosaic of disturbance and recovery. The most extensive and intensive stress and mortality are received from human activity, fire, and mountain pine beetles (MPB; Dendroctonus ponderosae ). Understanding disturbance interactions and disturbance-succession pathways are crucial for adapting management strategies to mitigate their impacts and anticipate future ecosystem change. Driven by this goal, we assessed the forest disturbance and recovery history in the Southern Rocky Mountains Ecoregion using a 13-year time series of Landsat image stacks. An automated classification workflow that integrates temporal segmentation techniques and a random forest classifier was used to examine disturbance patterns. To enhance efficiency in selecting representative samples at the ecoregion scale, a new sampling strategy that takes advantage of the scene-overlap among adjacent Landsat images was designed. The segment-based assessment revealed that the overall accuracy for all 14 scenes varied from 73.6% to 92.5%, with a mean of 83.1%. A design-based inference indicated the average producer’s and user’s accuracies for MPB mortality were 85.4% and 82.5% respectively. We found that burn severity was largely unrelated to the severity of pre-fire beetle outbreaks in this region, where the severity of post-fire beetle outbreaks generally decreased in relation to burn severity. Approximately half the clear-cut and burned areas were in various stages of recovery, but the regeneration rate was much slower for MPB-disturbed sites. Pre-fire beetle outbreaks and subsequent fire produced positive compound effects on seedling reestablishment in this ecoregion. Taken together, these results emphasize that although multiple disturbances do play a role in the resilience mechanism of the serotinous lodgepole pine, the overall recovery could be slow due to the vast area of beetle mortality.

Colorado, New Mexico, Wyoming

Post-fire reference densities for giant sequoia seedlings in a new era of high-severity wildfires

Many forests globally are experiencing increases in large, high-severity wildfires, often with increasingly inadequate post-fire tree regeneration. To identify areas that might need post-fire planting, forest managers have a growing need for seedling reference densities – the natural seedling densities expected to be adequate to regenerate a forest – to compare with observed post-fire seedling densities. The most useful reference densities will meet five criteria: they will (1) be specific to natural post-fire reproduction rather than planted seedlings (because planted seedlings can have substantially greater survival than natural seedlings, thus underestimating adequate natural reproduction), (2) apply to the first few years following fire (when management decisions and actions are most likely), (3) be specific to each of those post-fire years (because post-fire seedling densities can change rapidly with time since fire), (4) be associated with estimates of uncertainty, and (5) include consideration of novel environmental conditions during management applications (because most reference densities will be based on data collected under more environmentally benign conditions). The world’s most massive tree species, the giant sequoia ( Sequoiadendron giganteum ) of California’s Sierra Nevada, recently experienced historically unprecedented wildfires that killed an estimated 13–19% of mature sequoias across their native range. Seedlings germinating after these fires then experienced exceptional summer heat and the two most severe summer droughts of the 121-year historical record. To help inform management responses to these events, we used seedling censuses from past fires (mostly prescribed fires) to calculate sequoia seedling reference densities meeting the five criteria. The reference densities had three striking features, which are partly attributable to giant sequoia’s status as a pioneer species. First, despite being inherently conservative, the reference densities were quite high. For example, mean first-year reference density was 172,599 seedlings ha −1 . Second, reference densities declined precipitously with time since fire: the mean fifth-year reference density was only 5% of the mean first-year density. Third, the reference densities were associated with relatively substantial uncertainty, a consequence of density variations among seedling plots; for example, the 95% credible interval for first-year reference density was 64,377 to 313,438 seedlings ha −1 . Despite this uncertainty, a case-study sequoia grove that recently burned in a high-severity wildfire had second-year post-fire seedling densities that were significantly (and dramatically) lower than the corresponding second-year reference density, suggesting inadequate post-fire reproduction. Our results highlight the value of the five criteria for reference densities – criteria that, in current practice, are rarely all met.

