Effects of crown release on basal area growth rates of some broad-leaved tree species with different shade-tolerance

We compare three or four years effects of crown release on stem growth rates among four deciduous broad-leaved tree species,Fagus crenata, Quercus crispula, Magnolia obovata andAcer mono, in an even-aged coppice forest, central Japan. The crown release significantly improved relative growth rates for basal-area (RGR_BA) of the four species. However the effects of neighboring gap area (GAPA) on the RGR_BA differed among the species. The effects were significant forQ. crispula andM. obovata, indicating that growth rates of these two species increase with intensity of the crown release. In contrast, the RGR_BA ofF. crenata andA. mono were not correlated with the GAPA, suggesting that their growth rates are unrelated to the intensity. We considered that such differences among species were closely related to their photosynthetic characteristics of light use; less-tolerant species (Q. crispula andM. obovata) exhibit greater growth rates relative to the intensity of crown release than tolerant species (F. crenata andA. mono). Based on these results, we proposed implications for thinning practices in mixed forests of species with different shade-tolerance.     

Changes in size of soil seed bank in Robinia pseudoacacia L. (Leguminosae), an exotic tall tree species in Japan: Impacts of stand growth and apicultural utilization

This study examined the factors that influence the size of a soil seed bank in Robinia pseudoacacia stands. We proposed two hypotheses: the amount of buried seeds of R. pseudoacacia is proportional to stand biomass rather than stand age, and apicultural utilization influences the size of the soil seed bank. R. pseudoacacia generally produces seeds with various degrees of physical dormancy. In addition, this tree is short-lived, and fecundity declines after 30–40 years in relation to tree vigor. However, R. pseudoacacia is intensively used in apiculture as an important honey source, and supply of honeybees by beekeepers could influence seed crops. We investigated 25 plantations in Hokkaido (Central, Hidaka, and Oshima) and in Nagano, Japan, where many naturally regenerated R. pseudoacacia stands occur. We found great variation in the size of the soil seed bank among stands, ranging from 13,757 to 6.4 seeds m⁻² per plot. A generalized linear mixed model revealed that both basal area (BA_Rp) and apicultural utilization best explained the size of the soil seed bank. Both the positive effect of BA_Rp and small contribution of stand age to the model implied that the soil seed bank is transitory, and will not persist for very long time. The large contribution of apicultural utilization to the size of the soil seed bank implied pollinator limitation under natural conditions. R. pseudoacacia often has large flower crops, and native pollinators can not keep up with the demand for pollination. Thus, the supply of honeybees by beekeepers should improve the pollination success of R. pseudoacacia. From these results, we conclude that we must consider stand history with respect to apicultural utilization if the land is harvested or a stand declines.     

Differences in leafminer (Phyllonorycter, Gracillariidae, Lepidoptera) and aphid (Tuberculatus, Aphididae, Hemiptera) composition among Quercusdentata, Q. crispula, Q. serrata, and their hybrids

Leafminer (Phyllonorycter, Gracillariidae, Lepidoptera) and aphid (Tuberculatus, Aphididae, Hemiptera) composition were studied in three deciduous oak species, Quercus dentata, Q. crispula, and Q. serrata, and their hybrids in Tomakomai Experimental Forest of Hokkaido University, Hokkaido, northern Japan. Identification of trees in this forest was done mainly on the basis of discriminant analysis on leaf morphology with reference to trees in pure Q. dentata and Q. crispula stands and a Q. serrata stand mixed with Q. crispula. The results suggested that hybridization occurred in all combinations (i.e. Q. dentata–Q. crispula, Q. crispula–Q. serrata, and Q. serrata–Q. dentata) and the frequency of hybrids was approximately 10%. The composition of Phyllonorycter and Tuberculatus species differed between Q. dentata and Q. crispula or Q. serrata, but did not differ between Q. crispula and Q. serrata. Thus, Q. dentata could differ from Q. crispula and Q. serrata in chemical properties that determine herbivore host selection, survival, and performance, possibly reflecting eco-physiological differences or phylogenetic distances. The study insects were divided into three groups: species specialized to Q. dentata (three Phyllonorycter and one Tuberculatus species), those to Q. crispula and Q. serrata (six Phyllonorycter and two Tuberculatus species), and a species collected at least from Q. dentata and Q. crispula (one Tuberculatus species). Putative hybrid trees of Q. dentata and Q. crispula harbored both Q. dentata-specific and Q. crispula-specific insects.     

