Crown plasticity in mixed forests—Quantifying asymmetry as a measure of competition using terrestrial laser scanning

Interspecific competition is a key process determining the dynamics of mixed forest stands and influencing the yield of multispecies tree plantations. Trees can respond to competitive pressure from neighbors by crown plasticity, thereby avoiding competition. We employed a high-resolution ground-based laser scanner to analyze the 3-dimensional extensions and shape of the tree crowns in a near-natural broad-leaved mixed forest in order to quantify the direction and degree of crown asymmetry of 15 trees (Fagus sylvatica, Fraxinus excelsior, Carpinus betulus) in detail. We also scanned the direct neighbors and analyzed the distance of their crown centres and the crown shape with the aim to predict the crown asymmetry of the focal tree from competition-relevant attributes of its neighbors. It was found that the combination of two parameters, one summarizing the size of the neighbor (DBH) and one describing the distance to the neighbor tree (HD), was most suitable for characterizing the strength of the competitive interaction exerted on a target tree by a given neighbor. By summing up the virtual competitive pressure of all neighbors in a single competitive pressure vector, we were able to predict the direction of crown asymmetry of the focal tree with an accuracy of 96° on the full circle (360°).The competitive pressure model was equally applicable to beech, ash and hornbeam trees and may generate valuable insight into competitive interactions among tree crowns in mixed stands, provided that sufficiently precise data on the shape and position of the tree crowns is available. Multiple-aspect laser-scanning proved to be an accurate and practicable approach for analyzing the complex 3-dimensional shape of the tree crowns, needed to quantify the plasticity of growth processes in the canopy. We conclude that the laser-based analysis of crown plasticity offers the opportunity to achieve a better understanding of the dynamics of canopy space exploration and also may produce valuable advice for the silvicultural management of mixed stands.     

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.     

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.     

Growth responses of three coexisting conifer species to climate across wide geographic and climate ranges in Yukon and British Columbia

This work aimed to compare radial growth–climate relationships among three coexisting coniferous tree species across a wide geographic and climate range from southern British Columbia (BC) to central Yukon, Canada. Tree-ring data were collected from 20 mature stands of white spruce (Picea glauca), lodgepole pine (Pinus contorta var. latifolia), and subalpine fir (Abies lasiocarpa). Linear relationships between annual growth variation and monthly and seasonal climate were quantified with correlation and regression analyses, and variation in climate–growth responses over a climatic gradient were quantified by regressing growth responses against local mean climatic conditions. Temperatures had more consistent and stronger correlations with growth for all three species than precipitation, but growth–climate responses varied among species and among sites. In particular, pine and fir populations showed different responses between BC and Yukon, whereas spruce showed a more consistent response across the study domain. Results indicate that (1) the response and sensitivity of trees to seasonal climate variables vary among species and sites and (2) winter temperatures prior to growth may have significant impacts on pine and fir growth at some sites. The capacity to adapt to climate change will likely vary among the study species and across climatic gradients, which will have implications for the future management of mixed-species forests in Yukon and BC.     

The effects of belowground resources on aboveground allometric growth in Bornean tree species

Tree height and crown allometries reflect adaptations for resource acquisition and structural stability, as well as plastic responses to a heterogeneous environment. While both light and soil resources limit growth and influence competitive responses in tropical forests, the effects of belowground resources on allometries are less understood, especially within the understory. To characterize outcomes of tree competition along an edaphic resource gradient, we quantified variation in height and crown allometries of six Bornean tree species from contrasting regeneration niches (light-demanding vs. shade-tolerant) on two soil habitats (clay-fine loam and sandy loam) within a 52-ha forest dynamics plot. Using empirically-fit allometric parameters and diameter growth rates from plot census data, we modeled tree height and crown area growth over the projected life span of each species. Based on resource competition theory, we hypothesized that tree species specializing on and populations of generalist species growing on the relatively moister, more fertile clay-fine loam soil habitat would have faster height and crown growth rates, compared to those on the sandy loam habitat, regardless of regeneration niche. Among soil specialists and within generalists of both genera, trees growing on clay-fine loam had taller stems and larger crowns at a given age and faster height and crown area growth rates at most sizes than trees on sandy loam. Differences in height and crown growth were driven by the faster diameter growth rates of trees on clay-fine loam, not by differences in height- and crown-diameter allometries, as trees on sandy loam were significantly taller at a given diameter, and differences in crown allometry were not consistent across soil habitats. Characterizing the height and crown growth responses of trees along resource gradients provides insight into the mechanisms that maintain diversity in tropical forests. Our results point to the importance of adaptive and plastic responses to both above and belowground resource availability in determining the allometric growth of trees and suggest that this diversity of responses may contribute tree species coexistence through competition-based trade-off mechanisms and variation in growth among individuals. Additionally, as the importance estimating natural carbon sequestration increases with the escalating effects of anthropogenic climate change, differences in tree growth and architecture across soil habitats also have implications for the approximation of forest carbon storage on heterogeneous tropical soils.     

