林窗对川西亚高山岷江冷杉幼苗生物量及其分配格局的影响

采用样带法调查了川西亚高山原始岷江冷杉林林窗和林冠下岷江冷杉幼苗(H≤100 cm)的密度、高度、基径和各构件生物量,分析了各构件生物量间的相关性.结果表明：林窗与林冠下幼苗密度分别为12 903和2 017株·hm^-2;平均高度分别是26.6和24.3 cm,差异显著;平均基径分别为4.97和5.13 mm,差异不显著.林窗内各龄级幼苗的生物量与林冠下同龄级幼苗的生物量之间差异不显著.林窗对不同龄级幼苗各构件生物量积累的影响不同,林窗内幼苗枝干生物量比在12年达到最大(1.54),随后其比值维持在0.69左右;林冠下幼苗在15年后(含15年),枝条生物量大于主干生物量.生长在林窗和林冠环境的幼苗单株总生物量、叶生物量、茎生物量、地上生物量和根生物量与幼苗基径的平方(D²)和苗高(H)的乘积有显著的线性关系.林窗与林冠下幼苗各构件生物量之间有显著的正相关关系.     

林隙对小兴安岭阔叶红松林树种更新及物种多样性的影响

研究了小兴安岭阔叶红松林不同林隙梯度(林隙中心、林隙近中心、林隙边缘)中主要树种的数量特征,以及林隙大小对树种更新的影响.结果表明:不同梯度林隙内灌木树种的密度均明显高于非林隙,同种灌木密度的比值在1.08～18.15之间;随林隙面积增加,乔木幼苗更新密度逐渐增大,幼树Ⅰ(高度H≥1 m,胸径DBH≤2 cm)和幼树Ⅱ(H≥1 m,2 cm＜DBH≤5 cm)的更新密度呈多峰曲线.林隙灌木总体更新密度主要随幼苗和幼树Ⅰ数量而变化.林隙内不同位置幼苗的平均树高、平均基径、种密度和个体密度有所差异.从林隙中心到非林隙,更新层乔木幼苗重要值的大小顺序为:林隙中心＞林隙近中心＞林隙边缘＞非林隙;物种均匀度呈高-低-高的变化,物种多样性的变化为早期林隙＞中期林隙＞晚期林隙.     

岷江冷杉林林窗小气候及其对不同龄级岷江冷杉幼苗生长的影响

对川西亚高山原始岷江冷杉(Abies faxoniana)林林窗内和林冠下小气候及岷江冷杉幼苗生长和生物量进行了两个生长季的连续观测。结果表明:6月林窗内与林冠下太阳辐射的日积累量没有显著性差异,而7~8月的日积累量则有显著性差异。整个生长季节,林窗内太阳辐射的平均积累量为8.10×MJ·m^-2,而林冠下太阳辐射的平均积累量为5.02×MJ·m^-2,两个位点太阳辐射积累量的显著差异主要来自7~8月日积累量的不同;林窗内5和15 cm层土壤的日平均温度比林冠下相应深度分别高2.1和2.7℃,差异显著。林窗内和林冠下3~8年岷江冷杉幼苗高增长率分别为1.2±0.3 cm·a^-1和1.1±0.3 cm·a^-1,差异不显著;9~20年岷江冷杉幼苗高增长率分别为6.2±2.4 cm·a^-1和3.0±0.9 cm·a^-1,差异显著。林窗内岷江冷杉幼苗根、主茎和总生物量与林冠下幼苗根、主茎和总生物量没有显著差异。不同年龄的岷江冷杉幼苗叶和侧枝生物量积累对林窗微环境的响应不同。     

