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.     

Assessing the impacts of species composition, top height and density on individual tree height prediction of quaking aspen in boreal mixedwoods

In this study we developed an individual tree height prediction model for quaking aspen (Populus tremuloides Michx.) grown in boreal mixedwood forests in Alberta using the nonlinear mixed model (NLMM) approach. We examined the impacts of density, species composition, and top height on aspen height predictions. Statistically significant stand level variables were incorporated into the base height–diameter model to increase the predictive ability and accuracy of the model at both the population and subject-specific levels. Our analyses showed that top height and density impacted height growth, but species composition did not. More importantly, we found that the inclusion of additional variables into the base model, despite improving model fitting statistics on the modelling data, did not improve the model's predictive ability and accuracy when cross-validated and when tested on an independent testing data set. Under the NLMM framework the base model performed as well as or better than the expanded models that contained other stand level variables. This has important theoretical and practical implications because, other than for biological reasons, more accurate local tree height predictions for aspen can be achieved simply by using the base height–diameter model fitted with the NLMM approach without the inclusion of other variables.     

Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta

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

Leaf respiratory acclimation to climate: comparisons among boreal and temperate tree species along a latitudinal transect

In common gardens along an ～900 km latitudinal transect through Wisconsin and Illinois, U.S.A., tree species typical of boreal and temperate forests were compared with respect to the nature and magnitude of leaf respiratory acclimation to contrasting climates. The boreal representatives were trembling aspen (Populus tremuloides Michx.) and paper birch (Betula papyrifera Marsh.), while the temperate species were eastern cottonwood (Populus deltoides Bartr ex. Marsh var. deltoides) and sweetgum (Liquidambar styraciflua L.). Assessments were conducted on seedlings grown from seed sources collected near southern and northern range boundaries, respectively. Nighttime rates of leaf dark respiration (R(d)) at common temperatures, as well as R(d)'s short-term temperature sensitivity (energy of activation, E(o)), were assessed for all species and gardens twice during a growing season. Little evidence of R(d) thermal acclimation was observed, despite a 12 °C range in average air temperature across gardens. Instead, R(d) variation at warm temperatures was linked most closely with prior leaf photosynthetic performance, while R(d) variation at cooler temperatures was most strongly related to leaf nitrogen concentration. Moreover, E(o) differences across species and gardens appeared to stem from the somewhat independent limitations on warm versus cool R(d). Based on this construct, an empirical model relying on R(d) estimates from leaf photosynthesis and nitrogen concentration explained 55% of the observed E(o) variation.     

Effects of soil temperature on biomass production and allocation in seedlings of four boreal tree species

One-year old seedlings of trembling aspen (Populus tremuloides Michx.), black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss), and jack pine (Pinus banksiana Lamb.) were subject to seven soil temperatures (5, 10, 15, 20, 25, 30 and 35 °C) for 4 months. All aspen seedlings, about 40% of jack pine, 20% of white spruce and black spruce survived the 35 °C treatment. The seedlings were harvested at the end of the fourth month to determine biomass and biomass allocation. It was found that soil temperature, species and interactions between soil temperature and species significantly affected root biomass, foliage biomass, stem biomass and total mass of the seedling. The relationship between biomass and soil temperature was modeled using third-order polynomials. The model showed that the optimum soil temperature for total biomass was 22.4, 19.4, 16.0 and 13.7 °C, respectively, for jack pine, aspen, black spruce and white spruce. The optimum soil temperature was higher for leaf than for root in jack pine, aspen and black spruce, but the trend was the opposite for white spruce. Among the species, aspen was the most sensitive to soil temperature: the maximum total biomass for aspen was about 7 times of the minimum value while the corresponding values were only 2.2, 2.4 and 2.3 times, respectively, for black spruce, jack pine and white spruce. Soil temperature did not significantly affect the shoot/root (S/R) ratio, root mass ratio (RMR), leaf mass ratio (LMR), or stem mass ratio (SMR) (P>0.05) with the exception of black spruce which had much higher S/R ratios at low (5 °C) and high (30 °C) soil temperatures. There were significant differences between species in all the above ratios (P<0.05). Aspen and white spruce had the smallest S/R ratio but highest RMR while black spruce had the highest S/R but lowest RMR. Jack pine had the highest LMR but lowest SMR while aspen had the smallest LMR but highest SMR. Both LMR and SMR were significantly higher for black spruce than for white spruce.     

