Leaf dynamics and shoot phenology of eleven warm-temperate evergreen broad-leaved trees near their northern limit in central Japan

Dynamic features of shoot phenology including leaf emergence and leaf fall, and leaf life span for eleven evergreen broad-leaved tree species were investigated in a warm-temperate rain forest in Mount Kiyosumi, central Japan. All species had periodic leaf emergence or flushing pattern, and were classified into two types; single and multiple flush and only one species, Eurya japonica, represented the latter type and the rest had single flush in spring. The single flush type can further be subdivided into two groups according to their duration of shoot growth; short and long flush. Seasonal patterns of leaf fall were categorized into four; unimodal, bimodal, broad unimodal, and multimodal type though they were not fixed pattern. The leaf emergence and leaf fall patterns were correlated for the eleven species, and five phenological types were categorized. Four of them were the single flush types, i.e., short flush of leaf emergence with unimodal leaffall (SSU) type of Castanopsis sieboldii and Quercus salicina, short flush with bimodal leaf fall (SSB) type of Quercus acuta, Machilus thunbergii, Neolitsea sericea, and Cinnamomum japonicum, long flush with bimodal leaffall (SLB) type of Myrsine seguinii, and long flush with broad unimodal leaffall (SLR) type of Symplocos prunifolia, Cleyera japonica, and Illicium anisatum. The multiple flush type is only one species, Eurya japonica, and it had multimodal leaffall pattern (MM type). The phenological pattern varied in relation to leaf life span, leaf size, and tree habit. Leaf life span ranged from 1.1 to 5.8 yr. The short flush species or SSU and SSB types were all canopy or subcanopy trees, and the former had short and the latter had long leaf life spans. The long flush species were all microphyllous small trees, and SLB type had a relatively long leaf life span in understory, SLR type had a long leaf life span in understory or in open habitat and/or forest gap as a pioneer tree. MM type had a long leaf life span and colonizing species in open habitat but they can survive in understory as well. The phenological attributes of evergreen trees were well corresponded to the ecological guild of the tree in both forest structure and successional stage, and were also constrained by phylogenetic groups.     

Winter photosynthesis by saplings of evergreen broad-leaved trees in a deciduous temperate forest

* Here we investigated photosynthetic traits of evergreen species under a deciduous canopy in a temperate forest and revealed the importance of CO2 assimilation during winter for annual CO2 assimilation. * Saplings were shaded by the canopy trees from spring through to autumn, but were less shaded during the winter months. Photosynthetic rates at light saturation (Aarea) were lower during winter than during the growing season. Aarea was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus during the winter, but differed little during summer and autumn. * Estimated daily CO2 assimilation (Aday) was higher during the winter than during the growing season in Camellia, Ilex and Photinia but was higher than that during the growing season only at the beginning and end of winter in Castanopsis, Cleyera and Quercus. Aday was higher in Camellia, Ilex and Photinia than in Castanopsis, Cleyera and Quercus but differed little among them during the growing season. * These results reveal the importance of winter CO2 assimilation for the growth of Camellia, Ilex and Photinia. Furthermore, differences in annual CO2 assimilation among species are strongly modified by species-specific photosynthetic traits during the winter under deciduous canopy trees.     

Shoot dynamics and architecture of saplings in Fagus crenata across its geographical range

 Relationships between leaf or shoot size, number, and arrangement in response to light were investigated to test the hypothesis that these characteristics are linked. In order to test this hypothesis, the divergence in allometry and shoot dynamics in saplings of Japanese beech (Fagus crenata) obtained from four populations and having different leaf sizes were examined in a nursery under both full sun and shade conditions. Trees with different leaf sizes also showed large differences in canopy structure, particularly when shade-grown saplings were compared. The final leaf mass distributions of the large-leaf populations were conical or “bottom - heavy”, while those of the small-leaf populations were planar or “top - heavy”. The slope of the allometric relations between leaf mass and shoot and branch mass in small-leaved populations were steeper than those in large-leaf populations. The four populations were classified into two growth types: populations producing a few large leaf and shoot modules corresponded to “stem growth type”, and those producing many small leaf and shoot modules corresponded to “leaf growth type”. These kinds of intra-specific variation in architecture and growth of F. crenata trees may influence the structure and dynamics of forests in accordance with differences in competitive ability or sensitivity to disturbances such as windstorm. Received: 18 March 1997 / Accepted: 21 October 1997     