Aerially applied methylcyclohexenone-releasing flakes protect Pseudotsuga menziesii stands from attack by Dendroctonus pseudotsugae

We tested methylcyclohexenone (MCH), an anti-aggregation pheromone for the Douglas-fir beetle (Dendroctonus pseudotsugae), for protection of Douglas-fir (Pseudotsuga menziesii) stands by applying MCH-releasing polymer flakes by helicopter twice during summer 2006 to five 4.05-ha plots in the State of Washington, USA. Five similar plots served as untreated controls. We assessed D. pseudotsugae flight into study plots using baited pheromone traps, and tallied D. pseudotsugae attack rates on all P. menziesii trees in 2005 and 2006. We also measured stand basal area and incorporated that as an explanatory variable in the analysis. Significantly fewer D. pseudotsugae were trapped in treated plots than in control plots, and significantly fewer P. menziesii trees were attacked in treated plots than in control plots. The attack rate in untreated stands was nearly 10 times that of treated plots, and stands with higher basal area were significantly more likely to be attacked by D. pseudotsugae than were stands of lower basal area. Attack rates in 2006 and 2005 were significantly correlated, regardless of treatment.     

Two-compartment models for describing temporal variation in the effective leaf area index of Robinia pseudoacacia stands on the Loess Plateau

Leaf area index (LAI) is a key parameter in many ecological models and its phenology significantly affects on net ecosystem production in deciduous forests. We examined trends in LAIe (effective leaf area index) using two-compartment models to test effects of stand age and slope aspect on seasonal rates of LAIe increase (k₁) and decrease (k₂). LAIe measurements were acquired from 24 sample plots over five months and used to develop robust predictive models for modeling LAIe dynamics. Generally, the stands’ k₁ values were substantially greater than their k₂ values. The LAIe increased rapidly between leaf emergence and saturation, and then lower slowly from leaf saturation to senescence. The strongest increase in LAIe was observed in 11- to 15-year-old stands on shady slopes (k₁ = 4.76) and in young stands (<10 years) on sunny slopes (k₁ = 3.94), but k₂ values were largest in mature stands. The trees on shady slopes generally had higher k₁ and lower k₂ values than those on sunny slopes, except in the youngest stand. The results showed that two-compartment models can robustly describe leaf growth and loss rates in black locust stands, and that both stand age and aspect both strongly affect the rate and magnitude of changes in LAIe during the growing season. Hence, the two-compartment model is recommended as an effective method when monitoring the LAIe quantitative dynamics of broad-leaved forest.     

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.     

Comparison of the growth traits of a commercial pioneer tree species,paper mulberry (Broussonetia papyrifera L. Vent.), with those of shade-tolerant tree species: investigation of the ecophysiological mechanisms underlying shade-intolerance

