Crown conductance in dwarf, medium, and tall pitch pines in the Long Island Pine Plains

The New York Pine Plains are a unique ecosystem with normal statured and a dwarfed variety of pitch pines (Pinus rigida Mill.). Growing interspersed with the dwarf pines are trees of intermediate height and features. Several hypotheses have been put forward as to why some of the trees are dwarfed, but none have been substantiated. In this study, we tested whether dwarf or medium trees are hydraulically limited compared to normally growing trees. Granier style sap flux sensors were installed in three to six trees of each tree type and sap flux was measured in early August 2004. Sap flux measurements were scaled to crown stomatal conductance using leaf area to sapwood area ratios for each tree. Contrary to expectation, dwarf and medium stature trees had very low leaf area to sapwood area ratios, but high crown stomatal conductances compared to normal trees. Analyses of leaf area, ring widths, and crown stomatal conductance indicate that differences between normal, and dwarf and medium pines are not due to hydraulic limitation, but that stunted growth may be due to other causes.     

Competition-dependent modelling of foliage biomass in forest stands

Currently, foliage biomass is estimated based on stem diameter or basal area. However, it is questionable whether the relations between foliage and stem observed from plantations of a single tree species can be applied to stands of different structure or species composition. In this paper, a procedure is presented to simulate foliage and branch biomass of tree crowns relative to crown size and light competition. Crowns are divided into layers and segments and each segment is divided into a foliated and an unfoliated fraction. Depending on the competitive status of the segment, leaf area density, specific leaf area and foliated branch fraction are determined. Based on this information, foliage biomass is calculated. The procedure requires a crown shape function and a measure to characterise competition for light and space of each individual segment within the canopy. Simple solutions are suggested for both requirements to enable an application with data that can be measured non-destructively in the field; these were stem position, tree height, crown base height, crown radii and some general crown shape information. The model was parameterised from single trees of Norway spruce and European beech and partly evaluated with independent data close to the investigation plot. Evaluations showed that the model can attribute the ecology of the different crown forms. Modelled foliage distribution for beech and spruce as well as total needle biomass of spruce agreed well with measurements but foliage biomass of beech was underestimated. The results are discussed in the context of a general model application in structured forests.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

Development of biomass proportions in Norway spruce (Picea abies [L.] Karst.)

This study tests the hypotheses that (1) the above-ground structure of Norway spruce (Picea abies [L] Karst.) is derivable from the functional balance theory, and that (2) crown ratio is a key source of structural variation in trees of different age and social position. Twenty-nine trees were measured in three stands (young, middle-aged, and mature), with three thinning treatments (unthinned, normal, and intensive) in the two older stands. There was a strong linear relationship between the total cross-sectional area of branches and that of stem at crown base. Foliage mass was linearly related with stem basal area at crown base. Also an allometric relationship was found between foliage mass and crown length. The mean length (weighted by basal area) of branches obeyed an exponential function of crown length. The parameters of most of these relationships were independent of slenderness (tree height/breast height diameter) and tree age However, total branch cross-sectional area per stem cross-sectional area in the young trees was greater than in the older trees. The young trees also had slightly shorter branches than predicted by the mean branch length equation. This was probably caused by branch senescence which had not yet started in the young stand. The older trees had a relatively long lower crown segment which was growing slowly and senescing. It was proposed that a segmented crown structure is characteristic of shade tolerant tree species, and that the structural model could be further developed by making the two segments explicit.     

落叶松人工林单木模型的研究

根据吉林省松江河林业局所实测的落叶松人工林(Larix olgensis)临时标准地66块、固定标准地18块以及8块团状枝解析样地资料,通过对林分中优势木生长及树冠结构与动态的分析,提出适于树木生长的Korf方程并用来构造林木的潜在生长函数。选择林分密度指数(SDI)作为反映林分中林木之间平均拥挤指标。在单木竞争指标的选择上,通过引进树冠因子,并在与传统的竞争指标相比较的基础上,淡化距离因子的作用,应用优势木相对树冠表面积构造了与距离无关的单木竞争指标,以此建立了落叶松人工林单木生长模型。     

