Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

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.     

Patterns of lodgepole pine regeneration following the 1988 Yellowstone fires

In 1988, fires killed extensive lodgepole pine (Pinus contorta Dougl. ex. Loud) in Yellowstone National Park. This species bears both serotinous and non-serotinous cones, with the former most common in fire-origin stands of an even-aged character. Reconnaissance of burned stands indicated that former even-aged communities regenerated effectively. Others did not. The larger and more uniformly-sized seedling under formerly even-aged communities suggests primarily a single wave of regeneration there. Seedlings appeared to initiate to some degree over multiple years under other stands, but not sufficiently to make them well stocked. Four different regeneration pathways seem to characterize the natural reforestation of lodgepole pine following the 1988 fires. These include: (1) a dense, uniformly distributed cohort that will develop as a single-storied stand; (2) lodgepole pine islands that form over long periods around isolated seedlings; (3) a moderate to low density cohort that will gradually fill with multiple age classes over a protracted period; and (4) a cohort of only widely scattered single seedlings that initially form as small nearby tree islands, and may eventually converge into a more continuous stand with multiple age classes.     

Nitrogen cycling following mountain pine beetle disturbance in lodgepole pine forests of Greater Yellowstone

Widespread bark beetle outbreaks are currently affecting multiple conifer forest types throughout western North America, yet many ecosystem-level consequences of this disturbance are poorly understood. We quantified the effect of mountain pine beetle (Dendroctonus ponderosae) outbreak on nitrogen (N) cycling through litter, soil, and vegetation in lodgepole pine (Pinus contorta var. latifolia) forests of the Greater Yellowstone Ecosystem (WY, USA) across a 0-30 year chronosequence of time-since-beetle disturbance. Recent (1-4 years) bark beetle disturbance increased total litter depth and N concentration in needle litter relative to undisturbed stands, and soils in recently disturbed stands were cooler with greater rates of net N mineralization and nitrification than undisturbed sites. Thirty years after beetle outbreak, needle litter N concentration remained elevated; however total litter N concentration, total litter mass, and soil N pools and fluxes were not different from undisturbed stands. Canopy N pool size declined 58% in recent outbreaks, and remained 48% lower than undisturbed in 30-year old outbreaks. Foliar N concentrations in unattacked lodgepole pine trees and an understory sedge were positively correlated with net N mineralization in soils across the chronosequence. Bark beetle disturbance altered N cycling through the litter, soil, and vegetation of lodgepole pine forests, but changes in soil N cycling were less severe than those observed following stand replacing fire. Several lines of evidence suggest the potential for N leaching is low following bark beetle disturbance in lodgepole pine.     

Effects of the overstory on the diversity of the herb and shrub layers of Anatolian black pine forests

Understory plants are important components of forests because they are responsible for the majority of the vascular plant diversity of forest ecosystems. The richness and composition of understory communities are closely related to the tree layer diversity, structure and composition. The aim of this study was to examine the understory diversity of Anatolian black pine (Pinus nigra Arnold subsp. pallasiana (Lamb.) Holmboe)-dominated forests on the Kazda?? Mountains of West Turkey. To describe the overstory structure and composition in a numerically and quantitatively well-defined manner, cumulative abundance profiles (CAPs) of the tree species were used. The resemblance of the sampling plots was classified into five stand types assessing the CAP through the Fuzzy C-Means clustering method. A permutational multivariate analysis of variance (PERMANOVA) was performed to test the variance of the community ecological distance between the five stand types, and the results showed significant differences in these clusters. Many shade-tolerant plants were associated with the mixed stands of Anatolian black pine–Kazda?? fir. The composition of the herb and shrub layer could not be explained by the environmental variables but by differences in the overstory structure of the stands. Pure or nearly pure Anatolian Black pine stands were more diverse than mixed oak–Anatolian black pine and Kazda?? fir–Anatolian black pine stands. However, although dense and young pure Anatolian black pine stands had the most diverse plant species in the shrub layer, they were ranked third in terms of the herb layer diversity. The Anatolian black pine–Kazda?? fir mixed stands had the lowest herb and shrub layer diversity. These results allow us to comprehend the relationship between the overstory structure and composition, and the understory diversity. Understanding this relationship is important for the conservation of understory plant diversity in the management of forest ecosystems.     