Trembling aspen root suckering and stump sprouting response to above ground disturbance on a reclaimed boreal oil sands site in Alberta,Canada

New Forests - Trembling aspen (Populus tremuloides Michx.) is an important early successional species in the boreal region that commonly regenerates via root suckering and, to a lesser extent,...     

Pests,climate and competition effects on survival and growth of trembling aspen in western Canada

New Forests - Trembling aspen (Populus tremuloides Michx.) is experiencing increased drought stress resulting from climate change together with increases in damage by forest tent caterpillar and...     

Effects of species and tree size diversity on recruitment in the Alaskan boreal forest: A geospatial approach

This study empirically evaluates and maps the relationships between recruitment and species and tree size diversity, as measured with the Shannon’s index, within mixed poplar/birch and mixed spruce stands across the boreal forest of Alaska. Data were collected from 438 permanent sample plots re-measured at a 5-year interval. Significant explanatory factors of recruitment, including species and tree size diversity were first identified using hierarchical partitioning. The effects of tree diversity on recruitment were then studied using generalized linear models and universal kriging to account for non-spatial factors and for spatial autocorrelation. We found a consistent positive relationship between recruitment and species diversity and a general negative relationship between recruitment and tree size diversity, indicating a tradeoff between species diversity and tree size diversity in affecting recruitment. These relationships however were not uniform across the landscape, presumably because they were subject to strong spatial autocorrelation attributable to natural disturbances and environmental stressors. In general, diversity had least effect on recruitment in stressful environments where stress, rather than competition, most likely governed recruitment.     

Resin pocket occurrence in Norway spruce depending on tree and climate variables

Resin or pitch pockets cause a significant reduction in timber quality of Norway spruce (Picea abies (L.) Karst). Their occurrence is very variable within the stem and between trees and stands. Researchers have suggested several causes of resin pocket formation with a majority arguing in favour of storms as the main causing factor.     

树高、胸径、树皮厚对厚朴产量的影响

为研究厚朴树高、胸径和树皮厚度对其产量的影响,从而为厚朴品种选优及丰产栽培提供理论指导。采用系统抽样方法抽取了10~14 a的90个厚朴单株样本,测量其单株树高、胸径、树皮厚和树皮产量,通过建立和分析厚朴产量回归方程可知,胸径、树高和树皮厚与产量均成正相关关系;胸径、树高、树皮厚对产量的直接影响从大到小依次为胸径、树高、树皮厚,其中胸径对产量的影响以直接影响为主,树高对产量的间接影响稍高于直接影响,其间接影响主要是通过胸径对产量产生影响,树皮厚主要通过胸径和树高对产量产生间接影响,直接影响较小。     

Relationships between climate and tree radial growth in interior British Columbia,Canada

Climate is a main driving factor of the formation of annual tree-rings, but which climatic variables are the most influential on radial growth may vary among species and sites. To explore these interactions, tree-ring chronologies along a major elevation gradient were examined for three tree species in southern interior British Columbia (Canada): Pseudotsuga menziesii, Pinus contorta, and Picea glauca × engelmannii. We used correlations and linear and multiple regressions to explore the relationships between tree-ring radial growth and climate variables in the area from 1922 to 1997. All correlation coefficients between ring chronologies and monthly climatic variables were medium to low (from −0.3 to 0.4); nevertheless, moderate but significant trends could be identified. Multivariate models explained up to 53%, 43% and 32% of radial growth variability for P. contorta, P. menziesii and P. glauca × engelmannii, respectively. All three species showed similar radial growth–climate patterns across the elevational gradient, but they had different details that made ring width–climate relationships species-specific. Precipitation-related variables were more related to radial growth at low-elevations, changing into temperature-related variables at high-elevations. Tree-ring width for all three species was primarily and significantly affected by climate variables from the year previous to the growing season and only secondly by current year conditions, but the critical months varied for different species and altitudes. Winter precipitation also affected radial growth, either as a source of water or as a possible agent of physical damage. Although our work showed significant climate influences on breast height tree radial growth, our results also indicated that other site factors such as microclimate or stand dynamics can be as or more important than climate variability.     