Shoot growth phenology of co-existing evergreen and deciduous species in an oak forest

Shoot growth phenology was compared for the saplings of evergreen and deciduous woody species sharing the same microsite. Growth initiation occurred earlier in evergreens (among co-stratal species) while deciduous species completed their growth earlier. Shoot growth rate was significantly greater (P<0.01) for deciduous trees than evergreen trees. The amount of shoot elongations and shoot diameter was also significantly greater (P<0.01) for deciduous trees than evergreens. On the other hand, among shrubs the amount of shoot elongation and shoot diameter was greater for evergreens but the rate of elongation and diameter was more or less similar for both. The duration of shoot elongation and shoot diameter was significantly longer in evergreens than the deciduous species. Leaf packing (number of leaves per shoot) was significantly more dense in evergreen trees (P<0.01) than in deciduous tree species. Leaf packing was more dense in evergreen than deciduous shrubs but the difference was not significant. Leaf area (per individual leaf) at full expansion was significantly greater (P<0.01) in deciduous species. Leaf dry mass and specific leaf mass in the initial stage was significantly greater for evergreen species than for deciduous species. The number of buds/10 cm of shoot was higher in evergreens. However, the per cent mortality was also higher in them.     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.     

Partitioning of soil water among canopy trees in a seasonally dry tropical forest

Little is known about partitioning of soil water resources in species-rich, seasonally dry tropical forests. We assessed spatial and temporal patterns of soil water utilization in several canopy tree species on Barro Colorado Island, Panama, during the 1997 dry season. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θ_v), and sap flow were measured concurrently. Evaporative fractionation near the soil surface caused soil water δD to decrease from about –15‰ at 0.1 m to –50 to –55‰ at 1.2 m depth. Groundwater sampled at the sources of nearby springs during this period yielded an average δD value of –60‰. θ_v increased sharply and nearly linearly with depth to 0.7 m, then increased more slowly between 0.7 and 1.05 m. Based on xylem δD values, water uptake in some individual plants appeared to be restricted largely to the upper 20 cm of the soil profile where θ_v dropped below 20% during the dry season. In contrast, other individuals appeared to have access to water at depths greater than 1 m where θ_v remained above 45% throughout the dry season. The depths of water sources for trees with intermediate xylem δD values were less certain because variation in soil water δD between 20 and 70 cm was relatively small. Xylem water δD was also strongly dependent on tree size (diameter at breast height), with smaller trees appearing to preferentially tap deeper sources of soil water than larger trees. This relationship appeared to be species independent. Trees able to exploit progressively deeper sources of soil water during the dry season, as indicated by increasingly negative xylem δD values, were also able to maintain constant or even increase rates of water use. Seasonal courses of water use and soil water partitioning were associated with leaf phenology. Species with the smallest seasonal variability in leaf fall were also able to tap increasingly deep sources of soil water as the dry season progressed. Comparison of xylem, soil, and groundwater δD values thus pointed to spatial and temporal partitioning of water resources among several tropical forest canopy tree species during the dry season. Received: 5 October 1998 / Accepted: 23 June 1999     

Plant species diversity in abandoned coppice forests in a temperate deciduous forest area of central Japan

