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• 1. [Article] Post-fire tree establishment and early cohort development in conifer forests of the western Cascades of Oregon, USA

  To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by the author(s) ...

  Citation × Citation Citation Abstract Copy Title: Post-fire tree establishment and early cohort development in conifer forests of the western Cascades of Oregon, USA Author: Tepley, Alan J., Swanson, Frederick J., Spies, Thomas A. To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by the author(s) and published by the Ecological Society of America. The published article can be found at: http://www.esajournals.org/loi/ecsp. Title: Post-fire tree establishment and early cohort development in conifer forests of the western Cascades of Oregon, USA Author: Tepley, Alan J., Swanson, Frederick J., Spies, Thomas A. To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by the author(s) and published by the Ecological Society of America. The published article can be found at: http://www.esajournals.org/loi/ecsp. Close

• 2. [Article] Exploration of statistical methods for synthesizing the effects of variable-retention harvesting on multiple taxa

  Variable-retention harvesting was proposed to reduce loss of biodiversity and ecosystem processes associated with late-seral Douglas-fir (Pseudotsuga menziesii) forests in the Pacific Northwest. The Demonstration ...

  Citation × Citation Citation Abstract Copy Title: Exploration of statistical methods for synthesizing the effects of variable-retention harvesting on multiple taxa Author: Lam, Tzeng Yih Variable-retention harvesting was proposed to reduce loss of biodiversity and ecosystem processes associated with late-seral Douglas-fir (Pseudotsuga menziesii) forests in the Pacific Northwest. The Demonstration of Ecosystem Management Options experiment was established to test this hypothesis. Analysis presents various challenges to drawing statistical inferences about treatment effects. This dissertation explored novel statistical methods for understanding the response of multiple forest taxa to variable-retention harvesting. Excessive zero counts are common among terrestrial small mammal species that are captured infrequently. Zero-inflated and hurdle models are appealing tools for analyzing these data. A simulation was performed to understand the properties and robustness of these models. When true mean abundance was low, the estimated parameters from these models were highly unstable. Goodness of fit criteria could not discern among the processes generating the data. The Poisson and negative binomial Generalized Linear Models (GLMs) were fitted to four small mammal species with different rates of capture. Predictors included several variables representing vegetation structure. These models and overdispersed Poisson were then specified as Generalized Linear Mixed Models (GLMMs) to account for nesting and blocking in the experimental design. The fitted GLMs indicated that predictors were not consistent among models for the infrequently captured species. Differences in estimated coefficients between GLMs and GLMMs were noticeable. The overdispersed Poisson GLMM was suggested to be most suitable. Structural Equation Modeling (SEM) is suitable for modeling interactions of many cause-and-effect relationships in forest ecosystems. SEM was applied to understand overstory-understory relationships of late-seral herb species under mature forest conditions and immediately after variable-retention harvesting. In undisturbed forests, light attenuation, belowground competition and stand age were the primary drivers of late-seral herb cover. After variable-retention harvesting, microclimatic stresses were inferred to primarily affect late-seral species diversity and composition. Logging debris had little discernible effect on the change in the late-seral herb community. The explored statistical models complement conventional methods for studying the effects of variable-retention harvesting. These models address distributional issues of response data and provide further insight into the complex processes driving managed forest ecosystems. Future analyses should apply a suite of statistical models to gain different perspectives. Title: Exploration of statistical methods for synthesizing the effects of variable-retention harvesting on multiple taxa Author: Lam, Tzeng Yih Variable-retention harvesting was proposed to reduce loss of biodiversity and ecosystem processes associated with late-seral Douglas-fir (Pseudotsuga menziesii) forests in the Pacific Northwest. The Demonstration of Ecosystem Management Options experiment was established to test this hypothesis. Analysis presents various challenges to drawing statistical inferences about treatment effects. This dissertation explored novel statistical methods for understanding the response of multiple forest taxa to variable-retention harvesting. Excessive zero counts are common among terrestrial small mammal species that are captured infrequently. Zero-inflated and hurdle models are appealing tools for analyzing these data. A simulation was performed to understand the properties and robustness of these models. When true mean abundance was low, the estimated parameters from these models were highly unstable. Goodness of fit criteria could not discern among the processes generating the data. The Poisson and negative binomial Generalized Linear Models (GLMs) were fitted to four small mammal species with different rates of capture. Predictors included several variables representing vegetation structure. These models and overdispersed Poisson were then specified as Generalized Linear Mixed Models (GLMMs) to account for nesting and blocking in the experimental design. The fitted GLMs indicated that predictors were not consistent among models for the infrequently captured species. Differences in estimated coefficients between GLMs and GLMMs were noticeable. The overdispersed Poisson GLMM was suggested to be most suitable. Structural Equation Modeling (SEM) is suitable for modeling interactions of many cause-and-effect relationships in forest ecosystems. SEM was applied to understand overstory-understory relationships of late-seral herb species under mature forest conditions and immediately after variable-retention harvesting. In undisturbed forests, light attenuation, belowground competition and stand age were the primary drivers of late-seral herb cover. After variable-retention harvesting, microclimatic stresses were inferred to primarily affect late-seral species diversity and composition. Logging debris had little discernible effect on the change in the late-seral herb community. The explored statistical models complement conventional methods for studying the effects of variable-retention harvesting. These models address distributional issues of response data and provide further insight into the complex processes driving managed forest ecosystems. Future analyses should apply a suite of statistical models to gain different perspectives. Close

