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1.
Crown shyness or canopy disengagement, the phenomenon wherein gaps around trees develop from swaying, whipping and shading, has been identified in the literature since the 1920s. Recent results by researchers at the University of Alberta have clearly described many of the processes involved for lodgepole pine [e.g. Rudnicki, M., Silins, U., Lieffers, V.J., Josi, G., 2001. Measure of simultaneous tree sways and estimation of crown interactions among a group of trees. Trees 15, 83–90; Rudnicki, M., Lieffers, V.J., Silins, U., 2003. Stand structure governs the crown collisions of lodgepole pine, Canadian Journal of Forestry Research 33, 1238–1244; Rudnicki, M., Silins, U., Lieffers, V.J., 2004. Crown cover is correlated with relative density, tree slenderness, and tree height in lodgepole pine. Forest Science 50, 356–363; Fish, H., Lieffers, V.J., Silins, U., Hall, R.J., 2006. Crown shyness in lodgepole pine stands of varying stand height, density, and site index in the upper foothills of Alberta. Canadian Journal of Forestry Research 9, 2104–2111]. However, explicit models of crown shyness are sparse in the literature. This paper describes the development of empirical models of crown shyness in lodgepole pine for British Columbia (BC). We measured crown area and neighbour locations on 60 trees growing in 13 stands in central BC. We estimated potential crown area (AV) using stem maps and Voronoi polygons constrained by estimates of maximum crown width, and then related observed crown area (AC) to AV and additional individual tree variables. One of the nine prediction equations was coded into a spatially explicit tree growth model modified to evaluate the effects of crown shyness at the stand level. Crown shyness models validated well against two independent sources and when linked with a light model tRAYci [Brunner, A., 1998. A light model for spatially explicit forest stand models. Forest Ecology and Management 107, 19–46], increased the below-canopy light by 0.07–0.11.  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.
Using tree data from permanent sample plots and climate data from the ClimateWNA model, mixed-effects height to live crown (HTC) models were developed for three boreal tree species in Alberta, Canada: trembling aspen (Populus tremuloides Michx.), lodgepole pine (Pinus contorta var. latifolia Engelm.) and white spruce (Picea glauca (Moench) Voss). Three model forms, the Wykoff model, a logistic model and an exponential model, were evaluated for each species. Tree height was the most significant predictor of HTC and was used in all models. In addition, we investigated the effects of competition and climatic variables on HTC modelling. Height–diameter ratio and either total stand basal area or basal area of coniferous trees were used as competition measures in the models. Different climate variables were evaluated, and spring degree-days below 0 °C, mean annual precipitation and summer heat–moisture index were incorporated into the aspen, lodgepole pine and white spruce models, respectively. Site index was only significant in lodgepole pine models. Residual variances were modelled as functions of tree height to account for heteroscedasticity still present in the mixed-effects models after the inclusion of random parameters. Based on model fitting and validation results as well as biological realism, the mixed-effects Wykoff models were the best for aspen and white spruce, and the mixed-effects logistic model was the best for lodgepole pine.  相似文献   

4.
We used field measurements and Monte Carlo simulations of canopy gap-size distribution and gap fraction to examine how beam radiation interacts with clumped boreal forest canopies of aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.) and jack pine (Pinus banksiana Lamb.). We demonstrate that the Beer-Lambert law can be modified to accommodate transmission of radiation through a clumped forest canopy as a function of path length or sun zenith angle. Multiband Vegetation Imager (MVI) measurements and Monte Carlo simulations showed that values of the zenith element clumping index (Omega(e)(0)) are typically between 0.4 and 0.5 in jack pine and black spruce and 0.65 in aspen. Estimates of LAI obtained from MVI measurements of the canopy gap fraction and adjusted for canopy clumping and branch architecture yielded LAI values of 3.0 in jack pine, 3.3 in aspen, and about 6.0 in black spruce. These LAI estimates were within 10-25% of direct measurements made at the same sites. Data obtained with the MVI, along with numerical simulations, demonstrated that assumptions of random foliage distributions in boreal forests are invalid and could yield erroneous values of LAI measured by indirect techniques and false characterizations of atmosphere-biosphere interactions. Monte Carlo simulations were used to develop a general equation for beam radiation penetration as a function of zenith angle in clumped canopies. The essential measurements included stem spacing, crown diameter, crown depth, and within-crown gap fraction.  相似文献   

