Long‐term recovery of the functional community assembly and carbon pools in an African tropical forest succession

On the African continent, the population is expected to expand fourfold in the next century, which will increasingly impact the global carbon cycle and biodiversity conservation. Therefore, it is of vital importance to understand how carbon stocks and community assembly recover after slash‐and‐burn events in tropical second growth forests. We inventoried a chronosequence of 15 1‐ha plots in lowland tropical forest of the central Congo Basin and evaluated changes in aboveground and soil organic carbon stocks and in tree species diversity, functional composition, and community‐weighted functional traits with succession. We aimed to track long‐term recovery trajectories of species and carbon stocks in secondary forests, comparing 5 to 200 + year old secondary forest with reference primary forest. Along the successional gradient, the functional composition followed a trajectory from resource acquisition to resource conservation, except for nitrogen‐related leaf traits. Despite a fast, initial recovery of species diversity and functional composition, there were still important structural and carbon stock differences between old growth secondary and pristine forest, which suggests that a full recovery of secondary forests might take much longer than currently shown. As such, the aboveground carbon stocks of 200 + year old forest were only 57% of those in the pristine reference forest, which suggests a slow recovery of aboveground carbon stocks, although more research is needed to confirm this observation. The results of this study highlight the need for more in‐depth studies on forest recovery in Central Africa, to gain insight into the processes that control biodiversity and carbon stock recovery.     

Secondary succession and effects of clear-logging on diversity in the subtropical forests on Okinawa Island,southern Japan

In order to clarify the recovery process of the subtropical forest on Okinawa Island, southern Japan, biomass accumulation and the successional trend of species diversity with time were investigated by comparing plots of old-growth and clear logged secondary forests. Self-thinning was an important factor in the development of young secondary forests, and the small variance in tree size within a stand was related to the stand not being fully stratified after clear-cutting. A large variance of size structure in old secondary and old-growth forests implies re-initiation of the understorey. Additionally, the trajectory of stand development indicated that the subtropical forest quickly recovered aboveground biomass, which reached its upper limit at about 50 years after disturbance. However, there was a large distinction in species diversity between the secondary forests and old-growth forests. The diversity of forest floor plants did not recover well after being clear-cut. This indicates that management of the subtropical forest should not only take timber-oriented tree species into account, but also the biodiversity in ground flora. The secondary forests were characterized by Castanopsis sieboldii and Schima wallichii, and the monopolization of C. sieboldii through secondary succession had a negative influence on species diversity. Distylium racemosum dominated at the late development stage and was considered a long-lived competing species that reduced the dominance of C. sieboldii and facilitated the co-occurrence of understorey species. Light-demanding pioneer tree species such as S. wallichii that regenerated after logging should be replaced by competitive effects of climax species, and thus relayed floristic change might increase species diversity along secondary succession.Nomenclature: Hatushima and Amano (1994).     

Phylogenetic clustering increases with succession for lianas in a Chinese tropical montane rain forest

Previous research found that phylogenetic clustering increased with disturbance for tropical trees, suggesting that community assembly is mainly influenced by abiotic factors during early succession. Lianas are an important additional component of tropical forests, but their phylogenetic community structure has never been investigated. Unlike tropical trees, liana abundance is often high in disturbed forests and diversity can peak in old secondary forest. Therefore, phylogenetic structure along a disturbance gradient might also differ from tropical tree communities. Here we determined phylogenetic community structure of lianas along a disturbance gradient in a tropical montane forest in China, using the net relatedness index (NRI) from 100 equivalent phylogenies with varying branch length that were constructed using DNA‐barcode sequences. Three additional phylogenetic indices were also considered for comparison. When NRI was used as index phylogenetic clustering of liana communities decreased with decreasing tree basal area, suggesting that liana competitive interactions dominate during early succession, which is in contrast to the pattern reported for trees. Liana communities in mature forests, on the other hand, were phylogenetic clustered, which could be caused by dispersal limitation and/or environmental filtering. The three additional phylogenetic indices identified different, sometimes contradicting predictors of phylogenetic community structure, indicating that caution is needed when generalizing interpretations of studies based on a single phylogenetic community structure index. Our study provides a more nuanced picture of non‐random assembly along disturbance gradients by focusing on a non‐tree forest component.     