Yellow pine regeneration as a function of fire severity and post-burn stand structure in the southern Appalachian Mountains

We used pre- and post-burn fire effects data from six prescribed burns to examine post-burn threshold effects of stand structure (understory density, overstory density, shrub cover, duff depth, and total fuel load) on the regeneration of yellow pine (Pinus subgenus Diploxylon) seedlings and cover of herbaceous vegetation in six prescribed-fire management units located within western Great Smoky Mountains National Park (GSMNP) in east Tennessee, USA. We also evaluated the utility of the Keetch-Byram Drought Index (KBDI) as a predictor of post-burn stand and fuel conditions by comparing post-burn stand variables for different ranges of KBDI (23-78; more wet, and 328-368; more dry). We found that yellow pine seedlings were effectively absent in post-burn forests until overstory density was reduced over 40%, understory density was reduced over 80%, and post-burn shrub cover was 10% or less. We also observed that a reduction in total fuels of 60% and a post-burn duff layer depth of less than four cm were required for successful regeneration of yellow pine. Total herbaceous species cover exhibited near identical responses with increased cover following an 80% reduction in understory density and a post-burn duff depth of less than 4 cm. We observed strong positive relationships between high KBDI values and burn severity, changes in forest structure, reductions in fuels, and post-burn yellow pine reproduction. We observed continuous recruitment of yellow pine seedlings 5 years after fire in high KBDI burns while low KBDI burns showed little change in yellow pine density through time. An intense outbreak of the southern pine beetle (SPB; Dendroctonus frontalis) occurred within 2 years of our high KBDI burns and reduced shading resulting from overstory mortality likely enhanced the survival of yellow pine seedlings. The results of this study provide targets for the application of prescribed fire to restore yellow pine in the southern Appalachians. Continued research and monitoring will help determine how prescribed fire can best be applied in combination with other disturbance agents such as SPB to perpetuate yellow pine forests.     

Risk rating for mountain pine beetle infestation of lodgepole pine forests over large areas with ordinal regression modelling

The mountain pine beetle Dendroctonus ponderosae Hopkins is endemic to lodgepole pine, Pinus contorta var. latifolia Engelmann, forests in western Canada. However, the current beetle epidemic in this area highlights the challenges faced by forest managers tasked with prioritizing stands for mitigation activities such as salvage harvesting and direct control methods. In western Canada, the operational risk rating system for mountain pine beetle is based on biological knowledge gained from a rich legacy of stand-scale field studies. Due to the large spatial (millions of hectares affected) and temporal (over 10 years) extents of the current epidemic, new research into large-area mountain pine beetle processes has revealed further insights into the landscape-scale characteristics of beetle infested forests. In this paper, we evaluated the potential for this new knowledge to augment an established system for rating the short-term risk of tree mortality in a stand due to mountain pine beetle. New variables explored for utility in risk rating include direct shortwave radiation, site index, diameter at breast height, the temporal trends in local beetle populations, Biogeoclimatic Ecosystem Classification and beetle–host interaction variables. Proportional odds ordinal regression was used to develop a model for the Vanderhoof Forest District in west-central British Columbia. Prediction on independent data was assessed with the area under the receiver operator curve (AUC), indicating good discriminatory power (AUC = 0.84) for predicting levels of mountain pine beetle-caused pine mortality.     

The ecology of tree regeneration in mature and old forests: combined knowledge for sustainable forest management

