Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.     

Response of bark beetles and their natural enemies to fire and fire surrogate treatments in mixed-conifer forests in western Montana

Four treatments (control, burn-only, thin-only, and thin-and-burn) were evaluated for their effects on bark beetle-caused mortality in both the short-term (one to four years) and the long-term (seven years) in mixed-conifer forests in western Montana, USA. In addition to assessing bark beetle responses to these treatments, we also measured natural enemy landing rates and resin flow of ponderosa pine (Pinus ponderosa) the season fire treatments were implemented. All bark beetles were present at low population levels (non-outbreak) for the duration of the study. Post-treatment mortality of trees due to bark beetles was lowest in the thin-only and control units and highest in the units receiving burns. Three tree-killing bark beetle species responded positively to fire treatments: Douglas-fir beetle (Dendroctonus pseudotsugae), pine engraver (Ips pini), and western pine beetle (Dendroctonus brevicomis). Red turpentine beetle (Dendroctonus valens) responded positively to fire treatments, but never caused mortality. Three fire damage variables tested (height of crown scorch, percent circumference of the tree bole scorched, or degree of ground char) were significant factors in predicting beetle attack on trees. Douglas-fir beetle and pine engraver responded rapidly to increased availability of resources (fire-damaged trees); however, successful attacks dropped rapidly once these resources were depleted. Movement to green trees by pine engraver was not observed in plots receiving fire treatments, or in thinned plots where slash supported substantial reproduction by this beetle. The fourth tree-killing beetle present at the site, the mountain pine beetle, did not exhibit responses to any treatment. Natural enemies generally arrived at trees the same time as host bark beetles. However, the landing rates of only one, Medetera spp., was affected by treatment. This predator responded positively to thinning treatments. This insect was present in very high numbers indicating a regulatory effect on beetles, at least in the short-term, in thinned stands. Resin flow decreased from June to August. However, resin flow was significantly higher in trees in August than in June in fire treatments. Increased flow in burned trees later in the season did not affect beetle attack success. Overall, responses by beetles to treatments were short-term and limited to fire-damaged trees. Expansions into green trees did not occur. This lack of spread was likely due to a combination of high tree vigor in residual stands and low background populations of bark beetles.     

Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Thinning and prescribed fire effects on dwarf mistletoe severity in an eastern Cascade Range dry forest,Washington

Forest thinning and prescribed fire practices are widely used, either separately or in combination, to address tree stocking, species composition, and wildland fire concerns in western US mixed conifer forests. We examined the effects of these fuel treatments alone and combined on dwarf mistletoe infection severity immediately after treatment and for the following 100 years. Thinning, burning, thin + burn, and control treatments were applied to 10 ha units; each treatment was replicated three times. Dwarf mistletoe was found in ponderosa pine and/or Douglas-fir in all units prior to treatment. Stand infection severity was low to moderate, and severely infected trees were the largest in the overstory. Thinning produced the greatest reductions in tree stocking and mistletoe severity. Burning reduced stocking somewhat less because spring burns were relatively cool with spotty fuel consumption and mortality. Burning effects on vegetation were enhanced when combined with thinning; thin + burn treatments also reduced mistletoe severity in all size classes. Stand growth simulations using the Forest Vegetation Simulator (FVS) showed a trend of reduced mistletoe spread and intensification over time for all active treatments. When thinned and unthinned treatments were compared, thinning reduced infected basal area and treatment effects were obvious, beginning in the second decade. The same was true with burned and unburned treatments. Treatment effects on infected tree density were similar to infected basal area; however, treatment effects diminished after 20 years, suggesting a re-treatment interval for dwarf mistletoe.     