We investigated plant species diversity as it related to stand structure and landscape parameters in abandoned coppice forests in a temperate, deciduous forest area of central Japan, where Fagus crenata was originally dominant. The species occurring in the study plots were classified into habitat types based on a statistical analysis of their occurrence bias in particular habitats (e.g., primary forest, coniferous plantation) in the landscape studied. The relationships between stand structure, which reflected the gradient of management, and forest floor plant species diversity (H and J) and richness (number of species per unit area) were not significant. However, these factors did influence the forest floor plant composition of the different types of habitat. According to the multiple regression analysis, species diversity and the richness of forest floor plants was affected by landscape parameters rather than by stand structure. For trees, species richness was mainly affected by the relative dominance of F. crenata, which is one of the stand structure parameters that decreases with intensive management. This is probably because many of the tree species that are characteristic of coppice forests increase after F. crenata have been eliminated by management; these species are not dominant in the original forest, where they are suppressed by F. crenata, the shade-tolerant dominant species. The species diversity (H and J) of trees was positively correlated with some landscape parameters, including the road density around the study plot, which may be associated with the intensity of management activity. The number of disturbance-tolerant species increased with increasing road density. Stand structure mainly affected disturbance-intolerant forest floor plant species and disturbance-tolerant tree species. Thus, the species diversity responses differed between forest floor plants and trees. The impact of forest management on species diversity was more prominent for forest floor plants.     

Leaf and bud demography and shoot growth in evergreen and deciduous trees of central Himalaya, India

Leaf and bud demography and shoot growth were studied in 10 evergreen (ES) and 15 deciduous (DS) tree species occurring between 600 and 2200 m elevation in the central Himalayan mountains in India. Results were analyzed to help explain why ES prevail in the vegetation of this region, even though the number of ES is no greater than for DS. Although each species had its own pattern with regard to leaf and bud demography and seasonality of shoot extension and radial growth, it was possible to group the species on the basis of shoot growth phenology. In most species, leaves emerged during March-April, at the onset of warm and dry summer season. The ES recruit leaves in shoots more rapidly than the DS. Across all species, peak number of leaves per shoot (5.8–20.7), peak leaf area per shoot (116.2–1559.2 cm²), peak number of vegetative buds per shoot (1.9–14.5), bud survival per shoot (23–84%), shoot extension growth (6.4–40.8 cm) and shoot extension period (13–30 weeks) varied considerably. The peak leaf area per shoot (587.7 vs. 246.7 cm²) and shoot extension growth (19.3 vs. 11.2 cm) were significantly greater for DS than for ES, and these two functional groups of species were clearly separable with regard to shoot growth characteristics.Results indicate that rapid recruitment of leaf crop in the shoots, longer leaf life-span, and access to ground water due to deep roots were some of the advantages, the ES had over the DS, that may have likely enable them to maintain growth for a longer period in this region of warm winters and longer winter day length as compared to temperate climates. In the shallow rooted DS, shoot growth seems to be much affected by a seasonal drought in winter and they are likely to be affected more in the event of failure of monsoon rains in this region.     

Riparian disturbance and community structure of a Quercus-Ulmus forest in central Japan

Population structure and ecological characteristics of component species of a riparian Ulmus-Quercus forest in central Japan were analyzed with special reference to riparian disturbance regime. Though the dbh distribution of the whole community was L-shaped, those of several component tree populations had several modes, suggesting intermittent regeneration periods in the past. Correlation of spatial distributions among tree populations and subpopulations showed 6 major groups reflecting riparian disturbances in the past and different establishment patterns among species. A cluster of small-sized tree populations (Salix sachalinensis, Alnus hirsuta and Populus maximowiczii <30 cm dbh) were distributed on the lower terrace along the active river channel, while large-sized subpopulations (dbh 60 cm) of Quercus crispula and Ulumus davidiana var. japonica occurred on the higher terrace. The Phellodendron amurense population also occurred on the higher terrace in small clumps though the trees were small (less than 55 cm dbh). Subpopulations of intermediate-sized individuals (30 dbh < 60 cm) of Q. crispula and U. davidiana var. japonica, together with Betula and Acer spp. populations occurred on the intermediate terrace. Dendrochronological analyses indicated that the large and intermediate-sized tree groups were established about 330 and 90 years ago, respectively, while the small-sized tree group established about 35 years ago. A survey of historical disturbances showed that these periods of establishment of the former two groups almost coincided with the historically major floods occurring in 1662 and 1902. However, the disturbance that resulted in the establishment of the youngest group could not be precisely identified. Thus, the forest is a mosaic of three differently-aged patches, which is closely related to the frequency and scale of riparian disturbances. Longevity of trees and the preferred conditions for seed germination and/or seedling establishment were particularly important for the guild structure in this forest community.     

