Changes in forest structure of a mixed conifer forest, southwestern Colorado, USA

We selected a warm/dry mixed conifer forest (ponderosa pine, white fir, Douglas-fir, and aspen) in southwestern Colorado to reconstruct historical conditions of fire regime and forest structure in preparation for an experiment in ecological restoration. Although mixed conifer forests are of high ecological and social value in the Southwest, they have been less studied than ponderosa pine forests. Fire-scar analysis on a 150-ha area showed recurring fires at mean intervals of 24 years (all fires with minimum of 2 sample trees scarred) to 32 years (fire scarring 25% or more of sample trees) from the 16th century until the abrupt cessation of fire after 1868, concurrent with European settlement. There was no evidence in age or species-specific data of severe burning at the scale of the study blocks (approximately 200 ha). The forest remained unharvested throughout most of the 20th century, until a cut in the early 1990s removed approximately equal basal areas of ponderosa pine and white fir. Forest structure had already changed substantially, however. Total basal area increased from an average of 11 m² ha⁻¹ in 1870 to 27 m² ha⁻¹ in 2003, despite harvesting of at least 8.4 m² ha⁻¹. Ponderosa pine declined from representing nearly two-thirds of basal area in 1870 to one-third in 2003. The other species increased dramatically, especially white fir, which went from 12% to 35% of basal area and dominated stand density with an average of 392 trees ha⁻¹. Total tree density increased from 142 trees ha⁻¹ in 1870 to 677 trees ha⁻¹ in 2003. The ecological changes that occurred here since the 19th century have been in exactly the opposite direction considering the warm, fire-favoring climate expected in the 21st century. If warm/dry mixed conifer forests of southern Colorado are to have a reasonable chance for persistence under the future climate regime, restoring conditions more similar to the frequently burned, open forests of the past is likely to be a useful starting point.     

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.     

Thinning reduces soil carbon dioxide but not methane flux from southwestern USA ponderosa pine forests

Forest soils are important components of the global carbon cycle because they both store and release carbon. Carbon dioxide is released from soil to the atmosphere as a result of plant root and microbial respiration. Additionally, soils in dry forests are often sinks of methane from the atmosphere. Both carbon dioxide and methane are greenhouse gases whose increasing concentration in the atmosphere contributes to climate warming. Thinning treatments are being implemented in ponderosa pine forests across the southwestern United States to restore historic forest structure and reduce the risk of severe wildfire. This study addresses how thinning alters fluxes of carbon dioxide and methane in ponderosa pine forest soils within one year of management and examines mechanisms of change. Carbon dioxide and methane fluxes, soil temperature, soil water content, forest floor mass, root mass, understory plant biomass, and soil microbial biomass carbon were measured before and after the implementation of a thinning and in an unthinned forest. Carbon dioxide efflux from soil decreased as a result of thinning in two of three summer months. Average summer carbon dioxide efflux declined by an average of 34 mg C m⁻² hr⁻¹ in the first year after thinning. Methane oxidation did not change in response to thinning. Thinning had no significant short-term effect on total forest floor mass, total root biomass, or microbial biomass carbon in the mineral soil. Understory plant biomass increased after thinning. Thinning increased carbon available for decomposition by killing tree roots, but our results suggest that thinning reduced carbon dioxide emissions from the soil because the reduction in belowground autotrophic respiration was larger than the stimulation of heterotrophic respiration. Methane oxidation was probably not affected by thinning because thinning did not alter the forest floor mass enough to affect methane diffusion from the atmosphere into the soil.     

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.     

