Stand structure and regeneration in a Kamchatka mixed boreal forest

Abstract. A 1‐ha plot was established in a Betula platyphylla‐Picea ajanensis mixed boreal forest in the central Kamchatka peninsula in Russia to investigate stand structure and regeneration. This forest was relatively sparse; total density and stand basal area were 1071/ha and 25.8 m²/ha, respectively, for trees > 2.0 cm in trunk diameter at breast height (DBH). 25% of Betula regenerated by sprouting, and its frequency distribution of DBH had a reverse J‐shaped pattern. In contrast, Picea had a bimodal distribution. The growth rates of both species were high, reaching 20 m in ca. 120 yr. The two species had clumped distributions, especially for saplings. Betula saplings were not distributed in canopy gaps. Small Picea saplings were distributed irrespective of the presence/absence of gaps, while larger saplings aggregated in gaps. At the examined spatial scales (6.25–400 m²) the spatial distribution of Betula saplings was positively correlated with living Betula canopy trees and negatively with dead Picea canopy trees. This suggests that Betula saplings regenerated under the crowns of Betula canopy trees and did not invade the gaps created by Picea canopy trees. The spatial distribution of Picea saplings was negatively correlated with living and dead Betula canopy trees and positively with dead Picea canopy trees. Most small Picea seedlings were distributed under the crowns of Picea trees but not under the crowns of Betula trees or in gaps. This suggests that Picea seedlings establish under the crowns of Picea canopy trees and can grow to large sizes after the death of overhead Picea canopy trees. Evidence of competitive exclusion between the two species was not found. At a 20 m × 20 m scale both skewness and the coefficient of variation of DBH frequency distribution of Picea decreased with an increase in total basal area of Picea while those of Betula were unchanged irrespective of the increase in total basal area of Betula. This indicates that the size structure of Picea is more variable with stand development than that of Betula on a small scale. This study suggests that Betula regenerates continuously by sprouting and Picea regenerates discontinuously after gap formation and that the species do not exclude each other.     

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.     

Crown Spread Patterns for Five Deciduous Broad-leaved Woody Species: Ecological Significance of the Retention Patterns of Larger Branches

Patterns of crown spread and branch retention of two shade-intolerantspecies (Betula platyphyllaandB. maximowicziana) were comparedwith three more tolerant species (Quercus mongolica,Acer sieboldianumandMagnoliaobovata). Branching height (height of the lowest living branch)rose more rapidly with age for the twoBetulaspecies than forthe shade tolerant species. Branching heights ofA. sieboldianumandM.obovatawere similar, irrespective of tree height and age, andlarger trees tended to produce wider crowns than theBetulaspecieswhen trees of similar height were compared. In all species,the branch basal area (cross-sectional area of a branch at itsbase) and the leaf area per branch generally increased as thebranch position on a stem became lower. Therefore, retaininglarger branches contributed significantly to the support ofa larger leaf area per tree. The number of larger branches (branchbasal area >80 cm²) for bothBetulaspecies was significantlysmaller than that of the shade tolerant species. The branchretention pattern ofBetulaspecies was probably a consequenceof intolerance of the leaves to shade. The decline ofBetulaspecieswith forest succession is likely to occur through their inabilityto retain branches with a large base area in closed forests.Copyright1997 Annals of Botany Company Shade tolerance; crown spread; branch retention; branch size; broad-leaved woody species; leaf area index per tree     

Light acquisition and use by individuals competing in a dense stand of an annual herb, Xanthium canadense

