Microsite specialization and spatial distribution of <Emphasis Type="Italic">Geonoma brevispatha</Emphasis>, a clonal palm in south-eastern Brazil

We tested the hypotheses that: (i) reproductive Geonoma brevispatha, an understorey clonal palm endemic to South American swamps, is most abundant in more brightly lit microsites but the abundance of juveniles is not responsive to light availability, (ii) the species is restricted to the transitional zones between flooded and well-drained microsites, (iii) if hypotheses (i) and/or (ii) are accepted, the microhabitat specialization they represent should be reflected in the spatial distribution of the individuals at distinct scales. The study was carried out in a swamp forest in south-eastern Brazil. Soil moisture was autocorrelated at distances up to approximately 1m, reflecting a fine-scale microtopographic pattern of flooded pits and channels delimited by drier mounds. The first hypothesis was rejected, but the second hypothesis was accepted. No genets occurred on flooded microsites. A juvenile emergence experiment showed that seeds were unable to develop into juveniles under flooded conditions, but canopy openness did not influence the number of juveniles emerging. The third hypothesis was accepted. Individual genets were randomly distributed at scales corresponding to soil moisture patches, but were aggregated at larger scales. Juvenile genets were positively associated with reproducer genets. Our results indicate that palms may specialize on narrow parts of moisture gradients in swamp forests, and that this specialization can exclude them from even subtly distinct microhabitats.     

Use of early and late successional forest patches by the endangered Lowland tapir Tapirus terrestris (Perissodactyla: Tapiridae)

Large herbivores play crucial ecological roles, affecting the structure and function of terrestrial ecosystems. Their effect, however, depend on how they select plants and vegetation patches for foraging. At the landscape scale, succession is one of the processes that should generate vegetation patches with different nutritional quality, affecting selection by herbivores. Earlier successional stages should be preferred as they are dominated by plants with nutritious and palatable leaves. Here, we investigate if the Lowland tapir prefers early compared to late successional forest patches, aiming at contributing to the understanding of the ecological role of the largest terrestrial South American herbivore, and to conservation strategies for this endangered species. We sampled 12 vegetation patches varying in successional stages across a 20.000-ha continuously-forested landscape in the Brazilian Atlantic Forest, recording tapirs through standardized camera trap and track surveys, and quantifying vegetation structure and treefall gaps. Whereas the number of individuals using each patch was not influenced by successional stage, intensity of use was higher in patches in earlier successional stages, in particular patches with higher density of smaller trees and higher cover of treefall gaps. Although inferences on the effects of tapir on plant community depends on future, smaller-scale studies, our results suggest herbivory by tapirs affects forest regeneration, potentially contributing to the maintenance of plant diversity. Results also point out to the potential of mosaics encompassing old-growth and secondary forests for the conservation of the Lowland tapir.     

Complex spatial structure in a population of <Emphasis Type="Italic">Didymopanax pittieri</Emphasis>, a tree of wind-exposed lower montane rain forest

Didymopanax pittieri is a common shade-intolerant tree colonizing treefall gaps in the elfin forests on windswept ridgecrests in the lower montane rain forests of the Cordillera de Tilarán, Costa Rica. All D. pittieri taller than >0.5 m in a 5.2-ha elfin forested portion of a gridded study watershed in the Monteverde Cloud Forest Preserve were located, mapped, and measured. This local population of D. pittieri is spatially inhomogeneous, in that density increases with increasing wind exposure; D. pittieri are more abundant near ridge crests than lower on windward slopes. The important and ubiquitous phenomenon of spatial inhomogeneity in population density is addressed and corrected for in spatial analyses by the application of the inhomogeneous version of Ripley’s K. The spatial patterns of four size classes of D. pittieri (<5 cm dbh, 5–10 cm dbh, 10–20 cm dbh, and >20 cm dbh) were investigated. Within the large-scale trend in density driven by wind exposure, D. pittieri saplings are clumped at the scale of treefall gaps and at the scale of patches of aggregated gaps. D. pittieri 5–10 cm dbh are randomly distributed, apparently due to competitive thinning of sapling clumps during the early stages of gap-phase regeneration. D. pittieri larger than 10 cm dbh are overdispersed at a scale larger than that of patches of gaps. Natural disturbance can influence the distribution of shade intolerant tree populations at several different spatial scales, and can have discordant effects at different life history stages.     