5.
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   

6.
Competition for canopy space is a fundamental structuring feature of forest ecosystems and remains an enduring focus of research attention. We used a spatial neighborhood approach to quantify the influence of local competition on the size of individual tree crowns in north-central British Columbia, where forests are dominated by subalpine fir (Abies lasiocarpa), lodgepole pine (Pinus contorta) and interior spruce (Picea glauca × engelmanii). Using maximum likelihood methods, we quantified crown radius and length as functions of tree size and competition, estimated by the species identity and spatial arrangement of neighboring trees. Tree crown size depended on tree bole size in all species. Given low levels of competition, pine displayed the widest, shortest tree crowns compared to the relatively long and narrow crowns found in spruce and fir. Sensitivity to crowding by neighbors declined with increasing tree height in all but the pine crown radius model. Five of the six selected best models included separate competition coefficients for each neighboring tree species, evidence that species generally differ in their competitive effects on neighboring tree crowns. The selected crown radius model for lodgepole pine, a shade-intolerant species, treated all neighbors as equivalent competitors. In all species, competition from neighbors exerted an important influence on crown size. Per-capita effects of competition across different sizes and species of neighbors and target trees varied, but subalpine fir generally displayed the strongest competitive effects on neighbors. Results from this study provide evidence that species differ both in their response to competition and in their competitive influence on neighbors, factors that may contribute to maintaining coexistence.  相似文献   

7.
A series of conventional distance-independent and distance-dependent competition indices, a highly flexible distance-dependent crowding index, and two light resource estimation indices were compared to predict individual tree diameter growth of five species of mature trees from natural-origin boreal mixed forests. The crowding index was the superior index for most species and ecosites. However, distance-independent indices, such as basal area of competing trees, were also effective. Distance-dependent light estimation indices, which estimate the fraction of seasonal photosynthetically-active radiation available to each tree, ranked intermediate to low. Determining separate competition indices for each competitor species accounted for more variation than ignoring species or classifying by ecological groups. Species’ competitive ability ranked (most competitive to least): paper birch ≈ white spruce ≈> trembling aspen > lodgepole pine > balsam poplar. Stratification by ecosite further improved model performance. However, the overall impact of competition on mature trees in these forests appears to be small.  相似文献   

8.
In boreal forests of western Canada, lodgepole pine (Pinus contorta Dougl. ex. Loud.) and white spruce (Picea glauca (Moench) Voss) often grow together with numerous tall shrubs such as green alder (Alnus crispa (Ait.) Pursh) and little-tree willow (Salix spp.). In an area south of Grande Prairie, Alberta, Canada, we examined the effects of shrubs, herbs and other trees on nutrient and light availability and growth of white spruce and lodgepole pine. For white spruce the best competition measure (tested against volume increments of the past 3 years) was visually estimated % ground cover times the height of the competitor (VCHT) with light (DIFN) ranking in third place. For lodgepole pine, DIFN was the best competition measure for predicting volume increment and the best competition index was again VCHT. Taller conifers had a stronger competitive effect than tall shrubs, with their effect on white spruce being larger than that on lodgepole pine.  相似文献   

