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.     

PLASTICITY OF TREE ARCHITECTURE: SPECIFIC AND ECOLOGICAL VARIATIONS FOUND IN AUBREVILLE'S MODEL

The crown form and branching pattern of sapling and adult trees of Terminalia catappa, T. latifolia, and an unidentified species of Manilkara are described. Leaf distribution (leaf positions within the crown) and leaf area index are given for the three species. Although they all exhibit the same architectural model, the two species of Terminalia maintain a tiered canopy in both sapling and large adult trees, while Manilkara has a sapling with poorly defined tiers and an adult with a hollow, hemispherical crown. Thus, the same architectural model (Aubréville's) can give rise to different crown shapes due to quantitative differences in branch geometry, degree of reiteration, and longevity of leaf-bearing branch units. Conversely, similar crown shapes can be produced by trees with different models. Manilkara conforms to the monolayer model of leaf arrangement at maturity, but Terminalia cannot easily be described as a monolayer or a multilayer. Adaptive geometry of a tree and its architectural model are not equivalent nor necessarily comparable.     

神农架巴山冷杉(Abies fargesii)林群落特征及其优势种群结构

采用样方法对神农架自然保护区巴山冷杉林的群落组成、优势种群结构与动态进行了研究.结果表明,该群落具有明显的温带常绿针叶林特征,以中小型革质、单叶的常绿高位芽植物为主.群落成层现象明显,可划分为乔木层、灌木层和草本层,林下苔藓层发达.植物的科属组成较为分散,乔木层主要优势科有松科(Pinaceae)、桦木科(Betulaceae)、杜鹃花科(Ericaceae)、槭树科(Aceraceae)等.区系成分复杂分散,与温带植物区系关系密切.巴山冷杉是该群落的建群种,重要值达54.30%,幼苗储备丰富,占更新层的90.4%,但幼树缺失.巴山冷杉中小径级个体占多数(78.8%),为进展型种群.高度结构基本呈倒金字塔形,集中分布于8～10级3个高度级上(62.2%),10级个体最多,占29.3%.植株个体数随冠幅级的递增呈倒"J"形分布,冠幅小于20m2的植株占87.4%,其中5m2以下最多(30.8%).冠幅与胸径呈指数相关,回归方程为y＝2.7118e0.0308x,R2＝0.520,冠幅与胸径的关系比与树高更密切.     

Community characteristics and population structure of dominant species of Abies fargesii forests in Shennongjia National Nature Reserve

In this paper, the community composition of Abies fargesii and the population structure of dominant species in this community were studied using a sampling plot method. The research plot was located in Shennongjia National Nature Reserve, China. Results showed that the appearance of the community was characterized by evergreen phanerophytes with microphyllous (or mesophyllous), coriaceous and single leaves. The vertical structure of the community can be clearly divided into 4 layers: arbor layer, shrub layer, herb layer and developed moss layer. Abies fargesii population was a developing population as there were more seedlings and young trees than old trees. Saplings were found more frequently in gaps than under the close canopy, which indicated a pattern of gap regeneration. The height structure of Abies fargesii population presented a reversed pyramid shape. Crown size structure of the population showed a typical reversed J shape. The relation between crown size and DBH (diameter at breast height) was closer than that between crown size and height.     

山西濒危植物翅果油树植冠的构型分析

应用分形理论及方法,从分枝格局和冠幅扩展2个方面对山西翼城和乡宁的35株翅果油树(Elaeagnus mollis Diels)个体的植冠构型进行统计分析。结果表明,幼树和成树的总体分枝率和逐步分枝率有显著变化;一级枝的平均枝长和枝径有显著差异,但枝和叶的方位角及叶倾角差异不明显。不同发育阶段个体的冠幅变化较大,幼树冠幅的分形维数(2.0026)小于成树(2.2694)。翅果油树在不同生长发育阶段对生境变化有不同的构型策略。     