Co-occurring species differ in tree-ring delta(18)O trends

The stable oxygen isotope ratio (delta(18)O) of tree-ring cellulose is jointly determined by the delta(18)O of xylem water, the delta(18)O of atmospheric water vapor, the humidity of the atmosphere and perhaps by species-specific differences in leaf structure and function. Atmospheric humidity and the delta(18)O of water vapor vary seasonally and annually, but if the canopy atmosphere is well mixed, atmospheric characteristics should be uniform among co-occurring trees. In contrast, xylem water delta(18)O is determined by the delta(18)O of water being drawn from the soil, which varies with depth. If co-occurring trees draw water from different soil depths, this soil-water delta(18)O signal would be manifest as differences in delta(18)O among the trees. We examined the variation in tree ring delta(18)O, over eight decades during the 20th Century, among three species co-occurring in natural forest stands of the northern Rocky Mountains in the USA. We sampled 10 Douglas-firs (Pseudotsuga menziesii (Mirb.) Franco var. glauca), 10 ponderosa pines (Pinus ponderosa Laws.) and seven western white pines (Pinus monticola Dougl.). As expected, variation in atmospheric conditions was recorded in the delta(18)O of the cellulose produced in a given year, but observed climatic correlations with delta(18)O were weak. Significant correlations with June climate data included: daily maximum temperature (r = 0.29), daily minimum temperature (r = -0.25), mean temperature (r = 0.20), mean daily precipitation (r = -0.54), vapor pressure deficit (r = 0.32) and solar radiation (r = 0.44). Lagged effects were observed in Douglas-fir and western white pine. In these species, the delta(18)O of a given annual ring was correlated with the delta(18)O of the previous ring. Ponderosa pine showed no significant autocorrelation. Although the species means were correlated among years (r = 0.67 to 0.76), ponderosa pine was consistently enriched in delta(18)O relative to the other species; differences were close to 2 per thousand and they are steadily increasing. Relative to the mean for the three species, ponderosa pine is becoming steadily more enriched (-1.0 per thousand). In contrast, Douglas-fir is being steadily depleted and western pine is intermediate, with an enrichment of 0.5 per thousand. Because all trees were exposed to the same atmospheric conditions, the differences in delta(18)O observed between species are likely due either to differences in the depth of water extraction or leaf function. If the former, presumably ponderosa pine has steadily taken up more water from near the soil surface and Douglas-fir has shifted uptake to a greater depth. If the latter, we suggest the pronounced changes in leaf-water delta(18)O are a result of changes in leaf structure and function with tree size and age.     

Reliance on stored water increases with tree size in three species in the Pacific Northwest

In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.     

Regulation of ponderosa pine foliar physiology and insect resistance mechanisms by basal area treatments

We compared foliar physiology and several measures of tree resistance to insect attack among ponderosa pine (Pinus ponderosa var. scopulorum Engelm.) trees growing in thinned stands. Measurements were made in a second-growth ponderosa pine forest in northern Arizona where the basal area treatments (6.9, 18.4, 27.6, 78.2 m(2) ha(-1)) have been experimentally maintained by frequent thinnings for 32 years before our measurements began in 1994. Most of the physiological characteristics measured were affected by the basal area treatments. As stand basal area increased from 6.9 to 78.2 m(2) ha(-1), predawn water potential, midday water potential, net photosynthetic rate, resin production, phloem thickness, and foliar toughness decreased. Foliar nitrogen concentration was greatest in trees in the intermediate basal area treatments. Our results show that the physiological condition of second-growth ponderosa pine can be manipulated by silvicultural control of stand basal area, and support the hypothesis that high stand basal area increases tree stress and decreases tree resistance to insect attack.     

Tree mortality in drought-stressed mixed-conifer and ponderosa pine forests, Arizona, USA

We monitored tree mortality in northern Arizona (USA) mixed-conifer and ponderosa pine (Pinus ponderosa Dougl. ex Laws) forests from 1997 to 2007, a period of severe drought in this area. Mortality was pervasive, occurring on 100 and 98% of 53 mixed-conifer and 60 ponderosa pine plots (1-ha each), respectively. Most mortality was attributable to a suite of forest insects, mediated by drought stress. The number of trees dying from 2002 to 2007 was more than 200% greater than the number dying from 1997 to 2002 in mixed-conifer forest and 74% greater in ponderosa pine forest. Extent of mortality was spatially variable in both forest types. Median cumulative mortality (the ratio of dead to live trees) increased by approximately 53 and 65% in mixed-conifer and ponderosa pine forests, respectively, from 2002 to 2007. Median mortality rates from 2002 to 2007 were approximately 2.0% year⁻¹ in mixed-conifer forest (range = 0-28.5%) and 0.4% year⁻¹ in ponderosa pine forest (range = 0-13.6%). Mortality rates generally were not strongly related to either elevation or stand density. Mortality was nonrandom with respect to tree size classes and species. Proportions of trees dying were greatest in the largest size classes, particularly in mixed-conifer forest, where mortality in the largest size class exceeded 22% from 2002 to 2007. Mortality in mixed-conifer forest was particularly pronounced for quaking aspen (85%) and white fir (28%), the least drought tolerant species present. These results provide an early glimpse of how these forest types are likely to respond to predicted climate changes in the southwestern USA. They suggest that these forests are not resilient to climate change, and that treatments to increase resilience to climate change may be appropriate. Research on causes of spatial heterogeneity in extent of mortality might suggest valuable approaches to aid in increasing resilience.     

