论滇南西双版纳的森林植被分类

本文基于多年研究成果的总结,对西双版纳森林植被的分类、主要植被类型及其特征进行了系统归纳,并讨论了它们与世界类似热带森林植被的关系。以群落的生态外貌与结构、种类组成和生境特征相结合作为植被分类的原则和依据,可以将西双版纳的热带森林植被分类为热带雨林、热带季节性湿润林、热带季雨林和热带山地常绿阔叶林四个主要的植被型,包括有至少二十个群系。热带雨林包括热带季节雨林和热带山地(低山)雨林二个植被亚型。热带季节雨林具有与赤道低地热带雨林几乎一样的群落结构和生态外貌特征,是亚洲热带雨林的一个类型,但由于发生在季风热带北缘纬度和海拔的极限条件下,受到季节性干旱和热量不足的影响,在其林冠层中有一定比例的落叶树种存在,大高位芽植物和附生植物较逊色而藤本植物和在叶级谱上的小叶型植物更丰富,这些特征又有别于赤道低地的热带雨林。热带山地雨林是热带雨林的山地亚型,是该地区热带山地较湿润生境的一种森林类型,它在植物区系组成和生态外貌特征上类似于热带亚洲的低山雨林,隶属于广义热带雨林植被型下的低山雨林亚型。热带季节性湿润林分布在石灰岩山坡中、上部,在群落外貌上类似热带山地常绿阔叶林但在植物区系组成上与后者不同,它是石灰岩山地垂直带上的一种植被类型。热带季雨林是分布在该地区开阔河谷盆地及河岸受季风影响强烈的生境的一种热带落叶森林,是介于热带雨林与萨王纳之间的植被类型。热带山地常绿阔叶林(季风常绿阔叶林)是西双版纳的主要山地植被类型,它分布在热带季节雨林带之上偏干的山地生境。它在植物区系组成上不同于该地区的热带季节雨林,在生态外貌特征上亦不同于热带山地雨林,是发育在受地区性季风气候强烈影响的热带山地的一种森林植被类型。     

西双版纳森林植被研究

西双版纳是世界生物学多样性保护的关键和热点地区,倍受国际学术界的关注。笔者依据30多年来对西双版纳植被的调查,结合植物群落生态学与植物区系地理学研究,并参考世界类似热带植被的研究成果,对西双版纳植被的分类、物种组成、群落生态表现和植物区系特征等作了系统探讨,还进一步分析比较了其与世界类似热带森林植被的关系。结果显示,西双版纳的森林植被共包括32个较为典型的群系,且分属于7个主要的植被型,即热带雨林、热带季节性湿润林、热带季雨林、热带山地(低山)常绿阔叶林、热带棕榈林、暖热性针叶林和竹林。本文对西双版纳植被进行的全面记录和系统归纳,可为科学研究、生物多样性保护和自然保护区的管理提供参考。     

Geological History,Flora, and Vegetation of Xishuangbanna,Southern Yunnan,China1

Xishuangbanna of southern Yunnan is biogeographically located at a transitional zone from tropical Southeast (SE) Asia to subtropical East Asia, and is at the junction of the Indian and Burmese plates of Gondwana and the Eurasian plate of Laurasia. The flora of the region consists of a recorded 3336 native seed plant species, belonging to 1140 genera in 197 families, among which 83.5 percent are tropical genera and 32.8 percent are endemic to tropical Asia, suggesting a strong affinity to tropical Asian flora. The vegetation of Xishuangbanna is organized into four forest types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad‐leaved forest, and tropical monsoon forest. The tropical rain forest in Xishuangbanna has the same floristic composition of families and genera as some lowland equatorial rain forests in SE Asia, and is dominated (with a few exceptions) by the same families both in species richness and stem dominance. The exceptions include some deciduous trees in the canopy layer, fewer megaphanerophytes and epiphytes, and a higher abundance of lianas and microphyllic plants. We consider the tropical rain forest of Xishuangbanna as a type of tropical Asian rain forest, based on their conspicuous similarities in ecological and floristic characteristics.     

Species Composition,Physiognomy and Plant Diversity of the Tropical Montane Evergreen Broad-leaved Forest in Southern Yunnan

Species composition, physiognomy and plant diversity of the less known tropical montane forests in southern Yunnan were studied based on the data from 15 sampling plots in three sites. These forests are mainly dominated by the families Theaceae, Fagaceae, Lauraceae and Euphorbiaceae in floristic composition, and dominated by evergreen phanerophytes with mesophyllous leaves. These forests are similar to lower montane rain forests in equatorial southeastern Asia in floristic composition and altitudinal distributions, but differ in physiognomy by having few epiphytes, but more lianas and more plants with compound leaves. These differences could be due to strongly seasonal climate and so-called mass elevation effect in southern Yunnan. They also differ from the tropical seasonal rain forests at lower altitudes in southern Yunnan by having conspicuously lower species richness, few epiphytes, fewer mega-mesophanerophytes, more abundant micro-nanophanerophytes and hemicryptophytes and more plants with microphyllous leaves. It is suggested that these forests could be termed tropical montane evergreen broad-leaved forests, and be a vegetation type from the northern margin of mainland southeastern Asia controlled by a strongly seasonal climate.     