In rural areas of northern Laos, a commercially valuable pioneer tree species, paper mulberry (Broussonetia papyrifera L. Vent.) has been recommended for intercropping under plantations of commercial trees. However, less is understood about the growth of this pioneer tree species in the understorey and the mechanism underlying the shade intolerance. We measured growth characteristics for seedlings of paper mulberry under four light intensities. We compared the relative growth rates in aboveground biomass and standing leaf area (RGR_mass and RGR_leaf), light-capture efficiency, and seeding-level mass-based daily photosynthetic rates (A _mass) with those of field-grown seedlings of eight shade-tolerant species to identify factors potentially responsible for shade-intolerance. Most growth traits of the paper mulberry seedlings did not differ consistently from those of the shade tolerant species. The ecophysiological–architectural model software showed higher A _mass and RGR_mass capacity in paper mulberry than in shade-tolerant species. Despite their higher RGR_mass, paper mulberry seedlings had negative RGR_leaf under shaded conditions due to short leaf lifespan. The linear RGR_mass–RGR_leaf relationship for paper mulberry had a high RGR_mass intercept, indicating that a high RGR_mass was required to provide positive RGR_leaf. Progressive decreases in standing leaf area with time, and possibly photosynthesis, appear to be responsible for the shade-intolerance of paper mulberry. Although intercropping of paper mulberry has been suggested in the species’ native region, understorey cultivation of paper mulberry would only be possible with relatively open canopies.     

Effect of thinning on LAI variance in heterogeneous forests

Leaf Area Index (LAI) is a main variable controlling carbon and water fluxes. This paper estimated the effect of thinning on the spatial distribution of leaf area in French forests. While many studies have focused on average LAI, we estimated clumping and measured both average LAI and the variation around it. LAI was derived from digital hemispherical photos at three sites: an unmanaged Fagus sylvatica forest in temperate area (control site), a mixed Mediterranean forest of Quercus ilex and Pinus halepensis, and regeneration of F. sylvatica under a mature stand of Pinus nigra in mountainous area. LAI measurements were also made with LAI 2000 devices over 5 years (from 1994 to 1998) within forest stands dominated by either beech (F. sylvatica L.), by oaks (Quercus petraea (Matus) Liebl., Quercus robur (Matus) Liebl.), or by Scots Pine (Pinus sylvestris L.). Thinning led to a variable decrease in LAI. The coefficient of variation of LAI (CV_LAI) provided a useful ecological index of the level and type of thinning. For undisturbed stands, CV_LAI varied from 10% to 20%, corresponding to the higher average LAI values. Disturbances created by thinning increase LAI spatial variability, resulting in larger CV_LAI values for all stands considered. Possible explanations of these results and use in remote sensing were discussed.     

Stand and landscape-level factors related to bird assemblages in exotic pine plantations: Implications for forest management

Plantations cover a substantial amount of Earth's terrestrial surface and this area is expected to increase dramatically in the coming decades. Pinus plantations make up approximately 32% of the global plantation estate. They are primarily managed for wood production, but have some capacity to support native fauna. This capacity likely varies with plantation management. We examined changes in the richness and frequency of occurrence of bird species at 32 plots within a Pinus radiata plantation (a management unit comprising multiple Pinus stands) in south-eastern Australia. Plots were stratified by distance to native forest, stand age class and thinning regime. We also assessed the landscape context of each plot to determine relationships between bird assemblages and stand and landscape-level factors. Bird species richness was significantly higher at plots ≥300 m from native forest and in mature (∼20 years since planting) and old (∼27 years since planting) thinned pine stands. We were able to separate the often confounding effects of stand age and thinning regime by including old stands that had never been thinned. These stands had significantly fewer species than thinned stands suggesting thinning regime, not age is a key factor to improving the capacity of pine plantations to support native species (although an age × thinning interaction may influence this result). At the landscape level, species richness increased in pine stands when they were closer to native riparian vegetation. There were no significant differences in species composition across plots. Our study indicates the importance of stand thinning and retention of native riparian vegetation in improving the value of pine plantations for the conservation of native fauna.     