茂兰喀斯特常绿落叶阔叶混交林林窗下木本植物更新组成

对茂兰喀斯特森林中不同特征林窗内木本植物更新进行调查,计算不同特征林窗内木本植物的重要值、密度和丰富度,分析不同林窗大小、年龄和形成原因与木本植物更新组成的关系。结果表明:林窗大小对木本植物密度和丰富度有显著影响,耐荫树种密度和丰富度在大林窗(>200 m²)分别为0.32株·m^-2和3.26,中性树种密度和丰富度分别为0.40株·m^-2和2.54,均比在小林窗(≤200 m²)中显著增大,表明大林窗(>200 m²)能为耐荫树种和中性树种提供适宜光照条件和生长环境。幼年期林窗中以耐荫树种为优势种,重要值为0.377;中年期林窗以中性树种为优势种,重要值为0.400;老年期林窗以喜光树种为优势种,重要值为0.325,表明随着林窗年龄的增加,林窗内喜光树种逐渐占据优势。林窗形成原因对木本植物密度有显著影响,中性树种和喜光树种的密度在藤本绞杀形成的林窗均为0.43株·m^-2,比在人为伐倒和病虫害形成的林窗中显著增大,表明藤本绞杀可为喜光树种和中性树...     

华北落叶松人工林林隙大小对其更新的影响

对关帝山华北落叶松人工林20个林隙的幼苗(高度<1 m)、幼树(高度≥1 m,胸径<5 cm)进行调查，分析4个面积林隙等级(<60 m²、60～120 m²、120～180 m²、≥180 m²)下华北落叶松幼苗和幼树的更新密度、生长指标和空间分布。结果表明：在不同面积的林隙等级中，幼苗、幼树的生长指标(基径、高度)和幼苗更新密度均在小林隙(14～60 m²)下最好，幼树更新密度在中林隙(60～120 m²)下最大，且同等级面积林隙下幼树密度均大于幼苗密度。华北落叶松幼苗、幼树在小林隙和中林隙下更新状况良好，大林隙(120～180 m²)和特大林隙(≥180 m²)则不利于华北落叶松更新。华北落叶松幼苗、幼树主要分布在林冠投影区域以及林冠空隙区域的边缘。通过补种或间伐等人工干扰手段将林隙面积控制在14～120 m²,可以促进华北落叶松的更新。     

林窗对热带雨林冠层树种绒毛番龙眼幼苗生长的影响

在林窗中央、林窗边缘和林冠下3种不同光照梯度的森林生境中,研究了西双版纳季节雨林冠层树种绒毛番龙眼幼苗的早期 (种子萌发后10周内) 生长和定居后 (实生苗生长3个月以上) 的生长特点.结果表明: 绒毛番龙眼幼苗在早期生长阶段,林窗中央的株高、基径、总干质量、单株叶面积和相对生长率最大,分别为24.45 cm、3.17 mm、0.79 g、122.45 cm²和14.78×10^-3 g·d^-1.林冠下根冠比 (0.87) 高于林窗中央 (0.20) ,可能是光照和水分共同作用的结果.林窗中央较强的光照有利于定居后幼苗的生长,株高、基径、总干质量、单株叶面积、相对生长率和净同化率均在林窗中央最大,实验结束时分别达到31.48 cm、3.80 mm、2.22 g、174.52 cm²、2.29×10^-3 g·d^-1和2.54×10^-5 g·cm^-2·d^-1.幼苗死亡可能与水分胁迫密切相关,由水分胁迫引起的幼苗死亡率在林冠下最高 (26.88%),但林冠下由脊椎动物捕食引起的幼苗死亡率较低(2.93%),从而使林冠下幼苗的最终存活率最高 (70.19%).光照是影响绒毛番龙眼幼苗形态学调节的重要因素, 林窗中央不同生长阶段幼苗的比叶面积最低,但相对生长率和净同化率最大.水分胁迫和光照在幼苗定居后仍是影响幼苗生物量分配的重要因素, 林窗边缘幼苗的根冠比最高 (0.33).     