应用结构方程模型解析影响黄连木果实产量 和种子命运的因素

果实(种子)产量和质量是影响植物种群更新的重要因素。为了探明影响黄连木果实产量和种子命运的因素以及这些影响因素之间的相互作用, 作者于2009年对河南省济源市45株黄连木(Pistacia chinensis)结果样树的植株特征、果实特征、果实产量和种子命运等进行了测定, 并用结构方程模型进行综合分析。结果表明: (1)黄连木果实产量与树高、树冠面积和果序大小等特征成正相关, 但与胸径、果实大小相关性不显著; (2)与捕食者饱和假说的预测不一致, 单株果实产量对黄连木广肩小蜂(Eurytoma plotnikovi)的种子捕食率(即虫蛀率)无显著直接负向效应; (3)树高和果实大小对虫蛀率为显著直接正向效应, 胸径对虫蛀率为显著直接负向效应, 显示黄连木广肩小蜂对植株特征和果实特征有一定的选择能力; (4)空壳率与虫蛀率成显著负相关, 空壳果实越多, 越易逃避黄连木广肩小蜂的寄生, 空壳果实的存在对完好种子起到了一定保护作用, 可能是黄连木防御昆虫寄生的重要机制; (5)空壳率和虫蛀率对种子完好率有显著直接负向效应, 而胸径、果序大小和果实产量对完好率为间接正向效应, 树高和果实大小为间接负向效应。可见, 黄连木植株特征和果实特征均不同程度地影响其果实产量和昆虫寄生, 从而影响黄连木的种子质量和种群更新。     

Quantitative characterization of clumping in Scots pine crowns

Background and Aims

Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect.

Methods

Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR_crown) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium.

Key Results

Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented.

Conclusions

Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.     

Measure of simultaneous tree sways and estimation of crown interactions among a group of trees

We present a technique to measure the simultaneous sway of a group of trees and reconstruct the frequency of crown collisions and sway dynamics of individual or groups of trees. We placed a biaxial clinometer (tiltmeter) at the live crown base in each of ten neighboring 15-m-tall lodgepole pine trees in Alberta, Canada. Tree bole rotation at tiltmeter mount height was recorded during windy conditions at a rate of 10 times/s for the cluster of trees. Rotation angles were used in a bole curve calculation to estimate tree displacement in 2-dimensional (x, y) space. Collision reconstruction was done in Arc/Info by assigning asymmetrical crown area dimensions (polygons) to calculated bole displacement for each tree. Reconstruction of each time step measured any overlaps between crown polygon areas. Crown polygon overlaps estimated in this manner allowed assessment of collision frequencies, area of crown overlap during collisions, and identification of the tree(s) that a subject tree contacted. Collision statistics are only given for trees interior to the sensored cluster (n=3). For 15.0 min of data with an average wind speed of 4.5 m/s and a maximum of 10.0 m/s there was an average of 65 collisions/min for each tree, and an average collision overlap area of 24%. This frequency and depth of collisions supports the notion that wind-induced crown interaction inhibits lateral shoot extension and is an important mechanism for the development of crown asymmetry and crown shyness. Insight into dynamic tree sway behavior and crown interactions will allow estimation and cultivation of a forest stand structure that is more resistant to damage from wind. The techniques of recording multiple simultaneous bole sway and their reconstruction are applicable to a broad range of wind-forest interaction research.     

Allometric relationships for Eucalyptus nitens (Deane and Maiden) Maiden plantations

Allometric relationships between stem, leaf area and crown dimensions were determined for Eucalyptus nitens (Deane and Maiden) Maiden using 81 trees sampled from 13 post-canopy closure sites and 34 trees sampled from 6 pre-canopy closure sites. These sites differed in site quality, stand age, fertiliser treatment, stand density and levels of weed infestation. Overall, tree age ranged from 2 to 13 years, tree height from 1.4 to 26.1 m and diameter at breast height from 0.6 to 38.7 cm. Pre-canopy closure trees exhibited site-specific relationships which were to some extent confounded with tree age. However, post-canopy closure trees had relationships which were independent of site, age and silvicultural treatments. Strong relationships between structural components were found for both stem and crown. Stem diameter at breast height was non-linearly related to tree height and crown length. Stem sapwood area (breast height or crown base) could be predicted from stem cross-sectional area. For post-canopy closure trees, a functional relationship between sapwood area (breast height and crown base) and leaf area was site-independent. The lack of specificity in terms of both site and management techniques enables these relationships to be applied generally to E. nitens plantations in Tasmania.     