Evidence that fungal pathogens inhibit recruitment of a shade-intolerant tree,white birch (<Emphasis Type="Italic">Betula papyrifera</Emphasis>), in understory habitats

Evidence from tropical forests suggests understory habitats are associated with a high risk of disease, which may prevent the establishment of vulnerable tree species; in contrast, canopy gaps can act as refuges from these pathogens. However, few studies have investigated the impacts of pathogens on regeneration in temperate forests. To determine whether losses to fungi of seeds of Betula papyrifera, a light-loving species, varied between habitats that differed in their degree of openness, we applied fungicide to seeds buried in old fields, treefall gaps, and forest understory sites. We found that the application of fungicide significantly reduced losses in all habitats, relative to control values. This effect was habitat-dependent: the benefit of fungicide was greater in forest understory than in openings. This suggests that B. papyrifera is prevented from establishing in understory environments in part by its susceptibility to pathogen attack, and not solely because of a high light requirement.     

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.     

Vegetation and environmental heterogeneity relationships in a Neotropical swamp forest in southeastern Brazil (Itirapina, SP)

Patterns in substrate, canopy openness, tree species composition and structure were studied in a swamp forest in southeastern Brazil (Itirapina, SP), using a fine spatial scale (∼0.05 km²). Sixty quadrats of 10 m × 10 m were divided into three sample plots, located in the centre and at both ends of the forest remnant (upstream and downstream), encompassing different environmental conditions. In each quadrat we quantified and identified individuals with CBH (circumference at breast height) ≥ 10 cm, the chemical properties of the soil, the drainage (flooded area) and the percentage of canopy openness. We keep 5124 individuals distributed over 37 species and 25 families. The downstream site presented a lower frequency of flooding, canopy openness, species richness (15 species) and density of individuals. In the central site we found different patterns of drainage and light incidence, the greatest species richness (31 species) and density of individuals, but smaller trees both in height and in diameter. The upstream site was subject to higher frequency of flooding, with intermediate values for density and species richness (27 species). Floristic and structural variation between and within sites reflects the different environmental conditions related to substrate and canopy openness. However, a larger number of species were correlated with drainage pattern (21 species) rather than with canopy openness (14 species), suggesting that the main factor responsible for the spatial organization of the plant community in swamp forests is soil drainage.     

Logging leaves a fingerprint on the number,size, spatial configuration and geometry of tropical forest canopy gaps

Recent advances in remote sensing such as airborne laser scanning have revolutionized our ability to accurately map forest canopy gaps, with huge implications for tracking forest dynamics at scale. However, few studies have explored how canopy gaps vary among forests at different successional stages following disturbances, such as those caused by logging. Moreover, most studies have focused exclusively on the size distribution of gaps, ignoring other key features such as their spatial distribution and shape. Here, we test a series of hypotheses about how the number, size, spatial configuration, and geometry of gaps vary across a logging disturbance gradient in Malaysian Borneo. As predicted, we found that recently logged forests had much higher gap fraction compared to old-growth forests, a result of having both a greater total number of gaps and a higher proportion of large gaps. Regrowing forests, on the other hand, fell at the opposite end of the spectrum, being characterized by both fewer and smaller gaps compared to nearby old-growth forests. Across all successional stages gaps were found to be spatially clustered. However, logging significantly diluted the degree of spatial aggregation and led to the formation of gaps with much more complex geometries. Our results showcase how logging and subsequent regrowth substantially alter not just the number and size of gaps in a forest, but also their spatial arrangement and shape. Linking these emergent patterns to their underlying processes is key to better understanding the impacts of human disturbance on the structure and function of tropical forests.     

Hyperspectral remote detection of niche partitioning among canopy trees driven by blowdown gap disturbances in the Central Amazon

Advanced recruitment and neutral processes play important roles in determining tree species composition in tropical forest canopy gaps, with few gaps experiencing clear secondary successional processes. However, most studies are limited to the relatively limited spatial scales provided by forest inventory plots, and investigations over the entire range of gap size are needed to better understand how ecological processes vary with tree mortality events. This study employed a landscape approach to test the hypothesis that tree species composition and forest structural attributes differ between large blowdown gaps and relatively undisturbed primary forest. Spectral mixture analysis on hyperspectral satellite imagery was employed to direct field sampling to widely distributed sites, and blowdown plots were compared with undisturbed primary forest plots. Tree species composition and forest structural attributes differed markedly between gap and non-gap sites, providing evidence of niche partitioning in response to disturbance across the region. Large gaps were dominated by classic Neotropical pioneer genera such as Cecropia and Vismia, and average tree size was significantly smaller. Mean wood density of trees recovering in large gaps (0.55 g cm⁻³) was significantly lower than in primary forest plots (0.71 g cm⁻³), a difference similar to that found when comparing less dynamic (i.e., tree recruitment, growth, and mortality) Central Amazon forests with more dynamic Western Amazon forests. Based on results, we hypothesize that the importance of neutral processes weaken, and niche processes strengthen, in determining community assembly along a gradient in gap size and tree mortality intensity. Over evolutionary time scales, pervasive dispersal among colonizers could result in the loss of tree diversity in the pioneer guild through competitive exclusion. Results also underscore the importance of considering disturbance processes across the landscape when addressing forest carbon balance.     

Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Liana Abundance,Diversity, and Distribution on Barro Colorado Island,Panama

Lianas are a key component of tropical forests; however, most surveys are too small to accurately quantify liana community composition, diversity, abundance, and spatial distribution – critical components for measuring the contribution of lianas to forest processes. In 2007, we tagged, mapped, measured the diameter, and identified all lianas ≥1 cm rooted in a 50-ha plot on Barro Colorado Island, Panama (BCI). We calculated liana density, basal area, and species richness for both independently rooted lianas and all rooted liana stems (genets plus clones). We compared spatial aggregation patterns of liana and tree species, and among liana species that varied in the amount of clonal reproduction. We also tested whether liana and tree densities have increased on BCI compared to surveys conducted 30-years earlier. This study represents the most comprehensive spatially contiguous sampling of lianas ever conducted and, over the 50 ha area, we found 67,447 rooted liana stems comprising 162 species. Rooted lianas composed nearly 25% of the woody stems (trees and lianas), 35% of woody species richness, and 3% of woody basal area. Lianas were spatially aggregated within the 50-ha plot and the liana species with the highest proportion of clonal stems more spatially aggregated than the least clonal species, possibly indicating clonal stem recruitment following canopy disturbance. Over the past 30 years, liana density increased by 75% for stems ≥1 cm diameter and nearly 140% for stems ≥5 cm diameter, while tree density on BCI decreased 11.5%; a finding consistent with other neotropical forests. Our data confirm that lianas contribute substantially to tropical forest stem density and diversity, they have highly clumped distributions that appear to be driven by clonal stem recruitment into treefall gaps, and they are increasing relative to trees, thus indicating that lianas will play a greater role in the future dynamics of BCI and other neotropical forests.     

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.     

Formation and closure of canopy gaps in the rain forest at Nouragues, French Guiana

Formation and closure of canopy gaps was monitored for three years in 12 ha of primary rain forest at Nouragues, French Guiana. At the first inventory, in April 1991, 74 openings in the canopy > 4 m² (sensu Brokaw 1982a) were located; 60 of these gaps were formed before January 1990. Between January 1990 and December 1993, 5 to 15 gaps were annually formed, opening 0.64–1.33% of the forest canopy each year. Of all gaps, 41% were created by a falling, snapped tree, 34% by a falling, uprooted tree, 22% by a falling branch, and 3% by a falling dead stem. A refined nearest neighbour analysis showed that gaps formed after January 1990 were clustered: uprooting of trees seemed to be related to shallow soils, and relatively many other trees fell when a tree uprooted, independent of the dbh of the uprooted tree. In 37 gaps, canopy openness in the gap centre (determined by hemispherical photographs) was monitored over three years. In 54% of the gaps, canopy openness increased in two successive years. It is reasoned that edges of especially large gaps may frequently be re-disturbed by falling trees or branches. Results suggest that gaps have closed after around 15 years. More data are needed to verify this.     

Patch structure and ramet demography of the clonal tree, Asimina triloba, under gap and closed-canopy

Clonal understory trees develop into patches of interconnected and genetically identical ramets that have the potential to persist for decades or centuries. These patches develop beneath forest canopies that are structurally heterogeneous in space and time. Canopy heterogeneity, in turn, is responsible for the highly variable understory light environment that is typically associated with deciduous forests. We investigated what aspects of patch structure (density, size structure, and reproductive frequency of ramets) of the clonal understory tree, Asimina triloba, were correlated with forest canopy conditions. Specifically, we compared A. triloba patches located beneath closed canopies and canopy gaps. We also conducted a three-year demographic study of individual ramets within patches distributed across a light gradient. The closed canopy-gap comparison demonstrated that the patches of A. triloba had a higher frequency of large and flowering ramets in gaps compared to closed-canopy stands, but total ramet density was lower in gaps than in closed canopy stands. In the demographic study, individual ramet growth was positively correlated with light availability, although the pattern was not consistent for all years. Neither ramet recruitment nor mortality was correlated with light conditions. Our results indicate that the structure of A. triloba patches was influenced by canopy condition, but does not necessarily depend on the responses of ramets to current light conditions. The lack of differences in ramet recruitment and mortality under varying canopy conditions is likely to be a primary reason for the long-term expansion and persistence of the patches. The primary benefit of a positive growth response to increasing light is the transition of relatively small ramets into flowering ramets within a short period of time.     