影响不同林型天然红松混交林林隙更新的土壤特征因子

采用野外调查、样品采集和统计分析等相结合的方法,对小兴安岭天然红松混交林3种不同林型(椴树红松混交林(TP)、枫桦红松混交林(BP)、云冷杉红松混交林(PAP))的林隙及其邻近郁闭林分的土壤特征因子和树木更新的相关性进行了研究,旨在阐明林隙土壤特征因子对树木更新的影响,从而为小兴安岭天然红松混交林植被更新、退化生态系统的恢复和可持续经营提供基础数据和实践参考。结果表明:郁闭林分土壤有机质、全氮质量分数显著高于3种不同林型的林隙。有效磷和速效钾含量在BP内与其他林型之间差异显著。3种林型林隙内p H值均略高于其郁闭林分,但与其差异均不显著。3种林型林隙内更新总密度、幼树更新密度与郁闭林分差异显著(P0.05),PAP林隙中更新总密度和幼树更新密度最高。BP林隙面积与更新密度相关不显著,乔木幼苗、幼树更新密度与有机质(r=-0.400,r=-0.475)、全氮均呈显著负相关(r=-0.519,r=-0.603)。TP林隙内全氮与乔木幼苗更新密度呈正相关(r=0.092),而与乔木幼树更新密度呈显著负相关(r=-0.585)。PAP林隙内全氮与乔木幼苗、幼树更新密度均呈负相关。郁闭林分幼苗更新密度分别与有机质、全氮、速效钾、p H值、脲酶和蛋白酶呈负相关。主成分分析表明,全氮是影响林隙和郁闭林分树木更新的关键因素。     

天童常绿阔叶林林窗的地形分布格局

为探究多维地形因素对林窗分布的影响,以天童20 hm²常绿阔叶林动态监测样地内的林窗为对象,结合地理信息系统软件,分析了林窗空隙率、林窗密度以及林窗面积在海拔、坡度、坡向、坡形、坡位5种地形因子下的分布格局.结果表明: 该样地林窗空隙率为13.1%,林窗密度为9.5个·hm^-2,林窗平均面积为137.82 m²;因高海拔台风干扰的强度较大,高海拔段(≥500 m)林窗的空隙率和密度显著大于中低海拔段(<500 m);台风带来的强降雨极易引起小范围滑坡,使得沟谷的林窗空隙率和密度显著大于侧坡,林窗平均面积显著大于侧坡和山脊.台风及其带来的强降雨是造成林窗在海拔及坡位梯度上具有显著性差异的主要原因.     

天童常绿阔叶林林窗的地形分布格局

为探究多维地形因素对林窗分布的影响,以天童20 hm²常绿阔叶林动态监测样地内的林窗为对象,结合地理信息系统软件,分析了林窗空隙率、林窗密度以及林窗面积在海拔、坡度、坡向、坡形、坡位5种地形因子下的分布格局.结果表明: 该样地林窗空隙率为13.1%,林窗密度为9.5个·hm^-2,林窗平均面积为137.82 m²;因高海拔台风干扰的强度较大,高海拔段(≥500 m)林窗的空隙率和密度显著大于中低海拔段(<500 m);台风带来的强降雨极易引起小范围滑坡,使得沟谷的林窗空隙率和密度显著大于侧坡,林窗平均面积显著大于侧坡和山脊.台风及其带来的强降雨是造成林窗在海拔及坡位梯度上具有显著性差异的主要原因.     

地形对阔叶红松林幼苗更新的影响

依托黑龙江凉水国家级自然保护区9 hm²典型阔叶红松林动态监测样地的900个2 m×2 m多年生幼苗(H≥30 cm,DBH<1 cm)样方,基于2006、2008和2010年3次调查数据,分析了地形对幼苗建立的影响.结果表明: 样地内共有乔木幼苗26种,2006、2008和2010年的乔木幼苗总数分别为4514、6464和5611株·hm^-2.其中个体数前10位树种的幼苗数量占幼苗总数的90%以上.地形对8个主要乔木幼苗的分布有显著的影响,其中暴马丁香、冷杉、色木槭、春榆、花楷槭、紫椴和青楷槭幼苗与其成树分布基本一致,而红松幼苗的空间分布与成树分布有所不同.暴马丁香、冷杉、裂叶榆、青楷槭和紫椴幼苗的死亡与地形显著相关.暴马丁香、春榆、红松、冷杉、水曲柳、色木槭和紫椴新增幼苗与地形显著相关.
     