“西双版纳热带山地雨林的植物多样性研究”的一些问题讨论

该文针对“西双版纳热带山地雨林的植物多样性研究”论文中存在的一些问题进行了讨论。原文所依据的6个调查样地, 从其分布海拔、生境、群落的生态外貌特征、植物区系组成及单位面积植物种数的统计上反映出它们并非都属于同样的植被类型,即原文所称的热带山地雨林,而可能分别属于《云南植被》中所应用的热带季节雨林的次生林群落(样地I和II)、季风常绿阔叶林群落(样地V和VI)及类似于苔藓常绿阔叶林(样地III和IV)的群落类型。由于这些样地代表了不同的植被类型, 导致在对这些样地的植物多样性特征的比较上出现较大差异。对原文在资料分析和与其它森林群落植物多样性的比较上存在的一些问题也作了讨论。原文的研究结果显示了西双版纳不同海拔高度上森林植物群落的物种多样性特征,但并不能全部运用于该地区的热带山地雨林这一特定植被类型。     

滇南热带季雨林的一些问题讨论

通过分析云南南部的水、热条件及植被分布,讨论了季雨林植被类型及其特征,认为云南南部的季雨林是介于热带雨林与萨王纳之间的,在干季基本上是落叶的一种森林植被类型,符合Schimper (1903)定义的热带季雨林植被,并考虑它是一种生态学意义上的经向地带性植被,与该地区的纬向地带性植被热带季节雨林一起共同构成云南南部的水平地带性植被。在云南南部的石灰岩山坡分布的过去被认为是季雨林的森林植被,尽管也受到季节性干旱的影响而不同程度地具有落叶成分,但它在群落外貌上与典型的季雨林不相同,在植物区系组成上也明显不同于该地区非石灰岩山地的季风常绿阔叶林,在分布上亦是在石灰岩低山沟谷的热带季节性雨林水平地带性植被带之上,根据其生态外貌、植物区系组成和生境特点,我们建议用“热带季节性湿润林”来称呼这类石灰岩山地森林类型,在性质上属东南亚热带北缘石灰岩山地垂直带上的一种植被类型。     

大陆东南亚(中南半岛)的主要森林植被类型介绍

大陆东南亚(中南半岛)的植被研究情况鲜为人知,至今仍无系统研究资料。该文依据数次对该地区的野外考察和资料收集,介绍了东南亚植被的研究情况和文献资料以及对该地区主要森林植被的分类和各主要植被类型的特征。大陆东南亚地区在植被分类上包括七个主要的陆生及湿地的森林植被类型:针叶林、针阔混交林、热带山地常绿阔叶林、热带雨林、热带季节性湿润林、热带季风林(季雨林)、干旱刺灌丛/萨王纳植被。其中,针叶林植被型包括温性针叶林和热性针叶林二个植被亚型;针阔混交林包括温性针阔混交林和暖温性针阔混交林二个亚型;热带雨林植被型包括热带低地常绿雨林、热带季节性雨林(热带低地半常绿雨林)、热带山地雨林及泥炭沼泽森林四个植被亚型。该文还对大陆东南亚地区植被研究历史、植被分类系统、类型特征及植物区系组成进行了讨论。     

滇南勐宋热带山地雨林的物种多样性与生态学特征

 研究了鲜为人知的滇南勐宋地区的原始山地雨林植被, 根据分布生境、群落结构和种类组成特征,可将该山地雨林区分为沟谷和山坡两个类型, 分别定义为八蕊单室茱萸(Mastixia euonymoides)-大萼楠(Phoebe megacalyx)林和云南拟单性木兰(Parachmeria yunnanensis)-云南裸花(Gymnanthes remota)林。该山地雨林的外貌仍以单叶、革质、全缘、中叶为主的常绿中、小高位芽植物组成为特征,层间木质藤本植物仍较丰富,草本高位芽植物和附生植物丰富,但板根和茎花现象少见,属于热带山地垂直带上低山雨林或山地雨林植被类型。与该地区的典型热带季节雨林和赤道热带雨林相比, 勐宋的山地雨林群落中的大、中高位芽植物和藤本高位芽植物比例相对减少, 小、矮高位芽植物和草本高位芽植物比例相对增加,单叶、革质、非全缘叶和小叶比例相对增加,板根现象少见。与中国热带北缘-南亚热带地区（季风）常绿阔叶林比较, 勐宋的山地雨林有较多的附生植物和草本高位芽植物,相对较少的小高位芽植物和矮高位芽植物,小叶比例亦较少,非全缘叶和革质叶比例相对较低。故勐宋山地雨林是滇南热带北缘山地的一种较湿润生境的植被类型, 与所谓的季风常绿阔叶林不同。在物种多样性上,勐宋热带山地雨林在单位面积植物种数上并不比该地区的热带季节雨林低, 物种多样性指数与低丘季节雨林相当,比沟谷季节雨林低, 明显高于季风常绿阔叶林。     