Forest Ecology and Management

Breeding birds in managed forests on public conservation lands in the Mississippi Alluvial Valley

Managers of public conservation lands in the Mississippi Alluvial Valley have implemented forest management strategies to improve bottomland hardwood habitat for target wildlife species. Through implementation of various silvicultural practices, forest managers have sought to attain forest structural conditions (e.g., canopy cover, basal area, etc.) within values postulated to benefit wildlife. We evaluated data from point count surveys of breeding birds on 180 silviculturally treated stands (1049 counts) that ranged from 1 to 20 years post-treatment and 134 control stands (676 counts) that had not been harvested for >20 years. Birds detected during 10-min counts were recorded within four distance classes and three time intervals. Avian diversity was greater on treated stands than on unharvested stands. Of 42 commonly detected species, six species including Prothonotary Warbler ( Prothonotaria citrea ) and Acadian Flycatcher ( Empidonax virescens ) were indicative of control stands. Similarly, six species including Indigo Bunting ( Passerina cyanea ) and Yellow-breasted Chat ( Icteria virens ) were indicative of treated stands. Using a removal model to assess probability of detection, we evaluated occupancy of bottomland forests at two spatial scales (stands and points within occupied stands). Wildlife-forestry treatment improved predictive models of species occupancy for 18 species. We found years post treatment (range = 1–20), total basal area, and overstory canopy were important species-specific predictors of occupancy, whereas variability in basal area was not. In addition, we used a removal model to estimate species-specific probability of availability for detection, and a distance model to estimate effective detection radius. We used these two estimated parameters to derive species densities and 95% confidence intervals for treated and unharvested stands. Avian densities differed between treated and control stands for 16 species, but only Common Yellowthroat ( Geothlypis trichas ) and Yellow-breasted Chat had greater densities on treated stands.

Mississippi Alluvial Valley

Thermal characteristics of amphibian microhabitats in a fire-disturbed landscape

Disturbance has long been a central issue in amphibian conservation, often regarding negative effects of logging or other forest management activities, but some amphibians seem to prefer disturbed habitats. After documenting increased use of recently burned forests by boreal toads (Bufo boreas), we hypothesized that burned habitats provided improved thermal opportunities in terrestrial habitats. We tested this hypothesis by conducting a radio telemetry study of habitat use (reported previously) and by using physical models that simulated the temperature of adult toads. We deployed 108 physical models in and adjacent to a 1-year old burn using a fully-replicated design with three burn severities (unburned, partial, high severity) and four microhabitats (open surface, under vegetation, under log, in burrow). Model temperatures were compared to a range of preferred temperatures in published studies. We found 70% more observations within the preferred temperature range of B. boreas in forests burned with high severity than in unburned areas. Burned forest was warmer than unburned forest across all microhabitats, but the largest relative difference was in burrows, which averaged 3 ??C warmer in high-severity burn areas and remained warmer though the night. More than twice as many observations were within the preferred temperature range in high-severity burrows than in unburned burrows. Areas burned with high severity were still warmer than unburned forest 3 years after the fire. Habitat use of toads during the concurrent radio telemetry study matched that predicted by the physical models. These results suggest there are fitness-linked benefits to toads using burned habitats, such as increased growth, fertility, and possibly disease resistance. However, increased soil temperatures that result from wildfire may be detrimental to other amphibian species that prefer cooler temperatures and stable environments. More broadly, our data illustrate the use of physical models to measure and interpret changes that amphibians may experience from disturbance, and highlight the need for research linking vital rates such as growth and survival to disturbance.

Forest Ecology and Management

Historical dominance of low-severity fire in dry and wet mixed-conifer forest habitats of the endangered terrestrial Jemez Mountains salamander ( Plethodon neomexicanus )

Anthropogenic alteration of ecosystem processes confounds forest management and conservation of rare, declining species. Restoration of forest structure and fire hazard reduction are central goals of forest management policy in the western United States, but restoration priorities and treatments have become increasingly contentious. Numerous studies have documented changes in fire regimes, forest stand structure and species composition following a century of fire exclusion in dry, frequent-fire forests of the western U.S. (e.g., ponderosa pine and dry mixed-conifer). In contrast, wet mixed-conifer forests are thought to have historically burned infrequently with mixed- or high-severity fire—resulting in reduced impacts from fire exclusion and low restoration need—but data are limited. In this study we quantified the current forest habitat of the federally endangered, terrestrial Jemez Mountains salamander (Plethodon neomexicanus) and compared it to dendroecological reconstructions of historical habitat (e.g., stand structure and composition), and fire regime parameters along a gradient from upper ponderosa pine to wet mixed-conifer forests. We found that current fire-free intervals in Jemez Mountains salamander habitat (116–165 years) are significantly longer than historical intervals, even in wet mixed-conifer forests. Historical mean fire intervals ranged from 10 to 42 years along the forest gradient. Low-severity fires were historically dominant across all forest types (92 of 102 fires). Although some mixed- or highseverity fire historically occurred at 67% of the plots over the last four centuries, complete mortality within 1.0 ha plots was rare, and asynchronous within and among sites. Climate was an important driver of temporal variability in fire severity, such that mixed- and high-severity fires were associated with more extreme drought than low-severity fires. Tree density in dry conifer forests historically ranged from open (90 trees/ha) to moderately dense (400 trees/ha), but has doubled on average since fire exclusion. Infill of fire-sensitive tree species has contributed to the conversion of historically dry mixedconifer to wet mixed-conifer forest. We conclude that low-severity fire, which has been absent for over a century, was a critical ecosystem process across the forest gradient in Jemez Mountains salamander habitat, and thus is an important element of ecosystem restoration, resilience, and rare species recovery.

