Canopy surface topography in a French Guiana forest and the folded forest theory

The canopy surface is an undulating surface that follows the irregular contours of the upper tree crowns and defines the inner and the outer limits of the forest volume. In French Guiana, the height of the canopy surface was surveyed in both a primary and a 20-years old clear-felled secondary forest plot. The topographic surface was displayed in a three-dimensional mesh, where X and Y are horizontal co-ordinates, and Z is the canopy height measured from the ground with an optical telemeter. The statistical dispersion of Z-data, and the spatial tree height variations, are interpreted at different levels of ecosystem organisation, from forest type (primary or secondary forest) to the trees themselves, following the folded forest model theory (Oldeman 1992, 1994). The vertical growth of trees creates a convex pattern in the relief of canopy surface, whereas gaps make concavities which delimit impact of perturbation on the forest structure. These events are either the result of the dynamic of single trees (emergent and decayed trees), or arise from the dynamic of a group of trees working together (group of emergent trees or complex gaps). At the plot scale, the elementary events, convexities and concavities, are gathered on similar topo-sequences, and form canopy units either higher or lower than the average canopy height. This study suggests that the topography of the canopy surface is defined by a complex nested system from trees, to groups of trees, to canopy units, within a delimited floristic and physical environment.     

Winning by a neck: tall giraffes avoid competing with shorter browsers

With their vertically elongated body form, giraffes generally feed above the level of other browsers within the savanna browsing guild, despite having access to foliage at lower levels. They ingest more leaf mass per bite when foraging high in the tree, perhaps because smaller, more selective browsers deplete shoots at lower levels or because trees differentially allocate resources to promote shoot growth in the upper canopy. We erected exclosures around individual Acacia nigrescens trees in the greater Kruger ecosystem, South Africa. After a complete growing season, we found no differences in leaf biomass per shoot across height zones in excluded trees but significant differences in control trees. We conclude that giraffes preferentially browse at high levels in the canopy to avoid competition with smaller browsers. Our findings are analogous with those from studies of grazing guilds and demonstrate that resource partitioning can be driven by competition when smaller foragers displace larger foragers from shared resources. This provides the first experimental support for the classic evolutionary hypothesis that vertical elongation of the giraffe body is an outcome of competition within the browsing ungulate guild.     

开放式空气CO2浓度增高对水稻冠层微气候的影响

利用位于江苏省无锡市安镇的我国唯一的农田开放式空气CO2 浓度增高 (FACE)系统平台 ,于2 0 0 1年 8月 2 6日至 10月 13日 (水稻抽穗至成熟期 )进行水稻作物冠层微气候连续观测 ,以研究FACE对水稻冠层微气候特征的影响 .结果表明 ,FACE降低了水稻叶片的气孔导度 ,FACE与对照水稻叶片气孔导度的差异上层叶片大于下层叶片 ,生长前期大于生长后期 .FACE使白天水稻冠层和叶片温度升高 ,这种差异生长前期大于生长后期 ;但FACE对夜间水稻冠层温度的影响不明显 .在水稻旺盛生长的抽穗开花期 ,晴天正午前后FACE水稻冠层温度比对照高 1.2℃ ;从开花至成熟期 ,FACE水稻冠层白天平均温度比对照高 0 .4 3℃ .FACE对冠层空气温度也有影响 ,白天水稻冠层空气温度FACE高于对照 ,这种差异随太阳辐射增强而增大且冠层中部大于冠层顶部 ;冠层中部空气温度FACE与对照的差异 (Tface-Tambient)日最大值在 0 .4 7～ 1.2℃之间 ,而冠层顶部的Tface-Tambient日最大值在 0 .37～ 0 .8℃之间 .夜间水稻冠层空气温度FACE与对照差别不大 ,变化在± 0 .3℃之内 .而FACE对水稻冠层空气湿度无显著影响 ,表明FACE使水稻叶片气孔导度降低 ,从而削弱了植株的蒸腾降温作用 ,导致水稻冠层温度和冠层空气温度升高 ,改变了整个水稻冠层的温度环     