Container seedlings outperform barefoot stock: Survival and growth after 10 years

Ponderosa pine, Jeffrey pine, and Douglas-fir seedlings were planted in container or bareroot form at three elevations in northern California. At the lowest elevation (762 m), container seedlings of ponderosa pine were significantly taller than bareroot seedlings at ages 4 and 10, and had breast-height diameters that were significantly larger than bareroot counterparts at age 10. Survival of Douglas-fir container seedlings was significantly greater than that of barefoot seedlings for all ages tested. At the mid-elevation site (1220 m), container seedlings of ponderosa pine and Douglas-fir were significantly larger in breast-height diameter than bareroot seedlings at age 10. Douglas-fir container seedlings survived significantly better at all ages than barefoot seedlings. At the highest elevation (1662 m), seedling height and diameter did not differ significantly, but survival of container seedlings was significantly higher than barefoot seedlings for both pine species at all ages tested.     

Can pine forest restoration promote a diverse and abundant understory and simultaneously resist nonnative invasion?

An important goal of forest restoration is to increase native plant diversity and abundance. Thinning and burning treatments are a common method of reducing fire risk while simultaneously promoting understory production in ponderosa pine (Pinus ponderosa) forests. In this study we examine the magnitude and direction of understory plant community recovery after thinning and burning restoration treatments in a ponderosa pine forest. Our objective was to determine if the post-treatment community was a diverse, abundant, and persistent assemblage of native species or if ecological restoration treatments resulted in nonnative species invasion. This project was initiated at the Grand Canyon-Parashant National Monument, Arizona, USA in 1997. We established four replicated blocks that spanned a gradient of soil types. Each block contained a control and a treated unit. Treated units were thinned to emulate pre-1870 forest stand conditions and prescribed-burned to reintroduce fire to a system that has not burned since ∼1870. We measured plant cover using the point-line intercept method and recorded species richness and composition on 0.05 ha belt transects. We examined the magnitude of treatment responses using Cohen's d effect size analysis. Changes in community composition were analyzed using nonmetric multidimensional scaling (NMS). Native plant species cover and richness increased in the thinned and burned areas compared to the controls. By the last year of the study, annual species comprised nearly 60% of the understory cover in the treatment units. Cheatgrass (Bromus tectorum), a nonnative annual grass, spread into large areas of the treated units and became the dominant understory species on the study site. The ecological restoration treatments did promote a more diverse and abundant understory community in ponderosa pine forests. The disturbances generated by such treatments also promoted an invasion by an undesirable nonnative species. Our results demonstrate the need to minimize disturbances generated by restoration treatments and argue for the need to proactively facilitate the recovery of native species after treatment.     

Predicting survival of planted Douglas-fir and ponderosa pine seedlings on dry,low-elevation sites in southwestern Oregon

Four equations were developed for predicting the probability of Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. ex Laws.) survival for the first (0–1) and first to third (1–3) growing seasons after applying mulching, scalping, or artificial shading (shade cards) treatments in plantations in southwestern Oregon, U.S.A. Variables describing conifer size, levels of competing vegetation, presence of silvicultural treatments, site factors, and climate factors were collected from 13 sites ranging from 0 to 6 years after planting and examined as potential predictors of survival. Age, stem diameter, a competition index for shrubs, severity of growing season at time of treatment, average annual precipitation, aspect, and slope angle were predictors of Douglas-fir survival during 0–1 and 1–3 growing seasons after treatment; the presence of silvicultural treatments was also a predictor only during the first growing season after treatment. Age, aspect, and slope angle were predictors of ponderosa pine survival over both 0–1 and 1–3 growing seasons after treatment; height-diameter ratio, competition indices for herbs, shrubs, and hardwoods, silvicultural treatment, severity of growing season at time of treatment, and average annual precipitation were also predictors only during the first growing season after treatment; crown width was a predictor of survival only during 1–3 growing seasons after treatment. When significant in the models, predicted probability of survival increases with treatments, less severe weather conditions, diameter, crown width, age, and precipitation; probability decreases with increasing height-diameter ratio and competition indices for herbs, shrubs, and hardwoods.     