云南热带雨林:特征、生物地理起源与演化北大核心CSCD

云南热带雨林具有与东南亚低地热带雨林类似的群落结构、生态外貌特征和物种多样性,是亚洲热带雨林的一个类型。它的植物区系组成中有90%的属和多于80%的种为热带分布成分,其中约40%的属和70%的种为热带亚洲分布型,它含属种较多的优势科和在群落中重要值较大的科也与亚洲热带雨林相似,是亚洲热带雨林和植物区系的热带北缘类型。云南西南部、南部与东南部的热带雨林在群落结构和生态外貌上类似,但在南部与东南部之间有明显的植物区系分异,它们经历了不同的起源背景和演化历程。云南的热带雨林在很大程度上由西南季风维持。喜马拉雅隆升导致西南季风气候形成和加强,在云南热带局部地区产生了湿润气候,发育了热带雨林植被。现在的云南热带雨林里或其分布地区有落叶物种或热带落叶林存在,这不仅是季节性气候的影响,推测在晚第三纪或第四纪更新世云南热带地区曾经历了干旱气候。云南热带雨林的分布主要受制于局部生境,并非地区性气候条件。     

Ecological and biogeographical studies on the tropical rain forest of south Yunnan,SW China with a special reference to its relation with rain forests of tropical Asia

Abstract. Ecological and biogeographic analyses of the tropical rain forest in south Yunnan were made using data from seventeen sample plots and floristic inventories of about 1000 species of seed plants. The rain forest is shown to be a type of true tropical rain forest because it has almost the same profile, physiognomic characteristics, species richness per unit area, numbers of individuals in each tree species and diameter classes of trees as classic lowland tropical rain forests. As the area is at the northern margin of monsoonal tropics, the rain forest differs from equatorial lowland rain forests in having some deciduous trees in the canopy layer, fewer megaphanaerophytes and epiphytes but more species of lianas as well as more species of microphylls. In its floristic composition, about 80% of total families. 94% of total genera and more than 90% of total species are tropical, of which about 38% of genera and 74% of species are tropical Asian. Furthermore, the rain forest has not only almost the same families and genera, but also the same families rank in the top ten both in species richness and in dominance of stems, as lowland forests in southeast Asia. It is indisputable that the flora of the rain forest is part of the tropical Asian flora. However, most of the tropical families and genera have their northern limits in south Yunnan and most have their centre of species diversity in Malesia. More strictly tropical families and genera have relatively lower species richness and importance compared with lowland rain forests in tropical southeast Asia. Thus, the flora also shows characteristics of being at the margin of the tropics. Based mainly on physiognomy and floristic composition the tropical rain forest of Yunnan is classified into two types, i.e. seasonal rain forest and wet seasonal rain forest, the latter is further divided into two subtypes, i.e. mixed rain forest and dipterocarp rain forest. From analysis of geographic elements it is also shown that the tropical rain forest of Yunnan occurs at a geographical nexus with its flora coming mainly from four sources, i.e. Malesia, south Himalayas, Indochina and China.     

滇东南马关古林箐热带雨林望天树群落的研究

云南东南部马关县古林箐的望天树群落以龙脑香科植物望天树为乔木层优势种,无患子科植物番龙眼为亚优势种,外貌以常绿大、中高位芽植物组成为特征,林内板根和茎花现象普遍,层间木质藤本和维管附生植物丰富,属于一种热带季节性雨林群落类型。在植物区系组成上,该群落以樟科、番荔枝科、楝科、大戟科、桑科、无患子科、橄榄科、柿树科等为主要组成科,在重要值统计上,以大戟科、无患子科、柿树科、龙脑香科、番荔枝科、樟科、楝科、橄榄科、桑科等为较优势的科,与东南亚热带雨林很接近,故为东南亚热带雨林的北缘类型。     

Structural comparison of tropical montane rain forests along latitudinal and altitudinal gradients in south and east Asia

