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.     

Biomass,stem basic density and expansion factor functions for five exotic conifers grown in Denmark

Adequate allometric equations are needed for estimating carbon pools of fast growing tree species in relation to international reporting of CO₂ emissions and for assessing their possible contribution to increasing forest biomass resources. We developed models for predicting biomass, stem basic density and expansion factors of stem to above-ground biomass for five fast growing conifers. Data included destructive measurements of 236 trees from 14 sites, covering a wide range of growth conditions. To ensure model efficiency, models for predicting stem, crown and total above-ground biomass for the five species were estimated simultaneously using a linear, mixed effects model that allowed contemporaneous correlations between the different tree components. Models differed among species and included dbh and tree height. The models explained more than 98% of the variation in above-ground biomass and reflected differences in the allometry between tree species. Stem density differed among species but generally declined with increasing site index and dbh. The overall model for predicting stem basic density included dbh, H₁₀₀ and site index and explained 66% of the total variation. Expansion factors decreased from 1.8–2.0 in small trees (dbh < 10 cm) to 1.1–1.2 for large trees (dbh > 25 cm), but differed among species. The overall model explained 86% of the variation and included quadratic mean diameter and dbh.     

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.     

北京栓皮栎林分的干形质量初步研究

探讨了划分林木干形质量的标准,将林木干形质量分为通直、轻度弯曲、二杈分枝、多杈分枝和重度弯曲5类干形。选择了胸径、树高、冠幅、冠长等6个因子,并研究了这6个调查因子随干形质量变化的关系。通过方差分析发现不同干形质量的林木胸径差异并不明显,而其它的调查因子却因干形不同而表现出显著差异。对表现显著的林分因子进行多重比较研究,发现通直干形生长得较高、有较小的冠径比;二杈分枝和多杈分枝林木的冠幅较大,且二杈分枝类型林木的高径比最小。可以得出结论:在近自然经营过程中,在选择以材质为目的的目标树时,首先考虑干形通直的林木。     

Derivation of stem taper from the pipe theory in a carbon balance framework

A dynamic tree growth model is described. The model derives the development of stem taper and vertical distribution of branch basal area from the pipe model, assuming that reuse of active pipes is regulated by foliage dynamics in a vertically explicit crown with a foliage distribution of constant shape. Based on empirical findings, the pipe model was modified slightly to allow the foliage/sapwood ratio to vary as a function of distance from the treetop. Growth was derived from carbon balance in a stand of different size trees that may shade each other. The model was applied to old and middle-aged trees growing in dense and sparse stands of Scots pine for which stand-level measurements are available as a chronosequence, but individual trees have been measured only once. Measured trees were compared with corresponding simulated trees for stem taper and vertical distribution of branch basal area. The results indicated that the pipe model assumptions, combined with a model of tree growth, are capable of producing realistic predictions of the vertical distribution of stem and branch diameter in trees of different sizes in the stand. A comparison of the results with a simple form of the uniform stress theory showed good agreement between the two models. However, a significant difference was found between the measured relative contribution of heartwood to total stem diameter and the predicted share of disused pipes in the stem. A possible explanation for this discrepancy is that the transition from sapwood to heartwood is gradual rather than abrupt as assumed in the model. A modification of the pipe model to incorporate a gradual transition is outlined.     

华南地区7种阔叶树种早期生长表现

在广东省西江林场的7种阔叶树种幼林内设置标准地,对其幼林期的生长表现进行研究.结果表明:米老排和灰木莲的地径、树高及冠幅与树高显著相关,地径与冠幅相关不显著;西南桦、火力楠和降香黄檀的地径、树高、冠幅三者间都显著相关;黑木相思的地径与树高及冠幅显著相关,冠幅与树高相关不显著;红花荷的地径、树高、冠幅的生长均无显著相关;...     

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate

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

Modelling dimensional growth of three street tree species in the urban forest of the City of Tshwane,South Africa

Tree height, crown height, crown diameter and stem diameter were measured for 282 trees of the indigenous species Combretum erythrophyllum, Searsia lancea and S. pendulina. Growth relationships were modelled using age as explanatory variable for stem diameter and subsequently stem diameter as explanatory variable for tree height, crown height and crown diameter. Coefficients are presented for predicting tree dimensions using a logarithmic function. There were strong correlations for stem diameter and age (r ² ≥ 0.75), and crown diameter and stem diameter (r ² ≥ 0.74) for all three of the species investigated. Correlations were weaker for tree height and stem diameter (r ² ≥ 0.63), and crown height and stem diameter (r ² ≥ 0.60) for S. pendulina but stronger for both C. erythrophyllum (r ² ≥ 0.83) and S. lancea (r ² ≥ 0.70) in both instances. The results can be used in forecasting the physical dimensions of these species as a function of time. The results could also be used in the process of modelling energy use reduction, air pollution uptake, rainfall interception, carbon sequestration and microclimate modification of urban forests such as those found in the City of Tshwane.     