广东南澳岛不同森林群落的林窗环境对3种移植树苗生长的影响

 先锋种和非先锋种可能对林窗产生不同的反应。在广东南澳岛4个不同的森林群落样地里移植了大叶相思（Acacia auriculaeformis A. Cunn.）、尾叶桉（Eucalyptus urophylla S. T. Blake）和荷木（Schima superba Gardn. et Champ）3种树苗,其中大叶相思和尾叶桉是先锋种,而荷木是非先锋种,测定了林窗和非林窗生境中的透光率和幼苗生长高度。结果表明,林窗的透光率大于非林窗的;各群落的透光率有季节性的变化,变化幅度各不同。3种幼苗在林窗内比     

闽北闽粤栲天然林林隙物种数量特征及更新规律分析

以福建省顺昌县的闽粤栲〔Castanopsis fissa(Champ.ex Benth.)Rehd.et Wils.〕天然林为研究对象,根据林隙面积(A)划定4个等级(A<50 m2、50 m2≤A<100 m2、100 m2≤A<150 m2和150 m2≤A<200 m2)的样地,并以非林隙样地为对照,对林隙和非林隙的乔木层、灌木层和灌草层中的物种分布、组成及生长状况进行调查和统计;在此基础上,对林隙和非林隙中主要物种的数量特征和更新响应规律进行比较分析.结果表明:闽粤栲天然林林隙和非林隙的乔木层分别有54和35种植物,共有种28种;灌木层分别有61和37种植物,共有种29种;灌草层分别有72和49种植物,共有种37种.从乔木层和灌木层各种类的重要值及灌草层各种类的密度看,林隙和非林隙中的优势种类均为闽粤栲和木荷(Schima superba Gardn.et Champ.)等,且林隙和非林隙间各优势种类的重要值差异不明显,其中,闽粤栲的重要值和密度均位居第一.从林隙和非林隙间各优势种类的重要值差值看,林隙的主要优势种类(如闽粤栲和木荷等)对林隙更新响应不明显,而其他种类对林隙则有强烈或中等的正、负更新响应,且在不同林层间有差异.从平均更新密度看,林隙面积对各优势树种的更新有一定影响,其中,林隙面积在150~200 m2时闽粤栲和丝栗栲(Castanopsis fargesii Franch.)的平均更新密度最大,林隙面积在100~150 m2时米槠〔Castanopsis carlesii(Hemsl.)Hay.〕、木荷、青冈〔Cyclobalanopsis glauca(Thunb.)Oerst.〕和黄毛润楠(Machilus chrysotricha H.W.Li)的平均更新密度最大,但在不同面积林隙间各树种的平均更新密度无显著差异(P=0.867).研究结果说明:在闽粤栲天然林中,对林隙更新响应不明显的种类较易成为林分的优势种或建群种,但林隙面积对树种的更新密度没有明显影响;从林隙到林冠的发育过程中,闽粤栲始终处于该林分的优势地位.     

秦岭山地锐齿栎次生林幼苗更新特征

分析了秦岭山地锐齿栎次生林群落乔木及木本幼苗物种特征值、更新生态位宽度及不同影响因子下（包括林分密度、坡向、海拔等）乔木更新动态。结果表明,乔木层物种有25种,锐齿栎占有明显地优势,重要值为149.18%。其次为华山松(Pinus armandii)和千斤榆(Carpinus cordata)等;更新层共有木本植物41种,其中乔木种有28种,占所有木本植物总数量的68%,优势种有青蛙皮槭(Acer grosseri)、木姜子(Litsea pungens)等。林下有丰富的幼苗库,径级及龄级比较小。更新方式以实生为主;分析了10种更新优势乔木种群的生态位宽度,同种幼树种群生态位宽度均大于幼苗种群生态位宽度。幼苗种群生态位宽度从大到小依次为榛子(Corylus heterophylla)、锐齿栎、青蛙皮槭等。幼树种群生态位宽度从大到小依次为榛子、青蛙皮槭、锐齿栎等;林分密度对林下乔木幼苗和幼树数量影响不同。当林分密度（株?hm-2）从720增加到1460时,幼苗密度逐渐增大。随后随着林分密度的增加,又呈现出减少的趋势。而幼树密度随着林分密度增加一直呈现出减少的趋势。坡向对锐齿栎次生林林下乔木幼苗和幼树的影响各异,阳坡（南偏西3）均利于幼苗和幼树的更新。南偏西45林分内,幼苗密度急剧减少。随着坡向转为阴坡,幼苗密度又逐渐增加。而幼树密度逐渐减少。当海拔从1083 m增加到1547 m时,幼树密度逐渐增加。随后,随着海拔增加到1882 m,幼树密度则逐渐减小。而幼苗密度随着海拔的增高一直呈现减小的趋势。     