The importance of light acquisition and utilization by individuals in intraspecific competition was evaluated by determining growth and photosynthesis of individual plants in a dense monospecific stand of an annual, Xanthium canadense. Photosynthesis of individual plants in the stand was calculated using a canopy photosynthesis model in which leaf photosynthesis was assumed to be function of leaf nitrogen content and light availability. The estimated photosynthetic rates of individuals were strongly correlated with the measured growth rates. Photosynthetic rates per unit aboveground mass (RPR, relative photosynthetic rate) increased with increasing aboveground mass, suggesting asymmetric (one-sided) competition in the stand. However, larger individuals had similar RPRs, suggesting symmetric (two-sided) competition. These results were consistent with the observation that size inequality over the whole stand increased with growth, but it remained stable among the larger individuals. The RPR of an individual was calculated as the product of absorbed photon flux per unit aboveground mass (Φ_mass) and light use efficiency (LUE, photosynthesis per unit absorbed photon flux). Φ_mass indicates the efficiency of light acquisition, and was higher in larger individuals in the stand, while LUE was highest in individuals with intermediate aboveground mass. LUE depends on leaf nitrogen content. At an early stage, leaf nitrogen contents of smaller individuals were similar to those that maximize LUE. Light availability to smaller individuals decreased as they grew, while their nitrogen contents did not change markedly, which decreased their LUE. We concluded that asymmetric competition among individuals in the stand resulted mainly from lower efficiencies in both light acquisition and light use by smaller individuals. Received: 31 January 1998 / Accepted: 12 November 1998     

The role of human disturbance in the local Late Holocene establishment of Fagus and Picea forests at Flahult, Western Småland, Southern Sweden

A pollen record from a small alder carr located in the centre of aFagus stand near the hamlet of Flahult in southern Småland has shown thatFagus became established in a semi open cultural landscape about 900 B.P. Human disturbance seems to have controlled the local establishment ofFagus at this site through an expansion of pastoral farming. TheFagus dominance in the present stand seems to be of recent origin, asFagus pollen percentages and influx values have increased considerably only during the last 50 years. The modern composition and structure of theFagus stand are probably an effect of changes in land-use and decreased human activity at the end of the last century. Today, only occasionalPicea individuals occur in the studied stand, andPicea does not appear to have been more abundant in the recent past. The regional expansion ofPicea has probably occurred during this century and has been favoured by modern forestry during the last 50 years.     

Plastic Response of Crown Architecture to Crowding in Understorey Trees of Two Co-dominating Conifers

Crown architecture and growth rate of trunk height, trunk diameterand lateral branches of understorey trees (5–10m tall)were compared between two co-dominating conifers,Abies sachalinensisandPiceaglehnii, in relation to the index of local crowding intensity,W,represented as a function of density, distance and basal areaof taller neighbours. For the two conifers, the growth of trunkheight and diameter was decreased and crowns became flat withincreasingW, keeping crown projection area. Self-pruning oflower branches was more intense inAbiesthan inPiceain crowdedconditions, while both conifers showed similar crown forms inless crowded conditions. These results suggest that the growthin lateral branches exceeded that in height in crowded conditions,especially inAbies. Tree age of both conifers increased withincreasingW, resulting from the low growth rate in crowded conditions.The age of the longest and lowest branch ofPicea, up to 150years, was positively correlated with the tree age ranging from70 to 250 years, whereas that ofAbieswas constant at around30 years irrespective of tree age varying from 40 to 140 years.This result agrees with the observation that agedAbieshad moreflat-shaped crowns than in agedPiceain crowded conditions. Theseresults suggest that each conifer adapted to crowding in differentways: high elongation of branches with high turnover rate forAbiesandviceversaforPicea. Abies; crown form; neighbourhood interference; Picea; plasticity     

Limitations on photosynthesis of competing individuals in stands and the consequences for canopy structure