Defensive traits of savanna trees – the role of shoot exposure to browsers

To investigate patterns of plant defence, a study was conducted on 13 tree species in Botswana. We tested the hypotheses that (1) shoots on small, young trees are more defended than shoots at the same height on tall, mature individuals and (2) shoots within browsing height are more defended than shoots above the reach of browsers, on mature trees. Plant traits studied were total phenolics, tannin activity, nitrogen (N), acid-detergent fibre (ADF), and degree of spinescence (in three species). We found some support for the first hypothesis. Overall young trees had a significantly higher concentration of total phenolics than mature trees and on species level these differences were significant for three of the 13 species. However, we found no differences in tannin activity, spinescence or ADF concentration between young and mature trees, and four species had higher levels of N in young trees, suggesting higher overall palatability. Contrary to our second hypothesis, shoots within browsing height were less chemically defended (total phenolics and tannin activity) than shoots above browsing height in six of 13 species. Two species had less ADF in lower shoots, and no significant differences were found in N or spinescence for any species. This study included a broad range of tree species at two sites and the results indicate that general theories of plant defence, originating from boreal and northern temperate forests, cannot be simply applied to savanna ecosystems.     

Liana cover in the canopies of rainforest trees is not predicted by local ground‐based measures

Lianas (woody climbers) are structural parasites of trees that compete with them for light and below‐ground resources. Most studies of liana–tree interactions are based on ground‐level observations of liana stem density and size, with these assessments generally assumed to reflect the amount of liana canopy cover and overall burden to the tree. We tested this assumption in a 1‐ha plot of lowland rainforest in tropical Australia. We recorded 1072 liana stems (≥1 cm diameter at breast height {dbh}) ha^?1 across all trees (≥10 cm dbh) on the plot and selected 58 trees for detailed study. We estimated liana canopy cover on selected trees that hosted 0–15 liana individuals, using a 47‐m‐tall canopy crane. Notably, we found no significant correlations between liana canopy cover and three commonly used ground‐based measurements of liana abundance as follows: liana stem counts per tree, liana above‐ground biomass per tree and liana basal area per tree. We also explored the role of tree size and liana infestation and found that larger trees (≥20 cm dbh) were more likely to support lianas and to host more liana stems than smaller trees (≤20 cm dbh). This pattern of liana stem density, however, did not correlate with greater liana coverage in the canopy. Tree family was also found to have a significant effect on likelihood of hosting lianas, with trees in some families 3–4 times more likely to host a liana than other families. We suggest that local ground‐based measures of liana–tree infestation may not accurately reflect liana canopy cover for individual trees because they were frequently observed spreading through neighbouring trees at our site. We believe that future liana research will benefit from new technologies such as high‐quality aerial photography taken from drones when the aim is to detect the relative burden of lianas on individual trees.     

Appropriate vegetation indices for measuring the impacts of deer on forest ecosystems

Increasing deer density can cause serious degradation of forests in the Americas, Europe, and Asia. To manage deer impacts, evaluating their current impacts on forest ecosystems is necessary, usually via vegetation indices. However, the relationship between vegetation indices and absolute deer density, while taking into account tree size, snow depth, light condition, and the type of understory vegetation, has never been investigated. We examined the relationship between various vegetation indices and absolute deer density in 344 study plots in the deciduous broad-leaved forest of Yamanashi Prefecture, central Japan. In each plot, debarking and browsing, along with the coverage and maximum height of understory vegetation, were surveyed. Estimated deer densities for 82 5 × 5-km mesh units ranged from 0.8 deer/km² to 32.7 deer/km². The percentages of debarked trees within a plot ranged from 0 to 84%. Debarking was promoted by high deer density, small tree size, and thick snow. The effect of tree size on debarking was stronger than that of deer density. Occurrence of browsing on understory vegetation was higher at higher deer densities, and where understory vegetation was dominated by evergreen dwarf bamboo. Coverage and maximum height of understory vegetation were unaffected by deer density but increased with canopy openness and the dominance of dwarf bamboo in the understory. Overall, we predict that debarking of small trees living in heavy snow areas should occur even at low deer densities (<10 deer/km²). Browsing on dwarf bamboo should occur at intermediate deer densities (10–30 deer/km²), while debarking of thick trees living in low snow areas should occur only at high deer densities (≥30 deer/km²). Our study shows that debarking and browsing on understory vegetation are appropriate indices for evaluating deer impacts on forest ecosystems, but that tree size, snow depth, and the type of understory vegetation should also be considered.     

Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

Responses of crown development to canopy openings by saplings of eight tropical submontane forest tree species in Indonesia: a comparison with cool-temperate trees

BACKGROUND AND AIMS: Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0.2-2.5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan. METHODS: The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance. KEY RESULTS: Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity. CONCLUSIONS: This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.     

开放式空气CO2浓度增高对水稻冠层微气候的影响

利用位于江苏省无锡市安镇的我国唯一的农田开放式空气CO₂浓度增高(FACE)系统平台,于2001年8月26日至10月13日(水稻抽穗至成熟期)进行水稻作物冠层微气候连续观测,以研究FACE对水稻冠层微气候特征的影响.结果表明,FACE降低了水稻叶片的气孔导度,FACE与对照水稻叶片气孔导度的差异上层叶片大于下层叶片,生长前期大于生长后期.FACE使白天水稻冠层和叶片温度升高,这种差异生长前期大于生长后期;但FACE对夜间水稻冠层温度的影响不明显.在水稻旺盛生长的抽穗开花期,晴天正午前后FACE水稻冠层温度比对照高1.2℃;从开花至成熟期,FACE水稻冠层白天平均温度比对照高0.43℃.FACE对冠层空气温度也有影响,白天水稻冠层空气温度FACE高于对照,这种差异随太阳辐射增强而增大且冠层中部大于冠层顶部;冠层中部空气温度FACE与对照的差异(T_face-T_ambient)日最大值在0.47～1.2℃之间,而冠层顶部的T_face-T_ambient日最大值在0.37～0.8℃之间.夜间水稻冠层空气温度FACE与对照差别不大,变化在±0.3℃之内.而FACE对水稻冠层空气湿度无显著影响,表明FACE使水稻叶片气孔导度降低,从而削弱了植株的蒸腾降温作用,导致水稻冠层温度和冠层空气温度升高,改变了整个水稻冠层的温度环     

Inter- and intra-plant variations in nitrogen,tannins and shoot growth of <Emphasis Type="Italic">Sclerocarya birrea</Emphasis> browsed by elephants

Impacts of elephants (Loxodonta africana africana) on woody vegetation has attracted substantial attention for decades, but plant-level responses remain a gap in the understanding of savanna ecology. Marula (Sclerocarya birrea caffra) forms an important part of elephant diets. We investigated the relationships between browsing intensity and shoot/leaf size, nitrogen (N) and condensed tannin (CT) concentrations in upper and lower canopies of male and female marula individuals in Hluhluwe-iMfolozi Park, South Africa. Browsing intensity (54%) did not differ between sexes, suggesting no preference by elephants for either sex. Females had higher [CT] than males and tannin decreased with increasing browsing intensity in both sexes. In lightly or moderately browsed trees, [CT] was controlled by unmeasured factors such that within-tree impacts of browsing were more variable in lightly/moderately browsed than heavily browsed trees. There was little change in [N] up to ~60% browsing intensity, but [N] increased dramatically at higher intensity. Shoots and leaves on broken branches in the lower canopy were larger (2.5 and 1.2 times, respectively) than those on unbroken branches in either upper or lower canopies. Chemical responses were systemic and potentially influence browsing among trees, while growth responses were strongly localised and potentially influence browsing within trees. Although marula trees are able to compensate vigorously for browsing at the scale of individual organs, trees may become progressively carbon-deficient and have their lives shortened if total plant growth is negatively affected by chronic browsing, e.g. near permanent water.     