Forest Ecology and Management

Nitrogen dynamics across silvicultural canopy gaps in young forests of western Oregon

Silvicultural canopy gaps are emerging as an alternative management tool to accelerate development of complex forest structure in young, even-aged forests of the Pacific Northwest. The effect of gap creation on available nitrogen (N) is of concern to managers because N is often a limiting nutrient in Pacific Northwest forests. We investigated patterns of N availability in the forest floor and upper mineral soil (0-10 cm) across 6-8-year-old silvicultural canopy gaps in three 50-70-year-old Douglas-fir forests spanning a wide range of soil N capital in the Coast Range and Cascade Mountains of western Oregon. We used extractable ammonium (NH 4 + ) and nitrate (NO 3 - ) pools, net N mineralization and nitrification rates, and NH 4 + and NO 3 - ion exchange resin (IER) concentrations to quantify N availability along north-south transects run through the centers of 0.4 and 0.1 ha gaps. In addition, we measured several factors known to influence N availability, including litterfall, moisture, temperature, and decomposition rates. In general, gap-forest differences in N availability were more pronounced in the mineral soil than in the forest floor. Mineral soil extractable NH 4 + and NO 3 - pools, net N mineralization and nitrification rates, and NH 4 + and NO 3 - IER concentrations were all significantly elevated in gaps relative to adjacent forest, and in several cases exhibited significantly greater spatial variability in gaps than forest. Nitrogen availability along the edges of gaps more often resembled levels in the adjacent forest than in gap centers. For the majority of response variables, there were no significant differences between northern and southern transect positions, nor between 0.4 and 0.1 ha gaps. Forest floor and mineral soil gravimetric percent moisture and temperature showed few differences along transects, while litterfall carbon (C) inputs and litterfall C:N ratios in gaps were significantly lower than in the adjacent forest. Reciprocal transfer incubations of mineral soil samples between gap and forest positions revealed that soil originating from gaps had greater net nitrification rates than forest samples, regardless of incubation environment. Overall, our results suggest that increased N availability in 6-8-year-old silvicultural gaps in young western Oregon forests may be due more to the quality and quantity of litterfall inputs resulting from early-seral species colonizing gaps than by changes in temperature and moisture conditions caused by gap creation.

Forest Ecology and Management

Tree and stand transpiration in a Midwestern bur oak savanna after elm encroachment and restoration thinning

Oak savannas, once common in the Midwest, are now isolated remnants within agricultural landscapes. Savanna remnants are frequently encroached by invasive trees to become woodlands. Thinning and prescribed burning can restore savanna structure, but the ecohydrological effects of managing these remnants are poorly understood. In this study, we measured sap flow (Js) to quantify transpiration in an Iowa bur oak (Quercus macrocarpa) savanna woodland encroached by elms (Ulmus americana), and in an adjacent restored savanna after thinning to remove elms, during summer 2004. Savanna oaks had greater mean daily Js (35.9 L dm-2 day-1) than woodland oaks (20.7 L dm-2 day-1) and elms (12.4 L dm-2 day-1). The response of Js to vapor pressure deficit (D) was unexpectedly weak, although oaks in both stands showed negative correlation between daily Js and D for D > 0.4 kPa. An earlier daily peak in Js in the elm trees showed a possible advantage for water uptake. As anticipated, the woodland's stand transpiration was greater (1.23 mm day-1) than the savanna's (0.35 mm day-1), yet the savanna achieved 30% of the woodland's transpiration with only 11% of its sapwood area. The difference in transpiration influenced water table depths, which were 2 m in the savanna and 6.5 m in the woodland. Regionally, row-crop agriculture has increased groundwater recharge and raised water tables, providing surplus water that perhaps facilitated elm encroachment. This has implications for restoration of savanna remnants. If achieving a savanna ecohydrology is an aim of restoration, then restoration strategies may require buffers, or targeting of large or hydrologically isolated remnants. ?? 2007.