Leaf traits and photosynthetic parameters of saplings and adult trees of co-existing species in a temperate broad-leaved forest

In Central European forestry the establishment of broad-leaved mixed forests is attaining increasing importance, but little information exists about gas exchange characteristics of some of the tree species involved, which are less abundant today. In an old-growth forest in Central Germany (Hainich, Thuringia), (i) I compared morphological and chemical leaf traits that are indicative of leaf gas exchange characteristics among eight co-existing species, and (ii) analysed photosynthetic parameters of saplings and adult trees (lower and upper canopy level) in four of these species (Acer pseudoplatanus L., Carpinus betulus L., Fraxinus excelsior L. and Tilia platyphyllos Scop.).Leaves from the upper canopy in the eight species studied varied significantly in their specific leaf area (12.9–19.4 m² kg⁻¹), stomatal density (125–313 stomata mm⁻²), leaf nitrogen concentration (95–157 mmol N m⁻²) and δ¹³C content (–27.81 to –25.85‰). F. excelsior and C. betulus were largely contrasting species, which suggests that the species, which were studied in more detail, include the widest difference in leaf gas exchange among the co-existing species. The saplings of the four selected species exhibited shade acclimated leaves with net photosynthesis rates at saturating irradiance (A_max) between 5.0 and 6.4 μmol m⁻² s⁻¹. In adult trees A_max of fully sunlit leaves was more variable and ranged from 10.5 (C. betulus) to 16.3 μmol m⁻² s⁻¹ (F. excelsior). However, less negative δ¹³C values in F. excelsior sun leaves point to a strong limitation in gas exchange. In the lower canopy of adult trees A_max of F. excelsior (12.0 μmol m⁻² s⁻¹) was also greater than that of A. pseudoplatanus, C. betulus and T. platyphyllos (5.0–5.6 μmol m⁻² s⁻¹). This can be explained by the small leaf area and the absence of shade leaves in mature F. excelsior trees. Thus, a considerable variation in leaf traits and gas exchange was found among the co-existing tree species. The results suggest that species-specific characteristics increase the spatial heterogeneity of canopy gas exchange and should be taken into account in the interpretation and prediction of gas flux from mixed stands.In der Forstwirtschaft Mitteleuropas gewinnt die Begründung von Laubmischwäldern zunehmend an Bedeutung, aber über Eigenschaften im Gasaustausch einiger beteiligter Baumarten, die heute nicht so häufig sind, ist wenig bekannt. In einem Altbestand in Mitteldeutschland (Hainich, Thüringen) habe ich (i) morphologische und chemische Eigenschaften von Sonnenblättern, die Hinweise auf Charakteristika im Blattgaswechsel geben, an acht koexistierenden Baumarten untersucht, und (ii) Photosyntheseparameter von juvenilen und adulten Bäumen (unteres und oberes Kronenniveau) von vier dieser Arten (Acer pseudoplatanus L., Carpinus betulus L., Fraxinus excelsior L. and Tilia platyphyllos Scop.) erhoben.Blätter aus dem oberen Kronenraum der acht untersuchten Arten variierten signifikant in der spezifischen Blattfläche (12.9–19.4 m² kg⁻¹), der Stomatadichte (125–313 Stomata mm⁻²), dem Blattstickstoffgehalt (95–157 mmol N m⁻²) und den δ¹³C-Werten (–27.81 bis –25.85‰). In diesem Kollektiv zeigten F. excelsior und C. betulus groβe Unterschiede, was darauf hindeutet, dass die Arten, die genauer untersucht wurden, die Spannweite an Gaswechseleigenschaften unter den koexistierenden Baumarten umfassen. Die Jungpflanzen der vier ausgewählten Arten besaßen Schattenblätter, deren Netto-Photosyntheserate bei hoher Lichtintensität (A_max) zwischen 5.0 and 6.4 μmol m⁻² s⁻¹ variierte. An Sonnenblättern von Altbäumen war A_max variabler und lag zwischen 10.5 (C. betulus) und 16.3 μmol m⁻² s⁻¹ (F. excelsior). Allerdings weisen hohe δ¹³C-Werte in Sonnenblättern von F. excelsior auf eine starke Limitierung des Gasaustauschs hin. Auch in der unteren Krone der Altbäume war A_max von F. excelsior (12.0 μmol m⁻² s⁻¹) höher als A_max von A. pseudoplatanus, C. betulus und T. platyphyllos (5.0–5.6 μmol m⁻² s⁻¹). Dies kann durch die geringe Blattfläche und die Abwesenheit von Schattenblättern in der Krone adulter Bäume von F. excelsior erklärt werden. Zwischen den koexistierenden Baumarten wurde somit in Bezug auf Blatteigenschaften und Photosyntheseparameter eine erhebliche Variation festgestellt. Die Ergebnisse legen nahe, dass artspezifische Eigenschaften die räumliche Heterogenität des Gaswechsels im Kronenraum erhöhen und bei der Interpretation und Vorhersage von Gasflüssen über Mischbeständen berücksichtigt werden sollten.     