The canopy structure of a stand of vegetation is determined by the growth patterns of the individual plants within the stand and the competitive interactions among them. We analyzed the carbon gain of individuals in two dense monospecific stands of Xanthium canadense and evaluated the consequences for intra-specific competition and whole-stand canopy structure. The stands differed in productivity, and this was associated with differences in nitrogen availability. Canopy structure, aboveground mass, and nitrogen contents per unit leaf area (N_area) were determined for individuals, and leaf photosynthesis was measured as a function of N_area. These data were used to calculate the daily carbon gain of individuals. Within stands, photosynthesis per unit aboveground mass (P_mass) of individual plants increased with plant height, despite the lower leaf area ratios of taller plants. The differences in P_mass between the tallest most dominant and shortest most subordinate plants were greater in the high-nitrogen than in the low-nitrogen stand. This indicated that competition was asymmetric and that this asymmetry increased with nitrogen availability. In the high-nitrogen stand, taller plants had a higher P_mass than shorter ones, because they captured more light per unit mass and because they had higher photosynthesis per unit of absorbed light. Conversely, in the low-nitrogen stand, the differences in P_mass between plants of different heights resulted only from differences in their light capture per unit mass. Sensitivity analyses revealed that an increase in N_area, keeping leaf area of plants constant, increased whole-plant carbon gain for the taller more dominant plants but reduced carbon gain in the shorter more subordinate ones, which implies that the N_area values of shorter plants were greater than the optimal values for maximum photosynthesis. On the other hand, the carbon gain of all individual plants, keeping their total canopy N constant, was positively related to an increase in their individual leaf area. At the same time, however, increasing the leaf area for all plants simultaneously reduced the carbon gain of the whole stand. This result shows that the optimal leaf area index (LAI), which maximizes photosynthesis of a stand, is not evolutionarily stable because at this LAI, any individual can increase its carbon gain by increasing its leaf area.     

Protein induction process and stochastic nature of cell commitment to proliferation and differentiation

A differential equation model is constructed to describe competition between trees with overlapping crowns. The model is based upon Thornley's mechanistic plant growth model which divides plant biomass into three components corresponding to storage material, degradable structural tissue and non-degradable structural tissue. The available incident radiation is partitioned among trees according to their sizes, with large trees intercepting more light than smaller neighbours. Analysis of the dynamic stability of the model reveals that suppression occurs over a wide range of parameter space. Typically, as canopy overlap increases and competition for light becomes intense, some trees are suppressed by their neighbours. The suppression-dominance phenomenon occurs even in stands of trees with identical parameter values. Model simulations are compared with data on the growth of Pinus radiata.     

A protocol for measuring abundance and size of a neotropical liana,Desmoncus polyacanthos (Palmae), in relation to forest structure

This study investigated the relationship between the abundance and size ofD. polyacanthos and measures of forest canopy structure. Várzea and terra firme forest were selected for study at the mouth of the Amazon estuary, Pará, Brazil. The forest canopy characteristics investigated included individual tree height and height to base of the live portion of the crown, forest canopy depth (the distance from the lowest foliage in the understory to the top of the canopy including all free space between top and bottom), forest canopy thickness (the portion of the forest canopy depth occupied only by foliage) and forest canopy density (the sum of lengths of each of the individual crowns that make up forest canopy depth) (see Fig. 2b). The attributes ofD. polyacanthos measured included leaf number, stem height and percent coverage within the stands. Results from this study suggest thatD. polyacanthos can be encouraged to grow through selective cutting of canopy trees and promotion of regenerating forest stands on terra firme fallow. The protocol used to relateD. polyacanthos abundance and size to kind of forest canopy structure can be used for evaluation of other climbing palms.     

Regeneration in subalpine coniferous forests

Death of canopy trees when gaps are formed was studied in a subalpine coniferous forest, central Japan, which was composed ofAbies, Tsuga, Picea, Betula, andSorbus. Typhoons were considered to be the most important cause of the death of canopy trees. The degree of disturbance in each of 16 plots (20 m×20 m) was represented by the percentage of the total basal area of dead trees to that of living and dead canopy trees (disturbance magnitude; MAG). The mortality of canopy trees increased as their dbh increase in the plots of lower MAG than 90%. The mortality varied among genera, andTsuga was characterized as having lower mortality than that of the other conifers. 418 dead trees were observed. The standing dead trees made up 10.7% of the trees, the stem broken trees 46.7%, and the uprooted trees 42.2%. The stem breaking was most frequent inAbies, and the uprooting was most frequent inTsuga, Picea, andBetula. Undeveloped forests, which have the L-shaped dbh distribution, were destroyed only in high degree (70%<MAG), while developed forests were destroyed in various degrees (30%<MAG<100%). The percentage of uprooted trees in basal area decreased with the development of the forest, from 60% to 10%.     

