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

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

Predicting mortality for five California conifers following wildfire

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

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.     

A crown profile model for Pinus radiata D. Don in northwestern Spain

A crown profile model was developed for radiata pine (Pinus radiata D. Don) in Galicia (northwestern Spain). Data from 443 trees located in 56 permanent plots, established by the Unidade de Xestión Forestal Sostible (UXFS) of the University of Santiago de Compostela in plantations of this species in the region, were used. The crowns of the trees were measured by a visual method based on similar triangles. Both simple geometric shapes and mathematical equations were used to describe the crown profiles. As crown profile models usually require variables that are expensive to measure, equations to estimate the maximum crown radius and the height to the maximum crown radius were also developed, using other easily measured tree and stand variables. Several models were fitted using a system of equations approach and accounting for an autocorrelated, heteroscedastic error structure. The selected crown profile model consists of a system of two allometric equations for the crown below (primarily shade needles) and above (primarily sun needles) maximum crown radius. The model explained 88% of the variability in crown radius with a mean error of 0.24 m.     

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.     

Soil bacterial functional diversity is associated with the decline of Eucalyptus gomphocephala

This study investigates whether tree decline in Eucalyptus gomphocephala (tuart) is associated with the functional diversity of soil bacterial communities. We selected 12 sites with different stages of decline and assessed crown health [Crown density (CD), Foliage transparency (FT), Uncompacted live crown ratio (ULCR), Crown dieback ratio (CDR) and Epicormic index (EI)] and soil bacterial functional diversity based on Biolog EcoPlates™ incubation [Average well colour development (AWCD), Shannon diversity (H′), richness (S) and Shannon evenness (E)]. Crown health indices differed between sites with EI being the most robust indicator of decline in crown health followed by CDR and CD (P < 0.05). Soil bacterial indices collected at 0–10 and 20–30 cm soil depth between December (summer, dry season) and May (autumn, start of wet season) differed between sites (P < 0.05), and significant relationships between crown health indices, except ULCR, and all soil bacterial indices were observed. Principle component analysis (PCA) showed that a decrease in the utilization of carbohydrates, carboxylic acids, amino acids and amines by the soil bacterial communities correlated to sites with poor crown health, indicating some changes in physiological responses of bacterial groups with declining tree health. Using stepwise regression analyses, in the 0–10 cm soil layer in December, itaconic acid had a 46% contribution to the EI. Carboxylic acids, including itaconic acid, have a strong ability to solubilize soil minerals in calcareous soil, and these possibly increased the availability of soil mineral nutrients in the healthier sites compared to the declining sites, particularly in the dry season. In addition, lack of soil water in the declining sites limited soil bacterial diversity and was positively correlated with EI in the 0–10 cm soil layer in December. In conclusion, soil bacterial functional diversity has a strong relationship with tuart decline and the importance of soil microbes in tuart ecosystem health must be considered in the future.     

Snow damage in lodgepole pine stands brought into thinning and fertilization regimes

Several heavy wet snowfalls occurred during 2007-2009 across a broad-scale thinning and fertilization experiment to bring overstocked juvenile lodgepole pine (Pinus contorta var. latifolia) in the foothills of Alberta, Canada into an intensive management regime. We examined the bending and breakage of trees in relation to thinning and fertilization and used a multimodel information-theoretic approach to model stand and tree level predictors of snow damage. Fertilized stands suffered the greatest amount of snow damage, and this was most noteworthy when stands were also thinned; here 22% (17% broken stems) of trees were damaged compared to 8% (4% broken stems) in the thinned and unfertilized stands. At the stand level, needle weight and crown cover were reliable predictors of snow damage. At the tree level, separate models were developed for each combination of thinning and fertilization. All models used total tree volume; usually the smaller trees in the stands were more susceptible to damage but in the thinned and fertilized stands larger but slender trees with large asymmetrical crowns tended to be damaged. Also, trees with lower total stem volume were more susceptible to damage. Only in the thinned and fertilized stands were variables related to crown shape and asymmetry important predictors of snow damage. We conclude that snow damage is an important agent for self-thinning in unthinned stands and fertilization tends to exacerbate damage because of increase in foliage size. In areas with regular occurrence of heavy snow, we do not recommend fertilization at the same time as thinning, as the larger and more economically important trees in the stand are at risk.     