Changes in biotic and abiotic drivers of seedling species composition during forest recovery following shifting cultivation on Hainan Island,China

Seedlings play an important role in the processes of plant community succession. We compared seedling (dbh < 1 cm) species composition and diversity over a chronosequence (18‐, 30‐, 60‐year‐old second growth and old growth forest) after shifting cultivation in a tropical lowland rain forest area on Hainan Island, China. Seedling diversity reached a maximum in the 60‐year‐old second growth forest, which is consistent with the intermediate disturbance hypothesis. With the progression of secondary succession, canopy openness (CO), soil organic matter, soil phosphorus content, and tree abundance showed a general decreasing trend; soil water content and tree basal area showed a general trend of increase, while soil pH and other nutrients reached maximum values and tree richness reached a minimum value at intermediate stages of succession. Seedling composition and diversity were significantly affected by soil water, pH, soil nutrient content, and biotic factors in the 18‐year‐old second growth forests; by soil pH, soil nutrient content, and biotic factors in the 30‐year‐old second growth forests; by CO, soil nutrient content and tree abundance in the 60‐year‐old second growth forests; and by CO, soil pH, and soil nutrient content in the old growth forests. At earlier stages of succession, the effect of the proportion of old growth forest in the surrounding landscape on seedling diversity was greater than that of land‐use history, but the importance of these drivers was reversed at later stages of succession.     

南方红壤区3种典型森林恢复方式对植物群落多样性的影响

合理的森林恢复方式能提高植物多样性,进而提高生态系统服务功能.在我国南方红壤区研究了3种典型森林恢复方式(引进种恢复的湿地松(Pinus elliottii)人工林本地种恢复的马尾松(Pinus massoniana)人工林和自然恢复的天然次生林)的植物区系构成和植物群落多样性.结果表明:(1)湿地松人工林有155种植物,隶属66科118属,马尾松人工林有137种植物,隶属59科97属;天然次生林有226种植物,隶属86科160属,3种森林恢复方式的乔木层、灌木层和草本层的优势科属明显不同,马尾松人工林的优势物种和天然次生林更相似;(2)天然次生林的植物区系基本构成、植物区系类型种类高于马尾松人工林和湿地松人工林,并且天然次生林的温带成分比例高于湿地松人工林;(3)恢复方式对植物群落的多样性指数有显著影响,天然次生林的物种丰富度、辛普森指数明显高于马尾松人工林和湿地松人工林,两种人工林之间差异不显著;(4)3种森林恢复方式的植物群落结构存在显著差异,相比湿地松人工林,马尾松人工林的植物群落组成与天然次生林更相似.总之,自然恢复的天然次生林植物群落多样性高于人工恢复的马尾松人工林和湿地松人工林,本地种马尾松人工林在维持区域植物群落结构功能上优于引进种湿地松人工林.     

Tropical Secondary Forest Management Influences Frugivorous Bat Composition,Abundance and Fruit Consumption in Chiapas,Mexico

Most studies on frugivorous bat assemblages in secondary forests have concentrated on differences among successional stages, and have disregarded the effect of forest management. Secondary forest management practices alter the vegetation structure and fruit availability, important factors associated with differences in frugivorous bat assemblage structure, and fruit consumption and can therefore modify forest succession. Our objective was to elucidate factors (forest structural variables and fruit availability) determining bat diversity, abundance, composition and species-specific abundance of bats in (i) secondary forests managed by Lacandon farmers dominated by Ochroma pyramidale, in (ii) secondary forests without management, and in (iii) mature rain forests in Chiapas, Southern Mexico. Frugivorous bat species diversity (Shannon H’) was similar between forest types. However, bat abundance was highest in rain forest and O. pyramidale forests. Bat species composition was different among forest types with more Carollia sowelli and Sturnira lilium captures in O. pyramidale forests. Overall, bat fruit consumption was dominated by early-successional shrubs, highest late-successional fruit consumption was found in rain forests and more bats consumed early-successional shrub fruits in O. pyramidale forests. Ochroma pyramidale forests presented a higher canopy openness, tree height, lower tree density and diversity of fruit than secondary forests. Tree density and canopy openness were negatively correlated with bat species diversity and bat abundance, but bat abundance increased with fruit abundance and tree height. Hence, secondary forest management alters forests’ structural characteristics and resource availability, and shapes the frugivorous bat community structure, and thereby the fruit consumption by bats.     

