Vertical distribution of epiphytic bryophytes in an Indonesian rainforest

We studied species richness, composition and vertical distribution of epiphytic bryophytes in submontane rainforest of Central Sulawesi. Bryophytes were sampled on eight canopy trees and on eight trees in the forest understorey. Microclimate was measured at trunk bases and at crown bases. The total recorded number of 146 epiphytic bryophyte species is among the highest ever reported for tropical forests and underlines the importance of the Malesian region as a global biodiversity hotspot. Species composition differed significantly between understorey trees and canopy tree trunks on the one hand, and the forest canopy on the other. Fourty-five percent of the bryophyte species were restricted to canopy tree crowns, 12% to the understorey. Dendroid and fan-like species mainly occurred in the forest understorey whereas tufts were most species rich in the tree crowns. The findings reflect the different microclimatic regimes and substrates found in the understorey and in the forest canopy. The results indicate that assessments of the bryophyte diversity of tropical forests are inadequate when understorey trees and tree crowns are excluded.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Vertical stratification and effects of crown damage on maximum tree height in mixed conifer–broadleaf forests of Yakushima Island,southern Japan

We investigated vertical stratification and effects of crown damage on maximum tree height in two mixed conifer–broadleaf forests in Yakushima Island, southern Japan. In both research plots, the conifer trees dominated the upper canopy while the broadleaved trees dominated the middle to lower canopy. Most broadleaved trees were shorter than the median crown-base height (H _CB) of the conifer trees. Estimates of the maximum height (H _max) of the conifer trees were greater than those of the broadleaved trees. Crown damage had significant negative effects on maximum height of the conifer trees. Crown damage was observed for 72.8–88.7% of the conifer trees, and severe types of damage such as stem breakage and top die-back were the most predominant. The H _max of the damaged conifer trees was 16–17% shorter than that of the intact trees and as much as 16–28% shorter than the potential maximum height estimated from the diameter–height relationship of the tallest intact trees. We inferred that crown disturbance is an important factor determining the maximum height of the canopy of the two mixed forests. Our results suggested that vertical stratification between conifer and broadleaved trees may be an important mechanism contributing to their coexistence and additive basal area of mixed forests on Yakushima Island.     

Age‐related Crown Thinning in Tropical Forest Trees

Gap dynamics theory proposes that treefall gaps provide high light levels needed for regeneration in the understory, and by increasing heterogeneity in the light environment allow light‐demanding tree species to persist in the community. Recent studies have demonstrated age‐related declines in leaf area index of individual temperate trees, highlighting a mechanism for gradual changes in the forest canopy that may also be an important, but less obvious, driver of forest dynamics. We assessed the prevalence of age‐related crown thinning among 12 tropical canopy tree species sampled in lowland forests in Panama and Puerto Rico (total N = 881). Canopy gap fraction of individual canopy tree crowns was positively related to stem diameter at 1.3 m (diameter at breast height) in a pooled analysis, with 10 of 12 species showing a positive trend. Considered individually, a positive correlation between stem diameter and canopy gap fraction was statistically significant in 4 of 12 species, all of which were large‐statured canopy to emergent species: Beilschmiedia pendula, Ceiba pentandra, Jacaranda copaia, and Prioria copaifera. Pooled analyses also showed a negative relationship between liana abundance and canopy gap fraction, suggesting that lianas could be partially obscuring age‐related crown thinning. We conclude that age‐related crown thinning occurs in tropical forests, and could thus influence patterns of tree regeneration and tropical forest community dynamics.     

Variation in crown light utilization characteristics among tropical canopy trees

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.     

Flooding × tree fall gap interactive effects on blackwater forest floristics and physical structure in the Peruvian Amazon

To better understand how flooding and tree fall structure forests in the Amazon, I sampled vegetation plots within three blackwater forests (least flooded, medium flooded, highest flooded) and their tree fall gaps in the Peruvian Amazon. I found (1) increased flooding decreased family richness in the closed-canopy forests but increased it in their gaps, with no trends for no. of unique species, (2) flooding decreased stem size everywhere as did the number of stems as size increased especially for larger stems, (3) Green's index showed clumping only for least-flooded forests with the closed-canopy forest showing more than its tree fall gap, and (4) both flooding and tree fall gap creation decreased canopy coverage perhaps as an additive effect. Further among the stem size classes, only the smallest stems were significantly affected by openness and by type of forest, with a significant interaction term where flooding significantly decreased the number of these smaller stems in all forests and their gaps, except those with the highest level of flooding. Also tree fall gaps had significantly more, smaller stems than their forests in the two most-flooded forests, but not in the least-flooded forest. I conclude that flooding is a greater stressor and influence on the structure of these forests than tree fall, and so, in these forests, gradients and disturbances overlap in their traditional roles.     

