Stand development and regeneration during a 33-year period in a seral Picea glehnii forest, northern Japan

Stand development and regeneration were studied during a 33-year period (1965-1998) in a 1-ha plot in a seral Picea glehnii forest in northern Japan. P. glehnii was mono-dominant in the upper canopy layer, but its understory trees were rarely found in 1965. Other species were scarcely observed in 1965. Many recruited saplings of Abies sachalinensis which had grown to > 5 cm diameter at breast height (DBH) by 1998 had become dominant in the understory layer. Mortality of P. glehnii canopy trees was low. Therefore, the stand basal area increased during the census period due to the growth of surviving canopy trees. Stand development brought about intense competition among trees by increasing local crowding for each tree, and promoted dominance of larger trees and suppression of smaller trees. Although growth rates of understory trees of the two conifers decreased with the increase in local crowding, the growth rate of A. sachalinensis was consistently higher than that of P. glehnii at all extents of local crowding. The recruitment rate (growing to 5 cm DBH) of the two conifers was less affected by local crowding. However, the number of recruits of P. glehnii was only about a quarter of that of A. sachalinensis during the census period because the regeneration of P. glehnii was largely restricted to fallen logs and within 1 m of the base of any live tree > 20 cm DBH. Therefore, our long-term study suggests that A. sachalinensis will dominate over P. glehnii in the seral forest because of higher recruitment and growth rates of the former than the latter in the understory.An erratum to this article can be found at     

Effects of life history strategies and tree competition on species coexistence in a sub-boreal coniferous forest of Japan

We examined the effects of different life history strategies and tree competition on species coexistence in a northern coniferous forest. We investigated the growth and demography of trees with stems ≥1 cm dbh in a 2-ha study plot in the Taisetsu Mountains of northern Japan. Three species, Abies sachalinensis, Picea jezoensis, and Picea glehnii, were found to be dominant in the forest. A. sachalinensis was the most dominant species in the understory, while the two Picea spp. were more abundant in the larger dbh size classes. The turnover rate of A. sachalinensis was about twice that of the Picea spp. The relative growth rate of understory trees in each species did not differ between different canopy conditions (closed canopy or canopy gap). The competitive advantage between A. sachalinensis and P. glehnii switched as they grew from understory (A. sachalinensis superior competitor) to canopy trees (P. glehnii superior competitor). Meanwhile, A. sachalinensis and P. jezoensis exhibited different environmental preferences. We propose that reversal in competitive superiority between different growth stages and trade-off between longevity and turnover are more important factors to promote their coexistence than regeneration niche differentiation related to canopy gaps in this sub-boreal coniferous forest.     

Demographic traits of understory trees and population dynamics of aPicea-Abies forest in Taisetsuzan National Park, northern Japan

The size structure transition matrices ofPicea jezoensis, Picea glehnii andAbies sachalinensis of a sub-boreal forest in Hokkaido, northern Japan were constructed based on the demography of each species (Picea jezoensis andPicea glehnii were dealt with together asPicea) during a 4-year period. Two types of matrices, density-independent and density-dependent population dynamics models, were investigated for evaluating the ‘waiting pattern’ betweenPicea spp. andA. sachalinensis. For the density-dependent model, it was assumed that the demographic traits of understory trees, the recruitment rate, the understory mortality rate and the transition probability from the understory to canopy stages, were regulated by the one-sided competitive effect of canopy trees. The observed size structure ofPicea was almost consistent with the stationary size structure obtained in both the density-independent and the density-dependent models, whereas the observed size structure ofA. sachalinensis was not realized in the two models. The effects of both the transition probability from the understory to canopy stages and the recruitment rate on the dynamics of canopy trees were investigated. ForPicea, two parameters—recruitment rate (e _i ) and transition probability from the understory to canopy stages-exponentially affected the dynamics of canopy trees. In contrast, forAbies sachalinensis, the two parameters affected linearly the dynamics of canopy trees. In conclusion, the population dynamics ofPicea andA. sachalinensis was determined by the parameters of the recruitment rate and the transition probability from the understory to canopy stages, relating to waiting patterns of understory trees for future gap formation. InPicea, the demographic parameters of understory trees intensively regulated the dynamics of canopy trees if compared withA. sachalinensis, suggesting that the performance of understory trees plays a key role in the population dynamics ofPicea. This reflects the growth pattern of understory trees in the regeneration of the two species.     