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.     

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.     

An empirical model of crown shyness for lodgepole pine (Pinus contorta var. latifolia [Engl.] Critch.) in British Columbia

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 (A_V) using stem maps and Voronoi polygons constrained by estimates of maximum crown width, and then related observed crown area (A_C) to A_V 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.     

Five-year radial growth of red oaks in mixed bottomland hardwood stands

We studied the relationships among 5-year radial (diameter and basal area) growth of red oak (genus Quercus, subgenus Erythrobalanus) crop trees and predictor variables representing individual tree vigor, distance-dependant competition measures, and distance-independent competition measures. The red oaks we examined are representative of the commercially and ecologically important oak species of the bottomland hardwood forests of the southeastern US. The crown class score, a quantitative measure of crown class and tree vigor, performed best in accounting for the variability in tree diameter growth. Plot-level variables failed to account for a significant proportion of the variability in tree radial growth. The basal area of the first-order neighbors that were taller than the crop trees and located within 2.4 times the mean overstory crown radius had the highest negative correlation with crop tree 5-year radial growth. Red oaks were a major part of these competitors and likely exerted the greatest competitive pressure. However, crop tree radial growth was positively associated with the basal area of the red oaks which were indirect (second order) neighbors and which were taller than the crop trees. It is possible that indirect neighbors do not compete with the crop trees, but they likely compete with the direct competitors of the crop trees, thus having an indirect positive influence on crop tree growth. Such reasoning is consistent with previously observed spatial dependence up to four times the mean overstory crown radius. The findings may have implications for thinning hardwoods stands and crop tree management in that foresters need to take into account (1) oak intra-genus competition, (2) the negative competitive effect of direct neighbors, and (3) the potentially positive effect of the indirect neighbors, the competitors’ competitors.     

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.     

Evaluation of competition and light estimation indices for predicting diameter growth in mature boreal mixed forests

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.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

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.     

Evaluation of competitive effects of green alder,willow and other tall shrubs on white spruce and lodgepole pine in Northern Alberta

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.     

Height growth of planted conifer seedlings in relation to solar radiation and position in Scots pine shelterwood

Seedlings of different provenances of Scots pine (Pinus sylvestris L.), lodgepole pine (Pinus contorta Dougl., var. latifolia Engelm.) and Norway spruce (Picea abies (L.) Karst.) were planted in three Scots pine shelterwoods (125, 65 and 43 stems ha⁻¹) and a clear-cut, all in northern Sweden. The sites were mounded and planting took place during 2 consecutive years (1988 and 1989). The solar radiation experienced by the individual seedlings was determined using a simulation model. Height development of the seedlings was examined during their first 6 years after planting. During the final 3 years of the study, height growth of Norway spruce was relatively poor, both in the shelterwoods and the clear-cut area. Height growth of lodgepole pine was significantly greater than that of Scots pine, both in the shelterwoods and the clear-cut. In contrast to Norway spruce, Scots pine and lodgepole pine displayed significantly greater height growth in the clear-cut than in the shelterwoods. For all three species in the shelterwoods, regression analyses showed that height growth was more strongly correlated with the distance to the nearest tree than with the amount of radiation reaching the ground, i.e. growth was reduced in the vicinity of shelter trees. Therefore, we conclude that the significant reduction in height growth of seedlings of Scots pine and lodgepole pine in Scots pine shelterwoods was partially caused by factors associated with the distance to the nearest shelter tree. Because the substrate was a nitrogen-poor sandy soil, we suggest that root competition for mineral nutrients, especially nitrogen, accounts for the reduction in height growth.     

Species differences in total and vertical distribution of branch- and tree-level leaf area for the five primary conifer species in Maine, USA

Vertical distribution of leaf area largely governs both tree structure and function. Models of this important tree attribute have been constructed for several commercially important conifers. However, a limited number of studies have compared alternative modeling techniques and inherent species differences. This study used several existing datasets for the five primary conifer species in Maine, namely balsam fir [Abies balsamea (L.) Mill.], northern white-cedar [Thuja occidentalis (L.)], eastern hemlock [Tsuga canadensis (L.) Carr.], eastern white pine [Pinus strobus (L.)], and red spruce [Picea rubens (Sarg.)] to examine species variation in total and vertical distribution of projected leaf area at the individual branch- and tree-levels. In addition, multiple methods for modeling the vertical distribution of leaf area were examined across the species. For a given branch diameter and location within the crown, eastern hemlock branches held the greatest amount of leaf area, followed by balsam fir, northern white-cedar, white pine, and red spruce. At the tree-level, eastern white pine held the greatest amount of leaf area followed by eastern hemlock, balsam fir, red spruce, and northern white-cedar for a given tree size. Across species, the two-parameter, right-truncated Weibull distribution performed the best for predicting vertical distribution of leaf area when compared to the four-parameter beta and Johnson's S_B distributions (reduction of root mean square error of 1.7–21.1%). Northern white-cedar had a relative distribution of leaf area distinctly different than other species in this study with a mode shifted towards the upper crown. In contrast to red spruce and white pine, the mode of the relative distribution of leaf area for balsam fir and eastern hemlock occurred lower in the crown. Results of this study suggest that differences in total and vertical distribution of leaf area exist between species, but significant amounts of their variation are largely accounted for by bole and crown size.     