Dispersal process driving subtropical forest reassembly: evidence from functional and phylogenetic analysis

Understanding the mechanisms of secondary succession related to forest management practices is receiving increasing attention in community ecology and biodiversity conservation. Abiotic and biotic filtering are deterministic processes driving community reassembly. A functional trait or phylogeny-based approach predicts that environmental filtering induced by clearcut-logging results in functional/phylogenetic clustering in younger forests, while biotic filtering (competitive exclusion) promotes functional/phylogenetic overdispersion in old-growth forests. From this perspective, we examined the patterns of functional/phylogenetic structures using tree community data (147 species × 170 plots). These data were chronosequenced from clearcut secondary forests to old-growth subtropical forests in the Ryukyu Archipelago, with species’ trait data (leaf and stem) and species level phylogeny. To detect clustering or overdispersion in the functional and phylogenetic structures, we calculated the standardized effect size of mean nearest trait distance and mean nearest phylogenetic distance within the plots. Functional or phylogenetic clustering was relatively weak in secondary forests, and their directional change with increasing forest age was not generally detected. Mean nearest trait/phylogenetic distance for most plots fell within the range of random expectation. The results suggest that abiotic/biotic filtering related to functional traits or phylogenetic relatedness plays a diminished role in shaping species assembly during secondary succession in the subtropical forest. Our findings of functional and phylogenetic properties might shed light on the importance of dispersal (stochastic) processes in the regional species pool during community reassembly after anthropogenic disturbance. It will also contribute to the development of coordinated schemes that maintain potential species assembly processes in the subtropical forest.     

Rapid Liana Colonization along a Secondary Forest Chronosequence

Lianas (woody vines) can have profound effects on tree recruitment, growth, survival, and diversity in tropical forests. However, the dynamics of liana colonization soon after land abandonment are poorly understood, and thus it is unknown whether lianas alter tree regeneration early in succession. We examined the liana community in 43 forests that ranged from 1 to 31 yr old in central Panama to determine how fast lianas colonize young forests and how the liana community changes with forest succession. We found that lianas reached high densities early in succession, commonly exceeding 1000 stems/ha within the first 5 yr of forest regeneration. Lianas also increased rapidly during early succession in terms of basal area but did not show evidence of saturation within the 30 yr of our chronosequence. The relative contribution of lianas to total woody plant community in terms of basal area and density increased rapidly and reached a saturation point within 5 yr (basal area) to 15 yr (density) after land abandonment. Our data demonstrate that lianas recruit early and in high density in tropical forest regeneration, and thus lianas may have a large effect on the way in which secondary forests develop both early and throughout succession.     

Change in community phylogenetic structure during tropical forest succession: evidence from New Guinea

Ecologists have recently interpreted patterns of phylogenetic distance among coexisting species as indicative of processes affecting community assembly during forest succession. We investigated plant community phylogenetic structure along a successional gradient in New Guinean lowland rain forest. We surveyed all trees with diameter at breast height ≥ 5 cm in nineteen 0.25 ha plots representing younger secondary, older secondary, and primary forest. We estimated plant community phylogeny from rbcL gene sequences to quantify change in phylogenetic structure during succession. Mean phylogenetic distance among co‐occurring trees increased with total basal area per plot, a proxy for forest age. Significant phylogenetic clustering was detected in secondary forest whereas primary forest was significantly over‐dispersed relative to null expectations. We examined the sensitivity of these patterns to various methods of branch length estimation and phylogenetic uncertainty. Power to detect community phylogenetic patterns when equal branch lengths were assumed was weak in comparison to direct molecular and time‐calibrated measures of divergence. Inferred change during forest succession was also robust to phylogenetic uncertainty so long as temporal information was incorporated in estimates of divergence. The observed patterns are consistent with processes of environmental filtering during tropical forest succession giving way to other processes in primary forests including density‐dependence.     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Liana communities exhibit different species composition,diversity and community structure across forest types in the Congo Basin