Forest gap dynamics and the Ising model

The vegetation height in forest ecosystems is spatially heterogeneous. Canopy gaps (sites with low vegetation) are formed by treefalls, and they recover to canopy sites (with high vegetation) either by growth of small trees or by branch extension of surrounding trees. The dynamics of canopy gaps have been studied using a spatial Markov chain with nearest neighbor interaction. (1) If the canopy recovery rate is constant and if the gap formation rate for a site increases exponentially with the number of neighboring gap sites, the equilibrium distribution is the same as the one generated by the Ising model in statistical mechanics. Here, we extend the equivalence to the situation in which both the gap formation and canopy recovery depend on the neighborhood, as shown in recent forest data. (2) We develop a statistical test of whether a given spatial pattern is a random sample from the Ising model. The test is based on the conditional probability of configurations on a partial lattice. We apply the method to vegetation height data from the Ogawa forest reserve, Japan, measured on a 5x5 m grid in 1976, 1981, 1986, and 1991. The spatial pattern of the original forest data deviates significantly from the Ising model. We examine whether a larger sampling distance or the removal of the effects of the topography can reduce this deviation.     

樟子松人工林分枝结构的分析

基于对6块樟子松(Pinus sylvestris var. mongolica)人工林固定标准地中的30株样木枝解析调查数据,通过分析不同林分、不同大小林木1级枝和2级枝的分枝概率、分枝格局和分枝角度,揭示了樟子松人工林树冠的分枝结构特点。研究结果表明：樟子松人工林1级枝和2级枝的平均分枝数量分别为3.84个和2.80个,两者分枝概率均呈正态分布;1级和2级枝条在光照条件好的几个区间（方位角46°～225°）分布较多,1级枝条的水平分布遵从均匀分布,而2级枝条则不遵从均匀分布;树冠上层枝条的分枝角度略小于树冠中、下层,上层平均分枝角度为45.6°,而中下层平均分枝角度都为49.4°。不同大小林木的1级枝分枝结构规律表明：Ⅰ级木和Ⅴ级木的每轮平均分枝数非常接近,分别为3.89和3.94个,比Ⅲ级木每轮分枝数大0.5个左右;1级枝水平分布在各区间内(45°间隔)相差在0.24%～2.81%之间,方差分析结果表明枝条水平分布与林木大小无关;不同大小林木的分枝角度有所差别,Ⅰ级木、Ⅲ级木和Ⅴ级木的平均分枝角度分别为48.5°、42.2°和50.7°。     

Performance of trees in forest canopies: explorations with a bottom-up functional-structural plant growth model

Here we present a functional-structural plant model that integrates the growth of metamers into a growing, three-dimensional tree structure, and study the effects of different constraints and strategies on tree performance in different canopies. The tree is a three-dimensional system of connected metamers, and growth is defined by the flush probability of metamers. Tree growth was simulated for different canopy light environments. The result suggest that: the constraints result in an exponential, logistic and decay phase; a mono-layered-leaf crown results from self-shading in a closed canopy; a strong apical control results in slender trees like tall stature species; the interaction between weak apical control and light response results in a crown architecture and performance known from short stature species in closed forest; correlated leaf traits explain interspecific differences in growth, survival and adult stature. The model successfully unravels the interaction effects of different constraints and strategies on tree growth in different canopy light environments.     

Crown asymmetry in high latitude forests: disentangling the directional effects of tree competition and solar radiation

