基于地形因子的天童地区常绿树种和落叶树种共存机制研究

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种,那么它们与常绿树种的共存机制是什么?常绿树种和落叶树种生态习性差异较大,二者对生境的选择应有所不同,我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设,我们以天童20ha动态样地调查数据为依托,选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象,用典范对应分析(CCA)研究了地形因子对二者分布的影响差异,用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下:(1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%,对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明,与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%,成负关联的比例分别为40%和7%;与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%,成负关联的比例分别为10.9%和47.6%;与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种;两个植物类群对生境的选择多呈现相反格局,尤其是在沟谷生境和山脊生境,这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一,生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。     

地形异质性对云南普洱季风常绿阔叶林物种多样性的影响

地形异质性通过调控树木生长所需的养分、水分和光照等而成为亚热带森林结构与物种组成的重要驱动因子。但是, 地形异质性对季风常绿阔叶林物种多样性及其分布影响的研究还相对较少。该文基于云南普洱30 hm²森林动态监测样地(大样地) 750个20 m × 20 m的样方调查数据, 以海拔、坡度、凹凸度和坡向4个地形因子为变量, 采用C均值模糊聚类分析大样地的地形类型, 进而分析不同地形条件下的群落物种组成及群落物种多样性; 采用Torus转换检验法, 探讨物种与地形关联性, 为季风常绿阔叶林生物多样性保护提供科学依据。研究结果表明, 大样地可分为山脊、陡坡、缓坡、高谷和沟谷等5类地形, 地形面积分别是8.00、6.04、7.68、2.76和5.52 hm²。大样地中胸径(DBH) ≥ 1 cm的木本植物个体153 418株, 分属79科179属271种。5类地形中, 物种丰富度、不同径级的植株密度和比例明显不同, 多样性及优势物种多度分布具有较大差异。种-面积曲线表明, 同等面积条件下, 随着取样面积增加, 山脊的物种丰富度始终最小, 高谷次之, 沟谷的物种丰富度始终最大。种-个体数累积曲线表明, 随着个体数增加, 山脊物种丰富度的累积速率最小, 种丰富度增加缓慢, 高谷次之。在被检验的123个物种中, 与地形相关的物种有83个, 高达67.5%的物种与至少一类地形存在显著相关关系。山脊和缓坡中与地形具有显著负相关关系的物种数超过显著正相关的物种数; 而与陡坡、高谷和沟谷显著正相关的物种数高于显著负相关的物种数。普洱大样地地形异质性对物种多样性维持的贡献率为7.8%。     

Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species corre-spond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species correspond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

南亚热带常绿阔叶林地上碳储量空间分布特征及其影响因素

旨在探讨南亚热带常绿阔叶林地上碳储量空间分布特征及其影响因素,为了解该区域森林的碳汇功能提供理论依据。通过对鼎湖山南亚热带常绿阔叶林20 hm²固定森林样地调查数据,采用一元线性回归分析和主成分分析方法,划分优势种和非优势种,研究地上碳储量的空间分布和生物/非生物因素的影响,获取了以下结果:(1)优势种对鼎湖山南亚热带常绿阔叶林地上碳储量贡献更大(1533.85 Mg,74.72%),尤其是大径级物种(1389.68 Mg,67.69%)。优势种地上碳储量(CV=0.635)的空间分布较非优势种(CV=0.690)更加均匀。(2)物种多样性与优势种和总体地上碳储量负相关,而与非优势种正相关。(3)群落总体、优势种和非优势种的地上碳储量均与结构多样性显著正相关。然而,结构多样性对非优势种地上碳储量的影响程度高于优势种。(4)群落中的土壤营养度、凹凸度与地上碳储量正相关。综上所述,通过提升非优势种的物种多样性可以增加鼎湖山南亚热带常绿阔叶林地上碳储量。此外,改善土壤营养条件也有助于提升南亚热带森林的碳汇功能。     

Coexistence mechanisms of evergreen,deciduous and coniferous trees in a mid-montane mixed forest on Mt. Emei,Sichuan, China