Regeneration process of coniferous forests in northern Hokkaido I.Abies sachalinensis forest andPicea glehnii forest

Regeneration of natural forests was studied in the Nakagawa Experiment Forest of Hokkaido University using age distribution surveys made by the clear felling method. In Plot 1 (30 m × 65 m),Abies sachalinensis dominated the canopy layer but there were also a fewBetula ermanii trees.Sasa senanensis densely covered the forest floor. Most of the canopy trees were from 122 to 195 years old. Seedlings younger than 50 years old ofA. sachalinensis were found on fallen logs and root bases. There were, however, few trees from 50 to 120 years old. The present canopy trees seemed to have regenerated after competitive pressure from old canopy andSasa disappeared 180 years ago. Plot 2 (50 m × 100 m) on serpentinite soil was dominated byPicea glehnii. Sasa kulirensis covered the floor but not as densely asS. senanensis in Plot 1. The ages ofP. glehnii ranged from 1 to 586 years old, and the age distribution ofA. sachalinensis was L-shaped. A small gap in the canopy formed about 290 years ago, and it gradually extended. Conifers regenerated continuously in the extending gap butB. ermanii did not. One hundred thirty years ago, part of Plot 2 was again destroyed andA. sachalinensis andB. ermanii regenerated. Thus, two types of regeneration were found. One regenerated both conifers andBetula after a sudden disturbance of canopy layer or death ofSasa, and the other, under an extending gap, regenerated only conifers.     

Physiological and growth responses of switchgrass (Panicum virgatum L.) in native stands under passive air temperature manipulation

In the central Great Plains of North America, climate change predictions include increases in mean annual temperature of 1.5–5.5 °C by 2100. Ecosystem responses to increased temperatures are likely to be regulated by dominant plant species, such as the potential biofuel species Panicum virgatum (switchgrass) in the tallgrass prairie. To describe the potential physiological and whole‐plant responses of this species to future changes in air temperatures, we used louvered open‐sided chambers (louvered OSC; 1 × 1 m, adjustable height) to passively alter canopy temperature in native stands of P. virgatum growing in tallgrass prairie at varying topographic positions (upland/lowland). The altered temperature treatment decreased daily mean temperatures by 1 °C and maximum temperatures by 4 °C in May and June, lowered daytime stomatal conductance and transpiration, decreased tiller density, increased specific leaf area, and delayed flowering. Among topographic contrasts, aboveground biomass, flowering tiller density, and tiller weight were greater in lowland sites compared to upland sites, with no temperature treatment interactions. Differences in biomass production responded more to topography than the altered temperature treatment, as soil water status varied considerably between topographic positions. These results indicate that while water availability as a function of topography was a strong driver of plant biomass, many leaf‐level physiological processes were responsive to the small decreases in daily mean and maximum temperature, irrespective of landscape position. The varying responses of leaf‐level gas exchange and whole‐plant growth of P. virgatum in native stands to altered air temperature or topographic position illustrate that accurately forecasting yields for P. virgatum in mixed communities will require greater integration of physiological responses to simulated climate change (increased temperature) and resource availability over natural environmental gradients (soil moisture).     

