基于地形因子的天童地区常绿树种和落叶树种共存机制研究

位于亚热带的浙江天童和古田山常绿阔叶林大样地分布有较高比例的落叶树种,那么它们与常绿树种的共存机制是什么?常绿树种和落叶树种生态习性差异较大,二者对生境的选择应有所不同,我们推测生境分化可能是两类植物实现共存的主要机制。为检验该假设,我们以天童20ha动态样地调查数据为依托,选择个体数≥20的55个常绿树种和42个落叶树种作为分析对象,用典范对应分析(CCA)研究了地形因子对二者分布的影响差异,用torus转换检验来分析常绿树种和落叶树种与各类地形生境的关联。结果如下:(1)CCA分析表明地形因子对常绿树种分布的解释量为19.2%,对落叶树种分布的解释量为7.0%。(2)torus转换检验结果表明,与沟谷成正关联的常绿树种和落叶树种的比例分别为16.4%和28.6%,成负关联的比例分别为40%和7%;与山脊成正关联的常绿树种和落叶树种的比例分别为41.8%和4.8%,成负关联的比例分别为10.9%和47.6%;与受干扰生境成正关联的常绿树种和落叶树种的比例分别为16.4%和42.9%。上述结果说明地形对常绿树种分布的影响大于落叶树种;两个植物类群对生境的选择多呈现相反格局,尤其是在沟谷生境和山脊生境,这进一步表明生境分化是常绿树种和落叶树种共存的重要机制之一,生态位理论在一定程度上能较好地解释亚热带常绿阔叶林物种多样性的维持。     

Regeneration in subalpine coniferous forests

The regeneration process of a subalpine coniferous forest, a mixed forest ofTsuga diversifolia (dominant species),Abies veitchii, Abies mariessi, andPicea jezoensis var.hondoensis, was studied on the basis of annual ring data. The age class distribution was discontinuous and four age groups occurred in the study plot (30m×30m). The canopy layer was a mosaic of patches (83.8–133.7 m² patch area), which had different mean ages. The recruitment of canopy trees was carried out only by advance regeneration in the plot. The diameter growth ofAbies andPicea exceeded diameter growth ofTsuga in the gap.Abies lived for 200–300 years and their trunks were susceptible to heart rot.Picea lived for 300–400 years andTsuga for more than 400 years. The regeneration process derived from the analysis of the plot consisted of three phases leading to the development of a even-aged patch; (1) the establishment of saplings before a gap opening, (2) the opening of a gap in the canopy and repair of the canopy by advance regenerated saplings dominated by rapid growth species,Abies andPicea, and (3) the dying off of canopy trees as each species reached the end of its life-span, resulting in pure patches of long-livedTsuga.     

Tertiary relic deciduous forests on a humid subtropical mountain, Mt. Emei, Sichuan, China

The floristic characteristics, age structure and survival modes of Tertiary-relic deciduous forests were analyzed at 1600 m on Mt. Emei (3099 m), Sichuan, China. Three plots were selected to represent typical topographies: Plot 1 at 1620 m on a scree slope, Plot 2 at 1640 m on a slope with moderately rocky soils, and Plot 3 at 1616 m on a gentle slope with less rocky soils. At Plot 1, on the scree slope, the forest was rich in species and dominated by Tertiary remnants and other deciduous trees (Davidia involucrata, Styrax hemsleyana, Cercidiphyllum japonicum var.sinense, Pterocarya hupehensis, Prunus brachypoda, Prunus padus, Tetracentron sinense andStaphylea holocarpa). The relic deciduous tree taxaDavidia, Cercidiphyllum, Tetracentron andEuptelea occupied the unstable concave slopes, where evergreen broad-leaved trees (Castanopsis platycantha andMachilus pingii) were rarely able to survive. On the relatively stable convex slopes of this plot, evergreen trees with small diameter mainly appeared in the subcanopy and shrub layers.Davidia involucrata was the dominant species in this forest. On the slope with moderately rocky soils (Plot 2), the forest was co-dominated by relicDavidia and other deciduous (Styrax andPterocarya), and evergreen trees (Castanopsis andMachilus). On the gentle slope with less rocky soils (Plot 3),Davidia trees were found only in the subcanopy and shrub layers, and the forest was dominated byMachilus, Castanopsis, Styrax andPrunus trees. Regeneration ofDavidia occurs mainly on the scree slope where landslides are most common. The age structure of theDavidia stands indicates that this species is able to survive on the unstable scree habitat due to its strong sprouting ability. The Tertiary-relic deciduous forest on the scree slope is seen to be a topographic climax forest.     