Light foraging by trees is a fundamental process shaping forest communities. In heterogeneous light environments this behavior is expressed as plasticity of tree growth and the development of structural asymmetries. We studied the relative influence of neighborhood structure and directional solar radiation on horizontal asymmetry of tree crowns in late‐successional high latitude (67–68°N) forests in northern Fennoscandia. We described crown asymmetries as crown vectors (i.e. horizontal vectors from stem center to crown center), which we obtained from canopy maps based on crown perimeter measurements in the field. To disentangle the influence of the two main determinants, inter‐tree competition and directionality of above‐canopy solar radiation at high latitudes, we applied circular statistical models, utilizing cylindrical distributions, to these data consisting of orientations and intensities of crown asymmetry. At the individual tree level, our model predicted crown asymmetry vectors from the current stand structure, and the predictions became better when the intensity of asymmetry (i.e. crown vector length) was higher. Competition was the main determinant of crown asymmetry for 2/3 of trees, and the model predictions improved when we incorporated the directionality of solar radiation. At the stand‐level, these asymmetries had resulted in a small increment of the projected canopy area and an increased regularity of spatial structure. Our circular statistical modelling approach provided a quantitative evaluation of the relative importance of directionality of solar radiation and neighborhood stand structure, showing how both of these factors play a role in formation of crown asymmetries in high latitude forests. This approach further demonstrated the applicability of circular statistical modeling in ecological studies where the response variable has both orientation and intensity.     

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.     

Model simulations of spatial distributions and daily totals of photosynthesis in Eucalyptus grandis canopies

Summary A simulation model for radiation absorption and photosynthesis was used to test the hypothesis that observed nonuniform distributions of nitrogen concentrations in young Eucalyptus grandis trees result in greater amounts of daily assimilation than in hypothetical trees with uniform N distributions. Simulations were performed for trees aged 6, 9, 12 and 16 months which had been grown in plantations under a factorial combination of two levels of fertilization and irrigation. Observed leaf N distribution patterns yielded daily assimilation rates which were only marginally greater (<5%) than for hypothetical trees with uniform distributions. Patterns of assimilation distribution in individual tree crowns closely resembled those for absorbed radiation, rather than for N. These conclusions were unaffected by three choices of alternative leaf area density distributions. The simulation model was also used to calculate hourly and daily rates of canopy assimilation to investigate the relative importance of radiation absorption and total canopy nitrogen on assimilation. Simulated hourly rates of carbon assimilation were often lightsaturated, whereas daily carbon gain was directly proportional to radiation absorbed by the tree crown and to total mass of N in the leaves. Leaf nitrogen concentrations determined photosynthetic capacity, whereas total leaf area determined the amount of radiation absorbed and thus the degree to which capacity was realized. Observed total leaf area and total crown N were closely correlated. The model predicted that nitrogen use efficiences (NUE, mol CO₂ mol^–1 N) were 60% higher for unfertilized than for fertilized trees at low levels of absorbed photosynthetically active radiation (PAR). Nitrogen use efficiency was dependent on fertilizer treatment and on the amount of absorbed PAR; NUE declined with increasing absorbed PAR, but decreased more rapidly for unfertilized than for fertilized trees. Annual primary productivity was linearly related to both radiation absorbed and to mass of N in the canopy.     

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。     

Simple equations to estimate light interception by isolated trees from canopy structure features: assessment with three-dimensional digitized apple trees

* Simple models of light interception are useful to identify the key structural parameters involved in light capture. We developed such models for isolated trees and tested them with virtual experiments. Light interception was decomposed into the projection of the crown envelope and the crown porosity. The latter was related to tree structure parameters. * Virtual experiments were conducted with three-dimensional (3-D) digitized apple trees grown in Lebanon and Switzerland, with different cultivars and training. The digitized trees allowed actual values of canopy structure (total leaf area, crown volume, foliage inclination angle, variance of leaf area density) and light interception properties (projected leaf area, silhouette to total area ratio, porosity, dispersion parameters) to be computed, and relationships between structure and interception variables to be derived. * The projected envelope area was related to crown volume with a power function of exponent 2/3. Crown porosity was a negative exponential function of mean optical density, that is, the ratio between total leaf area and the projected envelope area. The leaf dispersion parameter was a negative linear function of the relative variance of leaf area density in the crown volume. * The resulting models were expressed as two single equations. After calibration, model outputs were very close to values computed from the 3-D digitized databases.     