9.
In southern British Columbia, juvenile trembling aspen is managed primarily as a competitor with conifers rather than for its ecological and economic value. As a result, brushing treatments have been applied on a widespread basis and this practice is likely to continue in the near future. Given the potential for climate change to affect our valuation of aspen, we require a better understanding of factors that affect its development, its competitive ability with conifers and its responses to brushing. We used data from 11 aspen management experiments to examine the influence of climate and site factors on aspen height, cover, and density in 17–24 year-old control stands and 9–16 years after manual cutting or girdling. Models explained 64% and 89% of the variation in aspen height in control and manually brushed stands, respectively, but were poor for girdling. Increasing length of the frost-free period was associated with increasing aspen height in control stands, whereas drier summer conditions on cool aspects favoured height growth of aspen suckers following manual cutting. We also examined the influence of climate and site factors on three simple competition indices that describe the height and density of aspen relative to conifer height, and then tested how well these indices predicted conifer growth. The density of aspen taller than conifers accounted for 39% of the variation in lodgepole pine diameter and the ratio of aspen/conifer height accounted for 33% of the variation in Douglas-fir height, suggesting that aspen competition was only moderately important to conifer growth. Our findings imply that aspen may become more productive with warmer summers provided it is not limited by summer moisture availability and that mixed stand management is a viable option in southern interior stands.  相似文献   

10.
Tree growth variables and more complex derived variables such as vigor index have all been linked to the ability of lodgepole pine (Pinus contorta var latifolia) trees to defend against insect herbivores, particularly mountain pine beetle (Dendroctonus ponderosae Hopkins, Coleoptera: Curculionidae). These variables are considered indirect measures of carbohydrate reserves. Trees with high vigor index values have high growth rates and are assumed to have high carbohydrate reserves. However, this critical assumption was untested for lodgepole pine. In this paper we evaluated the relationship between carbohydrate concentration and tree characteristics. We measured concentrations of root starch in 10 pure lodgepole pine stands of different ages in the Canadian Rockies which had been fertilized, thinned, fertilized and thinned or left untreated in a 2 × 2 factorial design. Both fertilization and thinning increased basal area increment while only fertilization increased lateral branch growth in the crown. Contrary to expectations, fertilization decreased root starch reserves although it increased basal area increment; thinning had no effect on root carbohydrates. Root starch reserves were positively related to basal area increment and vigor index in both fertilized and unfertilized plots, however, the best predictors of starch reserves were height-to-live-crown and cambial surface area below the live crown; starch reserves declined as both of these variables increased. These data suggest that large carbon sinks associated with long boles below the crown, and rapid growth of the crown and stem as a result of fertilization are detrimental to building starch reserves in the roots.  相似文献   

11.
We examined 65 lodgepole pine (Pinus contorta Dougl. ex Loud. var. latifolia Engelm.) sites in Alberta using a dendrochronological approach in order to examine the relationships between climate and growth of lodgepole pine across elevational ecoregions and diameter size classes. The 4 elevational ecoregions sampled included the Boreal Highlands (BH: 13 sites); the Foothills (FH: 36 sites); a grouping of the montane and subalpine zones of the southern Rocky Mountains (RM: 12 sites); and the montane zone of the Cypress Hills (CH: 4 sites). The first diameter size class was comprised of the three largest (top) diameter trees at each site. The tree list of each plot was ranked, then divided into three groups of equal basal area and the tree at the midpoint of each group (small, medium, and large) was selected for the other three size classes. Annual growth in basal area of lodgepole pine was generally sensitive to heat and moisture stress in late summer of the previous year, the degree of winter harshness, and the timing of the start of the growing season. Growth was inhibited by low temperature in all winter months at the most northern BH sites which had the coldest winters, but this effect was interrupted in some of the midwinter months in the more southerly sites in the RM, and we postulated this was due to the damaging influences of Chinook winds. Interannual growth patterns were strongly correlated between top diameter trees and the other classes, and trees of all diameter classes generally responded to climate in the same way, which indicated that it is sufficient to sample only the largest diameter trees in a stand to provide insight into growth–climate relationships. The forecasted growth estimates indicate that future climate warming will negatively impact the productivity of lodgepole pine in the FH, the heart of lodgepole distribution and productivity in Alberta.  相似文献   