We focused our attention on quantifying the factor complex of forest regeneration in 423 mature and old stands with contrasting environmental conditions. We recorded the microhabitat selection of tree recruits, the frequency of tree seedlings, and evaluated the drivers of sapling abundance and diversity. The majority of forest regeneration was established on undisturbed forest floor. Dead wood was a frequent substrate in spruce-(co)dominated forests. Seedling frequency within a stand was related to the site-type specific productivity gradient of stands—pine seedlings were common in low-productivity and spruce in high-productivity boreal forests. Seedlings of temperate broad-leaved trees dominated in productive boreonemoral forests, except for oak, which showed a uniform distribution of abundance in all forest site-types. Sapling abundance was dictated by forest site-type, and facilitated by stand diversity, variability in stand closure, lying dead wood, abundant moss, and a thick organic layer. Only in boreal forests was sapling abundance suppressed by the abundant spruce and younger trees. Upon considering the relationship between sapling abundance and species richness, sapling diversity was dependent on forest site-type, suppressed by stand density and dead wood (old gap) abundance, and facilitated by stand diversity. In addition, boreonemoral stands, competition from herbs, and facilitation by mosses occurred. The observed pattern of tree recruitment points to the importance of top-down effects of the overstory, competing or facilitating interactions with forest floor vegetation, and availability of regeneration microhabitats, which in complex make their ecology comparable with forest herbs. Natural forest regeneration can be enhanced if silvicultural methods support mixed stands and enhance field layer diversity. Oak can provide the universal tree species to improve stand structure over a wide range of habitats.     

The influence of mountain pine beetle outbreaks and drought on severe wildfires in northwestern Colorado and southern Wyoming: A look at the past century

Outbreaks of bark beetles and drought both lead to concerns about increased fire risk, but the relative importance of these two factors is the subject of much debate. We examined how mountain pine beetle (MPB) outbreaks and drought have contributed to the fire regime of lodgepole pine forests in northwestern Colorado and adjacent areas of southern Wyoming over the past century. We used dendroecological methods to reconstruct the pre-fire history of MPB outbreaks in twenty lodgepole pine stands that had burned between 1939 and 2006 and in 20 nearby lodgepole pine stands that were otherwise similar but that had not burned. Our data represent c. 80% of all large fires that had occurred in lodgepole pine forests in this study area over the past century. We also compared Palmer Drought Severity Index (PDSI) and actual evapotranspiration (AET) values between fire years and non-fire years. Burned stands were no more likely to have been affected by outbreak prior to fires than were nearby unburned stands. However, PDSI and AET values were both lower during fire years than during non-fire years. This work indicates that climate has been more important than outbreaks to the fire regime of lodgepole pine forests in this region over the past century. Indeed, we found no detectable increase in the occurrence of high-severity fires following MPB outbreaks. Dry conditions, rather than changes in fuels associated with outbreaks, appear to be most limiting to the occurrence of severe fires in these forests.     

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.     

Growth responses of three coexisting conifer species to climate across wide geographic and climate ranges in Yukon and British Columbia

This work aimed to compare radial growth–climate relationships among three coexisting coniferous tree species across a wide geographic and climate range from southern British Columbia (BC) to central Yukon, Canada. Tree-ring data were collected from 20 mature stands of white spruce (Picea glauca), lodgepole pine (Pinus contorta var. latifolia), and subalpine fir (Abies lasiocarpa). Linear relationships between annual growth variation and monthly and seasonal climate were quantified with correlation and regression analyses, and variation in climate–growth responses over a climatic gradient were quantified by regressing growth responses against local mean climatic conditions. Temperatures had more consistent and stronger correlations with growth for all three species than precipitation, but growth–climate responses varied among species and among sites. In particular, pine and fir populations showed different responses between BC and Yukon, whereas spruce showed a more consistent response across the study domain. Results indicate that (1) the response and sensitivity of trees to seasonal climate variables vary among species and sites and (2) winter temperatures prior to growth may have significant impacts on pine and fir growth at some sites. The capacity to adapt to climate change will likely vary among the study species and across climatic gradients, which will have implications for the future management of mixed-species forests in Yukon and BC.     