Differential photosynthetic characteristics between seedlings and saplings of Abies sachalinensis and Picea glehnii, in the field

Abies sachalinensis and Picea glehnii are co-dominant tree species and major components of the forests of Hokkaido, Japan. Recent work suggests that a reversal in potential competitive superiority at different developmental stages could be important to explain their coexistence. Such shifts in competitive advantage can be mechanistically understood by studying the corresponding physiological differences between distinct life stages. Accordingly, our objective was to investigate in the field the photosynthesis of shade-growing juveniles of these species from two different size-classes, seedlings and saplings. Our results show that seedlings of both species had higher concentrations of photoprotective xanthophylls than saplings, especially in spring, and suggest that seedlings have a lower threshold of stress tolerance than saplings. Photosynthetic capacity per needle area and lateral shoot extension rate decreased from the seedling to the sapling stage in A. sachalinensis, while in P. glehnii, both increased from the seedling to the sapling stage. Abies sachalinensis had higher photosynthetic rates at the seedling stage but lower rates at the sapling stage than P. glehnii. Nevertheless, A. sachalinensis had a higher lateral shoot extension rate than P. glehnii at both stages. Our physiological results support previous ecological observations that A. sachalinensis is a superior competitor to P. glehnii in the understory, and show that its competitive advantage is higher at the seedling stage than at the sapling stage.     

Growth and survival of <Emphasis Type="Italic">Abies sachalinensis</Emphasis> seedlings for three years after selection harvesting in northern Hokkaido,Japan

Previous studies have indicated that recruitment of Abies sachalinensis, a representative conifer species of northern Japan, decreased following single-tree selection harvesting in stands with dense dwarf bamboo understory. We tested the hypothesis that growth and survival of A. sachalinensis seedlings are reduced by canopy opening in that type of stand. A 0.75 ha study plot was examined, and all the seedlings (defined as trees with height 0.5–2 m) were identified and their shoot extensions measured for three years after single-tree selection harvesting (26% intensity in terms of basal area). The leader extensions of A. sachalinensis seedlings that experienced canopy opening were greatly improved. However, a negative effect on survival was also apparent; nearly 40% of seedlings died at the sites that experienced canopy opening. These results were supported by generalized linear models that examined variations of local harvesting intensity for individual seedlings. Despite the ability of A. sachalinensis to respond rapidly to exposure, some physiological stresses may have appeared, and presumably were amplified by co-occurring dwarf bamboos. With regard to the low seedling density (156 stems ha⁻¹), common in this type of stands, local harvesting intensity in the selection system should be reduced to maintain survival of advanced regeneration, thus sustaining stand structure and composition.     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Stand structure and growth patterns of understorey trees in a coniferous forest,Taisetsuzan National Park,northern Japan

Stand structure was studied with special reference to growth and mortality patterns of sapling and understorey trees in a coniferousPicea jezoensis andAbies sachalinensis forest in Taisetsuzan National Park, Hokkaido, northern Japan.Picea jezoensis was dominant in the basal area, whileA. sachalinensis was abundant in large numbers in the canopy. Estimated mortalities increased significantly with diameter at breast height (DBH) for bothP. jezoensis andA. sachalinensis in the canopy, but the tendency was different between the two species.Picea jezoensis had a lower mortality rate thanA. sachalinensis, especially at small DBH classes. The spatial distribution of understorey individuals ofA. sachalinensis did not show any significant correlation with the spatial distribution of canopy gaps, but that ofP. jezoensis showed a significant correlation.Abies sachalinensis can grow higher thanP. jezoensis under suppressed conditions; whileP. jezoensis requires canopy gaps for steady height growth. This growth pattern leads to a different waiting height in the understorey (≥2 m in height and 10 cm in diameter at breast height).Abies sachalinensis waited for an improvement in light conditions at higher strata (max. 7 m), whileP. jezoensis waited at lower strata (max. 3 m). The estimated mortality of understoreyA. sachalinensis increased with size, while that of understoreyP. jezoensis decreased. Therefore,P. jezoensis gives priority to survival whileA. sachalinensis gives priority to understorey growth. The difference in the ‘waiting pattern’ between the two species in the understorey was considered a significant feature for the canopy recruitment process ofP. jezoensis andA. sachalinensis.     