Lianas are poorly characterized for central African forests. We quantify variation in liana composition, diversity and community structure in different forest types in the Yangambi Man and Biosphere Reserve, Democratic Republic of Congo. These attributes of liana assemblages were examined in 12 1-ha plots, randomly demarcated within regrowth forest, old growth monodominant forest, old growth mixed forest and old growth edge forest. Using a combination of multivariate and univariate community analyses, we visualize the patterns of these liana assemblage attributes and/or test for their significant differences across forest types. The combined 12 1-ha area contains 2,638 lianas (≥2 cm diameter) representing 105 species, 49 genera and 22 families. Liana species composition differed significantly across forest types. Taxonomic diversity was higher in old growth mixed forests compared to old growth monodominant and regrowth forests. Trait diversity was higher than expected in the regrowth forest as opposed to the rest of forest types. Similarly, the regrowth forest differed from the rest of forest types in the pattern of liana species ecological traits and diameter frequency distribution. The regrowth forest was also less densely populated in lianas and had lower liana total basal area than the rest of forest types. We speculate that the mechanism of liana competitive exclusion by dominant tree species is mainly responsible for the lower liana species diversity in monodominant compared to mixed forests. We attribute variation in liana community structure between regrowth and old growth forests mostly to short development time of size hierarchies.     

The role of remnant trees in carbon sequestration,vegetation structure and tree diversity of early succession regrowing fallows in eastern Sierra Leone

Remnant tree presence affects forest recovery after slash‐and‐burn agriculture. However, little is known about its effect on above‐ground carbon stocks, especially in Africa. We focused our study on Sierra Leone, part of the Upper Guinean forests, an important centre of endemism threatened by encroachment and forest degradation. We studied 99 (20‐m‐radius) plots aged 2–10 years with and without remnant trees and compared their above‐ground carbon stocks, vegetation structure (stem density, basal area) and tree diversity. Above‐ground carbon stocks, stem density, basal area, species richness and tree diversity increased significantly with fallow age. Remnant tree presence affected significantly tree diversity, species dominance and above‐ground carbon stocks, but not vegetation structure (stem density, basal area). Number of remnant trees and number of species of remnant trees were also important explanatory variables. Although other factors should be considered in future studies, such as the size and dispersal modes of remnant trees, our results highlight that more strategic inclusion of remnant trees is likely to favour carbon stock and forest recovery in old fallows. To our knowledge, this is the first study on early succession regrowing fallows in West Africa.     

The Conservation Value of Secondary Forests for Vascular Epiphytes in Central Panama

Secondary forests that develop following land abandonment could compensate for the losses of diversity and structure that accompany deforestation of old‐growth forests in tropical regions. Whether secondary forests can harbor similar species richness, density, and composition of old‐growth forests for vascular epiphytes remains largely unknown for secondary forests older than 50 yr. We examined community structure (species richness, density, and species composition) of vascular epiphytes in older secondary forests between 35 and 115 yr after land abandonment and nearby old‐growth forests to determine if the community structure of epiphytes in secondary forests approaches that of old‐growth forests over time. The recovery of epiphyte species richness was rapid with 55‐year‐old forests containing 65 percent of old‐growth epiphyte species richness. Secondary forest epiphyte communities were found to be statistically nested within secondary forests older in age and within old‐growth forests. Similarity of epiphyte communities to old‐growth forests increased to 75 percent, 115 yr after abandonment. This study suggests that secondary forests will likely recover old‐growth epiphyte richness and composition given enough time. Epiphyte densities did not recover quickly with 55‐year‐old forests having 14 percent and 115‐year‐old forests having only 49 percent of the density of old‐growth forest epiphytes. The low density of epiphytes in secondary forests could impact rainforest diversity and function. We conclude that in less than 115 yr, although secondary moist forests have high conservation value for some aspects of community structure, they are unlikely to compensate biologically for the loss of diversity and ecosystem function that high epiphyte densities provide.     

