The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest

We assessed the composition of understory vascular plant communities in relation to the mosaic of canopy patch types, and their associated structure and environment, within unmanaged, mature boreal mixedwood forests in western Canada. Within a 30 km² area, we sampled patches of four different canopy types: conifer-dominated, broadleaf-dominated, mixed conifer-broadleaf, and canopy gaps (total n = 98). There were significant differences in understory composition among the four patch types (based on multi-response permutation procedure (MRPP)) and these were mainly due to differences in relative abundances of understory species. The understory communities of conifer patches were characterized by low abundances of shade intolerant species while shade-tolerant and evergreen species were indicators (based on an indicator species analysis (ISA)). Understory communities under gap and broadleaf patches were characterized by higher abundances of grasses and shade intolerant species. Gap, broadleaf, and mixed patches had higher abundances of certain shrub species than did conifer patches. The patch types also differed in terms of their environmental conditions. Conifer patches had drier, cooler soils and the lowest understory light. Broadleaf patches had the warmest soils while understory light during the leaf-off period was similar to that of canopy gaps. Gap patches had the lowest litter cover and PO₄⁻ availability and the highest light. Seven environmental variables (soil moisture, soil temperature, total light during the leaf-off period, cover of coarse and fine downed woody material, and availability of NH₄⁺ and Ca²⁺) were significantly related to understory species composition (in a constrained ordination by means of a distance-based redundancy analysis (db-RDA); 16.5% of variation in understory community data explained). Even within a single patch type, there was substantial environmental variation that was related to understory species composition. Our study suggests that the mosaic of canopy patches within mixedwood forests supports coexistence of both early and late successional understory plant species in mixedwood stands. Maintaining the mixture of canopy patch types within mixedwood stands will be important for conserving the natural patterns of understory plant composition in boreal mixedwood forests.     

Effects of aspen canopy removal and root trenching on understory microenvironment and soil moisture

Effects of three aspen (Populus tremuloides Michx.) canopy removal treatments and root trenching on understory microenvironment and moisture were tested at Parkland and Boreal sites in Alberta, Canada. Aspen canopies moderated air temperature by reducing maximums and increasing minimums, and increased the frost-free period in the understory by reducing radiative frosts. When daily differences were found among canopy treatments, maximum absolute humidity was greater with complete canopy removal. Maximum daily relative humidity was greater in openings at night than with either full or partial canopy cover. Predictably, increasing aspen cover reduced PAR reaching the understory. Soil moisture response was highly variable, changing with site, aspen density and precipitation patterns, but there were only marginal differences due to root trenching. In the Parkland site, soil moisture conservation from aspen canopy and leaf litter effects were masked by tree uptake in most periods, but a net increase in soil water (+5.2%) was observed during drought. Soil and microclimatic conditions in thinned aspen stands suggest potentially favourable production benefits from developing and adopting agroforestry systems in these northern ecosystems.     

Understory plant diversity and biomass in hybrid poplar riparian buffer strips in pastures

Understory plant biomass, species richness and canopy openness were measured in six-year old hybrid poplar riparian buffer strips, in the understory of two unrelated clones (MxB-915311 and DxN-3570), planted along headwater streams at three pasture sites of southern Quebec. Canopy openness was an important factor affecting understory biomass in hybrid poplar buffers, with lower understory biomass observed on sites and under the clone with lower canopy openness. Although tree size was an important factor affecting canopy openness, relationships between total stem volume and canopy openness, for each clone, also support the hypothesis of a clonal effect on canopy openness. Understory biomass and canopy openness as low as 3.6 g m⁻² and 7.6% in 1 m² microplots were measured under clone MxB-915311 at the most productive site. This reduction of understory plant growth could compromise important buffer functions for water quality protection (runoff control, sediment trapping and surface soil stabilisation), particularly were concentrated runoff flow paths enter the buffer. On the other hand, tree buffers that maintain relatively low canopy openness could be interesting to promote native and wetland plant diversity. Significant positive relationships between canopy openness and introduced species richness (R ² = 0.46, p < 0.001) and cover (R ² = 0.51, p < 0.001) were obtained, while no significant relationship was observed between canopy openness and native (wetland) species richness and cover. These results suggest that planting riparian buffer strips of fast-growing trees can rapidly lead to the exclusion of shade-intolerant introduced species, typical colonisers of disturbed habitats such as riparian areas of pastures, while having no significant effect on native (wetland) diversity. Forest canopy created by the poplars was probably an important physical barrier controlling introduced plant richness and abundance in agricultural riparian corridors. A strong linear relationship (R ² = 0.73) between mean total species richness and mean introduced species richness was also observed, supporting the hypothesis that the richest communities are the most invaded by introduced species, possibly because of higher canopy openness, as seen at the least productive site (low poplar growth). Finally, results of this study highlight the need for a better understanding of relationships between tree growth, canopy openness, understory biomass and plant diversity in narrow strips of planted trees. This would be useful in designing multifunctional riparian buffer systems in agricultural landscapes.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.     

