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1.
Chen Z  Kolb TE  Clancy KM 《Tree physiology》2001,21(16):1159-1169
We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.  相似文献   

2.
To facilitate ecosystem-specific management of juvenile mixtures of lodgepole pine (Pinus contorta Dougl. Ex Loud. Var. latifolia Engelm.) and trembling aspen (Populus tremuloides Michx.) in south-central British Columbia, we compared the characteristics of pine–aspen competition between a moist sub-boreal spruce and a dry interior Douglas-fir ecosystem. A total of 252 lodgepole pine and their neighbourhoods were examined across four untreated stands, each of which was sampled three times between ages 12 and 24 years. Pine diameter and height decreased with increasing density of trembling aspen at least as tall as the target pine (tall aspen) in both ecosystems. Regression analysis was used to examine the ability of tall aspen density and four competition indices (CIs) to predict pine size. Tall aspen density, which is easily assessed in the field, accounted for 63% and 69% of the variation in pine diameter and height in 20–24 year-old stands, respectively. The most successful competition index, based on the basal diameter ratio (BDR) of trembling aspen to pine accounted for, respectively, 78% and 73% of the variation. In the same stands, R2 values were 1–5% lower when tall aspen density and BDR at age 15–19 years were used to predict size of 20–24-year-old pine.  相似文献   

3.
Impact and recovery of juvenile western hemlock stands were measured 5 years after the end of an outbreak of western blackheaded budworm Acleris gloverana (Walsingham) (Lepidoptera: Tortricidae) and hemlock sawfly Neodiprion tsugae Middleton (Hymenoptera: Diprionidae) on Haida Gwaii, British Columbia, Canada. Overall, whole-tree mortality was infrequent (3.6–6.8%) but aggregated in a few, severely defoliated stands. Top-kill of the apical leader of surviving trees was more frequent (7.2–22.1%) and many of these trees had not recovered a growing dominant leader 5 years after the original damage. These severe impacts were most common in the largest trees in the youngest stands which had been spaced just before the outbreak. Radial growth was also reduced for 2 years following defoliation. However, mean annual ring width and basal area increment in surviving trees recovered quickly and was even enhanced in post-defoliation stands, particularly in spaced stands that had been moderately defoliated. Management recommendations include planning for likely outbreaks by avoiding creation of expanses of young regenerating stands of western hemlock during harvest and modifying spacing treatments when an insect outbreak is imminent to reflect the greater risk to trees <20 years of age.  相似文献   

4.
Overstory mortality, understory tree recruitment, and vegetation development were assessed in trembling aspen (Populus tremuloides Michx.) stands following two recent episodes of forest tent caterpillar defoliation (Malacosoma disstria Hbn.) in northeastern Ontario. The results suggest that poplar (aspen and balsam poplar (Populus balsamifera L.)) mortality increased with consecutive years of insect defoliation occurring from the mid-1980s to mid-2000s and the proportion of poplars in the overstory, but decreased with improved pre-defoliation tree vigour (DBH increment). The first outbreak, which lasted from the mid-1980s to early 1990s, was more severe in terms of insect defoliation and contributed more to poplar mortality and decline. The decline began in the late 1990s and peaked in early 2000s. Poplar regeneration and understory shrubs responded rapidly to foliage loss to insect defoliation and mortality of overstory poplars. The regenerated poplars were able to maintain their growth under developing shrubs and residual overstory canopy and numbers were sufficient to compensate for the poplar trees lost to insect infestation. The defoliation-induced overstory decline will accelerate the transition of aspen stands to conifer dominance through enhanced conifer recruitment and growth, and reduced hardwood overstory in aspen-dominated stands, while hardwood dominance will persist in pure aspen stands. From a timber supply perspective, the decline caused by forest tent caterpillar defoliation could delay the availability of aspen stands for harvesting by 40–50 years.  相似文献   

5.
Effective utilization of a patch or cluster approach to growing aspen-white spruce mixtures in the boreal forest requires an understanding of how the faster growing aspen patches influence microclimate and spruce growth in adjacent spruce patches. In this paper we examine, how young (11–13 year old) aspen (Populus tremuloides Michx) patches influence microclimatic conditions in adjacent openings and how these are reflected in the growth of white spruce (Picea glauca [Moench] Voss) on three boreal mixedwood sites in west-central Alberta, Canada. Light levels increased continuously across the aspen stand boundary and reached maximum levels within the opening, while soil moisture was highest near the edge of the opening and then decreased with distance into the aspen stand or into the adjacent opening north of the aspen stand. Light levels were reduced over a greater distance when moving north from aspen stand edges compared to openings located south of the young aspen. These young aspen stands provided growing season frost protection within one tree length from the edge. The growth of white spruce was positively related to initial crown size and either light levels or distance from the edge. Stem volume growth was best predicted by initial tree size and the light levels at the midcrown of seedlings.  相似文献   