Comparison between TanDEM-X- and ALS-based estimation of aboveground biomass and tree height in boreal forests

Interferometric Synthetic Aperture Radar (InSAR) data from TerraSAR-X add-on for Digital Elevation Measurement (TanDEM-X) were used to estimate aboveground biomass (AGB) and tree height with linear regression models. These were compared to models based on airborne laser scanning (ALS) data at two Swedish boreal forest test sites, Krycklan (64°N19°E) and Remningstorp (58°N13°E). The predictions were validated using field data at the stand-level (0.5–26.1 ha) and at the plot-level (10 m radius). Additionally, the ALS metrics percentile 99 (p99) and vegetation ratio, commonly used to estimate AGB and tree height, were estimated in order to investigate the feasibility of replacing ALS data with TanDEM-X InSAR data. Both AGB and tree height could be estimated with about the same accuracy at the stand-level from both TanDEM-X- and ALS-based data. The AGB was estimated with 17.2% and 14.6% root mean square error (RMSE) and the tree height with 7.6% and 4.1% RMSE from TanDEM-X data at the stand-level at the two test sites Krycklan and Remningstorp. The Pearson correlation coefficients between the TanDEM-X height and the ALS height p99 were r?=?.98 and r?=?.95 at the two test sites. The TanDEM-X height contains information related to both tree height and forest density, which was validated from several estimation models.     

基于树高和树冠因子的立木材积与地上生物量相容模型研究

【目的】无人机机载激光雷达能够准确地测定单木、林分乃至大尺度森林结构参数(树高和树冠因子)。为应用无人机激光雷达技术准确估测森林蓄积量、生物量和碳储量提供计量依据和技术支撑。【方法】以150株实测马尾松生物量样本数据为研究对象,采用非线性回归估计方法和度量误差联立方程组方法,分析立木材积和地上生物量与树高、树冠因子的相关性,并在此基础上研究建立基于树高和树冠因子的立木材积与地上生物量相容模型。【结果】单株材积和地上生物量与树高因子的相关性最为紧密,其次才是树冠因子;基于树高和冠幅因子的二元材积和地上生物量模型预估精度较高,达到92%以上,再考虑冠长因子的三元模型预估精度改进不大;基于树高和冠幅因子的二元立木材积与地上生物量相容模型估计效果更好,相对于一元相容模型系统而言,二元相容模型拟合效果有较大幅度提高,预估精度达到92%以上。【结论】采用度量误差联立方程组方法可以有效解决基于树高和树冠因子的立木材积与地上生物量相容问题,并且预估精度达到92%以上,所建二元立木材积与地上生物量相容模型可为应用激光雷达技术反演森林蓄积量和生物量提供计量依据。     