Influence of canopy tree size on stand basal area may reflect uncoupling of crown expansion and trunk diameter growth

Abstract A recent article by Midgley and colleagues suggests that large trees give rise to inordinately high stand basal areas because they pack canopy space more efficiently than smaller trees. We argue that this phenomenon bears more relation to the fact that diameter increment is not necessarily accompanied by significant crown expansion during all stages of a tree's life. Using data from a canopy tree population in an old‐growth temperate forest, we found that crown area scaled as roughly the 3/5 power of trunk basal area. Rather than reflecting fixed scaling laws, we suggest that this pattern arises because of limited opportunities for crown expansion in dense stands. Old canopy trees in dense stands can thus accumulate large basal areas without occupying a commensurately large canopy area.     

Crown alterations induced by decline: a study of relationships between growth rate and crown morphology in beech (Fagus sylvatica L.)

One of the first symptoms expressed by declining trees is reduced growth in stem diameter and length increment. The possibility of a relationship between length increment and crown thinning in beech (Fagus sylvatica L.) was investigated by developing a computer model to simulate first order branching patterns of the apical 2 m of monopodially branching beech trees, 70–100 years old, for a range of length increment rates. The model was based on values for branching angle, main axis and branch length increment, number of branches produced per year and branch mortality rates for six healthy and declining trees. Shoot growth rates in the apical 2 m of the sample trees ranged from about 5 cm/year (decline class 3) to 43 cm/ year (healthy). Simulations of branching patterns in the apical 2 m of trees growing at different rates indicated that, when growth rate exceeded about 20 cm/year, total first order branch length and area explored were independent of growth rate. When growth rates fell below this value there was a reduction in total area explored and first order branch length due primarily to the formation of fewer branches. More acute branching angles contributed to a reduction in the area explored. Growth rate-related crown thinning could increase the risk of bark necrosis and secondary pathogen infection during dry and/or hot spells.     

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.     

Influence of canopy tree phenology on understorey populations of Fagus crenata

Question: Is light available for subcanopy individuals of Fagus crenata spatiotemporally heterogeneous across patches with closed canopies of different foliage phenologies and gaps? Is local abundance of Sasa influenced by the composition of the canopy layer? If so, does the Sasa layer also affect the amount of light available to small F. crenata saplings? Is variation in F. crenata population structure consistent with the hypothesis that light is important? Location: Mt. Kurikoma, Japan 780 m a.s.l. Methods: Population structure of subcanopy individuals of Fagus crenata and importance of Sasa were examined in five patch types. The patch‐types were Fc (F. crenata only in the crown), Qm (Quercus mongolica var. grosseserrata only in the crown), Mo (Magnolia obovata only in the crown), Fc’ (periphery of F. crenata) and Gap. Seasonal changes in light availability above and below the Sasa layer was examined by using hemispherical photographs and quantum sensors. Results: Subcanopy individuals of F. crenata began unfolding their leaves approximately one month earlier than canopy trees of Q. mongolica var. grosseserrata and M. obovata, but a few days later than those of adult F. crenata. Accumulated photosynthetic photon flux density above the Sasa layer was greatest in Qm and Mo, and lowest in Fc. Importance of Sasa was highest in Gap. Maximum height and the number of subcanopy individuals of F. crenata were greatest in Qm, followed by Mo, and lowest in Fc. Conclusions: Differences in canopy layer composition probably influence the regeneration of F. crenata both directly through their foliage phenologies, and also indirectly by determining the importance of Sasa.     

