亚热带常绿阔叶林和暖温带落叶阔叶林叶片热值比较研究

植物干重热值(GCV)是衡量植物生命活动及组成成分的重要指标之一,反映了植物光合作用中固定太阳辐射的能力。利用氧弹量热仪测定了亚热带和暖温带两个典型森林生态系统常见的276种常见植物叶片的干重热值,探讨了亚热带和暖温带植物热值分布特征,以及不同生活型、乔木类型间植物热值的变化规律。实验结果发现:亚热带常绿阔叶林和暖温带落叶阔叶林叶片热值的平均值分别为17.83 k J/g(n=191)和17.21k J/g(n=85),整体表现为亚热带植物暖温带植物。不同地带性植被的植物叶片热值在不同生活型间表现出相似的规律,其中亚热带常绿阔叶林表现为:乔木(19.09 k J/g)灌木(17.87 k J/g)草本(16.65 k J/g);暖温带落叶阔叶林表现为:乔木(18.41 k J/g)灌木(17.94 k J/g)草本(16.53 k J/g);不同乔木类型间均呈现常绿乔木落叶乔木、针叶乔木阔叶乔木的趋势。落叶阔叶乔木表现为亚热带暖温带,而常绿针叶乔木则呈现亚热带暖温带的趋势。此外,我们对于两个分布区域内的4种针叶树种叶片热值进行了比较,发现华北落叶松(19.32 k J/g,暖温带)杉木(19.40 k J/g,亚热带)马尾松(19.82 k J/g,亚热带)油松(20.95 k J/g,暖温带)。亚热带常绿阔叶林和暖温带落叶阔叶林植物热值的特征及其变化规律,为森林生态系统的能量流动提供了理论基础。     

Stem growth and canopy dynamics in a world-wide range of Fagus forests

Abstract. Forests dominated by Fagus (beech) occur widely in the Northern Hemisphere. Tree species dominant together with beech vary in tolerance of understorey conditions. They are deciduous broad-leaved, evergreen broad-leaved or evergreen coniferous. The frequency and intensity of events that reduce the forest canopy cover are important determinants of the ratio of beech to other species in the canopy. For trees in the understorey and the canopy, stem diameter growth rate is determined by light regime and growing space which in turn are determined by canopy cover. We evaluated increase in stem diameter growth rate as an indicator of sudden reductions in canopy cover and canopy dynamics. We used tree-ring chronologies and calculated an index of growth rate increase (GI) to compare the canopy dynamics of 11 natural beech forests. Per site, the annual average value of GI poorly reflected the effects of dry or cool summers, and it clearly reflected events like tornados and hurricanes that removed substantial canopy cover. Among groups of sites average values of GI were significantly different. In the sites with a lower level of average GI, the establishment of the more shade tolerant tree and shrub species in the understorey was favoured, and subcanopy layers became more dense. On the other hand, higher levels of average GI allowed for more light demanding tree species to reach the canopy.     

中国4种典型森林中常见乔木叶片的非结构性碳水化合物研究

植物叶片的非结构性碳水化合物(non-structural carbohydrates,NSC)不仅为植物的代谢过程提供重要能量,还能一定程度上反映植物对外界环境的适应策略。以温带针阔混交林(长白山)、温带阔叶林(东灵山)、亚热带常绿阔叶林(神农架)和热带雨林(尖峰岭)4种森林类型的树种为研究对象,利用蒽酮比色法测定了163种常见乔木叶片可溶性糖、淀粉和NSC(可溶性糖+淀粉)含量,探讨了不同森林类型植物叶片NSC的差异及其地带性变化规律。结果显示：（1）从森林类型上看,植物叶片NSC含量从北到南递减,即温带针阔混交林(170.79 mg/g)>温带阔叶林(100.27 mg/g)>亚热带常绿阔叶林(91.24 mg/g)>热带雨林(80.13 mg/g)。（2）从生活型上看,无论是落叶树还是阔叶树,其叶片可溶性糖、淀粉和NSC含量均表现为：温带针阔混交林>温带阔叶林>亚热带常绿阔叶林>热带雨林;北方森林叶片可溶性糖、淀粉和NSC含量均表现为落叶树种>常绿树种,或阔叶树种>针叶树种。（3）森林植物叶片NSC含量、可溶性糖与淀粉含量比值与年均温和年均降水量均呈显著负相关。研究表明,森林植物叶片可溶性糖、淀粉和NSC含量以及可溶性糖与淀粉含量比值均具有明显的从北到南递减的地带性规律;其NSC含量以及可溶性糖与淀粉含量比值与温度和水分均呈显著负相关的变化规律可能是植物对外界环境适应的重要机制之一。该研究结果不仅为阐明中国主要森林树种碳代谢和生长适应对策提供了数据基础,而且为理解区域尺度森林植被对未来气候变化的响应机理提供新的视角。     

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).     