草本地下竞争对3种幼苗存活和生长的影响

为了研究不同土壤营养条件下草本地下竞争对华北落叶松、麦氏云杉、辽东栎3种1年生幼苗存活和生长的影响,以华北落叶松、麦氏云杉和辽东栎1年生幼苗为材料,以土壤营养条件(以施用肥料中的氮含量为基准:10 g·m^2和40 g·m^2)和草本地下竞争方式(完全地下竞争,部分地下竞争,无地下竞争)为处理组合,统计幼苗存活率并测定苗高。结果表明:1)完全和部分去除草本根系的地下竞争均可同样有效地提高华北落叶松幼苗的存活率,而不同土壤营养条件对其存活率和高生长均无影响,不同地下竞争方式不影响其高生长。2)对麦氏云杉幼苗而言,在40 g·m^2的土壤营养处理下其存活率显著降低;但高生长未受影响,而完全和部分去除草本根系的地下竞争均显著提高了其存活率,且高生长在完全去除草本地下竞争的处理下最高。3)辽东栎幼苗在所有处理组合下全部存活,高生长在40 g·m^2的土壤营养处理下以及完全和部分去除草本根系地下竞争的处理下显著较高。华北落叶松和麦氏云杉幼苗定居的关键并非是草本植物对土壤养分的争夺,而是草本植物地下部分对华北落叶松幼苗根系生长空间的争抢。在今后森林营造和恢复实践中,通过清除草本的地下部分,特别是宿根性草本,避免草本占据过多地下生长空间,协助华北落叶松和麦氏云杉成功渡过幼苗定居期,同时森林培育中的除草措施也在一定程度上有利于辽东栎幼苗的生长。     

Effects of nitrogen on the growth and competition between seedlings of two temperate forest tree species

To understand how coexisting temperate tree species react to nitrogen (N) addition, seedlings of two coexisting species, Phellodendron amurense Rupr. and Fraxinus mandshurica Rupr. were transplanted to a controlled environment. Seedlings were then planted in two competition scenarios, control and mixed, and then subjected to four N addition treatments. The seedling growth parameters and competition effect were monitored and analyzed. The height and stem-base diameter showed a positive response to the N addition in all competition scenarios except for mixed P. amurense, which responded negatively. Chlorophyll content of the seedlings showed a positive relationship to N addition while the chlorophyll a/b showed a complex trend. The addition of N showed a positive relationship to the competition effect of diameter growth in both species and height growth in P. amurense. These results indicate that F. mandshurica may be more sensitive to N addition and could benefit more from N addition than P. amurense. With an increasing N input from the atmosphere, forests may be more suitable for F. mandshurica reducing the abundance of P. amurense.     

Effects of climatic conditions on tree-ring widths of three deciduous broad-leaved tree species at their northern distribution limit in Mont St. Hilaire,eastern Canada

This study examined the effects of climatic conditions on tree-ring widths of red oak (Quercus rubra), sugar maple (Acer saccharum) and American beech (Fagus grandifolia) at their northern distribution limit in Mont St. Hilaire, eastern Canada. A residual chronology of tree-ring width index was developed for each species. The tree-ring width of Q. rubra showed negative and positive correlations with May temperature and June precipitation in the current year, respectively. A. saccharum also showed similar patterns with Q. rubra, i.e., negative and positive correlations with June temperature and precipitation in the current year, respectively. F. grandifolia showed neither positive nor negative correlations with temperature and precipitation. The expressed population signals of tree-ring width chronologies of the three species indicated that climatic factors more greatly influenced the variations in tree-ring widths in the order Q. rubra > A. saccharum > F. grandifolia. Therefore, the radial growth of Q. rubra was most restricted by drought stress in early summer among the three species. Global warming is suggested to affect the radial growth of the three species differently by increasing drought stress at Mont St. Hilaire in eastern Canada.     

不同施肥处理对3种海岛树种苗木生长及造林效果的影响

为了解不同施肥处理对普陀樟(Cinnamomum japonicum var.Chenii)、全缘冬青(Ilex integra Thunb.)、红楠(Machilus thunbergii Sieb.et Zucc)3种海岛树种苗木生长及造林效果的影响,设置了7个不同的施肥处理,即(1)N(对照),(2)N+P,(3)N+2P、(4)N+K、(5)N+2K、(6)N+1P+1K、(7)N+2P+2K,于2011年5月至2012年1月观测苗高和地径,并测定植物各部分的生物量。结果表明:3种苗木不同施肥处理下,通过测算苗木质量指数QI可知,在N+1K施肥组合下,全缘冬青的苗木质量最好,而红楠在N+2P+2K的施肥组合下苗木质量最高,普陀樟在N+1P+1K的施肥组合下,苗木质量要好于其他处理。     

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

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