基于边际回归的沈阳市绿化树种人工油松冠形模拟

构建城市典型绿化树种树冠外部轮廓预估模型,可以为城市绿化树种的空间配置优化奠定基础。本研究以辽宁省沈阳市典型绿化树种人工油松为对象,基于Crown Window装置获取60株样木树冠形状,利用幂函数、分段抛物线方程和修正Kozak方程选取基础模型,通过再参数化引入树冠结构因子(最大树冠半径)和相邻木竞争变量(相邻木平均树高、平均胸径、平均冠幅、相邻木株数,以及样木与其相邻木平均树冠接触高),构建耦合相邻木竞争及树冠结构因子的油松树冠外部轮廓预估模型。结果表明: 修正Kozak方程R_a²最大、均方根误差(RMSE)最小,模型稳定性良好,选取其为构建油松冠形模型的基础模型。通过再参数化在基础模型中引入最大树冠半径和相邻木平均胸径后,模型的R_a²提高0.0693,MSER为14.4%。分析最大树冠半径和相邻木竞争对油松树冠形状的影响发现,最大树冠半径对树冠形状影响较大,树冠半径随最大树冠半径增大而增大;相邻木平均胸径对树冠形状影响相比最大树冠半径较弱,树冠上半部分随相邻木竞争增强而增大,树冠下半部分随相邻木竞争增强而变小。本研究构建的耦合相邻木竞争及最大树冠半径的油松树冠外部轮廓边际回归模型具有良好的拟合优度,能够合理地模拟及预测沈阳市典型绿化树种人工油松的树冠形状。     

北京地区侧柏人工林密度效应

密度是影响森林尤其是人工林生长的重要因素,林冠层是森林生态系统与其他系统进行能量和物质交换的重要场所,树木及树冠生长对林分密度的响应关系可以看作是生物对环境变化产生的适应性现象。林分密度效应是生态学和森林培育学的重要研究内容之一。以23块8种不同密度梯度的北京山区侧柏人工幼龄林林分为研究对象分析其树木生长及树冠生长对密度的响应关系,其中树冠指标使用了参照了美国林务局(USDA)的树冠调查指标。研究结果表明:(1)林分平均胸径、平均树高和平均冠幅生长均随密度增大而减小,林分密度大于3000株/hm2时各指标减小的趋势变缓,使用异速生长模型可以很好地拟合这种变化关系;(2)随密度增加,树冠水平方向和垂直方向生长均到显著地抑制作用,树冠外形表现出由饱满冠型向狭长冠型变化的适应性现象;(3)使用树冠二维、三维指标与密度进行相关性分析可知树冠长度、树冠率等指标与林分密度呈负相关关系,树冠圆满度及树冠生产效率与密度表现出极显著正相关关系;(4)采用枝解析的方法研究了树枝长度、材积的平均生长量、连年生长量与密度的关系,结果表明幼龄期各生长量差异不大;(5)在建立冠幅模型时考虑了自变量间的多重共线性问题,所建的胸径单自变量二次方模型能够很好地预测侧柏人工幼龄林冠幅生长过程,模型相关系数R2为0.961。     

Responses of crown development to canopy openings by saplings of eight tropical submontane forest tree species in Indonesia: a comparison with cool-temperate trees

BACKGROUND AND AIMS: Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0.2-2.5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan. METHODS: The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance. KEY RESULTS: Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity. CONCLUSIONS: This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.     