Effects of planting spacing and site quality on 25-year growth and mortality relationships of Douglas-fir (Pseudotsuga menziesii var. menziesii)

Growth and mortality of coast Douglas-fir (Pseudotsuga menziesii var. menziesii) were studied for 25 years after planting seedlings at 1–6-m spacings on a site of moderate quality in the western Cascade Mountains of Washington. Responses were compared to those from two other studies representing high and low site qualities. Third-year height did not differ among spacings (P = 0.80), providing no evidence that close spacing stimulated early growth. Piecewise regression identified the onset of competition-induced mortality when stand density index (SDI [Reineke, L.H. 1933. Perfecting a stand density index for even-aged forests. Journal of Agricultural Research 46, 627–638]) exceeded 52% (S.E. = 4.6) of the species’ maximum or when average crown ratio (CR) declined below 52% (S.E. = 0.9). For a range of SDI values, CR averaged 2–7% points greater at the high-quality site than at the moderate-quality site. In a regression analysis of combined data from the moderate- and high-quality sites, relative values of average stem diameter and stand volume (% of maximum values observed per site) 23–25 years after planting increased and decreased with planting spacing, respectively (R² = 0.97 and 0.91, respectively). Intersection of these relationships at 3-m spacing indicated a point of equivalent relative development of tree size and stand yield. For a range of site qualities, stands planted at 3-m spacing: (1) maintained tree vigor (CR ≥ 50%) and stability (average height:dbh ratio <90), (2) experienced little or no competition-induced mortality through age 25 years, and (3) allocated 25-year growth equitably to development of tree size and stand yield, thereby providing a desirable starting point for subsequent management.     

Tree regeneration and future stand development after bark beetle infestation and harvesting in Colorado lodgepole pine stands

In the southern Rocky Mountains, current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreaks and associated harvesting have set millions of hectares of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) forest onto new stand development trajectories. Information about immediate, post-disturbance tree regeneration will provide insight on dynamics of future stand composition and structure. We compared tree regeneration in eight paired harvested and untreated lodgepole pine stands in the Fraser Experimental Forest that experienced more than 70% overstory mortality due to beetles. New seedlings colonized both harvested and untreated stands in the first years after the beetle outbreak. In harvested areas the density of new seedlings, predominantly lodgepole pine and aspen, was four times higher than in untreated stands. Annual height growth of pine and fir advance regeneration (e.g., trees established prior to the onset of the outbreak) has doubled following overstory mortality in untreated stands. Growth simulations based on our regeneration data suggest that stand basal area and stem density will return to pre-beetle levels in untreated and harvested stands within 80-105 years. Furthermore, lodgepole pine will remain the dominant species in harvested stands over the next century, but subalpine fir will become the most abundant species in untreated areas. Owing to terrain, economic and administrative limitations, active management will treat a small fraction (<15%) of the forests killed by pine beetle. Our findings suggest that the long-term consequences of the outbreak will be most dramatic in untreated forests where the shift in tree species composition will influence timber and water production, wildfire behavior, wildlife habitat and other forest attributes.     

Lodgepole pine growth as a function of competition and canopy light environment within aspen dominated mixedwoods of central interior British Columbia

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 (DIFN_t) 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 DIFN_t versus height growth indicated the response to be linear across the observed range of available light. Implications for management are discussed.     

Quantifying branch,crown and bole development in Populus tremuloides Michx. from north-eastern British Columbia

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.     