Predicting mortality for five California conifers following wildfire

Fire injury was characterized and survival monitored for 5677 trees >25 cm DBH from five wildfires in California that occurred between 2000 and 2004. Logistic regression models for predicting the probability of mortality 5-years after fire were developed for incense cedar (Calocedrus decurrens (Torr.) Florin), white fir (Abies concolor (Gord. & Glend.) Lindl. ex Hildebr.), sugar pine (Pinus lambertiana Douglas), Jeffrey pine (P. jeffreyi Balf.), and ponderosa pine (P. ponderosa C. Lawson). Differences in crown injury variables were also compared for Jeffrey and ponderosa pine. Most mortality (70–88% depending on species) occurred within 2 years post-wildfire and had stabilized by year 3. Crown length and crown volume injury variables predicted tree mortality equally well; however, the variables were not interchangeable. Crown injury and cambium kill rating was significant in predicting mortality in all models. DBH was only a significant predictor of mortality for white fir and the combined ponderosa and Jeffrey pine models developed from the McNally Fire; these models all predicted increasing mortality with increasing tree size. Red turpentine beetle (Dendroctonus valens) was a significant predictor variable for sugar pine, ponderosa pine, and Jeffrey pine; ambrosia beetle (Trypodendron and Gnathotrichus spp.) was a significant predictor variable for white fir. The mortality models and post-fire tree survival characteristics provide improved prediction of 5-year post-wildfire tree mortality for several California conifers. The models confirm the overall importance of crown injury in predicting post-fire mortality compared to other injury variables for all species. Additional variables such as cambium kill, bark beetles, and tree size improved model accuracies, but likely not enough to justify the added expense of data collection.     

Bark Beetle Demography in a Jeffrey Pine Stand as Influenced by Mechanized Thinning and Prescribed Fire

Forest thinnings implemented with cut-to-length and whole-tree harvesting systems followed by underburning were evaluated for their effects on bark beetle prevalence in pure, uneven-aged Jeffrey pine (Pinus jeffreyi Grev. & Balf.) interspersed with isolated California white fir (Abies concolor var. lowiana [Gord.] Lemm.). Based on pitch tube counts in a stand with a moderate bark beetle population in its pine component, the Jeffrey pine beetle (Dendroctonus jeffreyi Hopkins) generally preferred larger trees before treatment implementation, but after exhibiting mixed pretreatment tendencies concerning stand density demonstrated a posttreatment proclivity toward higher density. Cut-to-length thinning followed by underburning increased the pine beetle population while whole-tree thinning unaccompanied by burning reduced it. Tree mortality was induced by the bark beetle infestation but was not its sole cause. Pitch tube abundance on white fir far exceeded that on Jeffrey pine, and the greatest influence on the fir engraver (Scolytus ventralis LeConte) population was the prevalence of its host tree. The responses presented herein to these thinning and burning practices, which are being increasingly utilized in forest restoration efforts in the western USA, provide natural resource managers insight into potential forest health outcomes when implemented in Jeffrey pine and similar dry site forest types.     

Development of Leptographium wageneri on root surfaces and other substrata

Conidia of Leptographium wageneri var. ponderosum, Leptographium wageneri var. pseudotsugae and Leptographium wageneri var. wageneri germinated on wounded roots of ponderosa pine (Pinus ponderosa), Douglas fir (Pseudotsuga menziesii), and pinyon (Pinus edulis), but not on unwounded roots. On wounds, germination occurred regardless of the combination of pathogen variety and plant species. When wounded roots of ponderosa pine and Douglas fir seedlings were inoculated with conidia of L. w. var. ponderosum and L. w. var. pseudotsugae, respectively, 87-100% of the seedlings became infected. When unwounded roots were inoculated with wooden blocks colonized with L. wageneri, 60-66% of the seedlings became infected. The development of L. w. var. ponderosum and L. w. var. pseudotsugae on agar was compared with their development on agar overlaid with either cellophane or Nucleopore membranes. Undulating hyphae were found commonly in and on the agar, but less frequently on the membranes. On both membranes, mycelia formed mats which were tightly appressed to the surface; mats were not formed on the agar. On both agar and the membranes, two or more hyphae often formed a strand. When L. w. var. ponderosum mycelia were allowed to colonize the surface of an unwounded pine root, strands, undulating hyphae and poorly formed mats were seen.     

Effects of uneven-aged management on diameter distribution and species composition of northern hardwoods in Upper Michigan