热带次生林刀耕火种过程中土壤节肢动物群落结构及多样性的变化

通过模拟刀耕火种过程,对热带次生林土壤节肢运动群薄结构及多样性在刀耕火种过程中变化进行了初步研究,结果表明,火烧1周后,土壤节肢运行类群数和个体数从次生林地的14类,564头降为7类,164头,并且在群落组成上发生了较大变化,膜翅目的烧后1天成为突出优势类群,蜱蟥目在烧后1周成为唯一的优势类群,而在多样性方面,土壤节肢运动群落类群数（丰度）,个体数（多度）和DG多样性指数在刀耕火种过程中均降低,土壤节肢运动的垂直分布因刀耕火种干扰发生了很大变化,尚存土壤节肢运动随干扰强度的加剧,出现逐步由土壤表层向深层迁移的现象,一些土壤节肢运动在土层间做垂直迁移活动,说明土壤层在一定程度上减轻了刀耕火种对它们的破坏作用,维持了某些类群数量及群落多样性。     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

Restoration of the understorey layer of recent forest bordering ancient forest

Abstract. Recently established forests are commonly characterized by an impoverished understorey. Restoration is mostly based on spontaneous secondary succession, but little is known about the time period needed to achieve a community species pool with species composition equal to that of ancient forests. Vegetation in transects of 197 plots in 13 recent forest stands contiguous to the Meerdaalwoud ancient forest complex was surveyed. The recent forest stands ranged in age from 36 to 132 yr. The community species pool was described with an ecological, functional and phytosociological approach and based on groups derived from a CCA. Differences in community species pool between age classes of recent forest stands were analysed. During establishment of a new forest competitive species, forest edge species and species with high Ellenberg values for light and nitrogen and a more persistent seed bank will dominate the understorey. After 90 yr of succession the cover by these species decreases and reaches equal values to ancient forest after ca. 105 yr. A large number of forest species will be able to colonize the forest in less than 90 yr. Some typical forest species, however, have very low colonization rates and still have low cover in recent forest more than 105 yr old, so that complete restoration of the understorey requires a time period of over a century. Anthropogenic introduction of forest plant species may reduce the time required for ancient forest vegetation equality.     

次生林经营基础研究进展

次生林是中国森林的主体,在结构组成,林木生长,生产力,林分环境等诸多方面与原始林和人工林有着显著的不同,因此,经营好次生林对中国天然林保护等林业工程实施及国家生态安全建设具有重大意义。本文在广泛收集国内外有关次生林研究结果的基础上,探讨了次生林领域中基础研究所涉内容,揭示出次生林经营研究中所关注的基础问题,提出了次生林经营基础研究的主要内容与方向,对今后次生林经营研究和中国天然林保护等林业工程建设具有参考价值。     

From Forest to Farmland: Species Richness Patterns of Trees and Understorey Plants along a Gradient of Forest Conversion in Southwestern Cameroon

Vegetation surveys were carried out at 24 sampling stations distributed over four land use types, namely near-primary forest, secondary forest, agroforestry systems and annual crop lands in the northeastern part of the Korup region, Cameroon, to assess the impact of forest conversion on trees and understorey plants. Tree species richness decreased significantly with increasing level of habitat modification, being highest and almost equal in secondary and near-primary forests. Understorey plant species richness was significantly higher in annual crop lands than in other land use types. The four land use types differed in tree and understorey plant species composition, the difference being smaller among natural forests. Tree and understorey plant density differed significantly between habitat types. Density was strongly correlated with species richness, both for trees and understorey plants. Five tree and 15 understorey plant species showed significant responses to habitat. A 90% average drop in tree basal area from forest to farmland was registered. Our findings support the view that agroforestry systems with natural shade trees can serve to protect many forest species, but that especially annual crop lands could be redesigned to improve biodiversity conservation in agricultural landscapes of tropical rainforest regions.     

Mycorrhiza of plants in different vegetation types in tropical ecosystems of Xishuangbanna,southwest China

We examined plants growing in four tropical vegetation types (primary forest, secondary forest, limestone forest and a slash and burn field) in Xishuangbanna, southwest China for mycorrhizal associations. Of the 103 plant species examined (belonging to 47 families), 81 had arbuscular mycorrhizal (AM) associations, while three species possessed orchid mycorrhiza. AM colonization levels ranged between 6% and 91% and spore numbers ranged between 1.36 spores and 25.71 spores per 10 g soil. Mean AM colonization level was higher in primary and secondary forest species than in plant species from limestone forests and a slash and burn field. In contrast, mean AM fungal spore numbers of the primary and limestone forest were lower than in the secondary forest or the slash and burn field. AM fungal spores belonging to Glomus and Acaulospora were the most frequent in soils of Xishuangbanna. AM fungal colonization and spore numbers were significantly correlated to each other and were significantly influenced by vegetation type.     

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.