Forest Ecology and Management

Duration of fuels reduction following prescribed fire in coniferous forests of U.S. national parks in California and the Colorado Plateau

Prescribed fire is a widely used forest management tool, yet the long-term effectiveness of prescribed fire in reducing fuels and fire hazards in many vegetation types is not well documented. We assessed the magnitude and duration of reductions in surface fuels and modeled fire hazards in coniferous forests across nine U.S. national parks in California and the Colorado Plateau. We used observations from a prescribed fire effects monitoring program that feature standard forest and surface fuels inventories conducted pre-fire, immediately following an initial (first-entry) prescribed fire and at varying intervals up to >20 years post-fire. A subset of these plots was subjected to prescribed fire again (second-entry) with continued monitoring. Prescribed fire effects were highly variable among plots, but we found on average first-entry fires resulted in a significant post-fire reduction in surface fuels, with litter and duff fuels not returning to pre-fire levels over the length of our observations. Fine and coarse woody fuels often took a decade or longer to return to pre-fire levels. For second-entry fires we found continued fuels reductions, without strong evidence of fuel loads returning to levels observed immediately prior to second-entry fire. Following both first- and second-entry fire there were increases in estimated canopy base heights, along with reductions in estimated canopy bulk density and modeled flame lengths. We did not find evidence of return to pre-fire conditions during our observation intervals for these measures of fire hazard. Our results show that prescribed fire can be a valuable tool to reduce fire hazards and, depending on forest conditions and the measurement used, reductions in fire hazard can last for decades. Second-entry prescribed fire appeared to reinforce the reduction in fuels and fire hazard from first-entry fires.

Forest Ecology and Management

Canopy gap dynamics of second-growth red spruce-northern hardwood stands in West Virginia

Forest restoration requires an understanding of the natural disturbance regime of the target community and estimates of the historic range of variability of ecosystem components (composition, structure, and disturbance processes). Management prescriptions that support specific restoration activities should be consistent with these parameters. In this study, we describe gap-phase dynamics of even-aged, second-growth red spruce-northern hardwood stands in West Virginia that have been significantly degraded following early Twentieth Century harvesting and wildfire. In the current stage of stand development, gaps tended to be small, with mean canopy gap and extended canopy gap sizes of 53.4m2 and 199.3m2, respectively, and a canopy turnover rate of 1.4%year-1. The majority of gaps resulted from the death of one or two trees. American beech snags were the most frequent gap maker, partially due to the elevated presence of beech-bark disease in the study area. Gaps ranged in age from 1 to 28 years, had a mean of 13 years, and were unimodal in distribution. We projected red spruce to be the eventual gap filler in approximately 40% of the gaps. However, we estimated that most average-sized gaps will close within 15-20 years before red spruce canopy ascension is projected (30-60 years). Accordingly, many understory red spruce will require more than one overhead release - an observation verified by the tree-ring record and consistent with red spruce life history characteristics. Based on our observations, silvicultural prescriptions that include overhead release treatments such as thinning from above or small gap creation through selection harvesting could be an appropriate activity to foster red spruce restoration in the central Appalachians. ?? 2010 Elsevier B.V.

Forest Ecology and Management

Space use, forays, and habitat selection in California Spotted Owls (Strix occidentalis occidentalis): New insights from high resolution GPS tracking