Uneven-aged management using single-tree or group selection has been in practice for many decades, especially in northern hardwood forests. Use of stocking regulation tools is thought to produce and maintain specific stand structures that upon regulation, are sustainable over time. We evaluated stand structures in northern hardwoods in Upper Michigan across three ownerships that practice different approaches toward uneven-aged management. Industry land (MeadWestvaco—MWV) uses primarily diameter limit cutting combined with a sawyer-select harvest method, retaining a maximum diameter of 45.7 cm (18 in.) with a residual basal area of 16.1 m²/ha (70 ft²/ac) and a cutting cycle of 10 years. The Michigan Department of Natural Resources (DNR) uses a crop tree release technique with a residual basal area of 18.4 m²/ha (80 ft²/ac) and a cutting cycle of 20 years. Both ownerships view regeneration of new cohorts as inevitable given the intensity of disturbance and the forest type. The third ownership (the Ford Forestry Center School Forest at Michigan Technological University—FFC) employs strict stocking regulation using the BDq method (residual basal area of 16.1 m²/ha (70 ft²/ac), maximum diameter of 50.8–55.9 cm (20–22 in.), and a q-ratio of 1.3) with a cutting cycle ranging from 12 to 15 years.Stand structure on a total of 25 stands was characterized for these ownerships to assess the impact of management strategy on stand structure and species composition. Differences in species composition and lower diversity indices were found where increasing sugar maple dominance was an objective (FFC ownership). All ownerships showed reduced relative importance values of mid-tolerant species such as yellow birch in their stands as compared to values reported for old-growth or unmanaged stands. Diameter distributions were classified into one of three shape categories (negative exponential, increasing q-ratio, and rotated sigmoid) using the regression of DBH, DBH², and DBH³ on the log₁₀ of trees per hectare. The best model in terms of adjusted-R² and root mean square error (RMSE) was selected for each stand. All management strategies resulted in similar occurrences of distribution shapes, despite some evidence of a trend toward a rotated sigmoid distribution. These trends suggest that several different diameter distribution shapes in uneven-aged northern hardwoods in the Lake States are possible following a variety of management approaches, with sugar maple increasing in dominance with strict adherence to certain stocking regulation guidelines.     

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.     

The farm forest resource in Ireland: opportunities and challenges for rural development in Ireland

A critical mass of farm forestry and private forestry is now developing in Ireland with over 211,000 ha planted since 1980, with 190,000 ha of this planted since 1990. Currently, 90,000 ha of private forests are over 10 years of age, with 41,000 ha over 15 years of age. This paper examines the potential of the farm forest resource in Ireland and highlights some issues that need to be addressed in order that the full potential of farm forestry is realised. It is estimated that 876,000 m³ of timber is currently available from forests through removals from thinning operations from plantations that are at or have passed first thinning age. Cumulative volume output from thinning could be as high as 1.9 M m³ by 2010 increasing to 5.9 M m³ by 2015. Farm forestry has the potential to be a financially important farm enterprise with 15% of farmers (16,460) currently engaged in forestry in Ireland. The development of volume production forecasts will provide crucial supply information for gearing up of the wood-processing sector for additional volume from farms. However many enterprises face high harvesting and extraction costs. Solutions to overcoming economies of scale due to the small size of plantations (averaging 9.2 ha) are proposed using methods to group forestry operations together. The network of small sawmills throughout the country is favourable for private growers as analysis indicates that 82% of private forests are within 32 km of a sawmill and efforts to stimulate local processing are being encouraged. Financial analysis indicates that farmers can increase forestry returns by thinning their plantations. The risk of windthrow to plantations is one of the major factors that can lead to financial losses in farm forestry in Ireland and can be reduced by adequate management planning. A series of forestry extension programs will provide farmers with adequate information to help aid decision-making in managing their plantations.

Influences of Mechanized Thinning and Prescribed Fire on Natural Regeneration in an Uneven-Aged Jeffrey Pine Stand

Thinnings using cut-to-length or whole-tree harvesting systems followed by underburning were evaluated for their effects on seedling and sapling demography in a pure, uneven-aged Jeffrey pine (Pinus jeffreyi Grev. & Balf.) stand containing a minor component of California white fir (Abies concolor var. lowiana [Gord.] Lemm.). Depression of seedling counts due to forest floor disturbance associated with thinning was followed by a recovery largely confined to Jeffrey pine in the whole-tree treatment where final seedling counts exceeded those found initially. The postburn substrate was more favorable for establishment of Jeffrey pine than white fir seedlings, and the largest increase in seedling counts between the initial and final inventories occurred in the burned portion of the whole-tree treatment. Live sapling losses from thinning were greatest in the cut-to-length treatment, while underburning induced complete sapling mortality. Absent treatment, several stand and site variables influenced seedling and sapling abundance, prominent among them a propensity for mahala mat (Ceanothus prostratus Benth.) to elevate counts of white fir within both size classes. These results provide land managers insight into the impacts of six combinations of thinning and burning treatment on natural regeneration in eastern Sierra Nevada Jeffrey pine and similar dry site forest types.     