Structure and regeneration of a mid-montane (2200 m a.s.l.)mixed forest codominated by evergreen (Lithocarpuscleistocarpus), deciduous (Acer flabellatum)andconiferous (Tsuga chinensis, Abies fabri, andTaxus chinensis) trees were analyzed in a 40m× 60 m plot on Mt. Emei, Sichuan, China. Plant communitystructure and composition varied depending on topographic micro-habitat withinthe plot. Four topographic communities (topo-communities) were distinguishedwith dominant species corresponding to topography: (1)Abies – valley bank, (2) Acer– lower steep slope, (3) Lithocarpus – uppergentle slope, and (4) Tsuga – ridge. The coexistencemechanisms of the evergreen, deciduous, and coniferous trees were determined byidentifying the regeneration process characteristic of each dominant species,asrepresented by their seedling dispersion patterns and seedling establishmentalong the topographic gradients. The saplings and seedlings of the dominantswere distributed differently according to the topography:Lithocarpus under the canopy of parent trees and in gapsofthe upper gentle slope and ridge, Acer mainly in bothwell-lit and shady sites on the lower steep slope and on fallen logs,Abies on the valley bank but only in well-lit sites,Tsuga on the ridge, Taxus mostly onrock and well-lit sites on the slopes. Distribution of surviving saplings andseedlings was also related to the species of nearby canopy trees. We suggestthat Lithocarpus, Tsuga and Acer tendto be self-replacing in their own topographic habitats, andAbies survives as a fugitive by occupying occasionalsuitable gaps. The variation in soil conditions, particularly nutrients, withtopography affected seedling establishment and the growth of trees. Thesuccessional change of quantitative species composition, as predicted by theMarkovian model, shows the mixed forest to be in a sustained climax stage.     

Variation in tree growth,mortality and recruitment among topographic positions in a warm temperate forest

Questions: Do the population dynamics of trees differ among topographic positions and, if so, how does topographic position affect the population dynamics of species that are distributed in a topography‐specific manner? Which is the most important life stage in determining vegetation patterns? Location: Primary and secondary warm temperate evergreen broad‐leaved forest (40 ‐ 280 m a.s.l.) on the western part of Yakushima Island, Japan. Methods: Mortality, recruitment, DBH growth and distribution of stems (= 5 cm DBH) in a 2.62‐ha plot were surveyed in 1992 and 2002 to determine the relationships between population parameters and (1) topography and (2) distribution patterns of 17 common tree species. Results: Common species (n = 17) were classified into three distribution pattern groups: group A, distributed mainly on convex slopes; group B, on concave slopes, and group C, not aggregated with respect to topographic position. Stem mortality, recruitment and DBH growth were greater in group A than in group B within each topographic class. The hierarchy of stem mortality among topographic classes for groups A and B was convex > planar > concave. Stem recruitment density was relatively high on the convex and concave slopes, respectively, for groups A and B. Conclusions The topographical positions of adult trees were not always most suited for adult survival and growth. For group A, the distribution pattern of adults was determined in the juvenile stage, while this was not the case for group B. Studies of juvenile stages are important for understanding the demographic basis of vegetation distribution patterns.     

Functional diversity versus species diversity: relationships with habitat heterogeneity at multiple scales in a subtropical evergreen broad-leaved forest

Understanding the relationship between functional and species diversity as well as their association with habitat heterogeneity can help reveal the mechanisms of species coexistence in ecological communities. However, these interactions have been poorly studied in subtropical forests. In this paper, we evaluated functional diversity (as measured by Rao’s Q) and traditional species diversity (based on Simpson’s index) in a 24 ha forest plot in a subtropical evergreen broad-leaved forest (EBLF) in China. We compared the sensitivities of functional and species diversity to topographic variables (elevation, convexity, slope and aspect) at multiple spatial scales based on 10 × 10, 20 × 20, 40 × 40 and 50 × 50 m quadrats. Functional and species diversity were found to have different distribution patterns along a topographical gradient, with functional diversity better explained by topography than was species diversity using a spatial autocorrelation regression error model. Furthermore, functional diversity had a significantly greater association with topographic variables than species diversity in both adult and young trees; in both cases, the strength of the diversity-habitat association increased with quadrat size. We conclude that functional diversity reflects a greater diversity-habitat association in EBLF than does species diversity, and that the association depends on the spatial scale and life stages of the woody plants under evaluation.     