温带针阔混交林叶片性状随树冠垂直高度的变化规律

自然界中,森林植物叶片的生长随树冠高度呈现明显的垂直分布现象;然而,有关叶片性状随着树冠垂直高度增加的变化规律仍不清楚。为了更好地揭示植物叶片对光环境变化的适应策略以及对资源的利用能力,有必要深入探讨叶片性状与冠层高度的定量关系及其内在调控机制。以中国广泛分布的温带针阔混交林为对象,选取8种主要树种为研究对象(白桦、蒙古栎、水曲柳、大青杨、色木槭、千金榆、核桃楸和红松),通过测定这些物种9个冠层高度的叶片比叶面积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(N)、叶片磷含量(P)、氮磷比(N∶P)和叶绿素含量(Chl)等属性,探讨了针阔混交林叶片性状的差异以及各性状之间的相关关系,进而揭示叶片性状随树冠垂直高度的变化规律。实验结果表明:1)温带针阔混交林内优势树种的部分叶片性状在不同冠层高度之间差异显著。2)随着树冠垂直高度的增加,SLA、LDMC、N、P、N∶P和Chl呈现不同的变化趋势。其中,阔叶树种SLA随着树冠垂直高度的增加而减小;所有树种的LDMC随着树冠垂直高度的增加而增加;不同树种的N、P、N∶P和Chl随着树冠垂直高度的变化规律存在差异。3)对于温带针阔混交林冠层中,SLA与N、P、N∶P均存在显著的正相关关系,高SLA伴随着高的N、P、N∶P,表明植物通过SLA与N、P等性状的协同来提高叶片的光合作用(或对光热资源的利用效率)。本研究通过定量分析探讨温带针阔混交林叶片性状随冠层高度的变化规律,一定程度地揭示了树木对光、热和水资源竞争的适应机制,以及植物叶片的资源利用和分配策略,不仅拓展了传统性状研究的范畴,其相关研究结论也有助于树木生长模型的构建和优化。     

The functional trait space of tree species is influenced by the species richness of the canopy and the type of forest

Analysing how species modify their trait expression along a diversity gradient brings insight about the role that intraspecific variability plays over species interactions, e.g. competition versus complementarity. Here, we evaluated the functional trait space of nine tree species dominant in three types of European forests (a continental‐Mediterranean, a mountainous mixed temperate and a boreal) growing in communities with different species richness in the canopy, including pure stands. We compiled whole‐plant and leaf traits in 1719 individuals, and used them to quantify species trait hypervolumes in communities with different tree species richness. We investigated changes along the species richness gradient to disentangle species responses to the neighbouring environment, in terms of hypervolume size (trait variance), shape (trait relative importance) and centroid translation (shifts of mean trait values) using null models. Our main results showed differences in trait variance and shifts of mean values along the tree diversity gradient, with shorter trees but with larger crowns in mixed stands. We found constrained functional spaces (trait convergence) in pure stands, suggesting an important intraspecific competition, and expanded functional spaces (trait divergence) in two‐species admixtures, suggesting competition release due to interspecific complementarity. Nevertheless, further responses to increasing species richness were different for each forest type, waning species complementarity in sites with limiting conditions for growth. Our results demonstrate that tree species phenotypes respond to the species richness in the canopy in European forests, boosting species complementarity at low level of canopy diversity and with a site‐specific pattern at greater level of species richness. These outcomes evidence the limitation of functional diversity measures based only on traits from pure stands or general trait database values.     

Mapping Tropical Forest Trees Using High‐Resolution Aerial Digital Photographs

The spatial arrangement of tree species is a key aspect of community ecology. Because tree species in tropical forests occur at low densities, it is logistically challenging to measure distributions across large areas. In this study, we evaluated the potential use of canopy tree crown maps, derived from high‐resolution aerial digital photographs, as a relatively simple method for measuring large‐scale tree distributions. At Barro Colorado Island, Panama, we used high‐resolution aerial digital photographs (~0.129 m/pixel) to identify tree species and map crown distributions of four target tree species. We determined crown mapping accuracy by comparing aerial and ground‐mapped distributions and tested whether the spatial characteristics of the crown maps reflect those of the ground‐mapped trees. Nearly a quarter (22%) of the common canopy species had sufficiently distinctive crowns to be good candidates for reliable mapping. The errors of commission (crowns misidentified as a target species) were relatively low, but the errors of omission (missed canopy trees of the target species) were high. Only 40 percent of canopy individuals were mapped on the air photographs. Despite failing to accurately predict exact abundances of canopy trees, crown distributions accurately reproduced the clumping patterns and spatial autocorrelation features of three of four tree species and predicted areas of high and low abundance. We discuss a range of ecological and forest management applications for which this method can be useful.     