Pattern of changes in species diversity,structure and dynamics of forest ecosystems along latitudinal gradients in East Asia

We examined effects of seasonality of climate and dominant life form (evergreen/deciduous, broad-leaf/coniferous) together with energy condition on species diversity, forest structure, forest dynamics, and productivity of forest ecosystems by comparing the patterns of changes in these ecosystem attributes along altitudinal gradients in tropical regions without seasonality and along a latitudinal gradient from tropical to temperate regions in humid East Asia. We used warmth index (temperature sum during growing season, WI) as an index of energy condition common to both altitudinal and latitudinal gradients. There were apparent differences in patterns of changes in the ecosystem attributes in relation to WI among four forest formations that were classified according to dominant life form and climatic zone (tropical/temperate). Many of the ecosystem attributes—Fishers alpha of species-diversity indices, maximum tree height and stem density, productivity [increment rate of aboveground biomass (AGB)], and population and biomass turnover rates—changed sharply with WI in tropical and temperate evergreen broad-leaved forests, but did not change linearly or changed only loosely with WI in temperate deciduous broad-leaved and evergreen coniferous forests. Values of these ecosystem attributes in temperate deciduous broad-leaved and evergreen coniferous forests were higher (stem density was lower) than those in tropical and temperate evergreen broad-leaved forests under colder conditions (WI below 100°C). Present results indicate that seasonality of climate and resultant change in dominant life form work to buffer the effects of energy reduction on ecosystem attributes along latitudinal gradients.     

Contrasting patterns of phylogenetic diversity across climatic zones of Western Ghats: A biodiversity hotspot in peninsular India

This study attempts to understand the biogeographic history of the Western Ghats forests by investigating decoupling between phylogenetic and taxonomic diversity. We specifically test whether the deciduous forests have been recently established, whether the southern region was a refuge, and whether the deciduous and evergreen forest species have disparate evolutionary histories. We used species composition data from 23 forest types along the Western Ghats for all woody angiosperms above 10‐cm diameter at breast height. Forests were broadly grouped as either evergreen or deciduous. Mean phylogenetic distances corrected for species richness and mean phylogenetic beta diversity corrected for shared species were assessed using z‐scores from null distributions. Null distributions were generated by randomizing the species relationships on the phylogeny. We found that all evergreen forests showed a greater phylogenetic diversity as compared with null expectations. Deciduous forests showed the inverse pattern. Within the evergreen belt, there was a decreasing phylogenetic diversity from south to north, as predicted by the southern refuge hypothesis. The phylogenetic beta diversity across evergreen–deciduous forests was lesser than the null expectation, whereas it was much higher across forests within the evergreen belt. This study provides the first phylogenetic evidence for the antiquity of evergreen forests as well as the southern refuge hypothesis in the Western Ghats. The deciduous forests species have shared evolutionary histories with the evergreen forest species, suggesting multiple shifts between evergreen and deciduous states through the lineages. Conversely, the evergreen species exhibited a disparate evolutionary history across these forests, possibly owing to sharper ecological or climatic gradients.     

Variation of δ<Superscript>13</Superscript>C of wood and foliage with canopy height differs between evergreen and deciduous species in a temperate forest

We investigated post-photosynthetic fractionation and carbon transfer mechanisms of different plant functional types growing under the same climatic conditions in North-eastern China. The variations in δ¹³C of trunk and branches were compared with leaf δ¹³C at different canopy heights of Pinus koraiensis (evergreen coniferous species), Larix gmelinii (deciduous coniferous species) and Quercus mongolica (deciduous broad-leaved species). Results showed that δ¹³C of leaves increased (became more enriched) with increasing canopy height for both coniferous species (P. koraiensis, L. gmelinii) but not for Q. mongolica (a deciduous broad-leaved species). δ¹³C of both trunk and branches also increased with sampling height for the evergreen conifer P. koraiensis but did not significantly vary for either of the deciduous species (L. gmelinii or Q. mongolica), except a significant increase in branch δ¹³C for L. gmelinii. Similarly, δ¹³C of trunk and branches were strongly correlated with leaf δ¹³C only in the evergreen conifer, P. koraiensis. ¹³C was consistently more enriched in trunk, branches, and roots compared to leaves in all three species. Our findings suggest that, even under the same climatic conditions, different plant functional types may exhibit different carbon transfer mechanisms. This is contrary to the previous hypothesis that different carbon transfer mechanisms operate in forests of different climatic zones, especially in tropical and temperate forests. Particularly, the differences occur predominantly between evergreen and deciduous trees rather than between coniferous and broad-leaved trees. The significant difference in δ¹³C between leaves and wood tissues confirms a previous post-photosynthetic isotope fractionation in temperate forests.     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