Population structure and spatial patterns for trees in a temperate old-growth evergreen broad-leaved forest in Japan

The population structure and spatial pattern of major tree species in a warm-temperate old-growth evergreen broad-leaved forest in the Tatera Forest Reserve of Japan were investigated. All stems 5 cm in diameter at breast height (DBH) were mapped on a 4 ha plot and analyses were made of population structure and the spatial distribution and spatial association of stems in different vertical layers for nine species. This was done in the context of scale dependency. The plot was located on a very gentle slope and 17.1% of its canopy layer was in gaps. It contained 45 woody plant species and 4570 living stems with a basal area of 63.9 m² ha^–1. Castanopsis cuspidata var. sieboldii, the most dominant species for the basal area, had the maximum DBH among the species present, fewer smaller stems and a lower coefficient of statistical skewness of the DBH distribution. The second most dominant species, Dystilium racemosum, had the highest stem density (410 ha^–1), more abundant smaller stems and a relatively higher coefficient of skewness. Most stems in different vertical layers showed a weakly aggregated distribution with loose colonies as basic units. Gap dependency for the occurrence of stems under the canopy layer was weak. Maximum slope degree of the plot also weakly affected the occurrence of stems. Spatial associations varied among intra- and interspecific cohorts in the different layers and spatial scales examined, and positive associations among cohorts were found more frequently as the scales examined became larger. This tendency suggests that key factors forming observed spatial associations might vary with the spatial scales.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Intra- and inter-specific variation in canopy photosynthesis in a mixed deciduous forest