Impact of slash removal,drag scarification,and mounding on lodgepole pine cone distribution and seedling regeneration after cut-to-length harvesting on high elevation sites

Excessive slash loading could pose a problem for the regeneration of the serotinous lodgpole pine especially in forests at higher elevation where soil temperature is limiting. In the past, these forests have commonly been harvested using full-tree harvesting where trees are processed at roadside; however, recently cut-to-length harvesting has become a more frequent harvesting method. In cut-to-length harvesting the harvested trees are processed in the block, as a result slash accumulation is much higher on these cutblocks. In an experimental field trial, the cone distribution, natural lodgepole pine regeneration, and the growth and establishment of planted lodgepole pine were evaluated in response to slash load, drag scarification, and mounding after cut-to-length harvesting of high elevation lodgepole pine stands in the Rocky Mountains. Twelve sites were established, each contained six plots which were randomly assigned to six treatment combinations of two slash removal (slash and slash removed) and three mechanical soil preparation treatments (no soil preparation, drag scarifying, and mounding). The slash removal reduced slash volume by more than 50% but also reduced the number of lodgepole pine cones available for regeneration by over 33%. However, soil mechanical treatments offset this effect as fewer cones were necessary to achieve high natural pine regeneration densities. Drag scarification of plots resulted in 12 times the number of pine seedlings compared to the non-prepared plots. Although slash removal did not have an effect on the number of naturally regenerated lodgepole pine seedlings, it had a positive effect on their growth performance. Conversely, planted pine seedlings had lower mortality and better growth in soils that had been mechanically prepared and had the slash removed; however, the growth effects became only apparent 4 years after planting. While slash removal and mechanical soil preparation did increase soil temperatures; the slash removal treatment had a more transient effect on soil temperatures than soil preparation. Differences in soil temperature decreased over time which appeared to be mostly driven by a warming of the soils in the plots with no soil preparation, likely a result of the decomposition of the finer slash and feathermosses. Overall, it appears that surface disturbance on these high elevation sites had a far greater effect on lodgepole pine regeneration and growth than the increased accumulation of slash as a result of cut-to-length harvesting.     

Temporal dynamics of forage succession for elk at two scales: Implications of forest management

Rapid conversion of lodgepole pine (Pinus contorta) dominated landscapes in western Alberta to a mosaic of successional stand ages has long-term implications for elk populations in the region that have not been fully assessed. We developed stand-level models of forage succession using data from 159 cutblocks to simulate forage and cover availability within the home range of an elk under “even-flow” and “pulsed” timber cutting scenarios that are common in this area. We found forage biomass peaked approximately 9 years following felling for both herbaceous (graminoids and forbs) and palatable browse forage. Forage production was determined primarily by the age of the cutblock (time since felling) and to a lesser extent elevation, compound topographic index (a measure of site wetness), and the distance to the nearest deciduous forest patch. Forbs became increasingly abundant as stands matured. Browse composition shifted from palatable species to unpalatable species after approximately 30 years. Within an elk home range (100 km²), we found that simulating an “even-flow” cutting regime resulted in forage availability that was always higher than under the current conditions of 90% forests when more than 10 ha were harvested per year. Further, forest cover (>30$>30$-year-old stands) was never less than the guideline of 60% of the area needed to provide adequate thermal and hiding cover. In contrast, when a “pulsed” harvest scenario was simulated, similar to what occurs for mountain pine beetle control, forage availability changed dramatically over time as a large cohort of harvested timber aged. Without further harvesting the stand conditions in the home range eventually returned to a state of pre-harvest risk for mountain pine beetle outbreak. We demonstrated several additional harvesting scenarios that avoided the potential risk and produced a much higher level of forage than currently exists.     

Stand development patterns as a consequence of the mortality in Austrocedrus chilensis forests

The forests of Austrocedrus chilensis (D. Don) Pic. Sern. et Bizarri in Argentina suffer decline and mortality throughout their natural distribution known as ‘mal del ciprés’. While several aspects of this, to date, temporally unpredictable process of overstory tree mortality have been the focus of detailed studies, there has been little research on tree growth and stand dynamics in symptomatic forests nor stand development patterns and prediction of future stand structure. We studied 12 stands in northern Patagonia (Province of Río Negro) using stand reconstruction studies to examine the changes in stand structure over time as a consequence of overstory mortality and the implications of these structural changes on the establishment and growth of the residual overstory. Dendrochronological analyses were used to reconstruct stand establishment and structure over time, and to study past diameter growth patterns. Mortality in A. chilensis forests was variable in time among stands. As expected, overstory mortality led to the successful establishment of trees in the understory in all stands; however, the response of residual overstory trees was variable. Understory establishment was low in some stands and high in others depending on the density of the overstory. While overstory trees in almost all stands released after the onset of the mortality, the pattern was not distinctive, varying in time, number and magnitude. In some stands, growth releases occurred after single or multiple tree deaths suggesting a relationship between processes, while in others this was not the case. Even when the patterns of recruitment in the understory and the overstory response varied greatly among stands, when examined together, some general patterns emerged. This study is the first to intensively explore the dynamics of A. chilensis forests affected by ‘mal del ciprés’. Additionally, this study showed that arbitrarily categorizing disturbances as discrete or chronic masks the true process of release of growing space and the resulting stand dynamics.     