Growth Dynamics During Canopy Recruitment of Sprout-Origin Stems in Japanese Beech (Fagus japonicaMaxim.) Stools in Old Growth Forests of Central Japan

Abstract

In order to clarify mechanisms of canopy recruitment of Fagus japonicastems of sprout origin, age structure and growth dynamics were studied for stems within beech stools in two stands of old growth forests on the Pacific side of central Japan. The DBH vs. height and age relationships of the beech stems showed continuous distribution from small-young, understory stems to large-old, canopy ones. There were one to seven definable periods of suppression prior to canopy recruitment in F japonica.Prior to canopy recruitment number and length of release episodes were fewer and shorter than those of suppression. E japonicastems of sprout origin can endure suppression for a long time. Continuous emergence and the alternation of fast and slow growth in F. japonicastems suggest the sprouting ability for the beech canopy can play an important role in the successful closure of canopy openings. Canopy stems showed patterns of suppression and release that were often synchronous with other canopy stems within the same beech stool. Minor disturbances such as single crown falls can play an important role in the maintenance of the multi-stem structure of beech stools in old growth forests.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Carbon and nitrogen dynamics in topsoils along forest conversion sequences in the Ore Mountains and the Saxonian lowland, Germany

Carbon and nitrogen stocks and their medium-term and readily decomposable fractions in topsoils were compared in relation to soil microbial biomass and activity along sequences from coniferous to deciduous stands. The study was carried out in the Ore Mountains and the Saxonian lowland, representing two typical natural regions in Saxony, Germany. In accordance with current forest conversion practices, the investigation sites represent different stands: mature conifer stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) (type A); Norway Scots spruce and pine with advanced plantings of European beech (Fagus sylvatica L.) or European beech/Common oak (Quercus petreae Liebl.) (type B); and mature deciduous stands of European beech and European beech/Common oak (type C). The investigated forest sites can be grouped into three silvicultural situations according to the development from coniferous stands to advanced plantings and finally mature deciduous forests (chronosequence A–B–C). The organic layer (L, F and H horizons) and uppermost mineral soil (0–10 cm) were analysed for potential C mineralisation, microbial biomass, concentrations of total C and N (TOC and TN) and for medium-term and readily decomposable C and N fractions, obtained by hot- and cold-water extraction respectively. The results showed an increase in organic layer thickness and mass as well as TOC and TN stocks along the forest sequences in the lowland. Yet, underplanted sites with two storeys revealed higher organic layer mass as well as TOC and TN stocks as compared to coniferous and deciduous stands. Stocks of hot- and cold-water-extractable C and N in relation to microbial biomass and its activity revealed a high turnover activity in deeper organic horizons of deciduous forests compared to coniferous stands. The stand-specific differentiation is discussed in relation to microbial biomass, litter quantity and quality and forest structure, but also with respect to the site-specific climatic factors and water budget as well as liming and fly-ash impacts. Results indicate higher dynamics in deciduous stands in the lowland especially during the initial turnover phase. The elevated microbial activity in deeper organic horizons of deciduous litter-influenced sites in spring is discussed as a specific indicator for long-term C sequestration potential as besides C mineralisation organic compounds are humified and thus, can be stored in the organic layer or in deeper soil horizons. Due to liming activities, stand-specific effects on organic matter turnover dynamics have evened out today in the Ore mountain region, but will presumably occur again once base saturation decreases. Here, the stand-specific effect on microbial biomass can currently be seen again as C_mic in the L horizon increased from spruce to beech. Our study sites in the lowland revealed no significant fly-ash impact. Differences between sites were evaluated by calculating the discriminance function. TOC and TN as well as medium-term degradable C and N were defined in this study as indicators for turnover dynamics along forest conversion sites.     