多样化松林中昆虫群落多样性特征

马尾松和湿地松是我国南方的2种主要松树。通过对6种不同林分结构下的马尾松林和湿地松林内昆虫群落调查与多样性指数分析,表明2种松树内的昆虫种类和数量无显著差异,混交林中的昆虫群落的种类和数量比纯林多,尤其以捕食天敌类群的种类和数量更为明显。整个昆虫群落和植食类群多样性指数以湿地松林内较大,而天敌(捕食类群和寄生类群)多样性指数则以马尾松林较高。从不同林分结构下昆虫多样性的比较来看,混交林内昆虫群落多样性指数波动较小,明显地高于纯林。但不同林分结构下昆虫多样性随水平分布和垂直分层格局而变化,松树北面和东面各样地之间的昆虫群落多样性指数差异显著,而南、西面之间差异较小;树冠层各样地之间的差异达极显著水平,而枯枝落叶层和树干层之间差异不显著。由此,还进一步讨论了混交林中昆虫群落稳定性问题。     

吉林东部天然林林下光质分布特征

天然林林下光质对乔木幼苗以及灌草的组成与更新具有重要的生态学意义。但目前对于林下光质的研究仍然有限。以吉林东部地区天然林为例,通过调查乔木数据和林下光质数据,基于移动窗口法分析不同空间尺度森林冠层结构与林下光质的关系。结果表明：不同林型下红光光子通量密度(R)与蓝光光子通量密度(B)存在差异。其中沙松-千金榆-花楷槭混交林林下蓝光光子通量密度最小,而沙松-紫椴-臭冷杉混交林和长白落叶松纯林林下最大。随着尺度的增大,天然林乔木胸高断面积与R/PFD(红光/光子通量密度比值)和B/PFD(蓝光/光子通量密度比值)的比值呈显著正相关(P<0.05)。并且随着尺度的增加,相关系数总体逐渐增大,在35m处达到峰值。在此基础上在南向、东向和西向各延伸10m时呈现显著正相关(P<0.05)。在该尺度下分析优势树种对林下R/PFD和B/PFD比值的影响时发现：R/PFD与B/PFD比值随着针叶林胸高断面积的增加而增加。相对于阔叶林来说,多数林型针叶林下的冠层结构与林下R/PFD和B/PFD比值之间显著正相关(P<0.05)。在不同树种下,乔木冠层结构对R/PFD和B/PFD比值的影响...     

Confessions of solitary oaks: We grow fast but we fear the drought

Competition effects and management related disturbances can contribute to strong non-climatic signals in ring-width trends of trees from closed canopy forests. Removing this noise comes at the price of losing a considerable amount of (climatic) information on decadal and centennial time scales. Alternatively, open grown solitary trees, which never competed for light or other resources could function as less biased climate proxies. To explore this potential we analysed individual growth trends and climate-growth relationships of solitary oaks, potentially representative of former, open forests, and compared it to those of trees growing in adjacent closed canopy forests. Solitary oaks show differing tree allometry with bigger crowns and lower absolute heights. Their radial growth was significantly higher compared to forest trees. Contrary to our expectations, these solitary oaks show highly individualistic decadal growth trends. Their inter-annual variations are nevertheless in good agreement between individuals and stronger compared to trees from closed canopy forests. Climate –growth analysis revealed a significantly higher drought sensitivity of solitary oaks. Given the typical open medieval forest structure of large trees with extensive crowns growing in a mosaic of grazed woodlands and grasslands, it is likely that these trees contribute to the historical part of tree-ring collections. When trees from today’s denser forests forming the recent parts of the collections are used to calibrate reconstruction models, the differing drought sensitivity might therefore lead to an overestimation of historical droughts.     