Successional change of forest pattern along topographical gradients in warm-temperate mixed forests in Mt Kiyosumi,central Japan

Forest patterns along topographical gradients were compared between second- and old-growth forested watersheds in a warm-temperate zone of Mt Kiyosumi, central Japan. Three community types were distinguished depending on the topographical habitat type in each watershed, for example, conifer forest was dominated byAbies firma andTsuga sieboldii on ridge sites, evergreen broad-leaved forest was dominated byQuercus acuta, Q. salicina andCastanopsis cuspidata var.sieboldii on slope sites, and deciduous forest was dominated byEuptelea polyandra andCornus controversa in valley sites. Beta diversity and distinctiveness of each topographical community type increased with progression of secondary succession. Conifers and evergreen broad-leaved trees, which were intermingled with each other on ridges and slopes of the second-growth watershed, were in turn restricted to the ridge and slope habitat type, respectively, in the old-growth watershed. The process of this differentiation can be explained by the continuous regeneration of conifers on ridge sites, and its absence on slope sites due to different light conditions caused by progressive canopy closure of evergreen trees on the slope sites toward the old-growth watershed. In the valley type habitat, frequent soil disturbance, such as landslides and soil creep, hinder the continuous growth of late successional evergreen trees, and thus seral or pioneer deciduous trees can persist in the habitat.     

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.     

Forest succession in the subalpine region of Mt. Fuji,Japan

Subalpine forest succession was studied on Mt. Fuji, Japan, where various types of forests in different successional phases occur owing to volcanic action. Ninety stands were subjected to ordination using an index (SI) defined by the relative basal area and the life span of component woody species, and the cover of canopy layer of the sample stands. Two different sequences of sample stands were found. One was from deciduous scrubs, through Larix kaempferi forests and Abies forests, to Tsuga diversifolia forests, and the other from Abies-Tsuga thickets to Abies forests. Through analyses of the forest structure and composition, soil survey and identification of fallen logs, the former sequence was recognized as the primary sere and the latter as a regeneration sere following gap formation. During forest succession, basal area reached a maximum in the seral phase with a multi-layered structure. The Tsuga forests, whose understory is restricted to a moss layer, were regarded as the climax. The death or fall of Tsuga stems resulted in gaps, which were subsequently occupied by Abies-Tsuga thickets. The second Abies forests were distinguished from the ones in the primary sere by the occurrence of Dryopteris and Cacalia and the lack of Rhododendron in the understorey. Both Abies forest types included Tsuga saplings. Thus, a cyclic relation is supposed between Abies and Tsuga.Nomenclature follows Ohwi (1975) and Nakaike (1982) for vascular plants, Iwatsuki & Noguchi (1973) for mosses, Inoue (1981) for hepaticae, Kashiwadani (1981) for lichens, respectively. Abies veitchii, A. mariesii were lumped as Abies spp.I wish to express my sincere gratitude to Prof. Toshio Hamaya, Tokyo, for the cordial guidance and encouragement. I also thank Prof. M. Numata and Dr. M. Ohsawa, Chiba, Prof. K. Okutomi, Tokyo, Dr. K. Suzuki, Tokyo, Dr. M. Suzuki, Kanazawa, and Mr. H. Taoda, Kumamoto, for their valuable advice and discussions.     

Effect of bryophytes on survival of conifer seedlings in subalpine forests of central Japan

Bryophytes form the major seedbed for coniferous trees in the subalpine forests of central Japan. Field experiments were conducted on the floor of a closed stand dominated byTsuga on Mt Fuji to examine the seedling survival ofAbies veitchii andTsuga diversifolia in seven substrate types in relation to the morphological characteristics of the seeds and seedlings. NeitherAbies norTsuga seedlings could survive on dwarf-bamboo litter beyond 2 years after the seed rain.Abies seedlings survived not only in all types of bryophyte communities but also in larch litter. In contrast, the survival ofTsuga seedlings was restricted to communities of smaller-statured bryophytes found on logs. The seeds and 1st year seedlings ofA. veitchii were larger than those ofT. diversifolia. Abies seedlings produced hypocotyls taller than any bryophyte community and radicles which were long enough to penetrate into the moist humus layer, whereasTsuga seedling radicles presumably penetrate into humus only in bryophyte communities forming a thin mat on logs. The effect of bryophytes on survival of tree seedlings therefore depends on the relationships between the morphology of seeds and seedlings and the structure of bryophyte communities.     