Influence of herbicides and felling,fertilization, and prescribed fire on longleaf pine growth and understory vegetation through ten growing seasons and the outcome of an ensuing wildfire

Restoring longleaf pine (Pinus palustris Mill.) over much of its original range requires artificial regeneration. In central Louisiana, USA, two fertilization levels—No (NF) or Yes (F-36 kg/ha N and 40 kg/ha P) in combination with three vegetation treatments—Check, four prescribed fires (PF), or multi-year vegetation control by herbicidal and mechanical means (IVM) were applied to container-grown longleaf pine plantings in a grass savanna. After 10 years, P concentration in longleaf pine foliage was less on NF plots than F plots, but fertilization did not significantly affect tree stature. Survival was greater on NF plots than F plots, and so, NF plots were more productive (NF—63 m³/ha and F—45 m³/ha). Pine trees on IVM plots (37 dm³/tree) were significantly larger than on Check and PF plots, which averaged 17 dm³/tree. Survival was better on IVM plots (88%) than PF plots (78%), which was better than Checks (58%). Consequently, IVM plots were most productive (99 m³/ha), followed by PF plots (44 m³/ha), and lastly Checks (28 m³/ha). PF plots had greater grass cover (38%) than Check and IVM plots, which averaged 5%, whereas PF and IVM plots had less understory arborescent cover (an average of 25%) than Checks (91%). A wildfire in March 2007 reduced pine survival by 22, 14, and 1 percentage points on IVM, Check, and PF plots, respectively. Seven months later, longleaf pine production had decreased to 92 m³/ha on IVM plots while increasing to 55 m³/ha on PF plots and 30 m³/ha on Checks. Overall, the wildfire rejuvenated the herbaceous plant community. Grass cover on Check and IVM plots averaged 20% while grass cover on PF plots was 36%. Forb cover increased on all treatments from 2% before the wildfire to 13% seven months after the wildfire. Understory arborescent cover decreased on Checks to 62% but increased slightly on PF and IVM plots and averaged 30%.     

How well does Kirtland's warbler management emulate the effects of natural disturbance on stand structure in Michigan jack pine forests?

Jack pine (Pinus banksiana) forests in parts of northern Lower Michigan have been managed with 30 years of extensive clearcut harvesting followed by planted stand establishment in order to maintain habitat for the endangered Kirtland's warbler (Dendroica kirtlandii). We used two, parallel chronosequences to study how this management has affected the structural development of jack pine stands relative to the historically dominant disturbance regime of stand-replacing wildfire. Each chronosequence consisted of three young stands (age range 3–6 years), three intermediate-aged stands (age range of 12–17 years) and three mature stands (age range of 39–69 years). Average stem density in young plantations (2300 stems/ha) was lower than the average for young, fire-origin stands (11,000) and varied over a much narrower range among stands (1403–2667 for plantations and 1552–24,192 in fire stands). In addition, within-stand patchiness of stem density was also much higher in the wildfire sites for young and intermediate ages. Plantation sites possessed very little dead wood at young ages (averaging 3 snags/ha and 12 m³/ha CWD) compared to young fire-origin stands (averaging 252 snags/ha and 49 m³/ha CWD). In contrast, mature plantations had similar levels of dead wood (averaging 269 snags/ha and 22 m³/ha CWD) as mature fire-origin stands (averaging 557 snags/ha and 12 m³/ha CWD). Differences between the plantation and fire-origin chronosequences were driven mainly by young- and intermediate-aged stands, whereas mature stands were typically quite similar in all structural features. Our results show clearly that forest management aimed at preserving and enhancing the population of a single endangered species results in greatly simplified habitat structure at the stand level, and suggest that this simplification is perpetuated across the landscape as well. Of particular concern are the effects of extensive harvesting and planting on the availability of snags and CWD.     

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.     

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.     