Conserving genetic diversity in ecological restoration: a case study with ponderosa pine in northern Arizona,USA

Our study used allozyme analyses to evaluate potential impacts of ecological restoration treatments on genetic diversity of ponderosa pine (Pinus ponderosa var. scopulorum) populations within the Fort Valley Experimental Forest near Flagstaff, AZ. Allele frequencies varied among pre-settlement clumps, with trees more closely related to each other within clumps. This clumpy spatial stand structure typical of reference conditions thus represents “genetic neighborhoods” and suggests restoration of clumpy versus more evenly dispersed trees in naturally regenerating stands will protect evolutionary genetic patterns. Compared to pre-settlement populations, post-settlement trees had slightly greater heterozygosity, and allelic richness and allele frequencies varied between these two age groups. These genetic differences could have resulted from different selective conditions under which the two age groups became established. Genetic diversity of populations created using different selection criteria for residual post-settlement trees did not vary but simulated removal of 75% of post-settlement trees decreased allelic richness. Maintaining more clumps created from post-settlement populations with higher tree densities across the landscape would be more effective at conserving allelic richness. Overall results of this study indicate use of genetic structure and diversity can help guide restoration treatments to help ensure adaptive potential is conserved.     

Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

Fuel mass and stand structure after post-fire logging of a severely burned ponderosa pine forest in northeastern Oregon

Stand structure and fuel mass were measured before and after a post-fire logging operation conducted 2 years after the 1996 Summit Wildfire (Malheur National Forest), in a ponderosa pine-dominated forest in northeastern Oregon. Variables were measured both pre- and post-logging in four replicate units for each of three treatments [un-logged control, commercial harvest (most dead merchantable trees removed), fuel reduction harvest (most dead merchantable trees removed plus most dead trees >10 cm diameter)]. Post-fire logging resulted in a significant decrease in mean basal area, down to 46% pre-treatment level in commercial units, and down to 25% in fuel reduction units. Logging significantly reduced tree density, especially for the smallest (<22 cm diameter) and intermediate (23–41 cm) diameter classes. Fuel reduction units also had significantly fewer snags (dead trees >30 cm diameter—4 ha⁻¹), compared to both commercial (23 ha⁻¹) units and to un-logged controls (64 ha⁻¹) in the year following timber harvest. Logging did not change ladder height or tree species composition (% ponderosa pine, Douglas-fir and grand fir). Total woody fuel mass increased significantly in fuel reduction units when compared to controls, with the greatest difference among treatments occurring in the slash fuel (<7.6 cm diameter) component (mean of 6.2 Mg/ha for fuel reduction stands versus 1.3 Mg/ha for un-logged stands). Logging activity caused no change in the mass of the forest floor (litter or duff). Model projections of the fuel bed using the fire and fuels extension of the forest vegetation simulator (FVS–FFE) indicate that the disparity in slash fuel mass between fuel reduction and un-logged units would be sustained until about 15 years post-logging, but a re-burn of moderate intensity occurring during this time would likely kill all young trees, even in un-logged units, because of the influence of other components of the fuel bed, such as grasses and shrubs. Model projections of 1000-h fuels (woody fuels >7.6 cm diameter) indicate that standing structure in all stands would collapse quickly, with the result that un-logged stands would contain two- or three-fold greater masses at 25 and 50 years post-logging, leading to much higher consumption rates of fuel in the event of a re-burn in the same place. Variation in dead tree fall and decay rates did not change the relationship among treatments in 1000-h fuel loads, but changed the time at which treatment differences were projected to disappear. Despite treatment differences in heavy fuel accumulations over time however, FVS–FFE predicts no differences among treatments in mortality of young trees due to either moderate or high intensity fire occurring in the same place at 25, 50, or 100 years post-fire logging. The lack of a re-burn effect is in part due to the reliance on flame length as the primary mechanism leading to tree death in the fire effect models used by FVS–FFE. If tree death turns out to be caused more by root burning or cambial heating, the observed variations in 1000-h fuel loadings among treatments could be significant in the event of a future re-burn.     