楚雄花椒园半湿润常绿阔叶林的群落生态学特征

基于20个样方(各样方面积为25 m× 20 m,总面积1 hm2)调查,研究了楚雄花椒园自然保护区元江栲-米饭花林的群落生态学特征.该群落以壳斗科、越桔科、山茶科八角科、山矾科植物为乔木层主要成分,并以壳斗科植物占绝对优势,群落的物种多样性不高.木本植物以非全缘、革质、常绿、小叶等为其特征,反映该群落生境偏干,为典型...     

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。     

Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Climatic limits for the present distribution of beech (Fagus L.) species in the world

Aim Beech (Fagus L., Fagaceae) species are representative trees of temperate deciduous broadleaf forests in the Northern Hemisphere. We focus on the distributional limits of beech species, in particular on identifying climatic factors associated with their present range limits. Location Beech species occur in East Asia, Europe and West Asia, and North America. We collated information on both the southern and northern range limits and the lower and upper elevational limits for beech species in each region. Methods In total, 292 lower/southern limit and 310 upper/northern limit sites with available climatic data for all 11 extant beech species were collected by reviewing the literature, and 13 climatic variables were estimated for each site from climate normals at nearby stations. We used principal components analysis (PCA) to detect climatic variables most strongly associated with the distribution of beech species and to compare the climatic spaces for the different beech species. Results Statistics for thermal and moisture climatic conditions at the lower/southern and upper/northern limits of all world beech species are presented. The first two PCA components accounted for 70% and 68% of the overall variance in lower/southern and upper/northern range limits, respectively. The first PCA axis represented a thermal gradient, and the second a moisture gradient associated with the world‐wide distribution pattern of beech species. Among thermal variables, growing season warmth was most important for beech distribution, but winter low temperature (coldness and mean temperature for the coldest month) and climatic continentality were also coupled with beech occurrence. The moisture gradient, indicated by precipitation and moisture indices, showed regional differences. American beech had the widest thermal range, Japanese beeches the most narrow; European beeches occurred in the driest climate, Japanese beeches the most humid. Climatic spaces for Chinese beech species were between those of American and European species. Main conclusions The distributional limits of beech species were primarily associated with thermal factors, but moisture regime also played a role. There were some regional differences in the climatic correlates of distribution. The growing season temperature regime was most important in explaining distribution of Chinese beeches, whilst their northward distribution was mainly limited by shortage of precipitation. In Japan, distribution limits of beech species were correlated with summer temperature, but the local dominance of beech was likely to be dependent on snowfall and winter low temperature. High summer temperature was probably a limiting factor for southward extension of American beech, while growing season warmth seemed critical for its northward distribution. Although the present distribution of beech species corresponded well to the contemporary climate in most areas, climatic factors could not account for some distributions, e. g., that of F. mexicana compared to its close relative F. grandifolia. It is likely that historical factors play a secondary role in determining the present distribution of beech species. The lack of F. grandifolia on the island of Newfoundland, Canada, may be due to inadequate growing season warmth. Similarly, the northerly distribution of beech in Britain has not reached its potential limit, perhaps due to insufficient time since deglaciation to expand its range.     