• 3. [Article] Mesoscale Variation in Snag and Hardwood Densities and Sizes in Old-Growth Forests in Western Oregon

  To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by Society ...

  Citation × Citation Citation Abstract Copy Title: Mesoscale Variation in Snag and Hardwood Densities and Sizes in Old-Growth Forests in Western Oregon Author: Bright, Cheryl, Puettmann, Klaus, Ares, Adrian To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by Society of American Foresters and can be found at: http://www.safnet.org/. Title: Mesoscale Variation in Snag and Hardwood Densities and Sizes in Old-Growth Forests in Western Oregon Author: Bright, Cheryl, Puettmann, Klaus, Ares, Adrian To the best of our knowledge, one or more authors of this paper were federal employees when contributing to this work. This is the publisher’s final pdf. The published article is copyrighted by Society of American Foresters and can be found at: http://www.safnet.org/. Close

• 4. [Article] Approaches for Characterizing Plant Physiological Responses to Environmental Stress

  It is uncertain how predicted changes in climate will impact vegetation responses and plant species’ distributions because the physiological mechanisms underlying thresholds for damage are not well understood, ...

  Citation × Citation Citation Abstract Copy Title: Approaches for Characterizing Plant Physiological Responses to Environmental Stress Author: Marias, Danielle E. It is uncertain how predicted changes in climate will impact vegetation responses and plant species’ distributions because the physiological mechanisms underlying thresholds for damage are not well understood, and responses to stress vary by functional type and developmental stage. Thus, it is crucial to investigate physiological responses to heat and drought stress on multiple species, populations, and growth stages with diverse approaches. In this dissertation, I employ a suite of physiological and modeling methods to inform our knowledge of plant physiological responses to environmental stress in Coffea arabica saplings, Pseudotsuga mensizeii (PSME) and Pinus ponderosa (PIPO) seedlings, and old-growth PIPO. In Chapter 2, I evaluate the effect of leaf age and methodology on the thermotolerance or heat tolerance of C. arabica leaf discs using chlorophyll fluorescence and electrolyte leakage methods. I found that mature leaves were more heat tolerant than expanding leaves, longer time between temperature exposure and measurement yielded more accurate thermotolerance assessments, and photochemistry was more heat-sensitive than cell membranes. To complement the second chapter investigating heat stress responses on detached leaf discs, Chapter 3 examines the effect of leaf age and heat stress duration (45 min or 90 min) on whole-plant physiological responses and capacity to recover in C. arabica by monitoring chlorophyll fluorescence (F[subscript V]/F[subscript M]), gas exchange, and foliar non-structural carbohydrate (NSC) dynamics in situ in response to a simulated heat wave (49°C) in a growth chamber. I found that the 90 min treatment resulted in greater photosynthetic damage and slower recovery than the 45 min treatment, expanding leaves recovered more slowly than in mature leaves, and both heat treatments inhibited flowering. A leaf energy balance model demonstrated that heat stress would be exacerbated by drought-induced stomatal closure. Heat treatment duration significantly impacted NSC dynamics that were closely related to reproduction and repair. Because seedling establishment governs species’ distributions, and because seedlings are particularly threatened by high temperatures at the soil surface, in Chapter 4 I examined the thermotolerance and heat stress responses of PIPO and PSME seedling populations from contrasting climates. Unexpectedly, I found that PSME was more heat tolerant the PIPO. I also monitored physiological recovery after exposure to a simulated heat wave (45°C) by measuring photosynthesis, F[subscript V]/F[subscript M], foliar NSC, and carbon stable isotope ratios (proxy for intrinsic water use efficiency, iWUE). Heat stress responses were consistent with phenotypic plasticity and reflected the conditions under which the plants were grown, while iWUE, a measure of potential drought resistance, was consistent with ecotypic differentiation and the climates from which the seedlings originated. To investigate responses to environmental stress on larger temporal and spatial scales without the challenges of making repeated physiological measurements on old-growth trees, in Chapter 5 I used long-term trajectories of tree-ring growth and carbon and oxygen isotopes of tree-ring cellulose (δ¹³Ccell, and δ¹⁸Ocell) to successfully predict the stand