 Within the same forest, photosynthesis can vary greatly among species and within an individual tree. Quantifying the magnitude of variation in leaf-level photosynthesis in a forest canopy will improve our understanding of and ability to model forest carbon cycling. This information requires extensive sampling of photosynthesis in the canopy. We used a 22-m-tall, four-wheel-drive aerial lift to reach five to ten leaves from the tops of numerous individuals of several species of temperate deciduous trees in central Massachusetts. The goals of this study were to measure light-saturated photosynthesis in co-occurring canopy tree species under field conditions, and to identify sampling schemes appropriate for canopy tree studies with challenging logistics. Photosynthesis differed significantly among species. Even though all leaves measured were canopy-top, sun-acclimated foliage, the more shade-tolerant species tended to have lower light-saturated photosynthetic rates (P _max) than the shade-intolerant species. Likewise, leaf mass per area (LMA) and nitrogen content (N) varied significantly between species. With only one exception, the shade-tolerant species tended to have lower nitrogen content on an area basis than the intolerant species, although the LMA did not differ systematically between these ecological types. Light-saturated P _max rates and nitrogen content, both calculated on either an area or a mass basis, and the leaf mass to area ratio, significantly differed not only among species, but also among individuals within species (P<0.0001 for both). Differences among species accounted for a greater proportion of variance in the P _max rates and the nitrogen content than the differences among individuals within a species (58.5–78.8% of the total variance for the measured parameters was attributed to species-level differences versus 5.5–17.4% of the variance was attributed to differences between individual trees of a given species). Furthermore, more variation is accounted for by differences among leaves in a single individual tree, than by differences among individual trees of a given species (10.7–30.4% versus 5.5–17.4%). This result allows us to compare species-level photosynthesis, even if the sample size of the number of trees is low. This is important because studies of canopy-level photosynthesis are often limited by the difficulty of canopy access. As an alternative to direct canopy access measurements of photosynthesis, it would be useful to find an ”easy-to-measure” proxy for light-saturated photosynthetic rates to facilitate modeling forest carbon cycling. Across all species in this study, the strongest correlation was between nitrogen content expressed on an area basis (mmol m^–2, N _area) and light-saturated P _max rate (μmol m^–2 s^–1, P _maxarea) (r ²=0.511). However, within a given species, leaf nitrogen was not tightly correlated with photosynthesis. Our sampling design minimized intra-specific leaf-level variation (i.e., leaves were taken only from the top of the canopy and at only one point in the season). This implies that easy-to-measure trends in nitrogen content of leaves may be used to predict the species-specific light-saturated P _max rates. Received: 16 March 1996 / Accepted: 16 August 1996     

密度制约对宝天曼落叶阔叶林锐齿栎死亡前后分布格局的影响

为了在温带和亚热带的过渡带中验证森林树木死亡是否受密度制约的影响, 我们在宝天曼国家级自然保护区选择1个100 m×100 m的暖温带落叶阔叶林样地, 用双变量函数g(r)(the pair-correlation function)研究了锐齿栎(Quercus aliena var. acuteserrata)死亡前(活树和枯木统称为死亡前树木)和死亡后(活树为死亡后树木)的分布格局。把样地中的锐齿栎分为幼树(1 cm ≤ DBH<10 cm)、小树(10 cm ≤ DBH<20 cm)和成年树(DBH≥20 cm)3个不同的生长阶段来研究密度制约对空间分布格局的影响机制。结果表明: (1)死亡前锐齿栎在r>5 m尺度呈聚集分布, 死亡后幼树(1 cm≤DBH<10 cm)和成年树(DBH≥20 cm)在1-25 m尺度呈现随机分布, 死亡后小树(10 cm≤DBH< 20 cm)在r<1.5 m和2.5-4.5 m的尺度为随机分布, 在r>5 m的尺度呈聚集分布; (2)采用随机标签零模型和案例-对照设计的方法, 排除生境异质性影响后, 将幼树和小树的分布格局作为案例, 将成年树的分布格局作为对照, 并代表生境异质性的作用, 通过小径级树木与成年树分布格局的对比发现, 密度制约效应对死亡前后的锐齿栎分布格局均具有影响; (3)幼树和小树在成年树周围的分布死亡前为显著聚集分布格局, 死亡后剩余树木的聚集强度下降; 随着与成年树之间距离的增加, 死亡后的幼树(仅包括现存的活树)逐渐向随机分布格局演替。本研究初步表明锐齿栎空间格局受生境异质性的影响并呈现出显著的聚集效应, 排除生境异质性影响后, 锐齿栎死亡前后的空间格局受到密度制约的影响, 这一结果为Janzen-Connell假说提供了支持。     