南京灵谷寺森林50年来的动态变化研究

 为进一步揭示北亚热带森林次生演替规律,于2002年5月在定点样带上沿用前人的方法,对南京灵谷寺森林进行了群落调查。通过比较1951、1981、2002年的定点样带资料,对灵谷寺森林的物种组成、多度、频度、重要值、种群结构以及物种多样性的动态变化进行了研究。50多年来,群落中立木株数相对基本稳定,但空间分布异质性逐渐增加,而林下苗木和灌木个体数波动幅度极大,由1951年的4 712株大幅上升至1981年的44 130株,到2002年又回落至7 372株。群落中物种数和建群种构成等都存在明显变化,物种数由1951年的75种下降到1981年的50种,2002年又上升为73种。物种多样性指数也存在波动,但目前尚未发现物种多样性指数有明显的上升或下降趋势。历经50多年的次生演替,早期人工针叶林中的马尾松（Pinus massoniana）逐渐衰退,首先被阳性落叶阔叶树,如黄连木（Pistacia chinensis）、枫香（Liquidambar formosana）等取代,之后又被相对耐荫的栓皮栎（Quercus variabilis）等树种所替代,一些常绿树种成功入侵,人工针叶林经针阔混交林、落叶阔叶林,逐渐向地带性植被落叶 常绿阔叶混交林方向演替,其中立地条件较好地段的群落演替速度也较快。     

Impacts of land-use history on the diversity of a riparian forest landscape in warm-temperate Kyushu,southern Japan

We examined the impacts of land-use history on the species composition and diversity of a warm-temperate riparian forest landscape in Kyushu, southern Japan, focusing on the relationship between evergreen oaks and deciduous trees in natural and seminatural forests. The species composition of 59 plots was classified into four types (A to D). Type A, which showed a significant bias towards sites not subject to nonforest land use since 1947, had high species diversity consisting of (1) many lucidophyllous components of the region, including the rare indigenous oak Quercus hondae, and (2) summergreen tree species of varying dominance and number representing unique or locally rare elements of the riparian landscape in this warm-temperate region. Type B was dominated by a common species of oak, Q. glauca, and displayed less clear distribution bias with land-use history. In contrast to types A and B, types C and D, which were characterized by high dominance of deciduous trees, had negative bias away from sites that had been under forest land use in 1947. Presumably, intensive anthropogenic disturbances associated with nonforest land uses had expanded the habitats for deciduous trees. This phenomenon was represented by the establishment of forests (type D) dominated by Ulmus davidiana var. japonica (UDJ) after it had been released from the suppression of evergreen forest trees during a period of nonforest land use that prevents the successful recovery of evergreen trees. From these results we conclude that the impacts of land-use history on the diversity of warm-temperate riparian forest landscape are multiphased: a period of nonforest land use has a strong negative impact on lucidophyllous forest trees represented by the rare indigenous oak Q. hondae; release from the suppressive effects of the lucidophyllous species then encourages establishment of locally rare deciduous tree flora represented by UDJ, which continue to persist for decades after abandonment of nonforest land use.     

Zonal transition of evergreen,deciduous, and coniferous forests along the altitudinal gradient on a humid subtropical mountain,Mt. Emei,Sichuan, China

Altitudinal zonation of evergreen, deciduous and coniferous forests on Mt. Emei (3099 m asl, 29°34.5' N, 103°21.5' E), Sichuan, China was studied to understand the transition of vegetation zonation from tropical to temperate mountains in humid Asia. On the basis of quantitative data on floristic composition and community structure sampled at ten plots selected in different altitudes on the eastern slope of the mountain, forest zonation and the inter-relationships among different life-forms of trees in each zonal forest community were studied quantitatively. Three forest zones were identified physiognomically along the altitudinal gradient, viz. (i) the evergreen broad-leaved forest zone (660–1500 m asl), (ii) the mixed forest zone (1500–2500 m asl), and (iii) the coniferous forest zone (2500–3099 m asl). Great compositional changes were observed along elevation, and the zonal forest communities were characterized by their dominants and floristic composition. Maximum tree height decreased from 33 m at lower middle altitude (965 m asl) to 13 m near the summit (2945 m asl). There was no apparent deciduous forest zone along the altitudinal gradient, but true mixed forests of three life-forms (evergreen, deciduous, and coniferous) were formed around 2000–2500 m asl. Patches of deciduous forest were found in a lower part of the mixed forest zone, particularly on scree slopes, between 1450 m and 1900 m asl. These patches were dominated by the Tertiary relic deciduous trees, such as Davidia involucrata, Tetracentron sinense, and Cercidiphyllum japonicum var. sinense. High species diversity in the mixed forest zone resulted from the overlapping of different life-forms at middle altitudes, which is partly due to wider variety of temperature-altitude correlations. A comparison of the altitudinal zonation with the other east Asian mountain vegetation clarified that Mt. Emei is located exactly at the ecotone between tropical and temperate zonation types in eastern Asia.     