characteristics of two sets (upland, riparian) of old-growth PIPO using the Physiological Principles in Predicting Growth (3-PG) model, the δ¹³Ccell submodel, and a δ¹⁸Ocell submodel added by me. The expanded model helped to explain physiological drivers underlying the different tree-ring growth, δ¹³Ccell, and δ¹⁸Ocell trajectories measured at the upland and riparian sites. The combination of both δ¹⁸O and δ¹³Ccell submodels provided a useful and novel way to constrain 3-PG. This dissertation demonstrates an innovative strategy of applying diverse approaches to understand the physiological mechanisms behind vegetation responses to environmental stress. Title: Approaches for Characterizing Plant Physiological Responses to Environmental Stress Author: Marias, Danielle E. It is uncertain how predicted changes in climate will impact vegetation responses and plant species’ distributions because the physiological mechanisms underlying thresholds for damage are not well understood, and responses to stress vary by functional type and developmental stage. Thus, it is crucial to investigate physiological responses to heat and drought stress on multiple species, populations, and growth stages with diverse approaches. In this dissertation, I employ a suite of physiological and modeling methods to inform our knowledge of plant physiological responses to environmental stress in Coffea arabica saplings, Pseudotsuga mensizeii (PSME) and Pinus ponderosa (PIPO) seedlings, and old-growth PIPO. In Chapter 2, I evaluate the effect of leaf age and methodology on the thermotolerance or heat tolerance of C. arabica leaf discs using chlorophyll fluorescence and electrolyte leakage methods. I found that mature leaves were more heat tolerant than expanding leaves, longer time between temperature exposure and measurement yielded more accurate thermotolerance assessments, and photochemistry was more heat-sensitive than cell membranes. To complement the second chapter investigating heat stress responses on detached leaf discs, Chapter 3 examines the effect of leaf age and heat stress duration (45 min or 90 min) on whole-plant physiological responses and capacity to recover in C. arabica by monitoring chlorophyll fluorescence (F[subscript V]/F[subscript M]), gas exchange, and foliar non-structural carbohydrate (NSC) dynamics in situ in response to a simulated heat wave (49°C) in a growth chamber. I found that the 90 min treatment resulted in greater photosynthetic damage and slower recovery than the 45 min treatment, expanding leaves recovered more slowly than in mature leaves, and both heat treatments inhibited flowering. A leaf energy balance model demonstrated that heat stress would be exacerbated by drought-induced stomatal closure. Heat treatment duration significantly impacted NSC dynamics that were closely related to reproduction and repair. Because seedling establishment governs species’ distributions, and because seedlings are particularly threatened by high temperatures at the soil surface, in Chapter 4 I examined the thermotolerance and heat stress responses of PIPO and PSME seedling populations from contrasting climates. Unexpectedly, I found that PSME was more heat tolerant the PIPO. I also monitored physiological recovery after exposure to a simulated heat wave (45°C) by measuring photosynthesis, F[subscript V]/F[subscript M], foliar NSC, and carbon stable isotope ratios (proxy for intrinsic water use efficiency, iWUE). Heat stress responses were consistent with phenotypic plasticity and reflected the conditions under which the plants were grown, while iWUE, a measure of potential drought resistance, was consistent with ecotypic differentiation and the climates from which the seedlings originated. To investigate responses to environmental stress on larger temporal and spatial scales without the challenges of making repeated physiological measurements on old-growth trees, in Chapter 5 I used long-term trajectories of tree-ring growth and carbon and oxygen isotopes of tree-ring cellulose (δ¹³Ccell, and δ¹⁸Ocell) to successfully predict the stand characteristics of two sets (upland, riparian) of old-growth PIPO using the Physiological Principles in Predicting Growth (3-PG) model, the δ¹³Ccell submodel, and a δ¹⁸Ocell submodel added by me. The expanded model helped to explain physiological drivers underlying the different tree-ring growth, δ¹³Ccell, and δ¹⁸Ocell trajectories measured at the upland and riparian sites. The combination of both δ¹⁸O and δ¹³Ccell submodels provided a useful and novel way to constrain 3-PG. This dissertation demonstrates an innovative strategy of applying diverse approaches to understand the physiological mechanisms behind vegetation responses to environmental stress. Close