Succession after stand replacing disturbances by fire,wind throw,and insects in the dark Taiga of Central Siberia

The dark taiga of Siberia is a boreal vegetation dominated by Picea obovata, Abies sibirica, and Pinus sibirica during the late succession. This paper investigates the population and age structure of 18 stands representing different stages after fire, wind throw, and insect damage. To our knowledge, this is the first time that the forest dynamics of the Siberian dark taiga is described quantitatively in terms of succession, and age after disturbance, stand density, and basal area. The basis for the curve–linear age/diameter relation of trees is being analyzed. (1) After a stand-replacing fire Betula dominates (4,000 trees) for about 70 years. Although tree density of Betula decreases rapidly, basal area (BA) reached >30 m²/ha after 40 years. (2) After fire, Abies, Picea, and Pinus establish at the same time as Betula, but grow slower, continue to gain height and eventually replace Betula. Abies has the highest seedling number (about 1,000 trees/ha) and the highest mortality. Picea establishes with 100–400 trees/ha, it has less mortality, but reached the highest age (>350 years, DBH 51 cm). Picea is the most important indicator for successional age after disturbance. Pinus sibirica is an accompanying species. The widely distributed “mixed boreal forest” is a stage about 120 years after fire reaching a BA of >40 m²/ha. (3) Wind throw and insect damage occur in old conifer stands. Betula does not establish. Abies initially dominates (2,000–6,000 trees/ha), but Picea becomes dominant after 150–200 years since Abies is shorter lived. (4) Without disturbance the forest develops into a pure coniferous canopy (BA 40–50 m²/ha) with a self-regenerating density of 1,000 coniferous canopy trees/ha. There is no collapse of old-growth stands. The dark taiga may serve as an example in which a limited set to tree species may gain dominance under certain disturbance conditions without ever getting monotypic.     

Stomatal activity within the crowns of tall deciduous trees under forest conditions

The variation in stomatal activity within the crowns ofAcer campestre, Carpinus betulus andQuercus cerris was measured by vapour exchange porometer on several summer days in an oak-hornbeam forest, in SW Slovakia, Czechoslovakia. Variation resulted from crown position in the forest stand and from leaf position within the canopy. The highest stomatal conductance was in sunlit sun leaves in the upper part of the canopy. Stomatal conductance decreased with increasing depth in the canopy. The steepest decrease was in the upper canopy, in the intermediate zone between fully sunlit and fully shaded leaves, and was caused by the decline in leaf irradiance and in stomatal density. In codominant trees, the conductance in shade leaves at the base of the crown was significantly lower than in the sun leaves at the top of the crown. In a dominant tree,Q. cerris, the differences in stomatal conductance were small and most frequently insignificant. Variation in incident light also determined the diurnal variation of stomatal conductance with respect to crown aspect. Differences between sun leaves on the east and west facing aspects of the overstory crown ofQ. cerris were demonstrated for several days.     

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.     

Effects of Physiological and Environmental Variations on Size-Structure Dynamics in Plant Populations

Sensitivity analysis was conducted, based on the canopy photosynthesisand continuity equation models which were developed in a previouspaper (Yokozawa and Hara, 1992), to investigate effects of variationin physiological parameters (maximal photosynthetic rate perunit leaf area, respiration rate per unit leaf area, maintenancerespiration rate per unit weight, growth respiration rate perunit weight, light extinction coefficient of the canopy, etc.)on the size-structure dynamics in plant populations. As thedegree of asymmetry in competition between individuals increased,effects of variation in physiological parameters diminished.Therefore, a population undergoing one-sided competition (mostasymmetric competition) is a stable system, little affectedby temporal and spatial variations in the environmental conditionswhich lead to variation in physiological parameters, whereasa population undergoing symmetric two-sided competition is sensitiveto these fluctuations. It was also shown by simulation thatthe degree of asymmetry in competition decreases (through effectson canopy photosynthesis) as nutrient level in the soil is reduced.It is suggested that symmetric two-sided competition is associatedwith non-transitivity of competition between species (i.e. frequentreversals of rank order of species), and hence with speciesdiversity. Several other ecological phenomena are discussedin relation to allometry (i.e. allocation-growth pattern) andthe degree of asymmetry in competition.Copyright 1994, 1999Academic Press Allometry, canopy photosynthesis, competition mode, continuity equation, parameter sensitivity, stability of stand structure     