Bark beetle-caused mortality in a drought-affected ponderosa pine landscape in Arizona,USA

Extensive ponderosa pine (Pinus ponderosa Dougl. ex Laws.) mortality associated with a widespread severe drought and increased bark beetle (Coleoptera: Curculionidae, Scolytinae) populations occurred in Arizona from 2001 to 2004. A complex of Ips beetles including: the Arizona fivespined ips, Ips lecontei Swaine, the pine engraver beetle, Ips pini (Say), Ips calligraphus (Germar), Ips latidens (LeConte), Ips knausi Swaine and Ips integer (Eichhoff) were the primary bark beetle species associated with ponderosa pine mortality. In this study we examine stand conditions and physiographic factors associated with bark beetle-caused tree mortality in ponderosa pine forests across five National Forests in Arizona. A total of 633 fixed-radius plots were established across five National Forests in Arizona: Apache-Sitgreaves, Coconino, Kaibab, Prescott, and Tonto. Prior to the bark beetle outbreak, plots with mortality had higher tree and stocking compared with plots without pine mortality. Logistic regression modeling found that probability of ponderosa pine mortality caused by bark beetles was positively correlated with tree density and inversely related with elevation and tree diameter. Given the large geographical extent of this study resulting logistic models to estimate the likelihood of bark beetle attack should have wide applicability across similar ponderosa pine forests across the Southwest. This is particularly true of a model driven by tree density and elevation constructed by combining all forests. Tree mortality resulted in significant reductions in basal area, tree density, stand density index, and mean tree diameter for ponderosa pine and for all species combined in these forests. Most of the observed pine mortality was in the 10–35 cm diameter class, which comprise much of the increase in tree density over the past century as a result of fire suppression and grazing practices. Ecological implications of tree mortality are discussed.     

Modification of root IAA concentrations,tree growth,and survival by application of plant growth regulating substances to container-grown conifers

Tree survival after planting ispartially a function of the tree's capacity toproduce new roots. In a field trial we assessedthe potential to modify the IAA concentrationin roots, root growth responses, and plantsurvival by root application of plant growthregulators (PGRs) such as IBA, NAA, andethylene, or alginate, a moisture retentionmaterial. Container-grown Douglas-fir,Englemann spruce, and lodgepole pine werelifted before and during prescribed liftingwindows and treated with Stim-root®, Ethrel®,Hormogel®, or alginate before or aftercold-storage, then planted in a clearcut.Lifting trees outside of the prescribed liftingwindow decreased IAA concentrations in roots ofDouglas-fir, Englemann spruce, and lodgepolepine. Treating plants with different PGRs aftercold storage increased root IAA concentrationsand root growth after planting compared totreating plants prior to cold storage. Rootgrowth and above ground plant growth andsurvival were well correlated to IAAconcentrations in roots of Douglas-fir andEnglemann spruce. IAA concentrations in rootsof lodgepole pine correlated with root growth,but did not correlate with survival. A costanalysis of treatment effects on growth andsurvival showed that certain post-cold storagePGR treatments can decrease the cost necessaryto attain target stocking and increase the sizeof the trees. Our results suggest thatapplication of PGRs or other root-promotingmaterials to tree roots before planting has thepotential to be a cost-beneficial method forincreasing root growth and tree survival.     

Response of old-growth conifers to reduction in stand density in western Oregon forests

The positive growth response of healthy young trees to density reduction is well known. In contrast, large old trees are usually thought to be intrinsically limited in their ability to respond to increased growing space; therefore, density reduction is seldom used in stands of old-growth trees. We tested the null hypothesis that old-growth trees are incapable of responding with increased growth following density reduction. The diameter growth response of 271 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), ponderosa pine (Pinus ponderosa Dougl. ex Laws) and sugar pine (Pinus lambertiana Dougl.) trees ranging in age from 158 to 650 years was examined 20 to 50 years after density reduction. Density reduction involved either light thinning with removal of less vigorous trees, or shelterwood treatments in which overstory trees were not removed. Ratios of basal area growth after treatment to basal area growth before treatment, and several other measures of growth, all indicated that the old trees sometimes benefited and were not harmed by density reduction. Growth increased by 10% or more for 68% of the trees in treated stands, and nearly 30% of trees increased growth by over 50%. This growth response persisted for at least 20 years. During this 20-year period, only three trees in treated stands (1.5%) exhibited a rapid decrease in growth, whereas growth decreased in 64% of trees in untreated stands. The length of time before a growth response to density reduction occurred varied from 5 to 25 years, with the greatest growth response often occurring 20 to 25 years after treatment. These results have important implications both for the basic biology of aging in woody plants as well as for silvicultural practices in forests with old-growth trees.     

Reprint of: Lessons from long-term studies of harvest methods in southwestern ponderosa pine-Gambel oak forests on the Fort Valley Experimental Forest, Arizona, U.S.A.