Temporal patterns of water flux in trees and lianas in a Panamanian moist forest

Using constant heat sap flow sensors, xylem water fluxes in ten tree species and two liana species were monitored for 5–10 days during the beginning of the wet season in May, 1993. For a subset of the trees, a branch was also monitored at the top of the crown for 5 days. Xylem flux (J _S) was related diurnally in all plants to vapor pressure deficit (D) measured within the upper-third of the canopy, and to incoming shortwave radiation R _S above the canopy. Cross-correlation analysis was used to estimate time lags between diurnal patterns of J _S and D or R _S, and between J _S in stems and branches. The maximum correlation coefficient from cross-correlation of J _S with R _S (range=0.57–0.92) was often higher than the maximum of J _S with D (range=0.43–0.89), indicating that diurnal J _S was more dependent on R _S than D. Time lags (lag corresponding to maximum correlation) of J _S at stem-base with D was shorter (0–45 min) than with radiation (5–115 min), highly variable within a species, and uncorrelated to the height or exposure of tree crowns or liana in the canopy. On a stand level, not accounting for the diel lag between stem sap flux and canopy flux resulted in errors in estimated canopy transpiration of up to 30%. Received: 19 October 1998 / Accepted: 8 June 1999     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

Density-dependence and co-existence of conifer and broad-leaved trees in a Japanese northern mixed forest

Abstract. The growth and survival of coniferous trees (particularly Abies sachalinensis) and broad-leaved trees (particularly Quercus crispula) were followed over a 15-yr period in a 15.5-ha area in a northern mixed forest in Japan, and the coexistence of the two groups was simulated by a density-dependent projection matrix model. The density-dependent model assumes that the density effect of mother trees due to one-sided competition for light on smaller-sized tree regulates the demographic functions. The mother tree densities of conifers and broad-leaved trees have stronger negative effects on the recruitment and survival of seedlings of their own group than of the other group. These results support the idea of reciprocal replacement for conifer and broad-leaved trees. Simulations using the density-dependent model showed that the two groups will co-exist within a particular range of recruitment rates. However, the density of both groups did not affect the growth rate of any tree, and equilibrium DBH-distributions from density-dependent matrices were quite different from present distributions both for conifers and broad-leaved trees. On the other hand, equilibrium DBH-distributions of conifer and broad-leaved trees from density-independent matrices were quite distinct from each other, reflecting different survivorship curves of the two dominants. These results suggest that density-dependent processes are not so important for shaping population structures in this northern mixed forest.     

Adjustment of foliage structure and function to a canopy light gradient in two co-existing deciduous trees. Variability in leaf inclination angles in relation to petiole morphology

 Foliar inclination angles, petiole morphology and dry matter partitioning between assimilative and support biomass were studied in shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. along a natural light gradient across the canopy. The leaves of sub-canopy species T. cordata were on average exposed to lower irradiances, and they were also more horizontal with greater blade inclination angles (ϕ_B, defined as the angle between the leaf fall-line and the horizon; ϕ_B was positive for the leaves inclined upwards, and negative for the leaves inclined downwards) than those in P. tremula. Seasonal average daily integrated quantum flux density (Q _int, mol m^–2 day^–1) and ϕ_B were not related in T. cordata, and only a weak negative effect of Q _int on ϕ_B was detected in P. tremula. Nevertheless, when both species were pooled, there was a strong negative relationship between Q _int and ϕ_B, implying that the leaves became progressively vertical with increasing height in the canopy. Interspecific differences in foliage inclination were mainly related to petiole morphology, in particular to petiole length, rather than to contrasting biomass investment patterns between assimilative and support tissues within the leaf. It was suggested that more horizontal leaves, resulting from the species-specific structure of petioles, partly explain the superior performance of shade-tolerant T. cordata in the understory and the sub-canopy. Received: 13 November 1997 / Accepted: 6 March 1998     