Differential seed and seedling predation by crabs: impacts on tropical coastal forest composition

Recently, the importance of seed predation by crabs on mangrove species distributions and densities has been established by several studies. In a tropical coastal terrestrial forest in Costa Rica, we investigated the relative importance of predation by land crabs, Gecarcinus quadratus, and hermit crabs, Coenobita compressus, on measured forest composition through a series of seed removal and seedling establishment experiments. We also used natural light-gaps and adjacent non-gap sites to test how canopy cover affects crab predation (seed removal) and seedling establishment. We found fewer tree species (S=18) and lower densities (seedlings, saplings, and adults) in the coastal zone within 100 m of coastline, than in the inland zone (S=59). Land crab densities were higher in the coastal zone (3.03±1.44 crabs m^–2) than in the inland zone (0.76±0.78 crabs m^–2), and hermit crabs were not present in the inland zone. Seed removal and seedling mortality also were higher in the coastal zone than in the inland zone, and in the open controls than in the crab exclosures. Mortality of seeds and seedlings was two to six times higher in the controls than exclosures for four of the five experiments. Crabs preferred seeds and younger seedlings over older seedlings but showed no species preferences in the seed (Anacardium excelsum, Enterolobium cyclocarpum, and Terminalia oblonga) and seedling (Pachira quinata and E. cyclocarpum) stages. We conclude that the observed differences in tree densities were caused by differential crab predation pressure along the coastal gradient, while the differences in species composition were due to predator escape (satiation) by seed quantity. Canopy cover did not affect seed removal rates, but did affect seedling survival with higher mortality in the non-gap versus gap environments. In summary, crab predation of seeds and seedlings, and secondarily canopy cover, are important factors affecting tree establishment in terrestrial coastal forests.     

白皮松林林隙物种组成动态及更新规律

研究了白皮松林林隙物种组成和种间关系在林隙发育的各个阶段的动态变化,并对林隙内主要树种的更新反应进行了探讨。结果表明,在发育阶段为前期、中前期、中后期和后期的序列上,林隙中乔木层优势度最大的物种分别是白皮松（Pinus bungeana）、辽东栎（Quercus wutaishanica）、山桃（Prunus davidiana）和侧柏（Platy-cladus orientalis）;灌木层优势种包括白莲蒿（Artemisia gmelinii）、胡枝子（Lespedeza bicolor）、薄皮木（Lepto-dermis oblonga）和狼牙刺（Sophora viciifolia）;草本层优势种包括大披针苔草（Carex lanceolata）、糙隐子草（Cleistogenes squarrosa）、紫菀（Aster tataricus）和荩草（Arthraxon hispidus）;非林隙林分的优势种不明显。在林隙发育过程中乔木层、灌木层、草本层的平均相似性系数为0.32、0.38和0.27。可见林隙发育过程中群落各层次的相似性系数由小到大顺序为：草本〈乔木〈灌木;草本植物随着林隙环境条件的改变而出现较大的消亡和更新,显示出较大的波动性;乔木的稳定性和连续性介于灌木和草本之间。依据林隙中树种幼苗、幼树Ⅰ和幼树Ⅱ的数量差异,可将林隙更新反应分为单峰型、双峰型、差异不明显3种,并分别以白皮松、辽东栎和油松（Pinus tabu-laeformis）为代表树种。     