Effects of disturbance and size structure on the regeneration process in a sub-boreal coniferous forest,northern Japan

The stand structure and disturbance history in a sub-boreal coniferous forest dominated byPicea jezoensis, Picea glehnii andAbies sachalinensis were investigated in four study plots set up in Taisetsuzan National Park, Japan. The effect of stand characteristics on the growth and mortality rates of understory trees was examined. Although all the stands showed inverse J-shape d.b.h. (diameter at breast height) distributions, the age structure and disturbance history differed amongst the stands. The stands with wide d.b.h. distribution (i.e. large CV and skewness) were more uneven-aged than those with narrow d.b.h. distribution (i.e. small CV and skewness). The disturbance-return interval based on the model of Hett and Loucks was 31 to 65 years. The gap ratio in the canopy was also different among the stands. These suggest that the variations in stand structure represent different occurrences of natural disturbances. Furthermore, the structural features such as size structure, canopy gap ratio and density of canopy trees also affected the growth dynamics of understory trees (≥2 m in height and <10 cm in diameter at breast height). The growth and mortality rates of understory trees changed with the canopy gap ratio and canopy tree density. The understory trees of stands with wide canopy d.b.h. distribution had higher growth and canopy recruitment rates than those of stands with narrow canopy d.b.h. distribution, contributing to the maintenance of continuous stand stratification. The understory trees of stands with narrow canopy d.b.h. distribution showed lower growth and higher mortality rates than those of stands with narrow canopy d.b.h. distribution, leading to the formation of a single-canopy structure. It is suggested that natural disturbance governs the regeneration process in the future by affecting the growth and mortality patterns of understory trees through the stand structure (size and age structure, canopy tree density, canopy gap ratio).     

Relationship between plant cover type and soil properties on Syunkunitai coastal sand dune in eastern Hokkaido

A survey of soils and trees was conducted on Syunkunitai coastal sand dune in eastern Hokkaido to clarify the relationships between the soil properties and the plant cover type. A belt transect of 360m in length was established across the dune. Three community types, that is, a Picea glehnii forest, an Abies sachalinensis forest, and a salt marsh were recognized. Soil types at the study area were determined to be sandy immature soil and peat soil. Their horizon sequences were described as A₀–V–C or T–V–C layers (A₀, T, V, and C indicate layers of leaf litter, peat, volcanic deposit, and parent material, respectively). The Abies sachalinensis forest was characterized by a relatively high calcium concentration in the surface soil layer and a tendency for podzolization in the volcanic deposit layer. The Picea glehnii forest was characterized by peat accumulation because of the high ground water table, volcanic deposits in the soil profile, and the strong influence of sea salt on the soil chemistry. The roots in the Picea glehnii forest were distributed more shallowly than those in the Abies sachalinensis forest, thus avoiding the high water table level as well as the influence of seawater in the soil. The salt marsh showed an extremely high sodium concentration and base saturation, indicating that this area was directly affected by seawater. Recently, the periphery of the Picea glehnii forest on Syunkunitai sand dune has been declining because of seawater inundation caused by ground subsidence.     

Contrasting traits,contrasting environments,and considerations on population dynamics under a changing climate: an ecophysiological field study of two co‐dominant tree species

The increase in light availability resulting from canopy changes or opening is not always beneficial and can inhibit photosynthesis of tree seedlings already under other environmental stress. Tree seedlings' responses to compounded abiotic stress depend on their life‐history traits, and understanding the variations of such responses is important for understanding population dynamics under a changing climate. In this study we investigate how the photosynthesis of juveniles of two canopy tree species with different life‐history traits, Abies sachalinensis and Betula ermanii, differs in two contrasting sites at a sub‐boreal forest in northern Japan—one under a deciduous canopy (Closed site) and the other at a wide canopy opening (Open site). Seedlings at the Open site had low F_v/F_m (quantum yield of photosystem II) for a longer period than those at the Closed site. Abies sachalinensis at the Closed site showed lower F_v/F_m in spring than those at the Open site, but recovered after the canopy's new leaves flushed, indicating its acclimation to the shaded condition. Mean P_max (light‐saturated photosynthetic rate at ambient CO₂ levels) of A. sachalinensis seedlings was affected by site and air temperature, while B. ermanii seedlings were also affected by precipitation. Only B. ermanii's seedlings presented growth in the period studied, in spite of observed mid‐day drops to F_v/F_m attributed to water‐deficit‐related photoprotection. Results suggest that the climate change predicted for the Hokkaido area may increase the competitive advantage of broad‐leaved deciduous species, such as B. ermanii, in relation to evergreen conifers like A. sachalinensis.     