Prescribed fire effects on bark beetle activity and tree mortality in southwestern ponderosa pine forests

Prescribed fire is an important tool in the management of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests, yet effects on bark beetle (Coleoptera: Curculionidae, Scolytinae) activity and tree mortality are poorly understood in the southwestern U.S. We compared bark beetle attacks and tree mortality between paired prescribed-burned and unburned stands at each of four sites in Arizona and New Mexico for three growing seasons after burning (2004–2006). Prescribed burns increased bark beetle attacks on ponderosa pine over the first three post-fire years from 1.5 to 13% of all trees, increased successful, lethal attacks on ponderosa pine from 0.4 to 7.6%, increased mortality of ponderosa pine from all causes from 0.6 to 8.4%, and increased mortality of all tree species with diameter at breast height >13 cm from 0.6 to 9.6%. On a per year basis, prescribed burns increased ponderosa pine mortality from 0.2% per year in unburned stands to 2.8% per year in burned stands. Mortality of ponderosa pine 3 years after burning was best described by a logistic regression model with total crown damage (crown scorch + crown consumption) and bark beetle attack rating (no, partial, or mass attack by bark beetles) as independent variables. Attacks by Dendroctonus spp. did not differ significantly over bole heights, whereas attacks by Ips spp. were greater on the upper bole compared with the lower bole. Three previously published logistic regression models of tree mortality, developed from fires in 1995–1996 in northern Arizona, were moderately successful in predicting broad patterns of tree mortality in our data. The influence of bark beetle attack rating on tree mortality was stronger for our data than for data from the 1995–1996 fires. Our results highlight canopy damage from fire as a strong and consistent predictor of post-fire mortality of ponderosa pine, and bark beetle attacks and bole char rating as less consistent predictors because of temporal variability in their relationship to mortality. The small increase in tree mortality and bark beetle attacks caused by prescribed burning should be acceptable to many forest managers and the public given the resulting reduction in surface fuel and risk of severe wildfire.     

Modelling coarse woody debris dynamics in even-aged Scots pine forests

Coarse woody debris (CWD) has become an important component in the study of forest ecosystems, being a key factor in the nutrient cycle as well as a habitat for many species. CWD dynamics varies greatly from primeval to managed forests. To assess the CWD dynamics, a chronosequence trial was established in two Scots pine forests in the Central mountain range in Spain. Although, the shelterwood system has been applied in both forests, one has received more intensive silviculture, whereas in the other, regeneration has been much more gradual and the thinning regime has not been so intensive. In order to inventory CWD, five decay classes and four categories according to size and CWD type (stumps and fallen logs or branches) were defined. The volume of branches and logs (estimated from length and mean diameter) and the number of stumps by size class and decay class are used to characterise the CWD. The most notable differences between the two silvicultural systems can be appreciated in the graph as peaks for temporary distribution of larger logs and stumps when intensive silviculture is applied. The CWD observed in the forests studied is mainly produced by logging. The maximum volume of logs and branches above 5 cm in diameter is 43.25 m³/ha after regeneration felling in the first forest, whereas in the other, a maximum of 16.30 m³/ha is reached at 60 years, just after thinning. Large stumps (diameter equal or greater than 30 cm) make up an important part of CWD biomass in these forests just after the regeneration felling. A model was developed to predict the changes in CWD quantity and quality distribution over time as well as predicting the effect of different silviculture options on CWD dynamics. The model integrates two different processes: the CWD inputs (which may be continuous or instant), and the decay process, modelled through a Richards–Chapman function. The average lifetime of CWD obtained ranges from 30 years for stumps over 30 cm in diameter to 8 years for logs with a diameter less than 10 cm.     

Long-term trends in loblolly pine productivity and stand characteristics in response to thinning and fertilization in the West Gulf region

Two levels each of thinning and fertilization were applied to a 7-year-old loblolly pine (Pinus taeda L.) plantation on a nitrogen- and phosphorus-deficient West Gulf Coastal Plain site in Louisiana. Levels of thinning were no thinning, or thinning applied 7 and 14 years after stand initiation. Levels of fertilization were no fertilization or broadcast fertilization with diammonium phosphate at age 7 years plus refertilization with urea, monocalcium phosphate, and potash at age 14 years. Long-term measurements of climate, stand development and productivity, projected leaf area index, and foliar nutrition were initiated at age 11 years. We found that by age 17 years, thinning increased mean live-crown length from 4.2 to 7.8 m, and mean tree diameter from 15.0 to 21.8 cm compared to the unthinned treatment. After rethinning at age 14 years, stand basal area increased 1.2 and 19.2% between ages 15 and 17 years on the unthinned and thinned plots, respectively. Refertilization at age 14 years reestablished foliar N, P and K sufficiency, which increased leaf area index from 4.2 to 6.0 m² m⁻² on the unthinned plots and from 3.2 to 3.8 m² m⁻² on the thinned plots, and subsequently, increased gross stand biomass from 114 to 141 Mg ha⁻¹ on the unthinned plots and from 78 to 95 Mg ha⁻¹ on the thinned plots by age 17 years. Leaf area was an important factor controlling loblolly pine productivity. At our study site, however, competition for light and water and nutrition-limited foliage growth influenced the variability and scope of this relationship. Our results suggest that a positive and linear relationship between leaf area and loblolly pine productivity does not universally occur on loblolly pine sites.     