State of fallen logs and the occurrence of conifer seedlings and saplings in boreal and subalpine old-growth forests in Japan

Fallen logs on the forest floor play an important role in tree seedling establishment in boreal and subalpine old-growth forests. We examined the abundance and state (degree of decay, moss coverage on their surfaces) of fallen logs on the forest floor and compared the occurrence of conifer seedlings and saplings on soil and fallen logs in old-growth evergreen boreal (Taisetsu) and subalpine (Yatsugatake) coniferous forest stands in Japan. The forest floors of both stands were covered mostly by soil, and fallen logs covered only a small proportion (4.5% in Taisetsu and 2.5% in Yatsugatake) of the forest floor. In both stands, no seedlings were found on freshly fallen logs. Both moss coverage and the occurrence of seedlings or saplings on fallen logs increased with decay in both stands. In Taisetsu, the occurrence of all conifer seedlings and saplings was, in general, restricted to fallen logs. In contrast, in Yatsugatake, Picea and Tsuga seedlings occurred much more frequently on fallen logs than on soil, whereas Abies seedlings and saplings became established on both soil and fallen logs. Seedlings were taller on decayed fallen logs, but taller saplings were rare on fallen logs in Yatsugatake. Results indicate that species characteristics, the condition of the fallen logs and environmental factors determine the preponderance of seedling and sapling establishment on fallen logs. The condition of fallen logs varies with the degree of decay, and fallen logs that function as seedling establishment sites on the forest floor are thought to be limited by time and space.     

Forest encroachment into a Californian grassland: examining the simultaneous effects of facilitation and competition on tree seedling recruitment

Competition and facilitation are both considered major factors affecting the structure of plant assemblages, yet few studies have quantified positive, negative, and net effects simultaneously. In this study, we investigated the positive, negative, and net effects of tree saplings on the encroachment of two tree species, Douglas fir (Pseudotsuga menziesii) and tanoak (Lithocarpus densiflora), into a coastal California grassland. The study involved three components: sampling the spatial distributions of P. menziesii and L. densiflora in the grasslands, a field experiment examining seedling survival in different grassland environments, and a greenhouse experiment examining the effects of soil moisture on early seedling performance. The field experiment was conducted over a 2-year period, using Pseudotsuga in 2002 and both species in 2003. Seedlings were separated into four treatment groups: those planted in open grassland, in shaded grassland, under artificial (plastic) conifer saplings, and under natural Pseudotsuga saplings. Air temperature, relative humidity, soil moisture, incident radiation levels and fog water inputs were measured for each treatment group in 2003. In the greenhouse experiment, Pseudotsuga and Lithocarpus seedlings were grown for 13 weeks in watering treatments simulating the summer soil moisture conditions of the open grasslands and under Pseudotsuga saplings. Surveys of naturally established seedlings found that Lithocarpus occurred only under Pseudotsuga saplings, while most Pseudotsuga seedlings were located near but not directly under conspecific saplings. In the field experiment, positive effects of tree saplings were much larger than negative effects, resulting in strong net facilitation of seedling establishment. Survival for both species was always higher under the plastic and live trees than in the open or shade plots. The primary mechanism facilitating seedling survival appeared to be increased soil moisture caused by input of fog precipitation coupled with reduced microsite evaporation. The greenhouse experiment further showed that soil moisture strongly affected seedling performance, with both species having much higher photosynthetic rates in the higher moisture treatment. In the lower moisture treatment, Pseudotsuga seedlings had higher photosynthetic rates and stomatal conductance than Lithocarpus, suggesting they may be able to better tolerate the environmental conditions found in the open grasslands. Our combined results suggest that rate and patterning of woody plant encroachment can be strongly influenced by facilitation and that fog precipitation may play a key role in plant interactions.     