Modeling the vertical foliage distribution of an individual <Emphasis Type="Italic">Castanopsis cuspidata</Emphasis> (Thunb.) Schottky,a dominant broad-leaved tree in Japanese warm-temperate forest

The vertical foliage distribution of Castanopsis cuspidata (Thunb.) Schottky was examined in trees of various sizes to clarify its variation in relation to tree size and the light environment in a stand. As indices of these parameters, we analyzed crown social position (CSP: percent of stand height) and specific leaf area (SLA). The vertical foliage distribution of trees was expressed by a Weibull function. The variation in the vertical foliage distribution of C. cuspidata could be categorized into three types using crown social position and light environment. In the first type, leaves were concentrated to the top 20% of the tree; such trees are canopy trees that can receive full sunlight. The second type had a large relative crown depth and an asymmetric distribution with the maximum foliage located near the top of the tree; such trees are suppressed trees whose crowns do not receive sufficient light. The third type had a large relative crown depth and a symmetric distribution; such trees occur in high light environments, although their crowns are in the understory layer. The differences in the vertical foliage distribution are related to the strategies used to capture light. Multiple regression analysis showed that CSP and SLA at the top layer of the tree explained successive changes in the vertical foliage distribution. These results will contribute to scaling-up the vertical foliage distribution to the community level in pure stands of C. cuspidata using an individual-based model.     

Comparative height crown allometry and mechanical design in 22 tree species of Kuala Belalong rainforest, Brunei, Borneo

In rainforests, trunk size, strength, crown position, and geometry of a tree affect light interception and the likelihood of mechanical failure. Allometric relationships of tree diameter, wood density, and crown architecture vs. height are described for a diverse range of rainforest trees in Brunei, northern Borneo. The understory species follow a geometric model in their diameter-height relationship (slope, β = 1.08), while the stress-elasticity models prevail (β = 1.27-1.61) for the midcanopy and canopy/emergent species. These relationships changed with ontogeny, especially for the understory species. Within species, the tree stability safety factor (SSF) and relative crown width decreased exponentially with increasing tree height. These trends failed to emerge in across-species comparisons and were reversed at a common (low) height. Across species, the relative crown depth decreased with maximum potential height and was indistinguishable at a common (low) height. Crown architectural traits influence SSF more than structural property of wood density. These findings emphasize the importance of applying a common reference size in comparative studies and suggest that forest trees (especially the understory group) may adapt to low light by having deeper rather than wider crowns due to an efficient distribution and geometry of their foliage.     

Crown allometries are less responsive than stem allometry to tree size and habitat variations in an Indian monsoon forest

While theoretical allometric models postulate universal scaling exponents, empirical relationships between tree dimensions show marked variability that reflects changes in the biomass allocation pattern. As growth of the various tree compartments may be controlled by different functions, it is hypothesized that they may respond differently to factors of variation, resulting in variable tree morphologies and potentially in trade-offs between allometric relationships. We explore the variability of tree stem and crown allometries using a dataset of 1,729 trees located in an undisturbed wet evergreen forest of the Western Ghats, India. We specifically test whether species adult stature, terrain slope, tree size and crown light exposure affect the relationships between stem diameter and stem height (stem allometry), and between stem diameter and crown width, crown area and crown volume (crown allometries). Results show that both stem and crown allometries are subject to variations in relation to both endogenous (tree size, species adult stature) and exogenous (terrain slope, crown light exposure) factors. Stem allometry appears to be more affected by these factors than are crown allometries, including the stem diameter–crown volume relationship, which proved to be particularly stable. Our results support the idea that height is a prevailing adjustment factor for a tree facing variable growth (notably light) conditions, while stem diameter–crown volume allometry responds more to internal metabolic constraints. We ultimately discuss the various sources of variability in the stem and crown allometries of tropical trees that likely play an important role in forest community dynamics.     