Habitat fragmentation in forests affects relatedness and spatial genetic structure of a native rodent,Rattus lutreolus

Habitat fragmentation can have a range of negative demographic and genetic impacts on disturbed populations. Dispersal barriers can be created, reducing gene flow and increasing population differentiation and inbreeding in isolated habitat remnants. Aggregated retention is a form of forestry that retains patches of forests as isolated island or connected edge patches, with the aim of ‘lifeboating’ species and processes, retaining structural features and improving connectivity. Swamp rats (Rattus lutreolus) are a cover‐dependent species that are sensitive to habitat removal. We examined the effects of aggregated retention forestry and forestry roads in native wet Eucalyptus forests on swamp rat gene flow and population genetic structure. We characterized neighbourhood size in unlogged forest to provide a natural state for comparison, and examined population structure at a range of spatial scales, which provided context for our findings. Tests of pairwise relatedness indicated significant differentiation between island and edge populations in aggregated retention sites, and across roads in unlogged sites. Spatial autocorrelation suggested a neighbourhood size of 42–55 m and revealed male‐biased dispersal. We found no genetic isolation by geographical distance at larger (>2.3 km) scales and populations were all significantly differentiated. Our results suggest that removal of mature forest creates barriers for swamp rat dispersal. In particular, roads may have long‐term impacts, while harvesting of native forests is likely to create only short‐term dispersal barriers at the local scale, depending on the rate of regeneration.     

The ecology, silviculture and biogeography of mahogany (Swietenia macrophylla): a critical review of the evidence

Recent studies of Swietenia macrophylla King argue that natural regeneration of this highly valued tree is episodic and occurs only after infrequent catastrophic disturbance. An analysis of 14 population size-distributions in natural forests shows that population structure varies across this species' range. In forests that are in successional transition from open deciduous forest to evergreen forest there is often prolific regeneration. We show that in this type of forest in southern Pará (Brazil) patches of even-sized trees are small. Large-scale disturbance may be necessary for regeneration only in evergreen forests where little light penetrates through the canopy. The transition zone between evergreen and deciduous forest in Pará is moving south, and we propose that mahogany populations to the north are in the process of competitive exclusion. Sustainable harvesting of mahogany must take account of the successional stage of the forest.     

Treefall Gaps and Regeneration Composition in the Laurel Forest of Anaga (Tenerife): a Matter of Size?

The size of treefall gaps is an important determinant of regeneration composition in tropical and temperate forests. Preliminary studies in the laurel forest of Tenerife have shown that small gaps (<100 m²) were the most numerous. However, due to this small size, no significant differences were found between regeneration in gaps and regeneration below the canopy. Because infrequent large gaps (>100 m²) are present in the laurel forest, we analyzed the regeneration in these large uncommon gaps, considering their potentially important role in the dynamics of the system. Our main hypothesis is that large gaps are important disturbance to ensure the regeneration and stablishment of shade intolerant species. Only five gaps larger than 100 m² (ranging from 125–268 m²) were found in the study area. Data from a further 20 small gaps (<100 m²), analysed in a previous study, was also included. Control plots were examined close to the gaps in order to determine regeneration below the closed canopy. We did not find a significant difference between regeneration density in the gaps (<100 m²) and regeneration below the canopy in the control plots. Contrary to our expectations, regeneration was lower in the large gaps than under the canopy. The open canopy in the large gaps increases light intensity, and has a negative effect on the germination and growth of shade-tolerant tree species like Viburnum tinus (although non-statistically significant); however, the increase in light intensity is not sufficient to stimulate the germination of shade-intolerant tree species. The effects of treefall gaps in the dynamics of the laurel forest of Anaga should be not considered as significant in comparison to other factors such as human disturbances or infrequent disturbances (land slides or hurricanes).     

Spatial heterogeneity of post-dispersal survivorship of Queensland rainforest seeds

Short-term seed survivorship for several species of the Queensland rainforest was compared in forests and early successional fields, treefall gaps and forest interior, and an array of microhabitats (tree buttresses, logs, forest floor with and without leaf litter cover). Seed survivorship was lower in forest than in field at one site, but equal at another. No differences were found in and out of treefall gaps, in contrast to several other studies. Microsites had little consistent influence on short-term seed survivorship. A survey of the literature showed no consistent habitat or microhabitat patterns in seed survivorship.     