Firescape ecology: how topography determines the contrasting distribution of fire and rain forest in the south‐west of the Tasmanian Wilderness World Heritage Area

Aim To test the hypothesis that ‘islands’ of fire‐sensitive rain forest are restricted to topographic fire refugia and investigate the role of topography–fire interactions in fire‐mediated alternative stable state models. Location A vegetation mosaic of moorland, sclerophyll scrub, wet sclerophyll eucalypt forest and rain forest in the rugged, fire‐prone landscapes of south‐west Tasmania, Australia. Methods We used geospatial statistics to: (1) identify the topographic determinants of rain forest distribution on nutrient‐poor substrates, and (2) identify the vegetation and topographic variables that are important in controlling the spatial pattern of a series of very large fires (> 40,000 ha) that were mapped using Landsat Thematic Mapper (TM) satellite imagery. Results Rain forest was more likely to be found in valleys and on steep south‐facing slopes. Fires typically burned within highly flammable treeless moorland and stopped on boundaries with less flammable surrounding vegetation types such as wet sclerophyll forest and rain forest. Controlling for the effect of vegetation, fires were most likely to burn on flats, ridges and steep north‐facing slopes and least likely to burn in valleys and on steep south‐facing slopes. These results suggest an antagonism between fire and rain forest, in which rain forest preferentially occupies parts of the landscape where fire is least likely to burn. Main conclusions The distribution of rain forest on nutrient‐poor substrates was clearly related to parts of the landscape that are protected from fire (i.e. topographic fire refugia). The relative flammability of vegetation types at the landscape scale offers support to the proposed hierarchy of fire frequencies (moorland > scrub > wet sclerophyll > rain forest) that underpins the ecological models proposed for the region. The interaction between fire occurrence and a range of topographic variables suggests that topography plays an important role in mediating the fire–vegetation feedbacks thought to maintain vegetation mosaics in south‐west Tasmania. We suggest that these fire–topography interactions should be included in models of fire‐mediated alternative stable vegetation states in other fire‐prone landscapes.     

Occurrence and types of ectomycorrhizae present in seedlings ofPicea glehnii in a natural forest in Hokkaido

Picea glehnii seedlings whose ages ranged from 2 to 5 years and which were growing in aP. glehnii-Abies sachalinensis natural forest where the surface had been scarified 6 years before, were assessed for the occurrence and types of ectomycorrhizae. Based on the macro- and microscopic characteristics, 34 types of ectomycorrhizae were classified. The basidiomycetes were clearly dominant and it was common to observe more than one type of mycorrhiza occurring in the same root tip. The diversity of mycorrhizal types and the presence of at least one type of ectomycorrhiza in a single seedling, observed in this work, indicate a very dynamic system of mycorrhizal formation inP. glehnii seedlings, which allows them to survive and grow in a forest surface subjected to soil scarification.     