Our current understanding of the relationship between imperiled species and forest management can benefit from global positioning system (GPS) technologies. Fauna of lateseral stage forests have historically been difficult to detect and track in rugged terrain, leading to challenges in movement characterization and conservation. We investigated movement of California Spotted Owls ( Strix occidentalis occidentalis ) using automated GPS loggers affixed to 15 owls in the northern Sierra Nevada, California. We used >17,000 locations from individual owls to characterize homerange size, movement distances, and roosting and foraging habitat selection at four spatio-temporal scales (landscape, home range, foray, nightly) during the breeding season (April–August). Additionally, we assessed owl use of Protected Activity Centers (PACs), which are designated by the U.S.D.A. Forest Service to protect nesting and roosting habitat. Our results corroborated some previous findings about habitat requirements of California Spotted Owls, while also revealing new nuances in space use and habitat selection. Roosting and foraging owls selected stands with high canopy cover and large trees at multiple spatio-temporal scales, with foraging owls showing strongest selection at the largest (landscape) scale investigated. Although owls selected for PACs while foraging and roosting, PACs protected less than one quarter of foraging space use (volume of use) and fewer than half of observed roosts during the breeding season. Female owl home ranges were double the size of male home ranges, and distances travelled from the nest by females were 1.3 times greater than distances travelled by males, with non-breeding females travelling farthest and visiting up to six PACs during a single breeding season. Foraying behavior of this sort has not been documented previously in California Spotted Owls. Our findings support protection of later seral stage forest attributes for roosting and foraging California Spotted Owls. Given their selection for later seral forest attributes, strongest evidence of foraging habitat selection at the landscape scale, long distances travelled by owls and limited habitat protection afforded by PACs, habitat connectivity across the landscape is likely an important component for owl conservation, and distribution of current protected areas may be inadequate for this wide-ranging species.

California

Fuel deposition rates of montane and subalpine conifers in the central Sierra Nevada, California, USA

Fire managers and researchers need information on fuel deposition rates to estimate future changes in fuel bed characteristics, determine when forests transition to another fire behavior fuel model, estimate future changes in fuel bed characteristics, and parameterize and validate ecosystem process models. This information is lacking for many ecosystems including the Sierra Nevada in California, USA. We investigated fuel deposition rates and stand characteristics of seven montane and four subalpine conifers in the Sierra Nevada. We collected foliage, miscellaneous bark and crown fragments, cones, and woody fuel classes from four replicate plots each in four stem diameter size classes for each species, for a total of 176 sampling sites. We used these data to develop predictive equations for each fuel class and diameter size class of each species based on stem and crown characteristics. There were consistent species and diameter class differences in the annual amount of foliage and fragments deposited. Foliage deposition rates ranged from just over 50 g m -2 year -1 in small diameter mountain hemlock stands to ???300 g m -2 year -1 for the three largest diameter classes of giant sequoia. The deposition rate for most woody fuel classes increased from the smallest diameter class stands to the largest diameter class stands. Woody fuel deposition rates varied among species as well. The rates for the smallest woody fuels ranged from 0.8 g m -2 year -1 for small diameter stands of Jeffrey pine to 126.9 g m -2 year -1 for very large diameter stands of mountain hemlock. Crown height and live crown ratio were the best predictors of fuel deposition rates for most fuel classes and species. Both characteristics reflect the amount of crown biomass including foliage and woody fuels. Relationships established in this study allow predictions of fuel loads to be made on a stand basis for each of these species under current and possible future conditions. These predictions can be used to estimate fuel treatment longevity, assist in determining fuel model transitions, and predict future changes in fuel bed characteristics.

Forest Ecology and Management

Level and pattern of overstory retention shape the abundance and long-term dynamics of natural and created snags

Standing dead trees, or snags, serve myriad functions in natural forests, but are often scarce in forests managed for timber production. Variable retention (VR), the retention of live and dead trees through harvest, has been adopted globally as a less intensive form of regeneration harvest. In this study, we explore how two key elements of VR systems — level (amount) and spatial pattern of live-tree retention — affect the carryover and post-harvest dynamics of natural and artificially created snags. We present nearly two decades of data from the DEMO Study, a regional-scale experiment in VR harvests of Douglas-fir-dominated forests in the Pacific Northwest. Snag losses to harvest were greater at 15 than at 40% retention (67 vs. 47% declines in density) and greater in dispersed than in aggregated treatments (64 vs. 50% declines). Densities of hard and tall (≥5 m) snags were particularly sensitive to low-level dispersed retention, declining by 76 and 81%, respectively. Despite these losses, post-harvest densities correlated with pre-harvest densities for most snag size and decay classes. In contrast to initial harvest effects, snag densities changed minimally over the post-harvest period (years 1 to 18 or 19), with low rates of recruitment offsetting low rates of loss. Post-harvest survival of snags was greater at 15 than at 40% retention (79 vs. 69%), as were rates of decay (68 vs. 52% of hard snags transitioned to soft). However, pattern had no effect on either process. Snag recruitment did not vary with retention level or pattern at the scale of the 13-ha harvest unit, but was several-fold greater in the 1-ha aggregates (14.3–27.8 snags ha −1 ) than in the corresponding dispersed treatments (4.2–5.3 snags ha −1 ). Snag size (diameter) distributions showed greater change in dispersed than in aggregated treatments, reflecting greater loss of smaller snags and recruitment biased toward larger snags. Created snags showed uniformly high survival (97%), irrespective of treatment, but rates of decay were greater at lower retention. If a goal of VR is to sustain snag abundance and diversity through harvest, emphasis should be placed on minimizing initial losses, either by reducing the intensity of felling in areas of dispersed retention or locating forest aggregates in areas of greater initial snag density, diversity, or incipient decay.