Aspects of tissue water relations and seasonal changes of leaf water potential components of evergreen and deciduous species coexisting in tropical dry forests

Summary This study compared the tissue water relations and seasonal changes in leaf water potential components of an evergreen tree,Morisonia americana, and two evergreen shrubs,Capparis verrucosa andC. aristiquetae, with two deciduous trees,Humboltiella arborea andLonchocarpus dipteroneurus, and the deciduous vineMansoa verrucifera. All these species coexist in a tropical dry forest in Venezuela. Leaves of the evergreen species are sclerophyllous, while those of the deciduous species are mesophytic. Leaf area to leaf weight ratios of fully mature leaves were about 75 and 170 cm² g^–1 in evergreen and deciduous species, respectively. Seasonal fluctuations of leaf water content per unit of dry weight, water potential, and turgor pressure were smaller in evergreen than in deciduous species. The analysis of tissue water relations using pressurevolume curves showed that evergreen species could develop a higher leaf turgor and lose turgor at lower leaf water potentials than deciduous species. This was related to a lower osmotic potential at full turgor in evergreen (-3.0 MPa)_than in deciduous (-2.0 MPa) species, rather than to the elastic properties of leaf tissue. The volumetric modulus of elasticity was 14 MPa in evergreen compared with 7–10 MPa in deciduous species. Thus, leaf characteristics are important in determining the drought resistance of evergreen species of this tropical dry forest.     

Spatial pattern dynamics over 10 years in a conifer/broadleaved forest,northern Japan

The present study investigated stand dynamics during 10-year period in a conifer/broadleaved mixed forest in Hokkaido, northern Japan, focusing on spatially dependent recruitment, mortality and growth of two growth-form groups, deciduous broadleaved species and the dominant evergreen conifer Abies sachalinensis. The stand-level basal area was maintained over the 10-year period, while a compositional equilibrium at the individual species level was not confirmed. Univariate and bivariate spatial analyses revealed clustering of many of the constituent species. The absence of single-species patches suggested an ambiguous mosaic formed by co-occurrence of Abies and broadleaved trees. The trend towards an aggregated distribution of Abies and broadleaved trees was caused by spatially dependent recruitment rather than mortality. New recruits of broadleaved species were spatially associated with surviving broadleaved trees, while this was not the case for Abies. The degree of competitive effects on growth was not consistent over the 10-year period. Abies showed between-groups competition, but not within-group competition. In contrast, we found asymmetric competitions between the broadleaved trees. Our results suggest that Abies is not sufficiently competitively dominant to exclude broadleaved trees, and that the co-occurrence of the two growth-form groups might be maintained.     

Differentiation and maintenance of topo-community patterns with reference to regeneration dynamics in mixed cool temperate forests in the Chichibu Mountains,central Japan

Topo-community structure and dynamics were studied in mixed cool temperate forests, using the regeneration dynamics, to clarify the maintenance mechanisms of community patterns along a microtopographic gradient. The 76 stands studied were classified into two groups (i.e. convex slope and concave slope stands). The soil surface was more eroded on the concave slope than on the convex slope, while water potential was not significantly different between topographies. On the convex slope, even-aged patches alternated between young phase patches dominated by shade intolerant species, such asAcer rufinerve andBetula grossa, and mature phase patches, withTsuga sieboldii andFagus crenata. A slower lateral growth rate ofTsuga canopy trees and the absence of suppressed saplings in the mature phase may prolong the gap phase, which provides a favorable situation to shade-intolerant species. On the concave slope, patch structure was less clear, and process of replacement of canopy species by previously suppressed individuals of the same species was seen in the mature phase, which was mainly composed ofF. crenata, Fagus japonica, Acer sieboldianum andStewartia pseudo-camellia. Gaps on the concave slope were formed frequently but were generally closed within 10 years by lateral growth of deciduous canopy trees and by upgrowth of suppressed trees, and thus some individuals underwent recurrent periods of suppression until they reached the canopy. We concluded that soil surface stability and gap encroachment pattern are critical to the maintenance of the community pattern along a microtopographic gradient.     