6.
We used manual cutting to manipulate trembling aspen (Populus tremuloides Michx.) density and spatial arrangement in relation to crop lodgepole pine (Pinus contorta Dougl. Ex Loud. var. latifolia Engelm.) on two sites in contrasting dry, cool to cold ecosystems of south-central British Columbia. In the dry, cool interior Douglas-fir ecosystem (IDFdk3), we reduced the density of tall aspen (aspen at least as tall as target pine) to 0 (broadcast removal), 1000, 2500, or 4000 stems/ha when the planted lodgepole pine was 6 years old. Eight years later, pine height/diameter ratio (HDR) was significantly lower in the broadcast removal and 1000 stem/ha treatments than in the control. There were no other significant growth responses and pine survival and vigour were good regardless of treatment. In contrast, in a dry, cold sub-boreal pine spruce ecosystem (SBPSxc) where treatments were applied at a stand age of 11 years, naturally regenerated lodgepole pine stem diameter increased significantly in the broadcast removal treatment relative to the untreated control within 2 years. After 4 years, HDR had declined significantly relative to the control where tall aspen density was ≤1000 stems/ha. There were no significant pine responses where 2500 tall aspen stems/ha were retained or where tall aspen were removed only within a 1-m radius around pine. The greater difference in height (height differential) between aspen and pine at the SBPSxc than the IDFdk3 site may partly explain the differing response of lodgepole pine to treatment. Trends of decreasing sucker density with increasing aspen retention were evident at both sites, but differences were significant (p ≤ 0.05) only at the SBPSxc site.  相似文献   

7.
Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39-year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m2 ha−1. Thinning of overstory aspen to 1000 or 2000 stems ha−1 did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.  相似文献   

8.
Establishing white spruce (Picea glauca (Moench) Voss) by planting it under established aspen (Populus tremuloides Michx.), stands has substantial potential as a technique for regenerating boreal mixedwood stands. The presence of an aspen overstory serves to ameliorate frost and winter injury problems and suppresses understory vegetation that may compete with white spruce. In this study we examine the growth of white spruce during the first 10 years after being planted underneath a 39 year-old stand of trembling aspen following thinning and fertilization. Results indicate successful establishment and reasonable growth rates of white spruce planted under thinned and unthinned aspen stands, even with aspen basal area of 51 m2 ha−1. Thinning of overstory aspen to 1000 or 2000 stems ha−1 did not increase light reaching seedlings, but did result in improvements in light above the shrub layer and in diameter and height growth of the underplanted seedlings. However, these increases in growth of underplanted spruce may not justify the expense of thinnings. Fertilization of these stands prior to planting had no effect on spruce growth. Growth of spruce underplanted at this site near Fort Nelson was similar to that at two other stands near Dawson Creek, B.C.  相似文献   

9.
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10.
Trembling aspen (Populus tremuloides Michx.) density and growth were assessed 9–12 years after stand establishment to determine whether mechanical site preparation (MSP) affects crop tree quality. Study sites were either treated with disc trenching or ripper ploughing and planted with white spruce (Picea glauca (Moench) Voss) seedlings immediately after harvest (treated) or were undisturbed since harvest (control). Stands were surveyed during the summer of 2002 with standard regeneration survey plots. Results show that aspen stem density and height were lower in MSP-treated areas relative to untreated areas. Diameter growth rates were unaffected by treatment, however the percentage of stem discolouration was higher in untreated control stands compared to site prepared areas. The results of this study indicate that there are no long-term benefits to carry out MSP for aspen promotion. However, as MSP does not appear to seriously harm the aspen crop, we suggest that this treatment can still be used on sites where aspen densities may be low without treatment (e.g., sites with extremely low soil temperatures, poor soil aeration, or vigorous competitive vegetation) or where a mixture of aspen and planted spruce are desired.  相似文献   

11.
Aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench.) Voss) were planted 0.5 m apart in intimate mixtures in 5 × 4 m plots, with two moisture regimes—irrigation versus control—and five species compositions—pure aspen (Aw100), mixed aspen and spruce (Aw83Sw17, Aw50Sw50, Aw17Sw83), and pure spruce (Sw100), replicated six times. Fifth-year assessments indicated that irrigation increased individual tree growth (height, RCD, crown width), plot leaf area index (LAI), and wood biomass. Increased aspen composition reduced the availability of soil moisture and consequently the growth of individual trees. With increased aspen composition more growth was allocated to stem in aspen and to foliage in white spruce. Comparatively, aspen responded more to irrigation and thus their growth is more dependent on precipitation than that of spruce. Among the three growth variables assessed, height responded more to irrigation in both species. Equal mixtures and aspen-dominated mixtures in control plots had higher productivity in terms of total wood biomass in both absolute and relative terms. The implications of these findings are discussed in relation to managing aspen and white spruce mixedwood forests under increasing drought expected as a result of climate change.  相似文献   