Individual-scale responses of five dominant tree species to single-tree selection harvesting in a mixed forest in Hokkaido, northern Japan

We investigated the individual-scale responses of five dominant species (Abies sachalinensis, Acer mono, Tilia japonica, Quercus crispula, and Betula ermanii) to single-tree selection harvesting in a conifer–broadleaved mixed forest in Hokkaido, northern Japan. Using data from stems with a diameter at breast height of ≥12.5 cm, collected during 20 years of monitoring a 6.7-ha stand, we analyzed the effects of harvesting in the neighborhood on tree recruitment and the growth (diameter class transition) and mortality of the residual trees. The effects of harvesting varied considerably among tree sizes and species. Harvesting improved the recruitment of A. mono and B. ermanii, and moderated the negative effect of the initial basal area of the surrounding canopy trees on the recruitment of Q. crispula. Conversely, harvesting limited the recruitment of A. sachalinensis by offsetting the positive effect of the initial basal area of the surrounding canopy trees. The growth of A. sachalinensis and Q. crispula decreased with the initial basal area of the surrounding canopy trees. Harvesting in the neighborhood resulted in an improvement in the growth of the trees of these species only in the smaller size classes. With increasing local harvesting intensity, the mortality of smaller A. sachalinensis trees decreased, whereas the mortality of larger trees increased. These results suggest that differences in the local harvesting intensity, spatial patterns of harvesting, and initial stand structures influence the stand-scale dynamics in response to partial harvesting in the mixed forests of this region.     

Effects of previous forest types and site conditions on species composition and abundance of naturally regenerated trees in young <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> plantations in northern Japan

Naturally regenerated trees in young Cryptomeria japonica plantations were investigated in 141 quadrats of 10 × 50 m within a watershed of 1,000 ha, and factors affecting their composition and abundance were examined. The species composition of naturally regenerated trees was classified into four types. Dominant species were Swida controversa, Magnolia hypoleuca and Pterocarya rhoifolia in Type A1, Betula maximowicziana, M. hypoleuca, Quercus crispula and Castanea crenata in Type A2, Q. serrata and C. crenata in Type B1, and Pinus densiflora in Type B2. The results of path analysis showed significant influences of previous forest type and geology among the factors that correlated with the species composition of naturally regenerated trees; previous forest type in particular showed a higher absolute path coefficient value. Species composition types of naturally regenerated trees corresponded to the previous forest types: Types (A1 and A2) corresponded to the natural forests composed of Thujopsis dolabrata var. hondai, Fagus crenata, Aesculus turbinata, etc., Type B1 corresponded to the secondary Q. crispula and Q. serrata forest, and Type B2 corresponded to the secondary P. densiflora forest, respectively. The abundance of naturally regenerated trees was strongly affected by geology; i.e., plantations on soft-sedimentary dacitic tuff contained great amounts of colonizing P. densiflora trees.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

The effects of stand structure on specific needle area in closed-canopy Chinese pine plantations

The specific leaf area (SLA) is an important variable reflecting the carbon gain strategy of tree growth, but the relationships between SLA and other environmental factors have not been studied extensively at the stand level. The aim of this study was to define the relationships between stand structure and SLA in order to improve the predictive value of SLA for forest management models. Various parameters of stand structure and specific needle area (SNA) were measured in 14 different even-aged and closed-canopy stands of Chinese pine (Pinus tabulaeformis Carr.). Correlation and regression analysis revealed several significant relationships between stand structure and SNA. Stand density exerted a significant effect on mean-canopy SNA (SNA_mean). Stand density was also strongly correlated to mean-layer SNA (SNA_L), especially at lower canopy layers. SNA_L increased at lower canopy layers within each stand, and the increase was greater in higher density stands. Within the range of stand densities examined in this study, the SNA_mean initially increased sharply with increasing stand density, but the rate of rise declined as the density increased. Finally, it reached stability when the stand density was above about 3,000 trees ha^?1.     