Water relations of stem succulent trees in north-central Baja California

Summary Water relations of several stem succulent trees were measured in north-central Baja California in comparisons to other growth forms in the same habitat. Our research concentrated on three stem succulent species (Idria collumnaris, Pachycormus discolor and Bursera microphylla) each with a different succulent stem morphology. The stem succulent trees had 1 to 4 kg H₂O/m³ of trunk while the other trees and shrubs in the same habitat had 0.6 to 0.8 kg H₂O/m³. The diurnal and seasonal variation in leaf water potential was small for the stem succulent species in comparison to deciduous and evergreen species as a consequence of the stem-water, buffering capacity. In addition, the leaf conductance of the stem succulent species was low (60 mmol m^–2 s^–1) and yet, the leaf conductance decreased through the day similar to adjacent evergreen and deciduous species. The leaves of the stem succulent trees lost turgor at low saturated water deficits (0.06 to 0.14), had comparatively high osmotic potentials, and high values of elastic modulus in comparison to adjacent evergreen and deciduous species. The stem acts as an important buffering mechanism allowing for the maintenance of leaf turgor in these stem succulent trees. The low transpiration rates of the stem succulent trees may be a mechanism to minimize leaf saturated water deficit and extend leaf longevity.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

冰雪灾害对江西九连山常绿阔叶林的影响

以江西九连山国家自然保护区常绿阔叶林为对象,研究林分不同层次优势种受灾程度,分析物种海拔、坡度对受灾程度的影响。结果表明:乔木上层和中层树木以断稍为主,乔木下层树木以断稍和腰折为主,小树和幼树以压弯为主。在平均受损指数MDI(Mean Damage Index)方面,乔木上层米槠MDI值最高,拟赤杨最低; 乔木中层鸭公树MDI值最高,浙江新木姜子最低; 乔木下层米槠MDI值最高,浙江新木姜子最低; 小树米槠MDI值最高,细枝柃最低; 幼树二列叶柃MDI值最高,狗骨柴最低。海拔对不同林分层次树木的部分受损指标有显著影响(P<0.05); 整体上,不同林分层次树木的受损指标在700～760 m海拔生境高于640～700 m海拔生境。不同坡度生境下树木的受损指标均无显著差异(P>0.05)。     

Codominance of <Emphasis Type="Italic">Acer saccharum</Emphasis> and <Emphasis Type="Italic">Fagus grandifolia</Emphasis>: the role of <Emphasis Type="Italic">Fagus</Emphasis> root sprouts along a slope gradient in an old-growth forest

We studied how the unusual capacity of mature Fagus grandifolia to form clumps of clonal stems from root sprouts can contribute to its frequent codominance with Acer saccharum in southern Quebec, Canada. In an old-growth forest, the degree of dominance by the two species shifted along topographic gradients spanning a few hundreds of meters, with Fagus more frequent on lower slopes and Acer on upper slopes. The frequency distribution of Fagus stem diameter had an inverse J distribution at all slope positions, which is indicative of continuous recruitment. Acer stem diameter also had an inverse J pattern, except at lower slope positions where size structure was discontinuous. For stems <2 m tall, Fagus regenerated mainly by sprouts at the upper and mid-slopes, while regeneration from seed was more pronounced on the lower slope. This change of regeneration mode affected the spatial pattern of Fagus stems. Understory trees of Fagus were positively correlated with conspecific canopy trees on upper and mid-slopes, but not on lower slopes where Fagus regenerated mainly by seedlings. Understory trees of Acer were positively correlated with conspecific canopy trees only on the mid-slope. There were many Fagus seedlings around Acer canopy trees at the lower slope, suggesting the potential replacement of Acer canopy trees by Fagus. This study suggests that the regeneration traits of the two species changed with slope position and that Fagus patches originating from root sprouts can contribute to the maintenance of Acer–Fagus codominance at the scale of local landscapes.     

Topographical variations in a plant–soil system along a slope on Mt Ryuoh, Japan

The plant–soil system was studied at different topographic levels (i.e. ridge, backslope and footslope) along a slope in a Cryptomeria plantation. Soil solution chemistry at each representative topographic plot was investigated. Tree height and diameter of Cryptomeria decreased upslope. The understory species composition changed along the slope. The upper part of the slope with Oa horizon soil N transformation was characterized by ammonification, while most of the inorganic N in the lower part of the slope without Oa horizon was nitrified. The inorganic N form in the soil solution corresponded with soil N transformation. Ammonium was the dominant inorganic N at the ridge, while NO₃ predominated at the foot of the slope. Soil solution chemistry was similar to throughfall at the ridge. At the foot slope, the chemical composition of the soil solution was different from throughfall due to high NO_3– concentrations. This suggests that the inorganic N form regulated not only N concentration but also cation concentrations. The soil N transformation pattern is important in nutrient cycling.     