Recruitment limitation in forests: Lessons from an unprecedented mountain pine beetle epidemic

Since the mid-1990s the forests of central British Columbia have undergone an unprecedented Mountain Pine Beetle (Dendroctonus ponderosae Hopkins) (MPB) epidemic that has resulted in extensive mortality of canopy lodgepole pine (Pinus contorta var. latifolia Engelm.). This study investigated how seed-source availability, seedbed substrate, overstory structure, and time since MPB attack interact to affect post-MPB seedling recruitment of the dominant tree species of these forests. In addition to post-MPB recruitment, these forests may be regenerated by trees established in the understory prior to MPB disturbance. Accordingly, we examined abundance and patterns of all regeneration less than 130 cm tall. We found post-MPB recruitment was sparse. Subalpine fir (Abies laciocarpa (Hook.) Nutt.) comprised the majority of the post-MPB recruitment. It increased with local parent tree basal area and increased strongly with proximity to a major seed source. This resulted in a patchy distribution for subalpine fir post-MPB regeneration. Lodgepole pine post-MPB recruitment was limited by overstory shading. Recruitment of pine decreased as the total overstory basal area increased. Interior spruce (Picea glauca × engelmannii) post-MPB recruitment was similarly limited by total overstory basal area. Seedbed substrates were uniform and dominated by moss. Substrate type distribution did not change as time since MPB disturbance increased. The overall low post-MPB recruitment observed was likely due to a lack of disturbance to the moss-dominated forest floor. Moss is known to be a poor substrate in northern forests. The distribution of all regeneration less than 130 cm tall showed the same trends as the post-MPB regeneration. We believe the post-MPB seedling recruitment dynamics of these forests was not substantially changed from conditions prior to MPB disturbance. There was no pulse of regeneration up to 10 years post-MPB disturbance. Unless this changes, future stand structure will be dominated by the seedling bank established prior to the MPB epidemic. Subalpine fir dominated the seedling bank (68%) and post-MPB recruitment (94%). This suggests that MPB-disturbed forests are undergoing a substantial shift in landscape-level species composition.     

Modelling primary branch frequency and size for five conifer species in Maine,USA

Quantitative models of crown structure have been developed for several conifer species, but these studies have rarely simultaneously fit the models across multiple species. This study used extensive crown structure data for the five primary conifer species in Maine to test for species differences in maximum branch diameter profile, branch density, and relative branch diameter distribution. The species included balsam fir [Abies balsamea (L.) Mill], northern white-cedar [Thuja occidentalis (L.)], eastern hemlock [Tsuga canadensis (L.) Carr.], eastern white pine [Pinus strobus (L.)], and red spruce [Picea rubens (Sarg.)]. After accounting for key covariates, significant species differences were found in all crown structural attributes examined in this study. Profiles for the mean tree indicated that northern white-cedar had the smallest maximum branch diameters throughout the crown and white pine had the largest, except near the base of the crown where the species switched in rank. The density of live branches in a crown had the widest range of variation of the examined crown structural attributes. Red spruce had a significantly higher density of primary branches than the other conifers, particularly in the upper crown. The relative branch diameter distribution indicated that balsam fir had a distribution more skewed towards larger relative branch sizes, while eastern hemlock and red spruce had distributions shifted towards smaller relative branches. This study highlights the range of variability in key crown structural attributes due to inherent species differences, but indicates that models fit across multiple species can perform quite well as the amount of explained variation was relatively high.     

Pathogenicity of Phytophthora species and Pythium undulatum isolated from Abies procera Christmas trees in Ireland

Phytophthora cryptogea, Phytophthora cinnamomi, Phytophthora cambivora, Phytophthora megasperma and Pythium undulatum were isolated from diseased Noble fir (Abies procera) seedlings and soil associated with dead Noble fir in Ireland. Seedlings of four Christmas tree species (A. procera, Picea sitchensis, Picea abies and Pinus contorta) were inoculated with these oomycetes to test their pathogenicity and the susceptibility of the various tree species. Phytophthora spp. and Pythium undulatum caused root rot on all tree species. Disease symptoms included reddish brown cambial discoloration, crown symptoms, brown foliage, dark brown roots, root rot and seedling mortality. These symptoms were similar to those observed on Noble fir in naturally infested plantations. Pythium undulatum appeared as the most virulent pathogen followed by P. cinnamomi, P. cambivora, P. megasperma and P. cryptogea. Noble fir showed to be most susceptible and lodgepole pine most tolerant while Sitka spruce and Norway spruce were intermediate.