茂兰喀斯特常绿落叶阔叶混交林群落组成和优势种更新类型

为探讨茂兰喀斯特森林群落结构和优势种更新机制,对茂兰自然保护区常绿落叶阔叶混交林群落进行了调查。结果表明,群落中有木本植物37科58属70种,以樟科、蔷薇科、无患子科和漆树科为主。常绿树种41种,占总种数的58.6%;落叶树种29种,占总种数的41.4%,这体现了喀斯特常绿落叶阔叶混交林的群落学性质。群落的优势乔木树种有轮叶木姜子(Litseaverticillata)、青冈栎(Cyclobalanopsisglauca)、狭叶润楠(Machilusrehderi)和翅荚香槐(Cladrastisplatycarpa)等。灌木优势种有皱叶海桐(Pittosporumcrispulum)、裂果卫矛(Euonymusdielsianus)、革叶铁榄(Sinosideroxylonwightianum)和南天竹(Nandina domestica)等。优势种种群结构有5种类型:单峰型,如樱桃(Cerasus pseudocerasus)、南酸枣(Choerospondias axillaris)等阳性乔木,为林窗更新种,随林窗的郁闭将衰退消失;逆J字型,如轮叶木姜子、香港四照花(Dendrobenthamia hongkongensis)、皱叶海桐等耐荫性强的常绿阔叶树种,更新能力较强,为顶极群落优势种;间歇型,如青冈栎、狭叶润楠、齿叶黄皮(Clausena dunniana)等树种,群落中的个体多集中在小径级阶段,能长期存在于群落中;L字型,如桂皮(Cinnamomumtamala)、梓叶槭(Acer catalpifolium)、革叶铁榄等;单柱型,如石岩枫(Mallotus repandus)、南天竹等常绿灌木树种,个体集中于幼苗、幼树阶段,为顶极群落亚乔木层和灌木层的主要组成种。因此,这表明调查区域的植物群落处于相对稳定的顶极阶段。     

Distribution and structural-functional attributes of trees in the high altitude zone of Central Himalaya,India

Patterns of tree species distribution and their structural-functional features were studied along an altitudinal gradient in the Indian Central Himalaya. The forest vegetation falls within three formation types: low to mid-montane sclerophyllous, broad-leaved forests; mid-montane deciduous forests; and high-montane mixed stunted forests. Deciduous tree species formed a considerable (49%) portion of the high altitude forests between 1500–3300 m. The upper limit of forests ranges from 3000 to 3300 m, and three taxa are commonly found at tree line: an evergreen, needle-leaved gymnosperm (Abies pindrow Spach.), an evergreen, broad-leaved sclerophyllous oak (Quercus semecarpifolia Sm.), and a deciduous, broad-leaved birch (Betula utilis Don).The high altitude forests differed from the low altitude forests of Central Himalaya in their greater proportion of microphanerophytes. Vernal flowering is common at this elevation in Central Himalaya. In general, in their elevational distribution and structural-functional attributes, these high altitude forests are similar to other forests of cool temperate zones in Himalaya.     

哀牢山中山湿性常绿阔叶林木径向生长季节动态及其对气候因子的响应

研究采用树木生长环在哀牢山中山湿性常绿阔叶林持续9年(2009—2017年)监测了2个常绿树种(厚皮香,Ternstroemia gymnanthera;南亚枇杷,Eriobotrya bengalensis)和2个落叶树种(西桦,Betula alnoides;珍珠花,Lyonia ovalifolia)的树干月生长量,采用逻辑斯蒂生长模型(Logistic model)模拟树木径向生长量和物候参数,并分析了年、季尺度上径向生长与主要气候因子的关系。结果表明:1)4个树种年平均生长量为6.3 mm,落叶树种年平均生长量(10.6 mm/a)显著高于常绿树种(3.0 mm/a);2)雨季(5—10月)是哀牢山中山湿性常绿阔叶林树木生长的主要时期,4个树种雨季平均生长量为5.9 mm,占全年总生长量的93%,其中落叶树种雨季生长量占全年的96%,而常绿树种雨季生长量占全年的86%;3)常绿树种生长季长度为169天,长于落叶树种(137天),而落叶树种最大生长速率(0.14 mm/d)显著高于常绿树种(0.03 mm/d),最大径向生长速率能很好地预测树种年生长量;4)低温、雾日和光合有效辐射是影响哀牢山亚热带常绿阔叶林4个研究树种径向生长的重要环境因子,其中温度对常绿树种径向生长具有显著影响,而雨日、雾日与空气湿度等水分因子对落叶树种径向生长更为重要。常绿树种年生长量对旱季气候因子的响应相比落叶树种更为敏感,树木旱季生长量除了受低温限制外,也受到水分供给的影响。气候变化可能改变不同物候类型树种在哀牢山中山湿性常绿阔叶林中的生长状态与分布格局。     