• 5. [Article] Mixed-Severity Fire Effects on Biological Legacies and Vegetation Response in Pseudotsuga Forests of Western Oregon's Central Cascades, USA

  Mixed-severity fire occurrence is increasingly recognized in Pseudotsuga forests of the Pacific Northwest, but questions remain about how tree mortality varies, and forest structure is altered, across ...

  Citation × Citation Citation Abstract Copy Title: Mixed-Severity Fire Effects on Biological Legacies and Vegetation Response in Pseudotsuga Forests of Western Oregon's Central Cascades, USA Author: Dunn, Christopher J. Mixed-severity fire occurrence is increasingly recognized in Pseudotsuga forests of the Pacific Northwest, but questions remain about how tree mortality varies, and forest structure is altered, across the disturbance gradient observed in these fires. Therefore, we sampled live and dead biological legacies at 45 one ha plots, with four 0.10 ha nested plots, stratified across an unburned, low, moderate and high-severity fire gradient. We used severity estimates based on differenced Normalized Burn Ratio (dNBR), and captured a disturbance gradient, but plots in our low-severity class underestimated fire effects because of misclassification or delayed mortality. We estimated probability of mortality for shade-intolerant (Douglas-fir, incense-cedar, sugar pine) and shade-tolerant (western hemlock, western redcedar, true fir) trees from 5,079 sampled trees and snags. The probability of mortality was higher for shade-tolerant species across all fire-severity classes, and decreased with increasing DBH except for western hemlock. Only large, shade-intolerant trees survived high-severity fire. Post-fire snag fall and fragmentation were estimated from 2,746 sampled snags and logs. The probability of snag fall decreased with increasing DBH for all species, and was positively correlated with fire severity, except for Douglas-fir that had a higher probability following low-severity fire. Snag fragmentation was positively correlated with DBH and fire severity for all species. We also estimated the coefficient of variation within- and among-plots by fire severity class, as well as across all sampled conditions. Structural attributes varied more within- than among-plots, likely a result of increasing sub-hectare patchy mortality as fire intensity increased. Although vertical and horizontal structural diversity increased at sub-hectare scales, the coefficient of variation was highest for all structural attributes when compared across all fire severity classes. Therefore, the range of fire effects observed in mixed-severity fires may be functionally important in creating structural complexity across landscapes, which is an important attribute of old-growth forests in the Pacific Northwest. Understory vegetation response to mixed-severity fires has not been characterized for these forests even though the majority of vegetation diversity is found in these vegetation layers. Therefore, we sampled forest structure (1000 m² circular plots) and understory vegetation (100 m² plots) at 168 collocated plots stratified across unburned, low, moderate and high-severity conditions 10 years (Tiller Complex) and 22 years (Warner Fire) post-fire. We focused on shrub species, but sampled forbs, graminoids, ferns and moss as functional groups. Offsite colonization and fire stimulated soil seedbanks increased the total species richness from 23 to 46. The life-history strategies of residual and colonizing species resulted in three dominant species response-curves to the magnitude of disturbance: 1) 'disturbance-sensitive', when relative abundance was highest in unburned plots and continued to decline with increasing fire severity, 2) 'disturbance-stimulated', when relative abundance was highest following low or moderate-severity fire and 3) 'disturbance-amplified', when relative abundance increased with increasing fire severity. Residual and colonizing species assemblages promoted five or six distinct understory communities, dominantly driven by legacy tree basal area rather than the proportion of basal area killed. Understory communities were rarely associated with one disturbance severity class as fire refugia, variation in overstory and understory fire severity, and compensatory conditions offset fire effects. Early-seral habitats were the most different from unburned forests, but were not the only post-fire conditions important across these burned landscapes. Interactions among live and dead forest structures following low or moderate-severity fire, and the vegetation response to these conditions, are also unique to the post-fire landscape and likely important for various wildlife