Diversity of flowering and fruiting phenology of trees in a tropical deciduous forest in India

BACKGROUND AND AIMS: In the dry tropics, vegetative phenology varies widely with tree characteristics and soil conditions. The present work aims to document the phenological diversity of flowering and fruiting with reference to leafing events in Indian dry-tropical tree species. METHODS: Nine tree species, including one leaf-exchanging and eight deciduous showing varying leafless periods, were studied. Monthly counts of leaves, flowers and fruits were made on 160 tagged twigs on ten individuals of each species for initiation, completion and duration of different phenological events through two annual cycles. KEY RESULTS: Variation in flowering relative to leaf flushing (which occurred just prior to or during a hot, dry summer) revealed five flowering types: summer flowering (on foliated shoots), rainy-season flowering (on foliated shoots following significant rains), autumn flowering (on shoots with mature leaves), winter flowering (on shoots undergoing leaf fall) and dry-season flowering (on leafless shoots). Duration of the fruiting phenophase was shortest (3-4 months) in dry-season and winter-flowering species, 6-9 months in rainy-and autumn-flowering species, and maximum (11 months) in summer-flowering species. A wide range of time lag (<1 to >8 months) between the start of vegetative (first-leaf flush) and reproductive (first-visible flower) phases was recorded in deciduous species; this time lag was correlated with the extent of the leafless period. A synthesis of available phenological information on 119 Indian tropical trees showed that summer-flowering species were most abundant (56 % of total species) amongst the five types recognized. CONCLUSIONS: The wide diversity of seasonal flowering and fruiting with linkages to leaf flush time and leafless period reflect the fact that variable reproductive and survival strategies evolved in tree species under a monsoonic bioclimate. Flowering periodicity has evolved as an adaptation to an annual leafless period and the time required for the fruit to develop. The direct relationship between leafless period (inverse of growing period) and time lag between onset of vegetative and reproductive phases reflects the partitioning of resource use for supporting these phases. Predominance of summer flowering coupled with summer leaf flushing seems to be a unique adaptation in trees to survive under a strongly seasonal tropical climate.     

Dew and its effect on two heliophile understorey species of a tropical dry deciduous forest in Mexico

 A series of horizontal and vertical measurements of dew deposition and dew duration were carried out in a tropical dry deciduous forest in western Mexico (19° 30′ N, 105° 03′ W). The effect of dew on transpiration in heliophile species was also investigated. The amount of dew was very variable with no temporal or spatial pattern. The amount of dew measured at two horizontal transects (maximum and minimum) (at heights of 0.20 m and 1.30 m above ground level) was from 0.014 to 0.203 mm and from 0.013 to 0.061 mm in the middle and at the end of the dry season, respectively. Dew deposition at different vertical levels (0.50–12.5 m height) ranged from 0.04 to 0.36 mm. The duration of dew formation ranged between 60 and 129 min after sunrise above the canopy (a height of 11 m), and between 259 and 290 min after sunrise at a height of 2 m. Daily transpiration rates were 883 and 632 g m^–2 d^–1 in Coccoloba liebmannii and 538 and 864 g m^–2 d^–1 in Jacquinia pungens in January and April, respectively. Transpiration was restricted from sunrise to early afternoon in April, as a result of the pronounced midday closure of stomata in both species. The reduction of transpiration by dew ranged from 13.2 to 50.1 g m^–2 d^–1 and from 4.5 to 77.7 g m^–2 d^–1 for C. liebmannii and J. pungens, respectively. Dew can play an important role in enhancing the survival of heliophile species in the dry season by reducing transpiration rates during the morning. Received: 30 April 1998 / Accepted: 14 December 1998