Forest Ecology and Management

Patterns and correlates of giant sequoia foliage dieback during California’s 2012–2016 hotter drought

Hotter droughts – droughts in which unusually high temperatures exacerbate the effects of low precipitation – are expected to increase in frequency and severity in coming decades, challenging scientists and managers to identify which parts of forested landscapes may be most vulnerable. In 2014, in the middle of California’s historically unprecedented 2012–2016 hotter drought, we noticed apparently drought-induced foliage dieback in giant sequoias ( Sequoiadendron giganteum Lindl. [Buchholz]) in Sequoia and Kings Canyon national parks, California. Characteristics of the dieback were consistent with a controlled process of drought-induced senescence: younger (distal) shoots remained green while older (proximal) shoots were preferentially shed. As part of an ongoing interdisciplinary effort to understand and map sequoia vulnerability to hotter droughts, we reviewed historical records for evidence of previous foliage dieback events, surveyed dieback along trail corridors in eight sequoia groves, and analyzed tree-ring data from a high- and a low-foliage-dieback area. In sharp contrast to the greatly elevated mortality of other coniferous species found at low and middle elevations, we estimate that <1% of sequoias died during the drought. Foliage dieback was notably elevated in 2014 – the most severe single drought year in our 122-year record – but much lower in subsequent years. We found no historical records of similar foliage dieback during previous droughts. Dieback in 2014 was highly variable both within and among groves, ranging from virtually no dieback in some areas to nearly 50% in others. Dieback was highest (1) at low elevations, probably due to higher temperatures, reduced snowpack, and earlier snowmelt; (2) in areas of low adult sequoia densities, which likely reflect intrinsically more stressful sites; and (3) on steep slopes, probably reflecting reduced water availability. Average sequoia ring widths were narrower at the high-dieback than the low-dieback tree-ring site, but for reasons that remain unclear the sites did not differ in their proportional ring-width responses to past droughts. Collectively, our results suggest that giant sequoia vulnerability to hotter droughts may be spatially quite variable, and that at least some of that variability can be explained by metrics related to site water balance. Future research will focus on integrating our results with physiological and remote-sensing data, including tracking sequoias as they recover from the drought.

California

Resilience of a heavily logged grove of giant sequoia ( Sequoiadendron giganteum ) in Kings Canyon National Park, California

The Big Stump Grove of giant sequoia ( Sequoiadendron giganteum (Lindl.) Buchholz) was heavily logged between 1883 and 1889 and the stand naturally regenerated from seed following logging. In 1968, as part of a 100% sequoia tree inventory, all living sequoias ( n = 3587) and dead trees and stumps ( n =588) were measured (diameter at breast height, dbh) and mapped. A comparison of pre- to post-logging (85 years later in 1968) stand characteristics showed the estimated basal area of 56.7 m 2 ha −1 in the pre-cut 1883 Big Stump Grove was very similar to the population mean basal area of 30 other giant sequoia groves (with more than 30 trees) in Sequoia and Kings Canyon National Parks. Sequoia density in 1968 was 1.5 times higher than the population mean, and over 45% of the basal area had been recovered after only 85 years. Assuming most re-establishment occurred over roughly a 9 year period (1883–1892), the diameter growth rate of trees less than 1.95 m dbh, averaged 6.1–6.8 mm year −1 but greatly varied as the 24 trees in the 1.8 m size class had a mean diameter growth rate of 21–24 mm year −1 . Data generated by dividing the grove into 0.25 ha contiguous plots indicated that only about 3.3 ha of the pre-cut 1883 grove did not have sequoia regeneration whereas 16.5 ha of the 1968 grove had sequoia regeneration but no sign of logs or stumps. The proportion of only-regeneration plots was significantly greater ( P <0.05) in peripheral areas compared to core areas of the grove. A stage projection model showed that with typical natality, growth, and mortality rates in the current ( t =0; 1968) stand, overrepresentation of 0.3–1.2 m dbh trees may produce a bimodal size distribution lasting perhaps 800 years or more into the future. Giant sequoia stand characteristics such as age and size structure are not highly resilient and may take several centuries to approach the ‘domain’ of age or size structure typical of old-growth sequoia forests. Grove boundaries may be less stable following a major disturbance.