species. Therefore, if ecological forestry paradigms focus dominantly on creating old-growth structure or early-seral habitats, they might exclude important conditions that contribute to the landscape structural complexity created by mixed-severity fires. Additionally, tree regeneration response to mixed-severity fires has not been characterized for these forests even though they offer insight into one aspect of the resilience of these ecosystems to disturbance. Therefore, we sampled forest structure (1000 m² circular plots) and regeneration dynamics (100 m² plots) at 168 collocated plots stratified across unburned, low, moderate and high-severity conditions 10 years (Tiller Complex) and 22 years (Warner Fire) post-fire. The largest marginal increase in tree mortality (stems ha⁻¹) occurred between unburned and low-severity fires, given preferential mortality of small trees and shade-tolerant species, but basal area mortality had the largest marginal increase moving from moderate to high-severity. Pairwise comparisons of legacy tree basal area between low and moderate-severity weren’t as significant as other comparisons, but did capture a gradient of increasing fire effects. Quadratic mean diameter and canopy base height were positively correlated with fire severity as incrementally larger trees were killed and canopy ascension followed. Regeneration density increased regardless of severity, relative to unburned forests (median density of 1,384 trees ha⁻¹), but the highest median density (16,220 trees ha⁻¹) followed low-severity fire at the Tiller Complex and moderate-severity fire (14,472 trees ha⁻¹) at Warner Fire. Plot-level average species richness was highest following these same fire severity classes, supporting the Intermediate Disturbance Hypothesis. Statistically distinct regeneration communities occurred across the fire severity gradient at both fire sites. The relative abundance of shade-tolerant tree species decreased as fire severity increased, except for a divergent response following stand-initiation at the Warner Fire. While divergent successional pathways were evident within a couple decades following stand-initiation, low or moderate-severity fires also modified successional trajectories and may be the most functionally important disturbance magnitude because it has the greatest potential to increase compositional and structural diversity. Incorporating mixed-severity fire effects into landscape management of Pseudotsuga forests could increase structural complexity at stand and landscape-scales. Title: Mixed-Severity Fire Effects on Biological Legacies and Vegetation Response in Pseudotsuga Forests of Western Oregon's Central Cascades, USA Author: Dunn, Christopher J. Mixed-severity fire occurrence is increasingly recognized in Pseudotsuga forests of the Pacific Northwest, but questions remain about how tree mortality varies, and forest structure is altered, across the disturbance gradient observed in these fires. Therefore, we sampled live and dead biological legacies at 45 one ha plots, with four 0.10 ha nested plots, stratified across an unburned, low, moderate and high-severity fire gradient. We used severity estimates based on differenced Normalized Burn Ratio (dNBR), and captured a disturbance gradient, but plots in our low-severity class underestimated fire effects because of misclassification or delayed mortality. We estimated probability of mortality for shade-intolerant (Douglas-fir, incense-cedar, sugar pine) and shade-tolerant (western hemlock, western redcedar, true fir) trees from 5,079 sampled trees and snags. The probability of mortality was higher for shade-tolerant species across all fire-severity classes, and decreased with increasing DBH except for western hemlock. Only large, shade-intolerant trees survived high-severity fire. Post-fire snag fall and fragmentation were estimated from 2,746 sampled snags and logs. The probability of snag fall decreased with increasing DBH for all species, and was positively correlated with fire severity, except for Douglas-fir that had a higher probability following low-severity fire. Snag fragmentation was positively correlated with DBH and fire severity for all species. We also estimated the coefficient of variation within- and among-plots by fire severity class, as well as across all sampled conditions. Structural attributes varied more within- than among-plots, likely a result of increasing sub-hectare patchy mortality as fire intensity increased. Although vertical and horizontal structural diversity increased at sub-hectare scales, the coefficient of variation was highest for all structural attributes when compared across all fire severity