Wind speed and crown class influence the height–diameter relationship of lodgepole pine: Nonlinear mixed effects modeling

Wind-induced mechanical perturbation is one of the mechanisms determining the height–diameter relationship of trees, but its effects have largely been neglected in past height–diameter models. In this study, we examined the effects of including wind speed as a regressor in the height–diameter model for lodgepole pine (Pinus contorta var. latifolia Engelm.). We also tested the hypothesis that the height–diameter relationship differs between trees of different crown classes (dominant and subordinate trees), and that a mixed effects height–diameter model developed at the crown class level should achieve a better prediction than that developed at the plot level. The results showed that including wind speed significantly improved the fit of the height–diameter model. The results also showed that the height–diameter relationships differ significantly between the two crown classes. The mixed effects model developed based on crown classes of individual trees had better fit with reduced residual variance and decreased Akaike's information criterion and the Schwarz's Bayesian information criterion than the plot-based mixed model. Evaluation of the developed model suggests the crown class-based height–diameter model has better prediction in terms of prediction error and precision. The developed crown class-based mixed effects model can be used to provide more accurate prediction of tree heights for lodgepole pine from their diameters.     

Carbon accumulation along a stand development sequence of Nothofagus antarctica forests across a gradient in site quality in Southern Patagonia

Above- and below-ground C pools were measured in pure even-aged stands of Nothofagusantarctica (Forster f.) Oersted at different ages (5–220 years), crown and site classes in the Patagonian region. Mean tissue C concentration varied from 46.3% in medium sized roots of dominant trees to 56.1% in rotten wood for trees grown in low quality sites. Total C concentration was in the order of: heartwood > rotten wood > sapwood > bark > small branches > coarse roots > leaves > medium roots > fine roots. Sigmoid functions were fitted for total C accumulation and C root/shoot ratio of individual trees against age. Total C accumulated by mature dominant trees was six times greater than suppressed trees in the same stands, and total C accumulated by mature dominant trees grown on the best site quality was doubled that of those on the lowest site quality. Crown classes and site quality also affected the moment of maximum C accumulation, e.g. dominant trees growing on the worse site quality sequestered 0.73 kg C tree⁻¹ year⁻¹ at 139 years compared to the best site where 1.44 kg C tree⁻¹ year⁻¹ at 116 years was sequestered. C root/shoot ratio decreased over time from a maximum value of 1.3–2.2 at 5 years to a steady-state asymptote of 0.3–0.7 beyond 60 years of age depending on site quality. Thus, root C accumulation was greater during the regeneration phase and for trees growing on the poorest sites. The equations developed for individual trees have been used to estimate stand C accumulation from forest inventory data. Total stand C content ranged from 128.0 to 350.9 Mg C ha⁻¹, where the soil C pool represented 52–73% of total ecosystem C depending on age and site quality. Proposed equations can be used for practical purposes such as estimating the impact of silvicultural practices (e.g. thinning or silvopastoral systems) on forest C storage or evaluating the development of both above- and below-ground C over the forest life cycle for different site qualities for accurate quantification of C pools at regional scale.     

Potential effects of climate change on the growth of lodgepole pine across diameter size classes and ecological regions

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.     

Impacts of thinning on structure,growth and risk of crown fire in a Pinus sylvestris L. plantation in northern Spain

We studied the combined effects of thinning on stand structure, growth, and fire risk for a Scots pine thinning trial in northern Spain 4 years following treatment. The thinning treatments were: no thinning, heavy thinning (32–46% of basal area removed) and very heavy thinning (51–57% of basal area removed). Thinning was achieved via a combination of systematic and selective methods by removing every seventh row of trees and then by cutting suppressed and subdominant trees in the remaining rows (i.e., thinning from below). Four years after thinning, mean values and probability density distributions of stand structural indices showed that the heavier the thinning, the stronger the tendency towards random tree spatial positions. Height and diameter differentiation were initially low for these plantations and decreased after the 4-year period in both control and thinned plots. Mark variograms indicated low spatial autocorrelation in tree diameters at short distances. Diameter increment was significantly correlated with the inter-tree competition indices, and also with the mean directional stand structural index. Two mixed models were proposed for estimating diameter increment using a spatial index based on basal area of larger trees (BALMOD) in one model versus spatial competition index by Bella in the other model. As well, a model to estimate canopy bulk density (CBD) was developed, as this variable is important for fire risk assessment. Both heavy and very heavy thinning resulted in a decrease of crown fire risk over no thinning, because of the reduction in CBD. However, thinning had no effect on the height to crown base and thus on the flame length for torching. Overall, although thinning did not increase size differentiation between trees in the short term, the increase in diameter increment following thinning and the reduction of crown fire risks support the use of thinning. Also, thinning is a necessary first step towards converting Scots pine plantations to more natural mixed broadleaved woodlands. In particular, the very heavy thinning treatment could be considered a first step towards conversion of overstocked stands.     