Individual-tree diameter growth models for black spruce and jack pine plantations in northern Ontario

Individual-tree distance independent diameter growth models were developed for black spruce and jack pine plantations. Data used in this study came via stem analysis on 1170 black spruce (Picea mariana [Mill.] B.S.P.) and 800 jack pine (Pinus banksiana Lamb.) trees sampled from 75 stands of 25 even-aged monospecific plantations for each species in the Canadian boreal forest region of northern Ontario. Of the 75 stands, 50 were randomly selected for each species and all trees from these stands were used for model development. Trees from the remaining stands were used for model evaluation.A nonlinear mixed-effects approach was applied in fitting the diameter growth models. The predictive accuracy of the models was improved by including random effects coefficients. Four selection criteria - random, dominant or codominant, tree size close to quadratic mean diameter, and small sized - were evaluated for accuracy in predicting random effects for a new stand using the developed models. Random effects predicted based on trees selected using the random selection criterion provided more accurate diameter predictions than those using trees obtained via other selection criteria for both species. The models developed here are very important to forest managers as the diameters predicted by these models or, their stand-level summaries (i.e., basal area, average diameter), are used as inputs in any forest growth and yield models. In addition, individual-tree diameter growth models can be used to directly forecast changes in diameter distribution of stands.     

Sugar maple (Acer saccharum Marsh.) growth is influenced by close conspecifics and skid trail proximity following selection harvest

In this study, we quantified the effects of local neighbourhood competition, light availability, and proximity to skid trails on the growth of sugar maple (Acer saccharum Marsh.) trees following selection harvest. We hypothesized that growth would increase with decreasing competition and increasing light availability, but that proximity to skid trails would negatively affect growth. A total of 300 sugar maples were sampled ∼10 years after selection harvesting in 18 stands in Témiscamingue (Québec, Canada). Detailed tree and skid trail maps were obtained in one 0.4 ha plot per stand. Square-root transformed radial growth data were fitted to a linear mixed model that included tree diameter, crown position, a neighbourhood competition index, light availability (estimated using the SORTIE light model), and distance to the nearest skid trail as explanatory variables. We considered various distance-dependent or -independent indices based on neighbourhood radii ranging from 6 to 12 m. The competition index that provided the best fit to the data was a distance-dependent index computed in a 6 m search radius, but a distance-independent version of the competition index provided an almost equivalent fit to data. Models corresponding to all combinations of main effects were fit to data using maximum likelihood, and weighted averages of parameter estimates were obtained using multimodel inference. All predictors had an influence on growth, with the exception of light. Radial growth decreased with increasing tree diameter, level of competition and proximity to skid trails, and varied among crown positions with trees in suppressed and intermediate positions having lower growth rates than codominants and dominants. Our results indicate that in selection managed stands, the radial growth of sugar maple trees depends on competition from close (≤6 m) conspecific neighbours, and is still affected by proximity to skid trails ∼10 years after harvesting. Such results underscore the importance of minimizing the extent of skid trail networks by careful pre-harvest planning of trail layout. We also conclude that the impact of heterogeneity among individual-tree neighbourhoods, such as those resulting from alternative spatial patterns of harvest, can usefully be integrated into models of post-harvest tree growth.     

基于管道模型理论的长白落叶松叶生物量与径向生长关系研究

目的 为了解长白落叶松单木叶生物量与径向生长之间的关系。 方法 以小兴安岭地区长白落叶松为研究对象,结合管道模型理论,研究单木个体水平和单木内不同方位区间水平上的叶生物量与径向生长的关系,分析树冠分布与径向生长之间的相关性。 结果 叶生物量与胸高处和枝下高处树干直径、断面积、边材面积,近1、2、3、5年断面积生长量均呈显著正相关,使用胸高断面积作为预测因子的叶生物量模型拟合效果最好。单木株内不同方位区域的叶生物量与对应的胸径处和枝下高处的树干半径、断面积,近1、2、3、5年断面积生长量均呈显著正相关。 结论 估测单株叶生物量时,使用胸高断面积作为预测因子的预测精度最高。株内不同方位叶生物量与对应方位的树干半径、断面积及断面积生长量均呈显著正相关,树冠偏冠与髓心偏心具有一定的相关性。     