Canopy structure in a temperate old-growth evergreen forest analyzed by using aerial photographs

We used aerial photographs to create a digital elevation model of the canopy surface of a 10-ha study area in a temperate old-growth evergreen forest. A topographic map of the ground surface in a 4-ha permanent plot within the study area was also drawn from ground measurements. The difference between the two elevation values (i.e., canopy surface – ground surface) at each point in a 5-m grid was considered to be the canopy height, and a canopy height profile was constructed from these data. The canopy structure in the 4-ha plot that was estimated in this way was compared with that obtained by two ground observation methods, i.e., the canopy (vegetation) height profile method and the canopy coverage census method. Large gaps were adequately detected by the aerial photograph method, but small gaps were less often detected. Gap size distribution obtained by the aerial photograph method was similar to that observed on the ground, and was a function of gap depth. This study indicates that if a detailed topographic map can be made, the canopy height profile derived from aerial photography can be effective in analyzing the canopy structure of evergreen forests, such as tropical rain forests, over large areas.     

Effects of Deer Browsing, Fabric Mats, and Tree Shelters on Quercus rubra Seedlings

We examined the effects of deer browsing, fabric mats, and tree shelters on the growth and survival of Quercus rubra (northern red oak) seedlings planted as part of a reforestation project in southcentral Minnesota. Browsing by white-tailed deer occurred on 68.6% of the seedlings growing without tree shelters, whereas only 3.6% of the trees were browsed by mice or voles and 2.5% were browsed by rabbits. Fabric mats, used to control competition from herbaceous plants for roughly half of the seedlings included in the study, had a detrimental effect overall. Seedlings grown with mats had a greater frequency of deer browsing and a greater chance of dying than seedlings grown without mats. Stem height for seedlings browsed by deer was less overall than for nonbrowsed seedlings, although this pattern varied with use of fabric mats and plot location. The use of plastic tree shelters effectively prevented deer browsing and reduced the mortality rate from 34.6% to 3.2%. Our results indicate that fabric mats should not be used in restoration projects with large deer populations. They also suggest that planting seedlings away from existing forest edges and using seedling protection measures such as tree shelters will increase seedling survival and growth in future restoration projects.     

When do nurse plants stop nursing? Temporal changes in water stress levels in Austrocedrus chilensis growing within and outside shrubs

Question: Does the proximity of shrubs affect seasonal water stress of young Austrocedrus chilensis trees (a native conifer of the Austral Temperate Forest of South America) in xeric sites? Location: A. chilensis xeric forest in northwest Patagonia, Argentina. Methods: We examined the dependence of predawn twig water potential on tree development (seedling to adult) and proximity to nurse shrubs during spring and summer. We analysed spatial associations of seedlings, saplings and adult trees with nurse shrubs, and also evaluated if trees affected shrub canopy vitality. Results: Water stress in Austrocedrus trees was affected by shrub presence. Small trees (i.e.<0.5 m in height) growing in the open were most stressed, particularly in summer. Small trees growing within a shrub canopy had low water stress and little change between spring and summer. The opposite trend, however, was true for the medium‐height category (i.e. 0.5‐1.5 m in height); trees in this size category were more stressed when growing within the shrub canopy than in the open. Larger Austrocedrus trees (i.e.>2 m in height) were not affected by shrub presence. Austrocedrus trees were spatially associated with shrubs in all height classes; however, the percentage of living shrub canopy decreased with tree height. Conclusions: In xeric areas of northwest Patagonia, the strength and direction of interactions between A. chilensis and shrubs, in terms of tree water stress, are dynamic and modulated by tree size and environmental conditions. Overall, positive effects of shrubs on early developmental stages appear to be more important than subsequent negative interactions, since nursing effects could generate a spatial association of shrubs and Austrocedrus trees that persists through later successional stages. These findings shed light on mechanisms behind successional changes, and have important conservation and management implications.     