樟子松人工林树冠表面积及体积预估模型的研究

基于樟子松(Pinus sylvestris var. mongolica)人工林6块固定标准地30株枝解析数据,在分析树冠表面积和树冠体积与林分变量和林木变量的基础上,利用幂函数建立了树冠表面积（CSA）和树冠体积(CV)的预估模型,同时还对林木材积生长量与CSA和CV进行了相关分析。研究结果表明：樟子松人工林树冠表面积和树冠体积随着林木胸径、树高和冠长的增大而增大,林木材积生长量与树冠表面积和树冠体积均明显呈线性关系。不同林分条件的樟子松人工林CSA和CV随林分年龄和胸径的增大而增大,CSA随林分密度的增大而减小,而CV与林分密度相关不紧密。林分树冠表面积和树冠体积预估模型的检验结果表明,两个模型的平均相对误差都在±8%之内,预估精度均大于91%,说明所建模型可以很好地预估樟子松人工林不同林分条件下的林木树冠表面积和树冠体积。     

Estimating Canopy Structure in an Amazon Forest from Laser Range Finder and IKONOS Satellite Observations1

Canopy structural data can be used for biomass estimation and studies of carbon cycling, disturbance, energy balance, and hydrological processes in tropical forest ecosystems. Scarce information on canopy dimensions reflects the difficulties associated with measuring crown height, width, depth, and area in tall, humid tropical forests. New field and spaceborne observations provide an opportunity to acquire these measurements, but the accuracy and reliability of the methods are unknown. We used a handheld laser range finder to estimate tree crown height, diameter, and depth in a lowland tropical forest in the eastern Amazon, Brazil, for a sampling of 300 trees stratified by diameter at breast height (DBH). We found significant relationships between DBH and both tree height and crown diameter derived from the laser measurements. We also quantified changes in crown shape between tree height classes, finding a significant but weak positive trend between crown depth and width. We then compared the field‐based measurements of crown diameter and area to estimates derived manually from panchromatic 0.8 m spatial resolution IKONOS satellite imagery. Median crown diameter derived from satellite observations was 78 percent greater than that derived from field‐based laser measurements. The statistical distribution of crown diameters from IKONOS was biased toward larger trees, probably due to merging of smaller tree crowns, underestimation of understory trees, and overestimation of individual crown dimensions. The median crown area derived from IKONOS was 65 percent higher than the value modeled from field‐based measurements. We conclude that manual interpretation of IKONOS satellite data did not accurately estimate distributions of tree crown dimensions in a tall tropical forest of eastern Amazonia. Other methods will be needed to more accurately estimate crown dimensions from high spatial resolution satellite imagery.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Reprint of: Tree-tree interactions and crown complementarity: the role of functional diversity and branch traits for canopy packing

Previous studies have shown that tree species richness increases forest productivity by allowing for greater spatial complementarity of tree crowns (crown complementarity), which in turn results in more densely packed canopies. However, the mechanisms driving crown complementarity in tree species mixtures remain unclear. Here, we take advantage of a high-resolution, three-dimensional terrestrial laser scanning approach in the context of a large-scale biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) to quantify the extent to which functional dissimilarity and divergences in branch traits between neighbouring trees affect crown complementarity at the scale of tree species pairs (i.e., two adjacent trees). Overall, we found no support that functional dissimilarity (divergence in morphological flexibility, specific leaf area and wood density) promotes crown complementarity. However, we show that the effects of functional dissimilarity (the plasticity of the outer crown structure) on crown complementarity vary in their magnitude and importance depending on branch trait divergences. Firstly, crown complementarity tended to be highest for tree species pairs that strongly differed in their functional traits, but were similar in branch density. In contrast, heterospecific pairs with a low functional trait divergence benefitted the most from a large difference in branch density compared with pairs characterised by a large functional dissimilarity. Secondly, the positive effects of increasing divergence in branching intensity (the plasticity of the inner crown structure) on crown complementarity became most important at low levels of functional dissimilarity, i.e. when species pairs were similar in their branch packing and vice versa. This suggests that species mixing allows trees to occupy canopy space more efficiently mainly due to phenotypic changes associated with crown morphology and branch plasticity. Our findings highlight the importance of considering outer and inner crown structures (e.g. branching architecture) to deepen our understanding of tree-tree interactions in mixed-species communities.