Canopy disturbance and gap partitioning promote the persistence of a pioneer tree population in a near‐climax temperate forest of the Qinling Mountains,China

An unresolved question of temperate forests is how pioneer tree species persist in mature forests. In order to understand the responsible mechanisms, we investigated a near‐climax mixed temperate forest dominated by Betula albosinensis in the Qinling Mountains of China. Through establishing four 50 m × 50 m plots, we examined the canopy disturbance characteristics and its effects on tree recruitments. We further test the intra‐ and interspecific effects on the recruitment of B. albosinensis. The obtained data demonstrated canopy disturbance was frequent but most small‐sized. The canopy gaps are caused mainly by adult B. albosinensis by snapping. The regeneration of coexistent tree species shows a distinct preference for gap size. B. albosinensis were clumped at the juvenile stage and small scales. B. albosinensis juveniles were positively associated with B. utilis juveniles and negatively associated with the conspecific and B. utilis large trees. In addition, B. albosinensis juveniles showed negative associations with contemporary other tree species. Our results suggested that canopy disturbance caused by canopy trees and gap partitioning among the coexistent tree species are important for the persistence of the mixed forest. As a main gapmaker, B. albosinensis appear to develop a self‐perpetuating life‐history trait and allow them to persist.     

Foliage-canopy structure and height distribution of woody species in climax forests

The correlation between foliage-canopy structure and vertical woody species distribution was examined in seven climax forests ranging from alpine tree limit to tropical rain forest. Foliage density was measured by two-dimensional canopy tomography using photographs. Both foliage density and the vertical species density (the number of woody species having a maximum height within a vertical 1 m) were high in the upper canopy of warm-temperate and subtropical forests, but they were high at lower stratums in the tropical rain forest. Two variables correlated significantly despite the differences in foliage-canopy structures. In contrast to evergreen broad-leaved forests, a clear correlation could not be detected in northern cool-temperate and sub-alpine forests. A possible reason for species convergence in the foliage dense stratum is that species with maximum height in that stratum may be able to survive in the stratum due to symmetrical crown-to-crown interaction. If the maximum height of dwarf species is less than the foliage dense stratum, it may be difficult to survive in the community. The lack of correlation in northern forests may be due to poor canopy tree flora and a mixture of different life forms (non-sprouting trees and sprouting shrubs).     

Comparative study of additive basal area of conifers in forest ecosystems along elevational gradients

We examined the basal area of two life forms (conifers vs. broadleaf trees) along elevational gradients on Yakushima Island, Japan and on two series of geological substrate on Mount Kinabalu, Borneo. On Yakushima, total stand basal area abruptly increased from 700 to 1,050 m in accordance with the high dominance of conifers, indicating the presence of additive basal area of conifers in conifer–broadleaf mixed forests at higher elevations (1,050–1,300 m). Along two substrate series on Kinabalu, some forests at higher elevations (1,860–3,080 m) showed relatively high dominance of conifers, but conifer basal area did not appear to be additive. Conifers were emergents above the canopy of broadleaf trees in mixed forests on Yakushima, but two life forms usually coexisted in the single-story canopy in mixed forests on Kinabalu. Litterfall rate as a surrogate of productivity decreased with decreasing temperature along elevation on both the sites, but the rate of decrease was slower on Yakushima, where mixed forests at higher elevations showed relatively high rates. Thus, we suggest that additive basal area of conifers was linked to their emergent status, and that it enhanced productivity by complementary use of light by two life forms that occupy different stories. On Yakushima, typhoons are a major disturbance, but do not severely limit the height growth of conifers, allowing the development of two-story mixed forests. On Kinabalu, a major disturbance is El Niño-driven drought, and hydraulic limitation to tree height may explain the non-additive and non-emergent nature of conifers.     

Tamarind tree seed dispersal by ring-tailed lemurs

In Madagascar, the gallery forests of the south are among the most endangered. Tamarind trees (Tamarindus indica) dominate these riverine forests and are a keystone food resource for ring-tailed lemurs (Lemur catta). At Berenty Reserve, the presence of tamarind trees is declining, and there is little recruitment of young trees. Because mature tamarinds inhibit growth under their crowns, seeds must be dispersed away from adult trees if tree recruitment is to occur. Ring-tailed lemurs are likely seed dispersers; however, because they spend much of their feeding, siesta, and sleeping time in tamarinds, they may defecate a majority of the tamarind seeds under tamarind trees. To determine whether they disperse tamarind seeds away from overhanging tamarind tree crowns, we observed two troops for 10 days each, noted the locations of feeding and defecation, and collected seeds from feces and fruit for germination. We also collected additional data on tamarind seedling recruitment under natural conditions, in which seedling germination was abundant after extensive rain, including under the canopy. However, seedling survival to 1 year was lower when growing under mature tamarind tree crowns than when growing away from an overhanging crown. Despite low fruit abundance averaging two fruits/m³ in tamarind crowns, lemurs fed on tamarind fruit for 32% of their feeding samples. Daily path lengths averaged 1,266 m, and lemurs deposited seeds throughout their ranges. Fifty-eight percent of the 417 recorded lemur defecations were on the ground away from overhanging tamarind tree crowns. Tamarind seeds collected from both fruit and feces germinated. Because lemurs deposited viable seeds on the ground away from overhanging mature tamarind tree crowns, we conclude that ring-tailed lemurs provide tamarind tree seed dispersal services.     