classes. Therefore, the range of fire effects observed in mixed-severity fires may be functionally important in creating structural complexity across landscapes, which is an important attribute of old-growth forests in the Pacific Northwest. Understory vegetation response to mixed-severity fires has not been characterized for these forests even though the majority of vegetation diversity is found in these vegetation layers. Therefore, we sampled forest structure (1000 m² circular plots) and understory vegetation (100 m² plots) at 168 collocated plots stratified across unburned, low, moderate and high-severity conditions 10 years (Tiller Complex) and 22 years (Warner Fire) post-fire. We focused on shrub species, but sampled forbs, graminoids, ferns and moss as functional groups. Offsite colonization and fire stimulated soil seedbanks increased the total species richness from 23 to 46. The life-history strategies of residual and colonizing species resulted in three dominant species response-curves to the magnitude of disturbance: 1) 'disturbance-sensitive', when relative abundance was highest in unburned plots and continued to decline with increasing fire severity, 2) 'disturbance-stimulated', when relative abundance was highest following low or moderate-severity fire and 3) 'disturbance-amplified', when relative abundance increased with increasing fire severity. Residual and colonizing species assemblages promoted five or six distinct understory communities, dominantly driven by legacy tree basal area rather than the proportion of basal area killed. Understory communities were rarely associated with one disturbance severity class as fire refugia, variation in overstory and understory fire severity, and compensatory conditions offset fire effects. Early-seral habitats were the most different from unburned forests, but were not the only post-fire conditions important across these burned landscapes. Interactions among live and dead forest structures following low or moderate-severity fire, and the vegetation response to these conditions, are also unique to the post-fire landscape and likely important for various wildlife species. Therefore, if ecological forestry paradigms focus dominantly on creating old-growth structure or early-seral habitats, they might exclude important conditions that contribute to the landscape structural complexity created by mixed-severity fires. Additionally, tree regeneration response to mixed-severity fires has not been characterized for these forests even though they offer insight into one aspect of the resilience of these ecosystems to disturbance. Therefore, we sampled forest structure (1000 m² circular plots) and regeneration dynamics (100 m² plots) at 168 collocated plots stratified across unburned, low, moderate and high-severity conditions 10 years (Tiller Complex) and 22 years (Warner Fire) post-fire. The largest marginal increase in tree mortality (stems ha⁻¹) occurred between unburned and low-severity fires, given preferential mortality of small trees and shade-tolerant species, but basal area mortality had the largest marginal increase moving from moderate to high-severity. Pairwise comparisons of legacy tree basal area between low and moderate-severity weren’t as significant as other comparisons, but did capture a gradient of increasing fire effects. Quadratic mean diameter and canopy base height were positively correlated with fire severity as incrementally larger trees were killed and canopy ascension followed. Regeneration density increased regardless of severity, relative to unburned forests (median density of 1,384 trees ha⁻¹), but the highest median density (16,220 trees ha⁻¹) followed low-severity fire at the Tiller Complex and moderate-severity fire (14,472 trees ha⁻¹) at Warner Fire. Plot-level average species richness was highest following these same fire severity classes, supporting the Intermediate Disturbance Hypothesis. Statistically distinct regeneration communities occurred across the fire severity gradient at both fire sites. The relative abundance of shade-tolerant tree species decreased as fire severity increased, except for a divergent response following stand-initiation at the Warner Fire. While divergent successional pathways were evident within a couple decades following stand-initiation, low or moderate-severity fires also modified successional trajectories and may be the most functionally important disturbance magnitude because it has the greatest potential to increase compositional and structural diversity. Incorporating mixed-severity fire effects into landscape management of Pseudotsuga forests could increase structural complexity at stand and landscape-scales. Close