Crown release improves growth of 20-year-old Betula alleghaniensis in tolerant northern hardwood stands

Twenty-year-old overstocked naturally regenerated yellow birch (Betula alleghaniensis Britton) were thinned near Thessalon, Ontario. Treatments consisted of either a control, or the removal of all competing trees at 1, 2, 3, or 4 m around the boles of the crop trees. The size of the openings created around the crop tree crowns was found to be more strongly correlated to diameter, clear bole length, and crown increment than the removal of competing trees using fixed distance from the stem. Crop trees responded well to thinning. Five-year diameter increment and crown diameter growth were positively correlated to size of the opening created around the crop tree crown. However, the relationship between height growth and size of canopy opening was weak. Clear bole length was found to be negatively correlated to the size of canopy opening. Few epicormic sprouts developed on the stems when all competing trees were removed at 1, 2, or 3 m around the crop tree boles. However, release at 4 m greatly increased the number and size of epicormic sprouts. Providing about a 15-m² opening around the crop tree crown, corresponding to approximately a 3-m removal of competing trees around the bole, provided a good balance between diameter increment and stem quality maintenance.     

Modelling growth behaviour in monoculture in subtropical eastern Australia of two eucalypt species that differ in shade tolerance

Growth behaviour of the relatively shade-intolerant species Eucalyptus dunnii and the rather more shade-tolerant Corymbia citriodora was considered during several growth periods over 8–14 years of age in experimental plantation monoculture in subtropical New South Wales. Larger trees dominated growth in any stand; this was consistent with the principle that inter-tree competition in such forests is for light and is asymmetric in that larger trees shade smaller trees but the reverse does not occur. Once competition was established, competition-induced mortality of C. citriodora was less than that of E. dunnii. A model relating tree diameter growth rate to tree diameter, competitive position in the stand and stand basal area was developed. The model showed that smaller, hence partially shaded, trees in stands of C. citriodora were better able to maintain their growth rates than equivalent trees in stands of E. dunnii. It is hypothesised that this reflects the greater shade tolerance of C. citriodora. This pattern of growth led to a more even distribution of tree size classes across the stand in the more shade-tolerant species.     

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.     

Pipe-model ratio distributions and branch foliage biomass: differences between two sympatric spruce species

The foliage biomass–sapwood relationship (the pipe model) is critical for tree growth and is used in tree growth models for understanding the implications of this structural relationship on the allocation of resources. In this research, we compared this relationship for two commercially important and sympatric species, black spruce (Picea mariana (Mill.) B.S.P.) and white spruce (Picea glauca (Moench) Voss). At locations in eastern Canada, 57 black and 50 white spruce trees were destructively sampled to obtain foliage biomass, crown structure, and tree stem measures. Using a model-based approach, we compared foliage biomass–branch basal area and foliage biomass–sapwood relationships at the tree and disk (i.e. along the tree stem) levels (i.e. pipe-model ratios) between these two species. We found that (i) branch foliage biomass–branch basal area was greater for black spruce than white spruce and (ii) pipe-model ratios along the tree stem given tree size were greater for black spruce than for white spruce. We attributed these differences to: (i) greater shade tolerance and leaf longevity of black spruce; (ii) slower growth rates of black spruce; and (iii) differing hydraulic strategies and mechanical requirements.     

Branch mortality and potential litterfall from Douglas-fir trees in stands of varying density

Forest floor characteristics influence nutrient cycling and energy flow properties of forest ecosystems, and determine quality of habitat for many forest plants and animals. Differential crown recession and crown development among stands of differing density suggest that an opportunity may exist to control the input of fine woody litter into the system by manipulating stand density. The objective was to measure the rate of branch mortality among stands of differing density and to estimate the range in total per hectare necromass inputs. Although litter traps are reliable for estimating per hectare rates of litterfall, branch mortality dating on sectioned stems uniquely allows assessment of several other litterfall components: (1) individual tree contributions to total litterfall; (2) the amount of branch material released by mortality, regardless of whether the branches are shed to the forest floor; (3) the distribution of basal diameters characterizing the litterfall from a given tree and stand. Twenty-four trees were felled and sectioned on permanent plots that were part of a silvicultural study of stand density regimes in Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco.). Whorl branches were dissected out of bole sections to determine the dates of mortality, and a branch biomass equation was applied to estimate potential rate of litterfall. Periodic annual rates were expressed in four ways: (1) number of branches per tree; (2) mass of branches per tree; (3) mass of branches per unit of crown projection area; (4) mass of branches per hectare. For the growth periods investigated, larger trees and trees growing on denser plots tended to release a greater necromass through branch mortality. Average branch basal diameter generally decreased with increasing stand density. Annual branch mortality ranged from 33 to 430 g m⁻² crown projection area for individual trees, and from 236 to 1035 kg ha⁻¹ for individual plots. These rates approached the low end of the range of previously published fine litterfall rates for Douglas-fir. Rates on these plots were relatively low owing to the temporary delay in crown recession imposed by artificial thinning. A conceptual model of branch litter dynamics is presented to depict consistencies with crown development among stands managed under different density regimes.     