Dead trees and protected polypores in unmanaged north-temperate forest stands of Lithuania

The availability of coarse woody debris (CWD) and distribution of dead trees into categories of mortality (dead standing, broken and uprooted) were investigated in north-temperate forests of central Europe (Lithuania). The studied area comprised 188.7 ha and included 18 different stands 40–130 years of age with a variety of tree species (spruce (Picea abies (L.) Karst.), pine (Pinus sylvestris L.), alder (Alnus glutinosa (L.) Gaertn.), birch (Betula pendula Roth and B. pubescens Ehrh.), aspen (Populus tremula L.), oak (Quercus robur L.), forest types (caricus-sphagnum, vaccinium-myrtillus, oxalis, myrtillus-oxalis, caricus-calamagrostis) and edaphic conditions (peaty, sandy, loamy soils of different moisture). The stands were excluded from wood harvesting for at least 30 years. A total of 11 365 dead trees (over 10 cm in DBH) or 6160.7 m³ of dead wood was found (60.2 trees/ha and 32.6 m³/ha). The volume of CWD per hectare was larger in older stands (r_S=0.78, P<0.01). Tree mortality during the last 2 years consisted of 482 trees and 381 m³, or 1.28 trees/ha×year and 1.01 m³/ha×year. In 25–33% of cases it was wind-related. Uprooted and broken trees were of larger DBH than dead standing. The distribution into the categories of mortality was strongly dependent on tree species (chi-square test, d.f.=10,P=0). Dead standing dominated in CWD of pine and alder. Broken trees comprised almost a half in CWD of aspen, and about one-third in birch, alder and oak. Uprooting most often occurred in spruce, aspen and birch. Edaphic conditions and stand age had a pronounced impact on distribution into mortality categories for spruce (chi-square test, d.f.=20, P<0.00001) and pine (d.f.=8, P≤0.0003). On peat soil, only a minority of trees of both pine and spruce was uprooted, and standing dead prevailed. In CWD of spruce and pine, the proportions of both dead standing and broken decreased and that of uprooted trees increased on mineral soils of higher moisture and bulk density in older stands. By contrast, uprooting in birch and alder occurred less often on more wet sites, where the proportions of standing snags were higher. A total of 41 species of wood-decomposing polypores were found in the study area. Among those, 10 (24%) were of conservation value.     

Diversity and spatio-temporal dynamics of dead wood in a temperate near-natural beech forest (<Emphasis Type="Italic">Fagus sylvatica</Emphasis>)

The diversity, spatial patterns and temporal dynamics of dead wood were examined within the near-natural beech forests (Fagus sylvatica) of Serrahn (North-eastern Germany). Data were collected in an 8 ha sample plot and in two permanent plots (0.36 and 0.25 ha) that had been established at the end of the 1960s. The mean volume of dead wood was 94 m³ ha⁻¹, amounting to 14% of the total volume of all trees. The dead wood displayed a large variation in dead wood type, tree size and decay class. Standing dead wood accounted for about one-third of the total dead wood volume. The densities of standing dead trees were about 10% of the densities of the living trees over a wide range of diameters. The overall spatial distribution of dead trees exhibited a random pattern. Among the different dead wood types, standing entire dead trees and uprooted trees deviated from this pattern by displaying a significantly aggregated pattern. In the permanent plots a high mortality of overstorey trees was observed (1.3% year⁻¹) and the average amount of dead wood increased greatly from 2.9 to 111.6 m³ ha⁻¹ over the 35-year observation period. The near-natural beech forests of Serrahn have experienced a long period of low human interference. Nevertheless, our results suggest that the structure and dynamics of dead wood are strongly affected by the last major disturbance events that took place at the end of the Middle Ages. Information about the forest history is therefore a basic requirement when interpreting the results obtained in near-natural forests.