Forest Ecology and Management

Redistribution of soil nitrogen, carbon and organic matter by mechanical disturbance during whole-tree harvesting in northern hardwoods

To investigate whether mechanical mixing during harvesting could account for losses observed from forest floor, we measured surface disturbance on a 22 ha watershed that was whole-tree harvested. Surface soil on each 10 cm interval along 81, randomly placed transects was classified immediately after harvesting as mineral or organic, and as undisturbed, depressed, rutted, mounded, scarified, or scalped (forest floor scraped away). We quantitatively sampled these surface categories to collect soil in which preharvest forest floor might reside after harvest. Mechanically mixed mineral and organic soil horizons were readily identified. Buried forest floor under mixed mineral soil occurred in 57% of mounds with mineral surface soil. Harvesting disturbed 65% of the watershed surface and removed forest floor from 25% of the area. Mechanically mixed soil under ruts with organic or mineral surface soil, and mounds with mineral surface soil contained organic carbon and nitrogen pools significantly greater than undisturbed forest floor. Mechanical mixing into underlying mineral soil could account for the loss of forest floor observed between the preharvest condition and the second growing season after whole-tree harvesting.

New Hampshire

Intra-specific competition (crowding) of giant sequoias ( Sequoiadendron giganteum )

Information on the size and location of 1916 giant sequoias (Sequoiadendron giganteum (Lindl.) Buchholz) in Muir Grove, Sequoia National Park, in the southern Sierra Nevada of California was used to assess intra-specific crowding. Study objectives were to: (1) determine which parameters associated with intra-specific competition (i.e. size and distance to nearest neighbor, crowding/root system area overlap, or number of neighbors) might be important in spatial pattern development, growth, and survivorship of established giant sequoias; (2) quantify the level of intra-specific crowding of different sized live sequoias based on a model of estimated overlapping root system areas (i.e. an index of relative crowding); (3) compare the level of intra-specific crowding of similarly sized live and dead giant sequoias (less than 30 cm diameter at breast height (dbh) at the time of inventory (1969). Mean distances to the nearest live giant sequoia neighbor were not significantly different (at α = 0.05) for live and dead sequoias in similar size classes. A zone of influence competition model (i.e. index of crowding) based on horizontal overlap of estimated root system areas was developed for 1753 live sequoias. The model, based only on the spatial arrangement of live sequoias, was then tested on dead sequoias of less than 30 cm dbh (n = 163 trees; also recorded in 1969). The dead sequoias had a significantly higher crowding index than 561 live trees of similar diameter. Results showed that dead sequoias of less than 16.6 cm dbh had a significantly greater mean number of live neighbors and mean crowding index than live sequoias of similar size. Intra-specific crowding may be an important mechanism in determining the spatial distribution of sequoias in old-growth forests.

California

Arboreal lichen in uncut and partially cut subalpine fir stands in woodland caribou habitat, northern Idaho and southeastern British Columbia

To better understand the effects of partial cutting on arboreal lichen production within woodland caribou ( Rangifer tarandus caribou ) habitat, lichen was hand picked from 1228 branches on 307 subalpine fir ( Abies lasiocarpa ) trees in Idaho (ID), and in British Columbia (BC). Lichen biomass from partially cut stands was compared with biomass on trees from adjacent uncut stands at each site. Arboreal lichen biomass did not differ significantly between uncut and partially cut stands. Total number of branches per tree did not differ significantly between uncut and partially cut stands. Live branches had more lichen than dead branches. Species composition of arboreal lichen changed in partially cut stands compared with adjacent uncut stands. The ratio of live to dead branches was substantially different within the BC partial cut.

Forest Ecology and Management