12.
Carbon isotope composition (delta13C) of branchlet tissue at nine canopy positions, and nitrogen concentration (N(mass)) at four canopy positions, were assessed in 8-year-old hoop pine (Araucaria cunninghamii Ait. ex D. Don) trees from 23 half-sib families, grown in six blocks of a progeny test in southeastern Queensland, Australia. There was considerable variation among sampling positions, families and blocks in both delta13C and N(mass). The delta13C was positively related to N(mass) only for samples from the upper outer crown (P < 0.005). Phenotypic correlations existed between tree growth and canopy delta13C. Branchlet delta13C of the inner and lower outer crown was positively related (P < 0.037) to tree height, but delta13C in branchlets of the upper outer crown was not related to tree height, or was related negatively (P < 0.045). There were significant differences in delta13C between hoop pine families for six canopy positions (upper canopy positions as well as lower canopy positions on the northern side), with heritabilities greater than 0.40. The significance of these findings is discussed in relation to water and light competition within the tree canopy of hoop pine.  相似文献   

13.
The community composition of Carabid beetles, some species ofwhich are known predators of pine beauty moth, was examinedwithin a Scottish plantation forest. Sites differing in soiltype and the species of trees planted were sampled with pitfalltraps in most weeks of a 3—year period. There were faunaldifferences between sites with lodgepole pine on deep peat andother sites in the study (lodgepole pine on iron-pan soil, speciesmixture of lodgepole and Scots pine, and pure stands of Scotspine). In general deep peat sites supported fewer species andindividuals of carabids. Three Carabus species were implicatedas likely predators of Panolis flammea pupae and each was lessabundant on the sites with lodgepole pine. It is suggested thatthe susceptibility to pine beauty moth of lodgepole pine growingon deep peat substrates is at least partly attributable to impoverishedpredator faunas.  相似文献   

14.
We investigated the effects of herbaceous and woody vegetation control on the survival and growth of planted eastern white pine (Pinus strobus L.) seedlings through six growing seasons. Herbaceous vegetation control involved the suppression of grasses, forbs, ferns, and low-shrubs, and was maintained for 0, 2, or 4 years after white pine seedlings were planted. Woody control involved the removal of all tall-shrub and deciduous trees, and was conducted at the time of planting, at the end of the second or fifth growing seasons, or not at all. Seedling height and basal diameter responded positively and proportionally to duration of herbaceous vegetation control. Gains associated with woody control were generally not significant unless some degree of herbaceous vegetation control was also conducted. Only herbaceous control increased pine crown closure and rate of crown closure. Herbaceous control and the presence of 5000–15,000 stems per ha of young overtopping aspen were associated with reduced weevil (Pissodes strobi Peck.) injury and increased pine height growth. The study suggests that white pine restoration strategies on clearcut sites should focus on the proactive, early management of understory vegetation and the gradual reduction of overtopping cover from woody vegetation to create a seedling light environment that supports acceptable growth with minimal weevil damage.  相似文献   

15.
Stemflow is a focused point source input of precipitation and nutrients at the base of a tree or plant and can have a significant impact on site hydrology. To date, no known studies have modelled stemflow production for juvenile lodgepole pine(Pinus contorta var. latifolia).Meteorological conditions, tree characteristics, and stemflow were sampled for two juvenile lodgepole pine stands over the course of the 2009 growing season. Step-wise multiple regression was used to assess which meteorological and tree architecture variables influenced stemflow production for each research plot. Once predictor variables were identified, models were produced for each stand and a generic model was produced that applied to both plots. A model employing precipitation depth and crown projection area successfully explained 71.3% of the variation in stemflow production from sampled trees. Stemflow was found to represent 1.8% of the study period rainfall and,although not a large component of the plot-scale canopy water balance, it is an order of magnitude greater than the fractioning of stemflow from mature lodgepole and lodgepole pine dominated forest. Additionally, stemflow funnelling ratios were found to average 22.2 and 24.3 from the two sample plots over the study period with a single tree, single event maximum of 111.7 recorded for a tree with a 3.3 cm bole diameter and a rain depth of 17.4 mm.  相似文献   