Tree species composition affects productivity and carbon dynamics of different site types in boreal forests

The objective was to analyse how differences in the initial proportions of tree species and site fertility affect carbon sequestration in living biomass and soil. We used the individual-based simulation model EFIMOD, which is able to simulate spatially explicit competition between trees for light and nutrients. Simulations were carried out for three site types with distinct initial stocks of soil nutrients. For each site, the 100-years undisturbed dynamics of monocultures and mixtures of three tree species (Betula pendula Roth, Pinus sylvestris L. and Picea abies (L.) H. Karst.) was predicted. Changes in the proportions of competing tree species were dependent on the fertility of the site: on poor sites, pine was the most competent species, while on rich sites, spruce increased its proportion during stand succession. Net primary production (NPP) and soil respiration were the highest in stands of two coniferous species and in stands with a high initial proportion of pine. Mixed stands were more productive than monocultures; the highest overyielding was observed with mixtures of two coniferous species. Simulated NPP and carbon stocks in all pools increased from poor to rich sites. The highest carbon stocks in standing biomass were observed for mixtures of conifer species and three-species mixtures; the greatest accumulation of forest floor occurred in stands with high proportions of pine.     

Stability of subalpine forest-meadow boundary inferred from size and age structure ofAbies mariesii thickets on a Japanese snowy mountain

Subalpine forest zone in the Japanese snowy mountains is characterized by a mosaic of coniferous forests and meadows.Abies mariesii thickets surround mesic meadows, forming a forest-meadow ecotone. Size and age structure of the thickets were studied to examine the stability of the forest-meadow boundaries on Mount Aizukomagatake in central Japan. The age structure indicates that the small height ofA. mariesii in the thickets was not due to the early stage of succession but due to adverse site conditions. In spite of their small stem height, they were as old as canopy trees in neighboringA. mariesii forests. Snow settling pressure madeA. mariesii thickets with broken main stems on the crest slopes, while snow glide pressure formed those with bending stems on leeward slopes. No evidence was found to show any major changes in the position of the forest-meadow boundaries in the size and age structure. The boundary position appears to have been almost steady for at least this century.     

Changes in soil properties of Abies holophylla and Quercus-dominated stands 4 years after trenching

Abstract

Few studies tried to isolate the influence of plant roots on the soil characteristics including soil carbon (C) and nitrogen (N) storage. We evaluated the impact of root trenching on soil characteristics of coniferous (Abies holophylla) and deciduous (Quercus spp.) stands 4 years after trenching. Trenching treatment significantly increased the soil water content and nitrate concentration in both stand types. Soil pH, cation exchange capacity, soil C and N pools and isotopic compositions of C and N were significantly different between two stands, but trenching was not found to have significant impact on these soil properties. Our results indicated that root trenching in coniferous and broad-leaved deciduous forests of temperate region could significantly alter soil moisture regime and inorganic N levels, but not C and N stabilization in soils.     

Effects of forest structure and microhabitat on the distribution and flowering of a rare understory plant, Actaea elata

Knowing which environmental characteristics constrain forest understory species’ distribution and reproduction can inform conservation decisions about habitat management and locations for reintroduction efforts. Differential plant performance is common in varying environmental conditions, such as levels of canopy closure. Actaea elata (tall bugbane) is a rare, perennial, forest understory herb found from Oregon to British Columbia. Populations throughout this species’ range commonly occur in managed forests. We assessed the importance of environmental variation on the distribution and flowering of A. elata at both microhabitat and site-level scales. At four sites encompassing a variety of forest types, we measured several biotic and abiotic variables at vegetative and reproductive plants, as well as at random points. We measured these same variables at three sites where A. elata has been extirpated. Within occupied sites, reproductive plants were more likely to be located in canopy gaps than were vegetative plants. Both vegetative and reproductive plants were more likely to be found in areas of high herb cover compared to random points. These results indicate that gaps in the canopy may be critical for this species’ ability to reproduce, but survival may be possible in a broader range of forest understory conditions (similar to other co-occurring herb species). Random points at extirpated and occupied sites both tended to be located in areas of lower canopy openness and moisture availability than points in occupied sites where A. elata was present. Extirpated sites were either densely forested or clearcut, while occupied sites contained stands of intermediate density. Circaea alpina, Geranium robertianum, and Lactuca muralis were indicator species for A. elata and normally indicate moist soils, with Circaea alpina being a strong indicator of the presence of flowering A. elata. Taken together, our results suggest that maintenance of canopy gaps is important to promote flowering, and understory species that indicate moist sites, particularly Circaea alpina, can be used as aids to locate new populations of A. elata.     