Tree seed germination and seedling establishment in treefall gaps and understorey in a subtropical forest of northeast India

Abstract Seed germination, and survival and growth of seedlings of four dominant tree species, Quercus dealbata, Quercus griffithii, Quercus glauca and Schima khasiana were studied in the treefall gaps and forest understorey of an undisturbed mature-phase humid subtropical broadleaved forest in northeast India. Three important microenvironmental factors namely photosynthetically active radiation (PAR), soil moisture and litter depth, were also measured in the forest understorey and gaps and correlated with seedling mortality. Seed germination of S. khasiana was significantly higher in the treefall gaps than in the understorey; among the tree species studied, it had the highest germination. Quercus seedlings were abundant in the understorey and small gaps, while S. khasiana seedlings were more numerous in the large gaps. The survivorship curves for the seedling populations revealed that the three Quercus species survived better in the understorey, while S. khasiana did so in the gaps. PAR and soil moisture were positively correlated with tree seedling mortality, which occurred mainly during the winter months. The Quercus seedlings grew better in the forest understorey and small gaps and S. khasiana seedlings in the large gaps. The differential performance of the tree seedlings to the conditions prevailing in the understorey and gaps of two sizes indicates that different species were adapted to different light environments depending upon their optimum requirements. This could be an effective mechanism for promoting species coexistence in the forest community.     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

Analysis of seedling banks of a climax beech forest: ecological importance of seedling sprouts

Most of the common and dominant tree species in gaps of a climax Japanese beech (Fagus crenata) forest have large seedling banks on the forest floor. These seedlings were classified into “true seedlings” and “seedling sprouts”, each of which was further classified into two and three subtypes, respectively. Each species had a characteristic seedling form spectrum. The seedling bank of beech was composed almost solely of true seedlings, whereas, seedling sprouts predominated in the seedling banks of lower tree species such as Prunus grayana Maxim. and some species of Acer. The age structures suggested that seedling sprouts had longer life spans and had been recruited more constantly than true seedlings. The habit of sprouting and creeping seemed to enable seedlings of these species to persist for long periods, thus increasing their chance for eventual success in canopy gaps.     

Study on canopy disturbance regime and mechanism of tree species diversity maintenance in the lower subtropical evergreen broad-leaved forest,South China

ABSTRACT

The canopy disturbance, the gap environment, gap regeneration and maintenance of tree species diversity in the lower subtropical evergreen broad-leaved forest (LSEBF) of South China were studied in this paper. The most common manner of gap formation in the forest was by stem breakage. Most gaps were formed by two gap makers. The sizes of most expanded gaps (EG) and canopy gaps (CG) were in the range of 100 – 300 m² and 50 – 100 m², respectively. The ecological factors in gaps were analysed on the basis of contrasting measurement of the microclimatic regimes in gaps of different sizes and in non-gap stands. Tree species in the LSEBF were classified into 5 ecological species groups on the bases of their changes in order of importance values in gaps and in non-gap stands. Most of the species reached their peak of regeneration density around the gap sizes of 100 m² and 500 m². The curves of regeneration density vs. gap age for major species revealed two types. Regeneration densities of most species, and species diversity indices in gaps were greater than those in non-gap stands.     

Spatial structure and regeneration of Tetramerista glabra in peat swamp rain forest in Indonesian Borneo

We examined the spatial structure and regeneration of Tetramerista glabra, a dominant canopy tree in peat swamp rain forest in Borneo (West Kalimantan, Indonesia). T. glabra has strong spatial structure that changes dramatically during the life cycle; seedlings were highly aggregated, saplings were random and trees were evenly distributed. Germination and seedling relative growth were highest within canopy gaps, but seedling densities were highest at gap edges. Concentration of seedlings in gap edges, rather than localized seed dispersal, was responsible for seedling patchiness. The slope of the relationship between relative growth rate and seedling height decreased from gap, to gap edge to understory habitats, suggesting that competition within the seedling layer is more important in gaps than in the understory. The processes that break down seedling aggregation, leading to over-dispersion of trees, must be density dependent, but remain unknown.