Size-structure dynamics of two conifers in relation to understorey dwarf bamboo: A simulation study

Abstract. Size-structure dynamics and co-existence of the conifers Abies sachalinensis and Picea glehnii in subalpine forests in Japan, in relation to the understorey dwarf bamboo Sasa, were examined using a simulation model. This model explicitly incorporated recruitment rate, size growth rate and mortality. Recruitment and growth rates were assumed to be suppressed by the cumulative basal area of taller trees and the density of Sasa. As for the one-sided crowding effect, two types of model were examined. The total basal area of the two species suppresses the growth and recruitment rates in an additive model, while the basal area of each species has a separate effect included in a specific model. Two types of recruitment process were examined, i.e. open and closed systems; recruitment rates of open and closed systems are independent of, and proportional to, the basal area of the same species within the plot, respectively. Parameters were estimated from data of plot censuses for four years. The parameters of the specific model showed that recruitment and size growth rates were more suppressed by the same species than the other species. Recruitment of A. sachalinensis was more sensitive to suppression by Sasa compared with P. glehnii. The stationary size structure of the two species, generated from both the models with the open system, fitted well to the observed size structures across various Sasa densities. A. sachalinensis and P. glehnii dominated at lower and higher densities of Sasa, respectively. However, the closed-system simulation showed that the stable co-existence of the two species with a good fit to the observed size structure occurred only in the specific model. These results suggest that within-species interference, which is more severe than between-species interference, is important for the co-existence of the two species and that the relative dominance of the two species readily explained the differential responses to Sasa abundance.     

Differential survival among life stages contributes to co‐dominance of Abies mariesii and Abies veitchii in a sub‐alpine old‐growth forest

Questions: Are there interspecific differences in mortality and recruitment rates across life stages between two shade‐tolerant dominant trees in a sub‐alpine old‐growth forest? Do such differences in demography contribute to the coexistence and co‐dominance of the two species? Location: Sub‐alpine, old‐growth forest on Mt. Ontake, central Honshu, Japan. Methods: From 1980 to 2005, we recorded DBH and status (alive or dead) of all Abies mariesii and A. veitchii individuals (DBH ≥ 5 cm) in a 0.44‐ha plot. Based on this 25 year census, we quantified mortality and recruitment rates of the two species in three life stages (small tree, 5 cm ≤ DBH < 10 cm; subcanopy tree, 10 cm ≤ DBH < 20 cm; canopy tree, DBH ≥ 20 cm). Results: Significant interspecific differences in mortality and recruitment rates were observed in both the small tree and sub‐canopy tree stages. In this forest, saplings (< 5 cm DBH) are mostly buried by snow‐pack during winter. As a consequence, saplings of A. mariesii, which is snow and shade tolerant, show higher rates of recruitment into the small tree stage than do those of A. veitchii. Above the snow‐pack, trees must tolerate dry, cold temperatures. A. veitchii, which can more readily endure such climate conditions, showed lower mortality rate at the subcanopy stage and a higher recruitment rate into the canopy tree stage. This differential mortality and recruitment among life‐stages determines relative dominance of the two species in the canopy. Conclusion: Differential growth conditions along a vertical gradient in this old forest determine survival of the two species prior to reaching the canopy, and consequently allow co‐dominance at the canopy stage.     

Growth trends and dynamics in sub‐alpine forest stands in the Varaita Valley (Piedmont,Italy) and their relationships with human activities and global change

Abstract. A study of the forest lines, tree lines and the structures of the sub‐alpine forest was performed in Vallone Vallanta and in Alevé forest in the Varaita Valley (Cottian Alps, Piedmont, Italy). Forest‐ and tree lines were analysed over 1728 ha while forest structures were studied on six 3000‐m² plots located at the tree line (2), at the forest line (2) and inside the sub‐alpine forest (2). Dendro‐ecological analysis of living plants and stumps showed that Larix decidua was more abundant in the past than today and that Pinus cembra has expanded, both upwards and within sub‐alpine forests. Age structure analysis revealed that the current sub‐alpine forest stands were established 200–220 yr ago, probably following a clearcut. At the forest lines the tree density decreases, and some trees are more than 500 yr old, whereas at the tree lines most of the trees (almost exclusively Pinus cembra) are younger than 100 yr. Growth dynamics were investigated both by observing Basal Area Increment (BAI) in the old and dominant trees, and by comparing the BAIs of classes of trees with a given cambial age range in different time periods. The results showed that the growth rates of mature Pinus cembra and Larix decidua had increased. These increments are more substantial for Pinus than for Larix. The growth rate of young trees (< 100 yr) of both species has decreased over recent decades. This could be due to competition caused by increased tree densities that have resulted from a decrease in grazing.     