Effects of thinning level and site productivity on age-related changes in stand volume growth can be explained by a single rescaled growth curve

In this study, I developed a nonlinear mixed-effects model based on the Richards curve that describes the effects of thinning and site productivity on stand volume growth. I fitted the model to data from 29 long-term experimental monitoring plots in Japanese cedar (Cryptomeria japonica D. Don) plantations in northeastern Japan. Simulations using the model on high-productivity sites and at moderate thinning intensities indicated that net yields after ca. 80 years were lower in unthinned stands than in thinned stands; in other cases, such as at younger ages and at sites with low and average productivities, net yields were similar for each thinning intensity (including no thinning). These results could be attributed to the fact that the net current annual increment in stand volume (CAI_net) in the thinned stands exceeded that of the unthinned stands after a certain age (the “inversion” age) and to the negative correlation between site productivity and the inversion age. In addition, I found that a single rescaled growth curve was capable of describing the growth in stand volume at sites with different thinning histories and site productivities. This rescaled curve simply and clearly explained the effects of thinning and site productivity on stand volume growth. The existence of a single curve suggests that the positive effect of thinning on stand volume growth does not depend on site productivity, but it will require patient measurements over longer periods to reveal a positive effect at sites with low productivity.     

Ectomycorrhizal fungal communities of Quercus liaotungensis along different successional stands on the Loess Plateau,China

Although Quercus liaotungensis forest is an important component of Loess Plateau forest ecosystems, little is known about the ectomycorrhizal fungal (EMF) symbiosis of Q. liaotungensis in this ecosystem. Here we investigated EMF communities of Q. liaotungensis along three successional stands on the central Loess Plateau (Yan’an, China): a seedling stand (1 to 3-year-old seedlings under a Populus davidiana forest stand), a young tree stand (20 to 30-year-old Q. liaotungensis forest stand), and a mature tree stand (50 to 70-year-old Q. liaotungensis forest stand). In each stand, the Q. liaotungensis roots were sampled, then EMF communities were examined on the basis of EMF morphotyping, PCR–RFLP, and DNA sequencing. In total, 70 EMF species were observed, and the total richness of estimated species exceeded 100 EMF species. The EMF community was composed of a few common taxa (approx. 7 %) and many rare taxa (approx. 61 %). EMF richness in young and mature trees was higher than in seedlings. EMF communities of young and mature trees were more similar than those of seedlings and trees. Nonmetric multidimensional scaling ordinations of the EMF community revealed more separation among Q. liaotungensis of different age. This study provides preliminary insight into the taxonomic diversity of EMF of Q. liaotungensis along different successional stands on the Loess Plateau.     

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.     

Discrimination among host tree species by the ambrosia beetle Platypus quercivorus

The ambrosia beetle Platypus quercivorus is a vector for the fungus that causes Japanese oak wilt, and susceptibility to infestation by P. quercivorus varies by tree species. We postulated that P. quercivorus discriminates among host tree species differing in susceptibility to attack. To test this postulate, we counted the number of flying male beetles (NFM), the number of holes bored by male beetles (NH), and the number of flying female beetles (NFF) per unit area of bark surface in three fagaceous tree species: Quercus crispula (with high susceptibility to infestation) and Q. serrata and Castanea crenata (both with low susceptibility). NFM and NH were used to calculate the proportion of male beetles that bored holes out of those that flew to the tree (PBM). We used generalized additive models to predict NFM, NFF, and PBM. The locations of trees, expressed as x and y coordinates, numbers of weeks after the first male beetle’s flying (WEEK), diameters of trees 130 cm above ground (DBH), and tree species (SP) were incorporated into the models as candidate explanatory variables. The best-fit models for NFM and NFF included WEEK and DBH and the effect of location; SP was not included in the models. For PBM, the best-fit model included WEEK, DBH, and SP. The results indicate that male P. quercivorus prefer Q. crispula to Q. serrata and C. crenata and that selection is made before boring holes on trees, but that P. quercivorus do not discriminate among host species when they fly to trees.     