浙江天童国家森林公园常绿阔叶林凋落叶养分含量的时空分布特征

植物凋落叶养分含量的时空分布格局在森林生态系统的物质循环和能量流动中发挥着关键作用,尤其是对森林土壤养分情况有重要的影响,而在群落结构复杂的亚热带森林中基于群落整体水平的凋落叶养分含量特征研究相对匮乏。在位于浙江天童国家森林公园中的20 hm²动态监测样地进行,收集凋落叶并测定其碳、氮、磷含量,结合不同生活型物种的凋落高峰,春季和初夏为常绿树种凋落高峰;秋季为落叶树种凋落高峰,以及地形特征对凋落叶养分含量的时空分布特征进行探究。结果表明：在时间尺度上,凋落叶养分含量表现为不同凋落高峰之间的差异,碳含量表现为秋季高峰 < 春季高峰 < 初夏季高峰,氮、磷含量为秋季高峰大于春季和初夏季高峰;在空间尺度上,凋落叶养分含量则表现为沟谷地形和山脊地形之间的差异性,在秋季凋落高峰时碳含量在沟谷地形显著低于山脊地形,氮、磷含量在山脊地形显著低于其他地形。研究结果表明群落物种组成及其生活型差异是造成森林群落水平凋落叶养分含量时空差异的主要原因。     

Habitat differentiation of Lauraceae species in a tropical lower montane forest in northern Thailand

Dependency on topographical habitat was examined for Lauraceae tree species in a lower montane forest using a large-scale research plot established at Doi Inthanon National Park, northern Thailand. Twenty species of 10 genera of Lauraceae were recorded in a 7.5-ha part of the plot; Lauraceae accounted for 18% of the total basal area. Lauraceae was the most species-rich and most abundant family in the plot. In a cluster analysis based on the matrix of spatial associations between species, two clusters were recognized. Members of one cluster seemed to associate with lower-elevation habitats, and members of the other associated with habitats on ridges. By subdividing the study plot into 20m×20m squares, a discriminant analysis could be applied to the presence–absence data for the 17 species that had sufficient population density. The predictor variables used were the relative elevation, slope inclination, slope direction (transformed to deviation from SSW) and slope convexity for each of the squares. The discriminant models were tested statistically by applying the random shift technique. The models were significant for 11 of the species (65% of the species examined) and were associated with the topographical condition of the habitat. Stepwise selection of the predictor variables for these 11 species revealed that relative elevation and slope convexity were the most important factors for predicting the presence or absence of the Lauraceae species. Both these variables were considered to indicate the hydrological condition of the habitat.     

Fire survival of lowland tropical rain forest trees in relation to stem diameter and topographic position

The objective of this study was to relate patterns in forest structure, tree species diversity, and tree species composition to stem diameters and topography in unburned, once burned and twice burned lowland dipterocarp rain forests in East Kalimantan, Indonesia. To do this four unburned old growth forests were compared with three forests that burned once (1997/1998) and three forests that burned twice (1982/1983 and 1997/1998). Fire resulted in a strong reduction of climax tree density which was negatively related to tree diameter. However, a disproportionate reduction in small diameter understorey climax tree species occurred only after repeated fires. Climax tree species in both burned forest types were most common in swamps, river valleys and on lower slopes, while their density was much lower on places higher along hillsides. In unburned forest the opposite was observed, with climax tree density increasing steadily from swamp and river valleys to upper slopes and ridges. In contrast to climax trees, pioneer trees were abundant throughout the burned forest, with highest numbers on hill sides and ridges. Our results indicate that both diameter and topographic position of trees strongly affect their fire survival chances in tropical lowland forests.     

Saplings distribution in relation to topography and canopy openness in an evergreen broad-leaved forest

We examined the spatial distribution of saplings in relation to topography and stand structure in a subtropical evergreen broad-leaved forest in the northern part of Okinawa Island, Japan. The distributions of most species were influenced mainly by topography but also by canopy openness. Species were arranged along a topographical gradient from concave areas with low canopy openness to convex areas with high canopy openness. Canopy species were arranged along a gradient from unstable fertile areas to stable infertile areas. Species occurring mostly in convex areas had a narrower distribution range than those in concave sites. Thus, habitat heterogeneity provided by topography and stand structure appears to be important factors for maintaining high species diversity in the understory of this subtropical evergreen broad-leaved forest.     