Successional trends 219 years after fire in an old Pinus sylvestris stand in northern Sweden

Abstract. We present results from repeated analyses (1962, 1993) of a permanent plot established in 1947, combined with retrospective stand age structure data, in an old Pinus sylvestris stand in Muddus National Park, northern Sweden. The study points towards a successional pathway governed by concurrent disturbance effects of climate variability, reindeer grazing and fire. This is intermediate to the two often advocated ideas on dynamics in boreal forests, that is, one of disturbance-related tree regeneration/mortality and one of continuous regeneration. When the plot was established in 1947 the tree layer (> 1.3 m) consisted of 300 individuals/ha of P. sylvestris and 62/ha of Betula pubescens. Subsequently the stand has become more dense and the species dominance has shifted. In 1993, 362 P. sylvestris and 62 Picea abies individuals were present per ha, while no Betula individuals were found. The number of dead trees increased from zero in 1947 to 200/ha (Pinus) in 1993. Pinus was also the most common species in the sapling layer (< 1.3 m) throughout the study period, though the number dropped from 8912/ha in 1947 to 51% in 1993. Dead saplings decreased from 2650/ha in 1947 to ca. 50% in 1962, and only 9% in 1993. Temporal variations in mortality and sapling mean height coincided with variations in snow depth, indicating a critical period in sapling development when saplings are exposed at the snow/atmosphere interface. The number of living Picea saplings increased slowly until 1993; no dead saplings were found. Most Pinus recruited shortly after the 1774-fire, and during the second half of the 1900s. The major part of the spruce regeneration took place during the later half of the 1900s. No successful Betula recruitment has occurred after the 1930s, and no live Betula were present in 1993, which might be explained as an effect of increased reindeer browsing – the reindeer stock has grown by 50% since 1961. Although subjected to high mortality, Pinus regenerated and maintained a seedling/sapling bank. In this way Pinus remained dominant in the tree layer after more than 200 post-fire years. The importance of the shade-tolerant Picea has slowly increased, while Betula has died off. Thus, even after 219 yr since fire there is an early successional trend in the stand. This suggests that an increased chronic disturbance (grazing/browsing by reindeer) has partly succeeded earlier discrete fire-disturbance events, and maintained a continuous seedbed favouring the shade-intolerant pine recruitment.     

Tree mortality and effects of release from competition in an old‐growth Fagus‐Abies‐Picea stand

Abstract. In a montane mixed Fagus‐Abies‐Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old‐growth stand were studied by combining an 8‐yr annual census of trees in a 1‐ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap‐dynamics paradigm.     

Revegetation process on a subalpine mudflow

The process of revegetation on a subalpine mudflow was analyzed on the basis of the vegetation-soil relationship of the revegetated stands, and of the age structure and height growth process of the trees. Based on evidence derived from five independent sources, it was concluded that this mudflow was caused by a typhoon in 1959. The revegetated stands were classified into three vegetation types. AnAlnus maximowiczii stand was established where the surface soil was unstable; aBetula ermanii stand was established where the degree of disturbance by the mudflow was small; anAlnus-Betula stand was the intermediate type and occupied most of the area on the mudflow.Alnus had colonized concentrically 4–6 yr after the mudflow, andBetula and conifers had followed 1–2 yr later. As revegetation proceeded, the trees became differentiated into two populations, a canopy population and a suppressed one. The former was dominated mainly byAlnus and the latter byBetula and conifers. This differentiation was caused by the faster growth ofAlnus during the initial years of colonization. SomeBetula trees have recently grown faster thanAlnus, suggesting future replacement in dominance. Conifers have grown slowly even during recent years; the highly oligotrophic condition of the soil appears to have been responsible for this delayed growth.     