Competition and tree crowns: A neighborhood analysis of three boreal tree species

Competition for canopy space is a fundamental structuring feature of forest ecosystems and remains an enduring focus of research attention. We used a spatial neighborhood approach to quantify the influence of local competition on the size of individual tree crowns in north-central British Columbia, where forests are dominated by subalpine fir (Abies lasiocarpa), lodgepole pine (Pinus contorta) and interior spruce (Picea glauca × engelmanii). Using maximum likelihood methods, we quantified crown radius and length as functions of tree size and competition, estimated by the species identity and spatial arrangement of neighboring trees. Tree crown size depended on tree bole size in all species. Given low levels of competition, pine displayed the widest, shortest tree crowns compared to the relatively long and narrow crowns found in spruce and fir. Sensitivity to crowding by neighbors declined with increasing tree height in all but the pine crown radius model. Five of the six selected best models included separate competition coefficients for each neighboring tree species, evidence that species generally differ in their competitive effects on neighboring tree crowns. The selected crown radius model for lodgepole pine, a shade-intolerant species, treated all neighbors as equivalent competitors. In all species, competition from neighbors exerted an important influence on crown size. Per-capita effects of competition across different sizes and species of neighbors and target trees varied, but subalpine fir generally displayed the strongest competitive effects on neighbors. Results from this study provide evidence that species differ both in their response to competition and in their competitive influence on neighbors, factors that may contribute to maintaining coexistence.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

Stand and landscape level effects of a major outbreak of spruce beetles on forest vegetation in the Copper River Basin,Alaska

From 1989 to 2003, a widespread outbreak of spruce beetles (Dendroctonus rufipennis) in the Copper River Basin, Alaska, infested over 275,000 ha of forests in the region. During 1997 and 1998, we measured forest vegetation structure and composition on one hundred and thirty-six 20-m × 20-m plots to assess both the immediate stand and landscape level effects of the spruce beetle infestation. A photo-interpreted vegetation and infestation map was produced using color-infrared aerial photography at a scale of 1:40,000. We used linear regression to quantify the effects of the outbreak on forest structure and composition. White spruce (Picea glauca) canopy cover and basal area of medium-to-large trees [≥15 cm diameter-at-breast height (1.3 m, dbh)] were reduced linearly as the number of trees attacked by spruce beetles increased. Black spruce (Picea mariana) and small diameter white spruce (<15 cm dbh) were infrequently attacked and killed by spruce beetles. This selective attack of mature white spruce reduced structural complexity of stands to earlier stages of succession and caused mixed tree species stands to lose their white spruce and become more homogeneous in overstory composition. Using the resulting regressions, we developed a transition matrix to describe changes in vegetation types under varying levels of spruce beetle infestations, and applied the model to the vegetation map. Prior to the outbreak, our study area was composed primarily of stands of mixed white and black spruce (29% of area) and pure white spruce (25%). However, the selective attack on white spruce caused many of these stands to transition to black spruce dominated stands (73% increase in area) or shrublands (26% increase in area). The post-infestation landscape was thereby composed of more even distributions of shrubland and white, black, and mixed spruce communities (17–22% of study area). Changes in the cover and composition of understory vegetation were less evident in this study. However, stands with the highest mortality due to spruce beetles had the lowest densities of white spruce seedlings suggesting a longer forest regeneration time without an increase in seedling germination, growth, or survival.