Sampling strategies for the assessment of tree species diversity

Abstract. This paper aims at proposing efficient vegetation sampling strategies. It describes how the estimation of species richness and diversity of moist evergreen forest is affected by (1) sampling design (simple random sampling, random cluster sampling, systematic cluster sampling, stratified cluster sampling); (2) choice of species richness estimators (number of observed species vs. non-parametric estimators) and (3) choice of diversity index (Simpson vs. Shannon). Two sites are studied: a 28-ha area situated in the Western Ghats of India and a 25-ha area located at Pasoh in Peninsular Malaysia. The results show that: (1) whatever the sampling strategy, estimates of species richness depend on sample size in these very diverse forest ecosystems which contain many rare species; (2) Simpson's diversity index reaches a stable value at low sample sizes while Shannon's index is affected more by the addition of rare species with increasing sample size; (3) cluster sampling strategies provide a good compromise between cost and statistical efficiency; (4) 300 - 400 sample trees grouped in small clusters (10–50 individuals) are enough to obtain unbiased and precise estimates of Simpson's index; (5) the local topography of the Western Ghats has a major influence on forest composition, the steep slopes being richer and more diverse than the ridges and gentle slopes; (6) stratified cluster sampling is thus an interesting alternative to systematic cluster sampling.     

鼎湖山南亚热带常绿阔叶林荷木种群动态变化

荷木(Schima superba)是鼎湖山南亚热带常绿阔叶林群落中的优势种,为了解南亚热带常绿阔叶林的群落构建机制,该研究分析了鼎湖山南亚热带常绿阔叶林20 hm2样地中荷木种群在2005—2017年间的动态变化,并从径级和生境两个方面分析了其死亡率的分布特征。结果表明:(1)荷木种群个体数量从2005—2017年呈现出减少的态势。(2)在径级方面,荷木的死亡率和相对生长率的变化在不同径级区间呈现出不同的变化规律。径级在1～10 cm范围的荷木相对生长率大并迅速下降,死亡率上升且维持在较高水平;径级在10～50 cm范围的荷木随着径级增大,相对生长率减缓,然后下降,而死亡率不断下降;在50 cm以上的径级范围内,荷木相对生长率小、死亡率大。(3)不同生境中,2010年各个生境死亡率相差不大,2015年高海拔生境(高坡、山脊、高谷)死亡率大于低海拔生境(低坡、低谷),2017年则为低海拔生境大于高海拔生境。(4)荷木的死亡率与土壤中酸碱度呈正相关关系,而与土壤有机质、总氮、总磷、有效钾、有效氮含量呈负相关关系。     

Species–topography association in a species-rich subtropical forest of China

Habitat specialization has been considered as a primary factor in determining the distribution of species. In this study, we investigated species–habitat associations while controlling for spatial neighbourhood effects in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. Habitat specialization was measured by topographic variation and its effects on species distributions were modelled at three different spatial scales (10×10, 20×20 and 25×25 m²) using log-linear regression models and randomization tests. Our results showed: (1) 83% of the species were related to at least one or more topographic variables. Among them, 66%, 60%, 65% and 70% were closely dependent on slope, aspect, elevation and convexity, respectively. (2) Topographic variables have much stronger non-linear (quadratic and cubic) effects on species distributions than linear effects. (3) The effects of topographic heterogeneity on the distribution of shrubs species are smaller than on the distribution of canopy species, and smaller effects were also found in less abundant species. (4) There was a strong neighbourhood effect on species distribution: In 85% of the species, abundance in a focal quadrat was significantly correlated with abundance in the neighbour quadrats. We conclude that habitat specialization plays an important role in maintaining the diversity of this species-rich subtropical forest.     