16.
Stemflow is a focused point source input of precipitation and nutrients at the base of a tree or plant and can have a significant impact on site hydrology. To date, no known studies have modelled stemflow production for juvenile lodgepole pine (Pinus contorta var. latifolia). Meteorological conditions, tree characteristics, and stemflow were sampled for two juvenile lodgepole pine stands over the course of the 2009 growing season. Step-wise multiple regression was used to assess which meteorological and tree architecture variables influenced stemflow production for each research plot. Once predictor variables were identified, models were produced for each stand and a generic model was produced that applied to both plots. A model employing precipitation depth and crown projection area successfully explained 71.3% of the variation in stemflow production from sampled trees. Stemflow was found to represent 1.8% of the study period rainfall and, although not a large component of the plot-scale canopy water balance, it is an order of magnitude greater than the fractioning of stemflow from mature lodgepole and lodgepole pine dominated forest. Additionally, stemflow funnelling ratios were found to average 22.2 and 24.3 from the two sample plots over the study period with a single tree, single event maximum of 111.7 recorded for a tree with a 3.3 cm bole diameter and a rain depth of 17.4 mm.  相似文献   

17.
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.  相似文献   

18.
Crown structure, absorbed photosynthetically active radiation (APAR) and growth were analyzed in 300 replicated loblolly (Pinus taeda L.) and slash pine (Pinus elliottii Engelm. var. elliotti) clones to: (1) quantify genetic variation in crown structural traits, growth and APAR at the species, family and clonal levels; and (2) estimate within-family genetic and environmental influences on measured variables. Species and family-within-species differences were found in some growth traits, crown size, leaf area, APAR and branch angle. Loblolly pine developed larger crowns, exposed more leaf area with an acute angle, and intercepted more radiation than slash pine. Significant differences among clones within-family were found for stem volume and crown architecture. Loblolly pine and slash pine within-family, individual-tree broad-sense heritabilities ranged from 0.00 to 0.41 for growth and crown structural traits and most were between 0.10 and 0.25 when estimated from a combined analysis across families. Genetic correlations of crown size, leaf area and APAR with volume increment generally ranged from 0.60 to 0.75. This knowledge of the genetic interactions among growth and crown structural traits improves our understanding of how crown morphology affects light interception and stand development, and ultimately how these attributes can be incorporated in the selection of families or clones for the development of new crop tree ideotypes.  相似文献   

19.
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   

20.

Rate of growth and wood properties of Sitka spruce (Picea sitchensis (Bong.) Carr.) were investigated in triplet mixtures with lodgepole pine (Pinus contorta Loud.) (Alaskan provenance) and Japanese larch (Larix kaempferi (Lamb.) Carr.), established on nitrogen-deficient, iron pan soils. These ?nursing? mixtures were compared with pure stands of Sitka spruce that had been either regularly or periodically fertilized with nitrogen. Japanese larch promoted a growth rate in Sitka spruce greater than that achieved in the lodgepole pine-nursed spruce and equivalent to the two pure spruce treatments over the duration of the experiment (current age 28 yrs). Growth of regularly fertilized pure Sitka spruce was not significantly greater than that of periodically fertilized pure spruce. Alaskan lodgepole pine controlled branch size on the lower part of the spruce stems more effectively than the other treatments, although this may have been a function of tree size. Branch characteristics of Japanese larch-nursed spruce, however, were similar to those of the pure spruce treatments. Japanese larch caused an imbalance in crown development in the spruce, although it is unclear from the present study whether this will have an influence on stem and wood quality by the end of the rotation. Overall, the evidence from this study suggests that Japanese larch is an effective nurse of Sitka spruce on nitrogen-deficient iron pans, maintaining a rate of growth similar to that of pure Sitka spruce periodically fertilized with nitrogen and higher than that observed in spruce nursed by Alaskan lodgepole pine.  相似文献   

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