12.
There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.  相似文献   

13.
Trembling aspen (Populus tremuloides Michx.) from 14 aspen stands ranging in age from 5 to 142 years were destructively sampled to provide branch, crown and bole growth information. Analysis of crown development processes such as branch angle, branch extensional growth, branch crookedness and annual height growth suggested a significant increase in relative branch growth with height and age which was linked to the temporal decline of height growth. Crown class dependent crown length (%) and crown area relationships were observed indicating crown size followed the sequence: open grown trees > dominants > suppressed trees. A significant shift in crown development from vertical to lateral crown expansion was observed as aspen mature.  相似文献   

14.

Assessing defoliation and measuring litterfall are two different ways of estimating the shedding of needles and leaves from the forest canopy. Both variables can be said to reflect the crown condition, but the two methods have rarely been compared. In this study the visual observations of defoliation of individual trees were compared with sampled litterfall data for the two main tree species in Denmark, Fagus sylvatica and Picea abies. Defoliation assessments and litterfall measurements were performed in seven level II plots. Six of the stands were 40 yrs old and the remaining stand was 81 yrs old. Both a positive and a negative correlation, which were not significant, were observed between defoliation and total leaf litterfall on the beech sites. Similarly, no significant correlation was observed between defoliation and the yearly needle litterfall on the younger Norway spruce plots. However, the defoliation and the yearly needle litterfall at the old stand at Klosterhede were positively correlated on a 10% significance level. A positive correlation was also apparent at the younger stands between the defoliation and the needle litterfall from the period April-July of the same year in which defoliation was assessed. The absence of a clear connection between the two assessments is discussed. The two assessments are apparently of widely different origin, which makes comparison difficult.  相似文献   

15.
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 (rS=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.  相似文献   

16.
Three lodgepole pine and aspen mixedwood sites located in the central interior of British Columbia within the Sub Boreal Spruce (SBS) biogeoclimatic zone were chosen to study the neighbourhood aspen competition and canopy light environment of 14–19-year-old lodgepole pine. All three sites had previously been established as separate research trials designed to explore various silviculture options for controlling aspen competition (aspen brushing, herbicide, thinning and untreated areas). For each site, 33–36 pine trees were selected to represent the observed range of light regimes under the influence of various aspen competition levels. At each sample pine, competition and stand measurements were made and a series of vertical canopy light measurements from the top to the base of the live crown. After an evaluation of a variety of competition indices, the index DRD; sum of the ratio of each of the three nearest neighbour's DBH to the subject pine divided by their distance and, amount of available light at the top of the crown (DIFNt) were found as the best overall predictors of pine stem volume growth. A site specific exponential relationship of relative pine stem volume growth to DRD was found and minimum growth response competition thresholds were determined, which could provide useful targets where maximizing pine volume is intended. Evaluation of both linear and non-linear models of DIFNt versus height growth indicated the response to be linear across the observed range of available light. Implications for management are discussed.  相似文献   

17.
This paper summarises the results from 35 years-observed thinning experiments on 256 permanent sample plots in 10–60 year-old stands of ash, aspen, birch, oak, pine and spruce in Lithuania. Thinning enhanced crown projection area increment of residual trees. The largest effect was observed in stands of aspen and birch (growth increase by 200%), followed by ash and oak (over 100%), and spruce and pine (about 80%). Thinning also promoted dbh increment, especially in younger stands, and the increase of dbh increment was positively correlated with the thinning intensity. The strongest reaction was exhibited by oak and aspen, while ash, birch and conifers reacted to a lower extent. Low and moderate intensities of thinning stimulated volume production in younger stands while the opposite was observed in older stands with increasing removals. Spruce stands exhibited relatively strongest increase of volume increment and pine, –the weakest, while the effect on deciduous species was intermediate. The results demonstrate that significant increase in volume increment is achievable with thinning of only young forest stands, e.g. 10–20 year-old pine, birch and ash, or 10–30 year-old oak, aspen and spruce.  相似文献   