Diversity of naturally-regenerated native woody species in forest plantations in the Ethiopian highlands

Woody species diversity and ground layervegetation cover were studied in plantations ofPinus patula, Cupressus lusitanica,Grevillea robusta, and Juniperusprocera, and in surrounding natural forests inWondo Genet, Ethiopia. In the understory of theplantations, a total of 53 naturallyregenerated tree and shrub species belonging to31 families were recorded. Important indigenoustimber species were also represented. Treesaccounted for 72% of all naturally-regeneratedwoody plant species. On the average, 78% ofthe woody species found in plantations werealso found in the natural forest. There was nosignificant variation in understory woodyspecies richness among plantations. C.lusitanica plantations had a significantlyhigher abundance of woody species than those ofP. patula, but the difference wasnot significant in relation to J. procera,G. robusta and the adjacent natural forest.The herbaceous ground cover percentage in G. robusta and P. patula stands wasconsiderably higher than in C. lusitanicaand J. procera stands. Relative toplantations of other species, plantations ofthe native tree species J. procera hadhigher understory species richness, mediumwoody species abundance and low ground cover.Although there was a marked variation inoverstory plantation density, its effect onunderstory species richness and abundance wasnot significant. The density ofnaturally-regenerated woody species inplantations was over three times the usualplanting density in Ethiopia, indicating a highpotential of forest plantations for restoringthe natural forest ecosystems on degraded landsat a comparatively low cost.     

Inter-crown versus under-crown area: contribution of local configuration of trees to variation in topsoil morphology, pH and moisture in Abies alba Mill. forests

This investigation of three Abies alba stands differing in stem density (338–715 per ha) and vertical structure (one-storeyed or multi-layered) explored the relations between distance from neighbouring tree stems and local canopy openness and selected topsoil properties. The null hypothesis was that in relatively densely stocked forests of close–random stem distribution topsoil morphology, pH and moisture do not differ in inter-crown and under-crown patches. In three plots 1.1 ha in area, soil samples were taken in a square grid 5.0 × 5.0 m and analysed using semivariogram estimation and spatial autocorrelation. The local configuration of trees around the sampled locations was characterised using hemispheral photography and a local stand density index based on tree locations and diameters. The largest portion of the total variation in the soil variables analysed (68–100%) was attributable to small-scale variation in scales <5 m. In all stands, irrespective of density and vertical structure, local stand density/canopy openness correlated positively/negatively with ectohumus layer thickness but negatively/positively with upper soil pH and moisture. Variation in the local configuration of trees explained up to 17% of the total variation in organic horizon thickness, up to 22–29% in topsoil pH (depending on the horizon) and up to 19–27% in topsoil moisture. The results indicate that even in stands of random tree patterning, stem neighbourhood and small-scale variation in canopy density may contribute significantly to topsoil heterogeneity and potentially affect the functioning and structure of forest floor vegetation.     

The influence of forest cover on mule deer habitat selection, diet, and nutrition during winter in a deep-snow ecosystem

Recent research in western North America suggests that open forage areas are a greater limiting factor to mule deer abundance than closed conifer forests. However, much of this work was conducted in ecosystems prone to fire and low snow depths compared to the limits of mule deer range such as the Columbia Mountains, British Columbia, where snow is deep and fires are rare. We used snow track surveys as a measure of habitat use and fecal nitrogen as an index of dietary quality to compare the relative value to mule deer of open deciduous canopies to closed coniferous canopies in a wet ecosystem with deep snow and few fires. Deciduous canopies contain higher levels of understory forage compared to dense coniferous canopies, which are better at intercepting snow. We also evaluated food habits across landscapes with contrasting forest canopies. Results corroborated previous work in that foraging areas such as deciduous stands were strongly selected by deer, despite deeper snow relative to closed coniferous stands. Deer consumed fewer understory shrubs in coniferous-dominated stands, suggesting lower nutritional intake in these stands. Finally, deer appeared to derive a nutritional benefit in landscapes that had a higher proportion of open deciduous canopies, as indexed by fecal nitrogen. However, not all open canopy stands were of equal value to deer – deciduous-dominated stands were selected, whereas clearcuts were avoided. Similarly, not all closed coniferous stands were equally selected: cedar–hemlock stands were avoided whereas Douglas-fir stands were selected and indeed contained the highest proportion of deer tracks. We suggest that winter foraging areas have been underrepresented in management policy in British Columbia, but that snow-interception cover provided by coniferous stands still plays a role in winter deer ecology in deep-snow ecosystems.     