Analysis of development of a subalpineAbies stand based on the growth processes of individual trees

The progress of growth of a subalpine youngAbies veitchii andA. mariesii forest during 25 years was analyzed on the basis of measurements of the processes of height growth of about 230 trees in a quadrat where the old canopy had been completely destroyed by a typhoon in 1959. The original forest floor sapling population had consisted of trees shorter than 2 m. Saplings grew faster after the breakage of the canopy than before,A. veitchii growing faster thanA. mariesii. During the 25 years of growth, a few well developed trees exceeded 6 m in height, while others remained around only 1 m or less. Some small trees, mostlyA. veitchii died at sites of high density. A bimodality in the distribution of tree height had developed with a trough at about 2.5 m, differentiating the trees into canopy and suppressed populations. Canopy trees grew with wide variation of rates, while most of the suppressed trees showed little recent growth. No difference was found in recent growth rates between the two canopyAbies species. Differences in height growth rates among individual canopy trees were analyzed on the basis of their horizontal crown overlapping. Competition models evaluating the difference in height between trees with overlapping crowns were shown to be effective. The height growth rate of a canopy tree appeared to be controlled by both the closely grown taller trees and the local density of trees including those shorter than the subject tree.     

Effect of trap position, habitat, and height on the capture of pepper maggot flies (Diptera: Tephritidae)

Trapping results indicate that pepper maggot, Zonosemata electa (Say), flies occupy tree canopies adjacent to fields when not on host plants. Several in- and near-field trap positions were used to find a reliable monitoring system for adult pepper maggots. Traps baited with liquid ammonium hydroxide (Stills-style trap), hung in the canopy of trees on the edges of pepper fields, caught significantly more Z. electa flies than when positioned lower along the treeline or in the field. In a second experiment, significantly more pepper maggot flies were captured in sugar maples compared with choke cherry trees, which indicates a pest preference for certain nonhost habitats. The lowest trap height tested (2.1 m) failed to capture Z. electa flies in either tree species when the pest population level was low. These studies demonstrated that pepper maggot flies can be reliably detected with Stills-style traps positioned at approximately 6.4 m height within the canopy of sugar maple trees adjacent to pepper fields. Fruit oviposition scars also are useful site-specific indicators of pepper maggot presence/absence and may aid in determining if insecticide applications are necessary and in timing sprays.     

Disentangling the causes of damage variation by deer browsing on young Thuja plicata

Long‐lived trees experience different levels of damage due to mammalian herbivores. To untangle the mechanisms that underlie this variation, we combined chemical with dendrochronological analyses to study variation in browsing on Western redcedars (Thuja plicata) on Haida Gwaii (British Columbia, Canada). Since the last glaciation, Haida Gwaii forests had lacked large herbivore browser until Sitka black‐tailed deer (Odocoileus hemionus sitkensis) were introduced at the beginning of the 20th century. Dendrochronology yielded information on radial growth and plant annual responses to environmental stresses including herbivory. Secondary metabolite content and plant nutritional quality provided insights into proximate causes of food choices made by herbivores. We sampled lightly‐ and heavily‐browsed young trees at four sites: three clear‐cut sites with high browsing pressure and one old‐growth forest site where browsing pressure had, until recently, been lower. Heavily‐browsed young trees had lower concentrations of secondary metabolites and were of lower nutritive value than lightly‐browsed trees at all sites. Under high browsing pressure, tree growth patterns suggested that all young trees were initially severely browsed until some trees, currently scored as lightly‐browsed, started to escape deer. At the old‐growth site, both lightly‐ and heavily‐browsed trees tended to have lower overall average secondary metabolite concentrations than those of all other sites, a trend possibly related to greater canopy closure. Lightly‐browsed trees were older than heavily‐browsed ones which resulted, during the period of lower browsing pressure, in higher growth rate and a same pattern of change in growth from one year to the next year. This suggests that, under low browsing pressure, selection of young trees related to chemical defense was weak and that growth differences due to other factors than browsing could be expressed. Under strong browsing pressure, however, all young trees had equally low growth rates until trees with better genetic potential to produce effective defenses were able to escape deer. This suggests that selection by deer could occur on a long‐lived tree.     

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.     

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.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.