18.
In the boreal forest of Alberta, fire and wind often open gaps in the canopy where late-successional species can establish and over time cause a shift in the species distribution from deciduous (e.g., trembling aspen) dominated to mixedwood, to shade-tolerant conifer (e.g., white spruce) dominated stands. This study attempted to understand the change of density-dependent competition in a boreal chronosequence and the role of tree competition in affecting stand structure and mortality. Four 1-ha stem-mapped plots were established to represent a chronosequence comprised of aspen dominated, mixedwood, and spruce dominated stands in Alberta. Second order spatial point-pattern analysis using Ripley's K(t) function showed that intraspecific competition is a prevailing force causing conspecific tree mortality and thus shaping the stand structure. The results of bivariate K(t) function analysis did not reveal sufficient evidence of interspecific competition. This suggested that competitive interaction among heterospecific trees was not strong enough to cause significant tree mortality, but the analysis of marked correlation function revealed that interspecific competition could have a negative impact on tree growth. This study highlights the importance of density-dependent competition in understanding stand dynamics of boreal forests over succession.  相似文献   

19.

Tree mortality and growth losses following insect defoliation are poorly documented in Scandinavia. In 1990-1991, Diprion pini (L.) caused extensive defoliation to Scots pine in Lauhanvuori national park and surrounding areas in south-western Finland. Most trees lost all their foliage in 1990. In 1991, the outbreak area was sprayed with diflubenzuron (Dimilin®), except in the national park, where trees were severely defoliated again. No further defoliation occurred in 1992. In spring 1993, sprayed trees had ca 30% foliage, whereas unsprayed trees on average carried less than 10% of full foliage. The latter trees were susceptible to attack by Tomicus piniperda (L.), whereas the former largely escaped beetle attack. Beetle attacks peaked in 1993, and depletion of suitable host trees probably terminated the beetle outbreak in the area. Two years of severe defoliation resulted in substantial tree mortality and growth losses. In spring 1997, these unsprayed stands had suffered a ca. 50% loss in basal area which was mainly because of mortality, and about half of the dead trees had been attacked by T. piniperda. Surviving trees had ca 50% of full foliage, and radial growth had still not recovered. Basal area growth was reduced by 40-70%, depending on the amount of foliage left after the second year of defoliation. In contrast, tree mortality and beetle attack in the sprayed stands were negligible, and these trees had recovered full foliage and radial growth by spring 1997. Thus, one year of total defoliation resulted in an estimated loss in basal area growth of approximately 30% during ca. 5 yrs. In conclusion, the spraying operation was economically justified, as it prevented substantial tree mortality and reduced growth losses.  相似文献   

20.
When spruce beetles (Dendroctonus rufipennis) thin a forest canopy, surviving trees grow more rapidly for decades until the canopy closes and growth is suppressed through competition. We used measurements of tree rings to detect such growth releases and reconstruct the history of spruce beetle outbreaks at 23 mature spruce (Picea spp.) forests on and near the Kenai Peninsula, Alaska and four mature white spruce (Picea glauca) forests in Kluane National Park and Reserve, Yukon Territory. On the Kenai Peninsula, all stands showed evidence of 1–5 thinning events with thinning occurring across several stands during the 1810s, 1850s, 1870–1880s, 1910s, and 1970–1980s, which we interpreted as regional spruce beetle outbreaks. However, in the Kluane region we only found evidence of substantial thinning in one stand from 1934 to 1942 and thinning was only detected across stands during this same time period. Over the last 250 years, spruce beetle outbreaks therefore occurred commonly among spruce forests on the Kenai Peninsula, at a mean return interval of 52 years, and rarely among spruce forests in the Kluane region where cold winter temperatures and fire appear to more strongly regulate spruce beetle population size. The massive 1990s outbreaks witnessed in both regions appeared to be related to extremely high summer temperatures. Recent outbreaks on the Kenai Peninsula (1971–1996) were positively associated with the 5-year backwards running average of summer temperature. We suggest that warm temperature influences spruce beetle population size through a combination of increased overwinter survival, a doubling of the maturation rate from 2 years to 1 year, and regional drought-induced stress of mature host trees. However, this relationship decoupled after 1996, presumably because spruce beetles had killed most of the susceptible mature spruce in the region. Thus sufficient numbers of mature spruce are needed in order for warm summer temperatures to trigger outbreaks on a regional scale. Following the sequential and large outbreaks of the 1850s, 1870–1880s, and 1910s, spruce beetle outbreaks did not occur widely again until the 1970s. This suggests that it may take decades before spruce forests on the Kenai Peninsula mature following the 1990s outbreak and again become susceptible to another large spruce beetle outbreak. However, if the recent warming trend continues, endemic levels of spruce beetles will likely be high enough to perennially thin the forests as soon as the trees reach susceptible size.  相似文献   

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