Crown Architecture and Species Coexistence in Plant Communities

The relationships between crown architecture and species coexistencewere studied using the diffusion model and the canopy photosynthesismodel for multi-species plant communities. The present paperdeals with two species having different crown shapes [conic-canopyplant (CCP) and spheroidal-canopy plant (SCP)], for variousinitial mean sizes at the establishment stage and physiologicalparameter values (photosynthetic rate, etc.). Recruitment processeswere not incorporated into the model, and thus simulations weremade for the effects on the pattern of species coexistence ofeither sapling competition starting from different sapling banksor competition in single-cohort stands with little continualestablishment of species until a stand-replacement disturbance.The following predictions were derived: (1) SCPs can establishlater/slowly in the lower canopy layer even if they are overtoppedby a CCP which established first/rapidly; (2) if SCPs establishedfirst/rapidly and occupy the upper canopy layer, a CCP can rarelyestablish later/slowly in the lower canopy layer; (3) smallest-sizedCCPs can persist well in the lowermost canopy layer overtoppedby a SCP, suggesting a waiting strategy of CCP's saplings inthe understorey of a crowded stand; (4) even if CCPs establishedfirst/rapidly and occupy the upper canopy layer, an SCP canestablish later/slowly in the lower canopy layer. Therefore,the species diversity of SCPs which established first/rapidlyand occupy the upper canopy layer limits the number of CCP specieswhich can establish later/slowly. In contrast, the species diversityof CCPs which established first/rapidly and occupy the uppercanopy layer does not affect the number of SCP species whichcan establish later/slowly. The combination of initial sizesof a CCP and an SCP at the establishment stage (i.e. establishmenttiming) affects the segregation of vertical positions in thecanopy between the two species with different crown shape, andnot only species-specific physiological traits but also crownarchitecture greatly affects the coexistence pattern betweenspecies with different crown architectures. The theoreticalpredictions obtained here can explain coexistence patterns foundin single-cohort conifer-hardwood boreal and sub-boreal forests,pointing to the significance of crown architecture for speciescoexistence. Diffusion equation model; canopy photosynthesis model; conifer-hardwood boreal/sub-boreal forest; sapling establishment; vertical foliage profile     

应用修正的Gash解析模型对岷江上游亚高山川滇 高山栎林林冠截留的模拟

修正的Gash解析模型具有较好的物理基础,是目前估算和预测林冠截留的有效工具。基于2007年6月到9月的降雨数据、气象和林分结构资料,采用修正的Gash解析模型对岷江上游川滇高山栎林冠截留进行了模拟。结果表明,林冠持水能力为0.23mm,树干持水能力为0.041mm,树干茎流系数为0.013,使林冠饱和的降雨为0.3mm;研究期间实测的林冠截留量为80.2mm,占总降雨量的16.5%,应用模型模拟的林冠截留总量为81.8mm,高于实测值1.6mm(高于实测值2.0%),林冠截留和树干茎流周累积量模拟值均与实测值有很好的一致性,敏感性分析结果显示,应用修正的Gash解析模型进行岷江上游川滇高山栎林冠截留模拟研究中,模型最易受林冠持水能力影响,其次为郁闭度、树干持水能力,影响最小的林分参数为树干茎流系数。     

A model to assess relationships between forest dynamics and spatial structure

Abstract. A spatially explicit model was developed to study the relationships between the dynamics and spatial structure of forest stands. The objective was to test whether tree spatial structure can be used as an indicator of stand dynamics. The model simulates the growth, mortality and recruitment of trees in a multi‐specific and uneven‐aged stand. It includes deterministic and stochastic processes so that repeated simulations do not lead to the same stand but provide several possible results for a given dynamic (defined by a set of parameters). Second‐order neighbourhood analyses were used to characterize the resulting spatial structures. They showed a high variability for a given set of parameters. Only the main trends in the spatial structure can be interpreted. Sensitivity analyses, concerning the influence of competition on spatial structure, showed that in heterogeneous stands confounding effects can hinder the interpretation of the spatial structure if all the trees are considered. The spatial structure of the canopy trees alone proved easier to interpret as it is directly linked to post recruitment competition. Inference on the dominant modality of competition (one‐sided or two‐sided) based on the spatial structure proved difficult.