Scales of aboveground and below-ground competition in a semi-arid woodland detected from spatial pattern

Abstract. Semi-arid woodlands are two-phase mosaics of canopy and inter-canopy patches. We hypothesized that both aboveground competition (within canopy patches), and below-ground competition (between canopy patches), would be important structuring processes in these communities. We investigated the spatial pattern of trees in a Pinus edulis-Juniperus monosperma woodland in New Mexico using Ripley's K-function. We found strong aggregation of trees at scales of 2 to 4 m, which indicates the scale of canopy patches. Canopy patches were composed of individuals of both species. Crown centers of both species were always less aggregated than stem centers at scales less than canopy patch size, indicating morphological plasticity of competing crowns. In the smallest size classes of both species, aggregation was most intense, and occurred over a larger range of scales; aggregation decreased with increasing size as is consistent with density-dependent mortality from intraspecific competition. Within canopy patches, younger trees were associated with older trees of the other species. At scales larger than canopy patches, younger trees showed repulsion from older conspecifics, indicating below-ground competition. Hence, intraspecific competition was stronger than interspecific competition, probably because the species differ in rooting depth. Woodland dynamics depend on the scale and composition of canopy patches, aggregated seed deposition and facilitation, above- and below-ground competition, and temporal changes in the spatial scale of interactions. This woodland is intermediate in a grassland-forest continuum (a gradient of increasing woody canopy cover) and hence we expected, and were able to detect, the effects of both above- and below-ground competition.     

Changes in Forest Structure and Species Composition during Secondary Forest Succession in the Bolivian Amazon1

Changes in forest structure and species diversity throughout secondary succession were studied using a chronosequence at two sites in the Bolivian Amazon. Secondary forests ranging in age from 2 to 40 years as well as mature forests were included, making a total of 14 stands. Fifty plants per forest layer (understory, subcanopy, and canopy) were sampled using the transect of variable area technique. Mean and maximum height, total stem density, basal area, and species number were calculated at the stand level. Species diversity was calculated for each stand and for each combination of forest layer and stand. A correspondence analysis was performed, and the relationship between relative abundance of the species and stand age was modeled using a set of hierarchical models. Canopy height and basal area increased with stand age, indicating that secondary forests rapidly attain a forest structure similar in many respects to mature forests. A total of 250 species were recorded of which ca 50 percent made up 87 percent of the sampled individuals. Species diversity increased with stand age and varied among the forest layers, with the lowest diversity in the canopy. The results of the correspondence analysis indicated that species composition varies with stand age, forest layer, and site. The species composition of mature forests recovered at different rates in the different forest layers, being the slowest in the canopy layer. Species showed different patterns of abundance in relation to stand age, supporting the current model of succession.     

Effect of Crown Asymmetry on Size-Structure Dynamics of Plant Populations

The effect of crown asymmetry on the size–structure dynamicsof populations was evaluated using a spatial competition modelincorporating crown asymmetry. Computer simulations were carriedout with various combinations of density levels, spatial patterns,and degrees of asymmetry in competition to assess how they modifythe effect of crown asymmetry on size–structure dynamics. In the model, crown asymmetry is expressed by the crown-vector,or the vector linking the stem base and the centre of the projectedarea of the crown on the horizontal plane. Crown-vectors areassumed to develop in the manner by which crowns repel eachother. As crown-vectors develop, the positions of the crown-centresmove. Competition between individuals is expressed by a neighbourhoodmodel, in which individual growth is determined by the distancefrom, and size of, the neighbours' crown-centres. Generally, populations of individuals which developed asymmetriccrowns had larger survivorship, larger mean size, smaller coefficientsof variation and skewness, and a more regular spatial patternthan populations of individuals which developed symmetric crowns.The effect of crown symmetry is generally stronger in populationswith high density and a clumped spatial pattern. The effectof mortality caused by one-sided competition on size-structuredynamics was similar to that of crown asymmetry; mortality increasedmean size, reduced size hierarchy, and made the spatial patternmore regular. Because mortality was heavier in populations withoutcrown asymmetry, its effect on size-structure dynamics cancelledout, or overwhelmed, the effect of crown asymmetry in latergrowth stages. If crown asymmetry is associated with a reductionin growth, the effect of crown asymmetry is reduced. Nevertheless,the resultant population structure is different from that ofpopulations without crown asymmetry. Competition; crown asymmetry; morphological plasticity; neighbourhood interference model; size-structure dynamics