The Fort Valley Experiment Station (now Fort Valley Experimental Forest) has contributed many long-term studies to forest research. This paper focuses on a “Methods of Cutting” study initiated in 1913 on the Coulter Ranch Unit of Fort Valley and how that long-term study yielded important ecological and management lessons. We quantified the historical and contemporary forest patterns at this ponderosa pine-Gambel oak site, which was harvested using three different harvesting systems in 1913 (seed tree, group selection, and light selection) and was partially excluded from livestock browsing in 1919. Using nine historically stem-mapped permanent plots for the following three stand structural scenarios: 1913 pre-harvest (modeled), post-harvest (actual), and 2003-2006 (actual) conditions, we examined the short- and long-term consequences of harvest and livestock grazing land-use and stand dynamics. We assessed changes in spatial pattern under each harvesting system and in each structural scenario, and lastly, we examined spatial and temporal tree recruitment patterns as observed in the contemporary (2003-2006) conditions. The seed tree harvests effectively converted the spatial patterns from aggregated to random and left few trees, while the group selection and light selection had varying effects, but consistently exaggerated the spatial patchiness of the stand. By 2003-2006, all plots were aggregated at all scales and were one large patch of predominately small trees. Sites that were harvested, but excluded from livestock browsing had 40% more trees in 2003-2006. Contemporary recruitment patterns were significantly aggregated under all harvesting systems, but were most strongly aggregated if the site received a group selection or light selection cut. For group and light selection, pine seedlings initially established in stump patches created by harvesting and then proceeded to fill-in the remaining area, with recruitment rarely found under the residual pine or oak trees. Long-term data sets, such as these established by the Fort Valley Experimental Forest in 1913, are essential for quantifying the impact of historical land-use practices on contemporary forest composition and structure. Ignoring land-use legacies may lead to the misinterpretation of stand dynamics and development, and therefore should be explicitly quantified and incorporated into future management and restoration activities.     

Sources of within- and between-stand variability in specific leaf area of three ecologically distinct conifer species

Specific leaf area (SLA) is an important ecophysiological variable, but its variability within and between stands has rarely been simultaneously examined and modeled across multiple species. Extensive datasets on SLA in coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco), hybrid spruce (Picea engelmannii Parry × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carr.), and ponderosa pine (Pinus ponderosa Dougl. ex P. & C. Laws.) were used to estimate variability of SLA within a canopy and its relationship to tree- and stand-level covariates, and to predict SLA at various locations in tree crowns. Also, in the case of hybrid spruce, variation in SLA due to different relative horizontal lengths from the bole was examined. In all species, SLA systematically increased from tree tip to crown base and decreased with foliage age class. Cardinal direction did not have a highly significant influence in either Douglas-fir or hybrid spruce, but SLA did significantly decrease from branch tip to bole in hybrid spruce. Tree- and stand-level (e.g. density, site index) factors had relatively little influence on SLA, but stand age did have a significant positive influence. For ponderosa pine, a significant relationship between canopy mean current-year SLA and carbon isotope discrimination was also found, suggesting the importance of water stress in this species. An equation was fitted to estimate SLA at various points in the canopy for each species and foliage age class using absolute height in the canopy, relative vertical height in the tree, and stand age.     

Spatial and temporal patterns in structure,regeneration, and mortality of an old-growth ponderosa pine forest in the Colorado Front Range

Effective management and restoration of ponderosa pine forests requires an understanding of the heterogeneity of contemporary and historical stand structures. We assessed spatial and temporal patterns of tree establishment, mortality and size structure over a 30-year period in an old-growth ponderosa pine stand in the mid-montane zone of the Colorado Front Range. We analyzed spatial patterns and spatial associations using Ripley's K(t) and K₁₂(t) and then modeled the patterns using point process models. Forest age structure was estimated by aging a sub-sample of trees in the stand. Climate appeared to play a significant role in the coarse-scale temporal pattern of regeneration events. Stand structure (distribution of patches, light availability, and seed trees) influenced the spatial and temporal pattern of more recent regeneration events. Patchy regeneration resulted in spatial independence and some segregation of size classes. Older trees in the stand (40–55 cm dbh) exhibited some regularity in their spatial distribution at short distances indicating that patterns of mortality had been historically patchy. Contemporary patterns of mortality were mostly patchy, and mountain pine beetles caused a significant amount of mortality in the 1970s and 1980s. Both establishment and mortality retained spatial patterns that were somewhat consistent with pre-settlement forests, despite changes in driving processes.