Comparison of climatic effects on radial growth of evergreen broad-leaved trees at their northern distribution limit and co-dominating deciduous broad-leaved trees and evergreen conifers

Using dendrochronological techniques, this study examined whether tree-ring width of two evergreen broad-leaved species (Cleyera japonica, Eurya japonica) at their inland northern distribution limit in central Japan is more limited by low temperature compared with two co-dominating deciduous broad-leaved species (Fagus japonica, Magnolia hypoleuca) and two evergreen conifer species (Chamaecyparis obtusa, Abies firma), whose distribution limits are further north. The two deciduous broad-leaved species and the two evergreen conifers are tall tree species. Evergreen broad-leaved Cleyera japonica is a sub-canopy species and Eurya japonica is a small tree species. The tree-ring widths of four of the six species (except for Eurya japonica and Magnolia hypoleuca) correlated positively with the March temperature just before the start of the growth period. For deciduous broad-leaved Magnolia hypoleuca, the tree-ring width was correlated positively and negatively with July temperature and precipitation, respectively. However, the other deciduous broad-leaved Fagus japonica showed no such relationships. For the evergreen broad-leaved Cleyera japonica and evergreen conifers Chamaecyparis obtusa and Abies firma, tree-ring widths correlated positively with winter temperatures, probably because evergreen species can assimilate during warm winters. The tree-ring width of Cleyera japonica also correlated positively with temperatures of many months of the growth period. By contrast, the tree-ring width of the other evergreen broad-leaved Eurya japonica showed no positive correlation with the temperature in any month. Most Eurya japonica trees were suppressed by tall trees, which might disguise any climate effect. Thus, there were species differences in response to climate for each life form, and the tree-ring width of Cleyera japonica at the northern distribution limit was more limited by low temperatures compared with co-dominating species. It is suggested that growth of Cleyera japonica is increased by global warming at the latitudinal ecotone.     

Gradational Forest Change along the Climatically Dry Valley Slopes of Bhutan in the Midst of Humid Eastern Himalaya

Altitudinal forest and climate changes from warm, dry valley bottom (1250 m a.s.l.) to cool, humid ridge top (3550 m a.s.l.) along the typical dry valley slopes of the Bhutan Himalaya were studied. Annual mean temperature decreased upslope with a lapse rate of 0.62 °C·100 m⁻¹ from 18.2 °C at the valley bottom to 4.3 °C at the ridge top. On the contrary volumetric soil moisture content increased from 14.7 to 75.0%. This inverse relationship is the major determinant factor for the distribution of different forest types along the altitudinal gradient. Based on the quantitative vegetation data from 15 plots arranged ca. 200 m in altitude interval (1520–3370 m a.s.l.), a total of 83 tree species belonging to 35 families were recorded. Three major formation types of lower and upper coniferous forests, and a mid-altitude evergreen and deciduous broad-leaved forest were contrasted. Including two transitional types, five forest zones were categorized based on cluster analysis, and each zone can be characterized by the dominants and their phytogeographical traits, viz. (1) west Himalayan warm, dry pine (1520–1760 m a.s.l.), (2) wide ranging east-west Himalayan mixed broad-leaved (1860–2540 m a.s.l.), (3) humid east Himalayan evergreen broad-leaved (2640–2820 m a.s.l.), (4) cool, humid east Himalayan conifer (2950–3210 m a.s.l.), and (5) wide ranging cold, humid conifer (3305–3370 m a.s.l.). Structurally, total basal area (biomass) increased from 15.2 m² ha⁻¹ in the pine forest (1520 m) to 101.7 m² ha⁻¹, in the conifer forest (3370 m a.s.l.). Similarly, soil organic carbon increased from 2.7 to 11.3% and nitrogen from 0.2 to 1.9% indicating dry, poor nutrient fragile ecosystem at the dry valley bottom. We concluded that low soil moisture content (<20%) limits downslope extension of broad-leaved species below 1650 m a.s.l. while coldest month’s mean temperature of −1 °C restricted the upslope extension of evergreen broad-leaved species above 3000 m a.s.l. Along the dry valley slopes, the transition from dry pine forest in the valley bottom, to a mixture of dry west Himalayan evergreen and deciduous east Himalayan broad-leaved, and to humid evergreen oak–laurel forests feature a unique pattern of forest type distribution.