Susceptibility of northern British Columbia forests to spruce budworm defoliation

Stand susceptibility to defoliation by spruce budworm, Choristoneura fumiferana (Clem.), was examined in the Fort Nelson area of the Prince George Forest Region of British Columbia. In a retrospective study, defoliation maps of the study area were overlaid onto British Columbia Ministry of Forests cover type maps using a geographic information system. Analysis of the combined data identified forest characteristics associated with increased susceptibility to defoliation by spruce budworm. These were stands where the leading species was white spruce (Picea glauca (Moench) Voss), or where spruce was associated with aspen (Populus tremuloides Michx. and P. balsamifera L.) in mixed stands. Susceptibility to defoliation also was related to site quality, level of crown closure and stand age. Spruce stands on medium quality sites (site index 15 to 25 m, at reference breast height age 50 years) were more susceptible than stands on both poor- and high-quality sites. When spruce was mixed with aspen, stands on higher quality sites were more susceptible to budworm attack than poor sites. Open stands, where crown closure was <50%, were more susceptible to attack by spruce budworm than closed canopy stands. Older stands (120–199 years) were more susceptible to budworm attack than younger stands (40–110 years). In defoliated plots monitored for 6 years, tree mortality and top-kill reached a maximum of 30.4 and 47.2%, respectively. The losses varied with level of defoliation and were reduced by applications of the biological pesticide Bacillus thuringiensis.     

Growth and productivity of black spruce in even- and uneven-aged stands at the limit of the closed boreal forest

The increasing commercial interest and advancing exploitation of new remote territories of the boreal forest require deeper knowledge of the productivity of these ecosystems. Canadian boreal forests are commonly assumed to be evenly aged, but recent studies show that frequent small-scale disturbances can lead to uneven-aged class distributions. However, how age distribution affects tree growth and stand productivity at high latitudes remains an unanswered question. Dynamics of tree growth in even- and uneven-aged stands at the limit of the closed black spruce (Picea mariana) forest in Quebec (Canada) were assessed on 18 plots with ages ranging from 77 to 340 years. Height, diameter and age of all trees were measured. Stem analysis was performed on the 10 dominant trees of each plot by measuring tree-ring widths on discs collected each meter from the stem, and the growth dynamics in height, diameter and volume were estimated according to tree age. Although growth followed a sigmoid pattern with similar shapes and asymptotes in even- and uneven-aged stands, trees in the latter showed curves more flattened and with increases delayed in time. Growth rates in even-aged plots were at least twice those of uneven-aged plots. The vigorous growth rates occurred earlier in trees of even-aged plots with a culmination of the mean annual increment in height, diameter and volume estimated at 40–80 years, 90–110 years earlier than in uneven-aged plots. Stand volume ranged between 30 and 238 m³ ha⁻¹ with 75% of stands showing values lower than 120 m³ ha⁻¹ and higher volumes occurring at greater dominant heights and stand densities. Results demonstrated the different growth dynamics of black spruce in single- and multi-cohort stands and suggested the need for information on the stand structure when estimating the effective or potential growth performance for forest management of this species.     

The effects of partial cutting on stand structure and growth of western hemlock–Sitka spruce stands in southeast Alaska

The effects of partial cutting on species composition, new and residual-tree cohorts, tree size distribution, and tree growth was evaluated on 73 plots in 18 stands throughout southeast Alaska. These partially cut stands were harvested 12–96 years ago, when 16–96% of the former stand basal area was removed.Partial cutting maintained stand structures similar to uncut old-growth stands, and the cutting had no significant effects on tree species composition. The establishment of new-tree cohorts was positively related to the proportion of basal-area cut. The current stand basal area, tree species composition, and stand growth were significantly related to trees left after harvest (p<0.001). Trees that were 20–80 cm dbh at the time of cutting had the greatest tree-diameter and basal-area growth and contributed the most to stand growth. Diameter growth of Sitka spruce and western hemlock was similar, and the proportion of stand basal-area growth between species was consistent for different cutting intensities.Concerns about changing tree species composition, lack of spruce regeneration, and greatly reduced stand growth and vigor with partial cuts were largely unsubstantiated. Silvicultural systems based on partial cutting can provide rapidly growing trees for timber production while maintaining complex stand structures with mixtures of spruce and hemlock trees similar to old-growth stands.