Comparison of terpenes in extracts from the resin and the bark of the resinous stem canker of Chamaecyparis obtusa and Thujopsis dolabrata var. hondae

A monoterpene and 15 diterpenes were isolated from the ethyl acetate extracts of the bark-glued resin from the resinous stem canker ofThujopsis dolabrata var.hondae Makino. A monoterpene (nezukone20) and 4 diterpenes (acetyl torulosol5, acetyl isocupressic acid8, acetyl abietinol11, and 7-methoxytotarol18) were characteristic constituents of the ethyl acetate extracts but were absent in then-hexane extracts from the resinous stem canker ofT. dolabrata var.hondae. These terpenes were first isolated fromT. dolabrata var.hondae andT. dolabrata. The available literature suggests that diterpene18 is a new compound. The resinous stem canker ofChamaecyparis obtusa Endlicher contained larger amounts of manool1,trans-communic acid6, and ferruginol12 and smaller amounts of isocupressic acid7 and abietinol10 than the resinous stem canker ofT. dolabrata var.hondae. The concentration of18 was less than 2% in the extracts, and the resinous stem canker ofC. obtusa lacked this compound. The resinous stem cankers ofC. obtusa andT. dolabrata var.hondae provided extracts 15.6 and 4.96 times, respectively, heavier than the healthy ones. Large differences in the ratios and compositions of terpenes were also observed between the resinous stem canker and the healthy trees. Terpenes isolated from the extracts contained many kinds of diterpene, especially the labdane-type diterpenes, in these diseased trees. These results suggest that the presence of labdane-type diterpenes is closely associated with the resinous stem canker or the causal fungi of this disease.Part of this report was presented at the 50th annual meeting of the Japan Wood Research Society, Kyoto, April 2000     

Horizontal distribution of stolons of a temperate lianaWisteria floribunda DC. and its ecological significance

The spatial distribution of stolons ofWisteria floribunda DC. was investigated, and the ecological significance of clonal growth of this temperate liana is discussed. The study plot (0.15 ha) was located in a secondary deciduous broad-leaved forest dominated byQuercus serrata in central Japan. The tree canopy was almost closed with a mean openness of 4.2%.W. floribunda was dominant among lianas accounting for 86% of their total basal area.W. floribunda stolons of three individuals branched numerous times and intersected among individuals, and showed a large horizontal “web-like” extension on the ground. The total length of stolons was 66.0, 260.2, and 310.6 m, respectively, for each individual. A mean of 25 ramets were established on stolons per individual, and one-third of the ramets obtained a host tree. These results suggested that the expansion of stolons was effective in obtaining host trees forW. floribunda individuals. On the other hand, clonal growth ofW. floribunda did not always contribute to vertical growth of ramets in the study forest, since few ramets reached the forest canopy. Lack of sub-canopy trees in this forest appeared to constrain the vertical growth ofW. floribunda ramets, since sub-canopy trees are thought to bridge lianas from smaller trees to canopy trees.     