Seedling establishment of deciduous trees in various topographic positions

Abstract. We investigated the effect of topography‐related environmental factors (i.e., ground‐surface stability and soil moisture) on seedling establishment of 8 deciduous tree species. A field experiment was carried out using canopy species, which were classified into 3 groups based on the spatial distribution of adult trees (ridge, slope and valley). Demographic parameters were compared among species during the early stage of seedling establishment among 3 topographic positions in combination with gap and canopy conditions. The percentage of emerging seedlings tended to be lower on the ridge irrespective of the adult topographical distribution patterns. There was no clear trend in seedling emergence among the species groups classified by their spatial distribution. Seedling survival during 2 growing seasons was significantly different among species, topographic positions and light conditions. On the ridge, seedlings of the species dominating ridge tops had greater survival than those of other species, probably due to differences in demand for soil moisture. On the slope, frequent physical damage caused by surface material movement was observed and some species showed greater adaptability to the disturbed slope habitat. Survival of all seedlings was highest in the valley plots. Light conditions were the critical factor for seedling survival in some species. The results of this study suggest that topography creates diverse habitats for the establishment of tree seedlings. In addition to soil moisture, surface material movement may be a significant factor affecting seedling establishment.     

Size-dependency in hydraulic and photosynthetic properties of three <Emphasis Type="Italic">Acer</Emphasis> species having different maximum sizes

Recent studies suggest that physiological traits can be affected by tree size due to stronger hydraulic limitation in taller trees. As trees vary greatly in size, both within and among species, the adaptive responses to hydraulic limitation may be different among species with different maximum sizes. To investigate this, we explored size-dependency in photosynthetic and hydraulic parameters of three Acer species (Acer mono Maxim., Acer amoenum Carr and Acer japonicum Thunb.) using trees of various sizes under well-lit conditions. Leaf stomatal conductance of the Acer species decreased with tree size, implying that water supply to leaves decreases as trees grow. In contrast, content of nitrogen increased with tree size, which may compensate for the decrease in stomatal conductance to maintain the photosynthetic rate. Although the increase in nitrogen and leaf mass per area were larger in species with larger statures, the size-dependency in stomatal conductance was not different among species, and photosynthetic rate and hydraulic conductance were maintained in the three Acer species. Therefore, we suggest that hydraulic limitation on gas exchange does not necessarily depend on the maximum height of the species and that maintenance of photosynthesis and hydraulic properties is a fundamental physiological process during tree growth.     

Vertical vegetation zones along 30° N latitude in humid East Asia

Structural changes in altitudinal vegetation zones along a 30° N parallel were studied based on vegetation data from 20 mountains in East Asia, from 85° E to 130° E longitude. The altitude of comparable vegetation zones showed a sharp increase of 1400–1900 m from east to west. Forest limit reached an altitude of 4400–4600 m in the eastern Tibetan Plateau, being the highest forest limit in the world. The limidng factor for the upper limit of a vegetation zone was different in the east and west. Low temperature in winter controlled the upward distribution of the evergreen broadleaf forest in the east, whereas the limiting factor was growing season warmth in the west. A close correlation was found between the climatic indices and annual range of monthly mean temperature (ART) at the upper limit of a vegetation zone.Component genera of each vegetation zone along the 30° N parallel were analyzed, and it was found that the alternation of component genera from east to west was much more apparent in cool-temperate forests, reflecting their response to macrotopography and air masses. The distribution of Fagus extended into winter-cold regions, whilst Tsuga occurred principally in oceanic and warm climates. The northern limit of Tsuga corresponded well to an ART isotherm of 23 °C and its southern limit coincided with that of Fagus. According to the distribution of Fagus and Tsuga, the cool-temperate forests in East Asia along the 30° N belt were divided into three types: deciduous broadleaf forest (represented by Fagus), mixed forest (dominated by Fagus, Tsuga and others), and mixed evergreen forest (consisting mainly of Tsuga and sclerophyll oaks).