The contribution of structural indices to the modelling of Sitka spruce (Picea sitchensis) and birch (Betula spp.) crowns

Crown dimensions are important for the quantification of tree interactions in some growth models. This study investigates the potential for structural indices and other spatial measures to improve the prediction of crown radius and crown length for birch (Betula spp.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) in forests in Wales. Crown dimensions were measured for 125 birch and 154 spruce in six fully stem-mapped research plots. These data were used to test the performance of a crown radius model and a crown length model which estimated crown dimensions on the basis of allometric relationships with stem dimensions. Spatial data from the six plots were used to calculate the structural indices mean directional index, diameter correlation index, species mingling, dbh and height dominance, and dbh differentiation, as well as the Hegyi competition index, and basal area of neighbours and larger neighbours, for each crown measurement sample tree, using various numbers of nearest neighbours. Two non-spatial indices, BAL and BALMOD, were also calculated for all sample trees for comparison. These spatial and non-spatial variables were then incorporated into modified crown dimension models. Model performances, in terms of efficiency and relative bias, were compared to determine whether the inclusion of spatial or non-spatial variables resulted in any improvements over models using tree dimensions alone. Crown length and radius were found to be correlated with most of the spatial measures studied. Models incorporating spatial variables gave improvements in performance over allometric models for every data set, and performed more consistently than models containing non-spatial variables. The greatest improvements were achieved for suppressed birch in unthinned forests which had irregularly shaped and strongly displaced crowns. The spatial variable contributing to the most efficient model for each data set varied widely. This points to the complexity of tree spatial interactions and indicates that there is a great deal of scope for investigating other structural indices and crown dimension model forms.     

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

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

Impact of stand density on water status and leaf gas exchange in Quercus ilex

Tree thinning reduces tree-to-tree competition and likely contributes to the improvement of tree water status and productivity in water-limited systems. In this study, we examined the importance of competition for water among Quercus ilex trees in open woodlands by comparing the water consumption and physiological status of trees located along stand density gradients which ranged from 10% (low density; LD) to 100% (high density; HD) of canopy cover. The study was carried out at two sites which differed in mean annual rainfall (506 and 816 L m⁻²; D_site and W_site, respectively). Predawn and midday leaf water potential (ψ_d and ψ_m, respectively) and CO₂ assimilation rate (A) were measured every two weeks from mid May to mid September, in eight trees located along a stand density gradient at each site. Sap flow and soil moisture were measured only at D_site. Sap flow was continuously recorded by sap flowmeters (constant heating method) installed in 12 trees along two stand density gradients. Soil moisture (?) was measured every 20 cm for the first meter and then every 50 cm up to 250 cm. Measurements were conducted in 18 soil profiles, 6 located in HD and 12 in LD (six beneath and six out the canopy). At W_site, differences among stand densities for ψ and A were very small and emerged only at the end of the dry season. At D_site, ψ (both predawn and midday), A, ?, and sap flow density were significantly higher in LD trees than in HD ones. At D_site, some water remained unused in the soil at the end of the dry season beyond the canopy in the LD areas, and trees did not experienced such an acute water deficit (ψ_d > −1 MPa) as the HD trees did (ψ_d < −3 MPa). Summer tree transpiration at the stand level (E_stand) tended to saturate with the increase of canopy cover. E_stand increases by 32% when canopy cover goes from 50% to 100%. Results confirmed that the increase of tree-to-tree competition with stand density was much more significant at dry sites. In these sites, tree thinning is recommended as a way to maintain tree functioning.     

Modelling stemflow production by juvenile lodgepole pine (<Emphasis Type="Italic">Pinus contorta</Emphasis> var. <Emphasis Type="Italic">latifolia</Emphasis>) trees

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.     

Effects of competition and climate variables on modelling height to live crown for three boreal tree species in Alberta,Canada

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.