地形对阔叶红松林物种多样性的影响

为了更好地了解阔叶红松林(Pinus koraiensis)物种多样性的维持机制,对不同地形下的阔叶红松林乔木和灌木种,进行了物种多样性指数比较、物种多度分布模型拟合及物种多样性指数与环境因子的排序分析。结果表明,乔木在阴坡、半阴坡的丰富度显著高于半阳坡;灌木在坡度25°处的均匀度显著高于坡度6°处,在谷地处的4个多样性指数显著低于其他坡位,在阴坡、半阴坡处的4个多样性指数显著高于平地。统计模型、生态位模型、中性模型在不同地形下均能够通过拟合优度检验,物种多度分布在不同地形间差异不明显;地形对乔木的多样性指数的影响:海拔凹凸度坡度坡向坡位,对灌木的多样性指数的影响:坡位海拔坡向坡度凹凸度,土壤对乔木的物种多样性指数的影响:土壤pH土壤容重土壤有效N含量土壤速效K含量,对灌木的物种多样性指数的影响:土壤pH土壤有机质含量土壤速效P含量土壤全N含量土壤全P含量。总体而言,物种多样性随着坡度的增加而增加,阴坡处最高,谷地处最低。     

古田山常绿阔叶林优势树种甜槠和木荷的空间格局分析

基于24 hm²古田山中亚热带常绿阔叶林长期监测样地调查资料, 采用Ripley的K函数点格局分析等方法, 具体分析了古田山常绿阔叶林优势树种甜槠(Castanopsis eyrei)与木荷(Schima superba)不同年龄阶段的空间分布格局以及它们之间的空间关联性。结果表明: 两种优势物种在总体上及不同年龄阶段主要呈聚集分布, 且随年龄阶段的增加, 聚集程度有降低的趋势。两物种在大尺度直观上有明显的生境偏好。两种优势物种的幼树、小树与大树和老树主要呈空间负相关或无空间关联性, 与中树呈空间正相关。甜槠中树与大树、大树与老树主要呈空间正相关, 而甜槠的中树与老树呈空间负相关。木荷中树与大树和老树、大树与老树均呈现空间负相关或无关联。我们发现古田山优势物种通过密度制约和Janzen-Connell效应释放空间, 为其他物种共存提供了条件, 密度制约和Janzen-Connell效应可能是古田山样地甜槠和木荷空间格局形成的重要原因。     

南岭国家级自然保护区林下植物分布的地形相关性

以南岭国家级自然保护区山地森林群落林下植物为研究对象,探讨地形因子对林下植物分布的影响。该区3个海拔高度下的山地常绿阔叶林林下植物组成丰富,但个体多度在样方中的分布存在很大的空间异质性。在9000m^2的样地中设置的450个4m^2小样方,共记录到林下维管植物255种,隶属于95科、181属,总的物种个体多度为18203株。多响应置换过程（MRPP）分析表明,坡向和海拔（p〈0．0001）及坡度（p〈0．05）这3个地形因子对林下植被分布有显著的影响,其影响程度大小排序为：海拔〉坡向〉坡度。指示种分析（ISA）进一步确定了不同地形条件下具有显著指示值（IV≥50）的指示种。本项研究表明林下植物的空间分布会同时受到多个地形因子的影响,因此需综合各因子间的相互作用全面考虑;同时,MRPP结合ISA对于揭示群落植物分布的空间异质性及其与环境因子作用的相互关系、生物多样性保育、森林恢复以及造林引种等方面均有着重要的理论意义和实践价值。     

Canopy structure in a temperate old-growth evergreen forest analyzed by using aerial photographs

We used aerial photographs to create a digital elevation model of the canopy surface of a 10-ha study area in a temperate old-growth evergreen forest. A topographic map of the ground surface in a 4-ha permanent plot within the study area was also drawn from ground measurements. The difference between the two elevation values (i.e., canopy surface – ground surface) at each point in a 5-m grid was considered to be the canopy height, and a canopy height profile was constructed from these data. The canopy structure in the 4-ha plot that was estimated in this way was compared with that obtained by two ground observation methods, i.e., the canopy (vegetation) height profile method and the canopy coverage census method. Large gaps were adequately detected by the aerial photograph method, but small gaps were less often detected. Gap size distribution obtained by the aerial photograph method was similar to that observed on the ground, and was a function of gap depth. This study indicates that if a detailed topographic map can be made, the canopy height profile derived from aerial photography can be effective in analyzing the canopy structure of evergreen forests, such as tropical rain forests, over large areas.