Tree morphology responds to neighbourhood competition and slope in species-rich forests of subtropical China

Trees are able to respond to their local biotic and abiotic environment with morphological adjustments which improve resource acquisition and, thus, growth. In forests, light is broadly recognised as one of the major factors determining growth, and morphological responses comprise changes in crown architecture and stem stature. On sloping terrain, the interplay of phototropism and gravitropism may further affect morphological growth characteristics. However, different tree species are expected to show species-specific responses. In this study, we analysed three growth characteristics of tree individuals belonging to four species of two functional groups (evergreen: Schima superba, Castanopsis eyrei, deciduous: Quercus serrata var. brevipetiolata, Castanea henryi) in a species-rich Chinese subtropical forest. Crown projection area, relative crown displacement and stem inclination were related to biotic (local species richness, functional richness, competition, stand age) and abiotic (slope aspect and inclination, soil depth) variables in the local neighbourhood of the tree individuals. We hypothesised that (i) there are species-specific differences in the morphological response of crown architecture and stem stature and (ii) that crown size and asymmetry as well as stem inclination are influenced by both, biotic and abiotic factors. In contrast to our expectations we were unable to reveal any species-specific differences in any of the three growth characteristics. The results of mixed effects models showed that crown area was mainly affected by the target tree's dbh and biotic variables related to neighbours (competition, functional diversity), whereas stem inclination was mainly influenced by slope. Relative crown displacement was influenced by both, biotic and abiotic variables. We conclude that growth responses resulting in crown displacement and stem inclination seem to be an important mechanism to ameliorate foraging for light in our study area, but that these responses appear to be species-independent. The interplay of stem inclination and crown displacement allows for a plastic response of tree individuals in biotically and abiotically heterogeneous environments. Our results indicate that forest management in this region should focus on functionally diverse stands which are promoting crown area positively resulting in increased growth rates of individual trees.     

Species-specific biomass allometric models and expansion factors for indigenous and planted forests of the Mozambique highlands

Secondary Miombo woodlands and forest plantations occupy increasing areas in Mozambique, the former due to anthropogenic activities. Plantations, mainly species of Eucalyptus and Pinus, are being established on sites previously covered by secondary Miombo woodlands. This affects the evolution, cycle and spatiotemporal patterns of carbon(C) storage and stocks in forest ecosystems. The estimation of C storage, which is indispensable for formulating climate change policies on sequestrating CO_2, requires tools such as biomass models and biomass conversion and expansion factors(BCEF). In Mozambique, these tools are needed for both indigenous forests and plantations. The objective of this study is to fit species-specific allometric biomass models and BCEF for exotic and indigenous tree species. To incorporate efficient inter-species variability, biomass equations were fitted using nonlinear mixed-effects models. All tree component biomass models had good predictability; however, better predictive accuracy and ability was observed for the 2-predictors biomass model with tree height as a second predictor. The majority of the variability in BCEF was explained by the variation in tree species. Miombo species had larger crown biomass per unit of stem diameter and stored larger amounts of biomass per stem volume. However, due to relatively rapid growth, larger stem diameters, heights, and stand density, the plantations stored more biomass per tree and per unit area.     

樟子松人工林一级枝条基径模型的研究

以东北林业大学帽儿山实验林场樟子松人工林为研究对象,采用树干解析、枝解析的方法,分别于2007年和2008年在老山施业区选择不同年龄、不同立地和不同密度的樟子松人工林有代表性的林分设置固定标准地15块（其中,2007年设置7块,2008年设置8块）,共获取解析样木53株,实测了2298个一级枝活枝条变量因子（包括总着枝深度（DINC）、方位角（φ）、着枝角度（θ）、基径（BD）、枝长（BL）、弦长（BCL）、弓高（BAH））数据资料。基于理论或经验生长方程,建立了樟子松人工林树冠结构一级枝条基径静态模型,包括平均基径模型和最大最小一级枝条基径模型,并对模型进行了检验,结果表明模型预估效果良好。