Diversity of canopy and understorey spiders in north-temperate hardwood forests

1 We characterized and compared diversity patterns of canopy and understorey spiders (Arachnida: Araneae) on sugar maple ( Acer saccharum Marsh.) and American beech ( Fagus grandifolia Ehrh.) in hardwood forests of southern Québec, Canada.
2 We sampled canopies of 45 sugar maple and 45 American beech trees and associated understorey saplings in mature protected forests near Montréal. Samples were obtained by beating the crown foliage at various heights and by beating saplings around each tree.
3 Eighty-two species were identified from 13 669 individuals. Forty-eight species and 3860 individuals and 72 species and 9809 individuals were collected from the canopy and the understorey, respectively.
4 Multivariate analyses (NMDS ordination and NPMANOVA) showed the composition of canopy and understorey assemblages differed significantly, and canopy assemblages differed between tree species. Rank-abundance distribution models fitted to the canopy and understorey data indicated that different mechanisms structure the assemblages in both habitats. Three abundant spider species were significantly more common in the canopy; ten species were collected significantly more often in the understorey.
5 The forest canopy was shown to be an important reservoir for spider diversity in north-temperate forests.     

Diversity patterns of eucalypt canopy arthropods in eastern and western Australia

1. As part of a larger study on canopy arthropods and birds, a 1‐year chemical knockdown study was carried out in one Western Australian forest, where jarrah Eucalyptus marginata and marri E. (Corymbia) calophylla were sampled, and one eastern Australian (New South Wales) forest, where narrow‐leaved ironbark E. crebra and grey box E. moluccana were sampled. 2. Ten trees of each species were sampled during each of the four seasons and the arthropods were sorted to morphospecies level. This paper documents the foliage‐associated component of arboreal arthropod communities and compares arthropod species richness within orders and families, between tree species, and between the two forest types. 3. Hymenoptera, Coleoptera, Diptera, and Araneae were the richest in species. Nine hundred and seventy‐six species in 173 families were found in the eastern Australian forest, while 687 species in 176 families were found in the western Australian forest. Only 53% of families were common to both forests, but almost half the families recorded were represented by fewer than five species. Species overlap between tree species in each region was 40–53%. 4. Analysis using nonparametric bootstrapping procedures showed that sampling of foliage was comprehensive and that only 4–9% more species would be expected with more intensive sampling of the canopy. Absolute richness, as well as differences between tree species and regions, therefore appear to be real and not the result of sampling errors. As a consequence, arthropod species richness in Australian eucalypt forests is shown to be substantially greater than previous estimates.     

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.     

Three-dimensional Structure of an Old-growth <Emphasis Type="Italic">Pseudotsuga-Tsuga</Emphasis> Canopy and Its Implications for Radiation Balance,Microclimate, and Gas Exchange

We describe the three-dimensional structure of an old-growth Douglas-fir/western hemlock forest in the central Cascades of southern Washington, USA. We concentrate on the vertical distribution of foliage, crowns, external surface area, wood biomass, and several components of canopy volume. In addition, we estimate the spatial variation of some aspects of structure, including the topography of the outer surface, and of microclimate, including the within-canopy transmittance of photosynthetically active radiation (PAR). The crowns of large stems, especially of Douglas-fir, dominate the structure and many aspects of spatial variation. The mean vertical profile of canopy surfaces, estimated by five methods, generally showed a single maximum in the lower to middle third of the canopy, although the height of that maximum varied by method. The stand leaf area index was around 9 m² m^–2, but also varied according to method (from 6.3 to 12.3). Because of the deep narrow crowns and numerous gaps, the outer canopy surface is extremely complex, with a surface area more than 12 times that of the ground below. The large volume included below the outer canopy surface is very porous, with spaces of several qualitatively distinct environments. Our measurements are consistent with emerging concepts about the structure of old-growth forests, where a high degree of complexity is generated by diverse structural features. These structural characteristics have implications for various ecosystem functions. The height and large volume of the stand indicate a large storage component for microclimatic variables. The high biomass influences the dynamics of those variables, retarding rates of change. The complexity of the canopy outer surface influences radiation balance, particularly in reducing short-wave reflectance. The bottom-heaviness of the foliage profile indicates much radiation absorption and gas exchange activity in the lower canopy. The high porosity contributes to flat gradients of most microclimate variables. Most stand respiration occurs within the canopy and is distributed over a broad vertical range.