Relative importance of climate vs local factors in shaping the regional patterns of forest plant richness across northeast China

Northeast (NE) China covers three climatic zones and contains all the major forest types of NE Asia. We sampled 108 forest plots in six nature reserves across NE China to examine the influence of climate and local factors (canopy seasonality, successional stage, topography and forest structure) on geographic patterns of plant richness. We analyzed the relative effects of different factors at two spatial scales: the regional scale (across both latitude and altitude) and the local scale (along the altitudinal gradient within site). Our results showed that the relative importance of climate vs local factors differed remarkably depending on scale and functional group. While total and tree species richness were mainly limited by climate, herb and shrub richness was more related to local factors (especially at the local scale). In the climatic factors, heat sum was the major correlate of tree, shrub and total species richness, while herb richness was more associated with winter coldness. Precipitation was not a limiting factor for forest plant richness in NE China. Climate accounted for 34–76% of variation in richness at the regional scale, but explained only 0–44% at the local scale. Among the local factors, shrub species richness was sensitive to seasonal canopy openness, with higher richness in deciduous forests than in the evergreen needle-leaf forest. On the other hand, herb richness was sensitive to forest successional stage, with higher richness in middle- successional forests than in the early and late-sucessional forests. Local topography (aspect and position on slope) and forest structure (tree density) also showed remarkable influence on species richness. Our results suggest the importance of including local factors when examining large scale diversity gradient (especially for understory species), and the necessity of comparing diversity patterns among functional groups at different spatial scales.     

Composition,structure and diversity of cove forest stands in the Great Smoky Mountains: a patch dynamics perspective

Abstract. Cove forests of the Great Smoky Mountains are North American examples of old-growth temperate forest. Ecological attributes of seven stands were studied using one 0.6 - 1.0 ha plot per stand. Stand basal area (39 - 55 m²/ha) and biomass (326 - 471 Mg/ha) were high for temperate deciduous forest. Density ranged from 577 to 1075 stems/ha. All stands had a mixture of deciduous canopy species. Only rarely did a single species comprise more than half of the stand by density, basal area or biomass. Shade-intolerant species were present at low levels (1 - 5 % of total stand density). A wide range of stem diameters was characteristic of most species. However, some species lacked small stems, indicating discontinuous regeneration. Stands tended to have 10 - 20 tree species per ha and at least five species had biomass levels > 10 Mg/ha, indicating high evenness. Canopy gaps covered 10 % of the total area (2 - 21 % by stand). Gaps and conspecific patches of canopy trees > 0.05 ha in size were infrequent. Spatial analyses revealed a variety of patterns among species at inter-tree distances of 1 to 25 m. When all species were combined, juveniles showed aggregation, and adults were often hyperdispersed. Analyses for individual species confirmed that the mosaic of canopy species is influenced by non-random spatial processes. Adults of several species were aggregated at distances > 10 m. Juveniles of all major species exhibited aggregation. Several species exhibited regeneration near conspecific adults. This pattern suggested limited mobility for such species within the shifting mosaic. A diverse patchwork resulted despite the fact that many species did not exhibit segregation of adults and juveniles. Further understanding of patch dynamics and the potential for compositional steady state in cove forests requires long-term study with spatial data.     

The spatial pattern and canopy-understory association of trees in a cool temperate,mixed forest in western Japan

The spatial distribution pattern of trees and the association between canopy and understory individuals were examined with reference to the distribution of tree crowns in a cool temperate, mixed forest in Ohdaigahara, western Japan. Line transect and contact sampling methods were used to examine the pattern over various spatial scales. These methods are useful to detect patterns over a large study area. The dominance ofChamaecyparis obtusa on steep slopes forming large patches suggested that the distribution of this species is a consequence of landslides. UnderstoryFagus crenata showed a clumped distribution, and the relative coverage of this species was larger in canopy gaps than under a closed canopy. Understory individuals ofAbies homolepis showed a positive association with canopy trees ofF. crenata but a negative association with conspecific canopy individuals. These patterns suggested thatF. crenata regenerates in canopy gaps and is replaced byA. homolepis. The dynamics of these two species are consistent with the process of gap dynamics. The effects of both small- and large-scale disturbance must be evaluated to understand the mechanisms of patch formation and the coexistence of forest tree species.