Microtopography and distribution of canopy trees in a subtropical evergreen broad-leaved forest in the northern part of Okinawa Island,Japan

The spatial distribution of trees in relation to topography was investigated using two topographic indices, slope steepness and slope configuration, in a 4-ha plot in a subtropical evergreen forest in the northern part of Okinawa Island, Japan. Most species showed an aggregated distribution and patch size was related to various microtopographical features, including small ridges and valleys. In a cluster analysis based on the dissimilarity of each species distribution, the species were divided into three major clusters. A significant correlation between the dissimilarity and the distance between each species on the axis of the two topographic indices indicated that species association was, in part, explained by the topographic indices. I suggest that species distributed on steep and concave slopes regenerate depending on disturbances such as landslides on unstable topography, whereas species distributed on ridges and upper slopes regenerate depending on the canopy gap. A number of species that were less abundant in the 4-ha plot occurred in the riparian area near a stream, where the density of more abundant species was low. The results of this study support the existence of habitat niche divergence related to topography in a subtropical evergreen broad-leaved forest.     

Comparative demography of three coexisting Acer species in gaps and under closed canopy

Questions: 1. Is there a trade‐off between gap dependency and shade tolerance in each of the life‐history stages of three closely related, coexisting species, Acer amoenum (Aa), A. mono (Am) and A. rufinerve (Ar)? 2. If not, what differences in life‐history traits contribute to the coexistence of these non‐pioneer species? Location: Ogawa Forest Reserve, a remnant (98 ha), species‐rich, temperate deciduous forest in central Japan (36°56’ N, 140°35’ E, 600 ‐ 660 m a.s.l.). Methods: We estimated the demographic parameters (survival, growth rate and fecundity) by stage of each species growing in gaps and under closed canopy through observations of a 6‐ha permanent plot over 12 years. Population dynamics were analysed with stage‐based matrix models including gap dynamics. Results : All of the species showed high seedling and sapling survival rates under closed canopies. However, demographic parameters for each growth stage in gaps and under closed canopies revealed inter‐specific differences and ontogenetic shifts. The trade‐off between survival in the shade and growth in gaps was detected only at the small sapling stage (height < 30 cm), and Ar had the highest growth rate both in the shade and in the gaps at most life stages. Conclusions: Inter‐specific differences and ontogenetic shifts in light requirements with life‐form differences may contribute to the coexistence of the Acer species in old‐growth forests, with Aa considered a long‐lived sub‐canopy tree, Am a long‐lived canopy tree, and Ar a short‐lived,‘gap‐phase’ sub‐canopy tree.     

Succession in sub‐boreal forests of West‐Central British Columbia

Abstract. Structural and compositional changes were analysed over the course of 400+ yr of post‐fire succession in the sub‐boreal forests of west‐central British Columbia. Using a chronosequence of 57 stands ranging from 11 to 438 yr in age, we examined changes in forest structure and composition with complementary PCA and DCA ordination techniques. To determine stand ages and timing of tree recruitment, approximately 1800 trees were aged. Most early successional forests were dominated by Pinus contorta, which established rapidly following fire. Abies lasiocarpa and Picea glauca × engel‐mannii were also able to establish quickly, but continued to establish throughout the sere. Few Pinus contorta survived beyond 200 yr, resulting in major changes in forest structure. In some stands P. contorta never established, which led to considerable variation among stands less than 200 yr old. The oldest forests converged on dominance by Abies lasiocarpa. Vascular plant diversity decreased during succession whereas canopy structure became more complex as gap dynamics developed. Although these sub‐boreal forests contain few tree species, successional changes were pronounced, with structure changing more than composition across the chronosequence.