Heavy thinning in hinoki plantations in Shikoku (southwestern Japan) has limited effects on recruitment of seedlings of other tree species

Heavy thinning may provide a feasible means to convert single-species coniferous plantations to broadleaved or mixed forests. To assess this possibility, we monitored tree seedling dynamics of other tree species, the understory microenvironment, and seed availability following heavy thinning (50 % by volume) in two hinoki (Chamaecyparis obtusa) plantations in Shikoku, southwestern Japan. Conventional (35 % by volume) and row thinning (50 % by volume) treatments were also applied at one of the sites. Diffuse transmittance in the understory dramatically increased after all thinning treatments then rapidly decreased, except after row thinning. Heavy thinning accelerated recruitment of small seedlings (with stems <30 cm high) at one of the sites. However, the densities of these seedlings declined sharply, and increases in the densities of larger seedlings recorded in subsequent surveys were very small. Furthermore, at the other site, recruitment of small seedlings under heavy thinning was equivalent to that under conventional thinning and lower than that under row thinning. Abundant species in the recruited seedlings were pioneer or subcanopy species found in the soil seed bank. Low seed rain density at both sites appeared to be responsible for the poor recruitment of canopy tree species. Our results suggest that heavy thinning had limited effects on the seedling recruitment of other tree species in these hinoki plantations due to insufficient seed rain and rapid canopy closure.     

Secondary Succession Following Catastrophic Windthrow in a Boreal Forest in Hokkaido,Japan

Abstract

In 1954, Picea-Abiesforests of central Hokkaido were leveled by a windstorm caused by a typhoon. Successional dynamics following disturbance on several forest habitats were monitored over a thirty-four year period. Differences in site conditions, such as soil, topography, and forest floor vegetation, influenced the direction of succession, but some patterns prevailing in this area were discernible. In the early stage of succession (up to year 10), all sites were immediately occupied by Rubus idaeusvar. aculeatissi-muswhich was an inconspicuous component of the stands before disturbance. Plants which dominated the understory of stands before disturbance mostly disappeared during secondary succession except for Sasa senanensis.Residual conifers also survived except for those at a moss-floor site. Betulaspp. and Salixspp. invaded within 6-8 years. At year 10-18, R. idaeussuddenly decreased in canopy cover, while S. senanensisreplaced R. idaeusas the dominant component. Trees, both residuals and invaders, were successful in growing over the shrub layer during this period and species diversity reached a peak. These observations suggest that the timing of tree establishment occurs during the transition from R. idaeuscommunity to S. senanensiscommunity in the shrub-layer, as a deterministic process founded in the life history traits of species.     

The role of gaps and tree regeneration in the transition from dense to open black spruce stands

Black spruce forests growing on clay soils in northwestern Quebec change structure from dense even-aged stands to open uneven-aged stands such that almost all forests older than 200 years have an open canopy. These forests become unproductive over time because they are prone to paludification. The main goal of our study was to document the transition between dense and open stands in terms of gap dynamics, with a focus on tree regeneration. Our objective was to determine whether forests remain open due to a lack of regeneration, a lack of growth or both. Nine stands along a 50–250-year-old time since fire gradient were sampled with the line intersect sampling method. Gap fraction increased with stand age and reached a maximum of 77% in the oldest site. In old-growth stands, gaps were interconnected due to the low density of these forests. Most of the gap makers were found with broken stems. Regeneration was dominated by black spruce layers and was relatively abundant (1.71 stems/m²). However, the majority of gap fillers were smaller than 1 m in height in stands of all ages. Instead of a lack of regeneration, the opening of the forests is due to a lack of growth associated with cold and wet organic deposits. Partial harvesting could be implemented on the most productive sites, while management techniques including soil disturbances will be required on low productivity sites to recreate good growth conditions.     

Stand structure and composition of a 60-year-old larch (Larix kaempferi) plantation with retained hardwoods

Tree retention is understood as a key practice in creating complexity, leading to heterogeneity in resources and habitats in managed stands. In this article, we clarify the long-term effects of tree retention on stand structure and tree-species composition in a 60-year-old Larix kaempferi plantation in central Japan. In our study plot (1.5 ha) there were 18 stems/ha of retained trees (determined by tree-ring analysis), mostly Quercus crispula. We conducted spatial analyses and tested the hypothesis that tree abundance, size structure, and species composition and diversity change with distance from the retained trees. Near the retained trees, L. kaempferi showed a reduction of 40%–60% in basal area, due presumably to the shading effect. In contrast, the nearby area showed greater species diversity in the canopy layer. The retained trees created patches of different species composition in the understory. The spatial gradient of shade and colonization opportunity provided by retained trees greatly affect the distribution of the colonized species, according to their shade tolerance and seed-dispersal ability, which resulted in the stand structure with a heterogeneous shrub-layer vegetation. Retention proved particularly important for the enhancement and long-term maintenance of structural and compositional complexity in L. kaempferi plantations.