Geographical patterns of altitudinal zonation, floristic composition, and structural features of tropical montane rain forests were examined along latitudinal gradients in south and east Asia. On equatorial mountains, the tropical montane rain forests occur above 1000 m. Toward middle latitudes, they come farther down and reach sea level at c. 35° N. Thus, the forests are equivalent to the subtropical rain forests of the latitudinal, horizontal zonation series. They exhibit gradual changes in floristic composition and structure along both altitudinal and latitudinal gradients. On equatorial mountains, they are divided into three types, i.e. tropical lower montane, upper montane, and subalpine forests. The three tree regeneration types, having emergent, sporadic and inverse-J type stem-diameter class frequency distributions, coexist in the lower montane forests, but the upper and subalpine forests display only the inverse-J type species with a few species of the sporadic type. Toward the northern latitudinal limit, the distinction between the three tropical montane forest zones in equatorial mountains becomes less clear. This can be explained by temperature conditions: on equatorial mountains, a temperature sum of 85° C months which controls the upper limit of the lower montane forests, and a coldest month mean temperature of-1° C which controls the evergreen broad-leaved trees, appear at c. 2500 and c. 4000 m respectively. The altitudinal range between 2500 m and 3800 m, which is the upper forest limit, is covered by upper montane and subalpine forests. On the other hand, at the latitudinal northern limit, the tropical upper montane and subalpine forests cannot exist because the above mentioned two temperature conditions occur at nearly the same point. Thus, at the northern latitudinal limit of the tropical montane forests, the three zones of equatorial mountains amalgamate into a single subtropical lowland forest community. This is due to the seasonal temperature climate in middle latitudes in, e.g., central Japan and central China.A part of this paper was presented as an oral presentation at the Vth International Congress of Ecology, Yokohama 23–30.8.1990.     

西双版纳龙脑香林与热带亚洲和中国热带北缘地区植物区系的关系

西双版纳龙脑香林只有与热带亚洲的龙脑香林完全一致的植物科的组成,绝大多数的属也与后者共有,特别是在乔木层的区系组成和数量特征上,二者很类似,表明西双版纳龙脑香林在植物区系性质上归属于热带亚洲龙脑香林,但为后者的一种北缘类型。与中国热带边缘地区的其它一些森林类型相比,西双版纳龙脑香林亦具有与它们基本一致的科的组成,但无论按在群落中的种数,按占科的世界区系百分比,还是按重要值指数,西双版纳龙脑香林中热带科均占有显著地位,并且包含有较高比例的热带亚洲分布属和种,故可认为它是中国热带北缘地区的森林类型中热带性强,在区系组成上接近热带亚洲龙脑香林而作为后者的一个部分的森林类型。     

西双版纳热带雨林与海南热带雨林的比较研究

西双版纳的热带雨林与海南低地热带雨林和热带季雨林有基本一致的植物区系组成,群落中优势科无论在种数百分比还是重要值排名上均较接近,显然属于同样性质的植物区系。在生态特征上,西双版纳热带雨林群落高大,分层不明显,B层为林冠层,散生巨树常见,在生活型谱上以高位芽植物占绝对优势,大、中高位芽植物相对较多,落叶树种比例小;以中叶、纸质,全缘和复叶比例较高为特征,具有最接近海南低地湿润雨林的群落垂直结构和生态外貌,其雨林特点虽不如湿润雨林浓厚,但明显强于海南的热带常绿季雨林和山地雨林,海南常绿季雨林群落高度明显较矮,小叶比例通常较高,革质叶比例亦较高,群落具遥明显的旱生特点,海南的山地雨林群落高度较矮,A层连续,成为林冠,无散生巨树,分层明显,在生活型谱上大高位芽植物比例减少,附生植物丰富,并具有相当比例的地面芽植物;叶级虽以中叶占优势,但通常革质,非全缘和单叶比例较高,明显由于热量不足的影响而带有亚热带森林特色,在物种多样性上,西双版纳热带雨林的乔木物种多样性指数似乎与海南的低地热带雨林相当,低于海南的山地雨林群落,海南的热带雨林群落种类丰富度不同人研究的结果差异较大,如果这些用于比较的数据可靠和具有可比性的话,西双版纳热带雨林的物种多样性要比海南的山地雨林低。     

云南热带季雨林及其与热带雨林植被的比较

在中国植物学文献中, 对热带季雨林的解释和运用是不一致的, 特别是易于把季雨林与热带雨林相混淆。季雨林是在具有明显干、湿季变化的热带季风气候下发育的一种热带落叶森林植被, 是介于热带雨林与热带稀树草原(savanna)之间的一个植被类型。云南的热带季雨林在分布生境、生态外貌特征、植物种类组成和地理成分构成上, 均与热带雨林有明显区别, 季雨林主要分布在海拔1 000 m以下的几大河流开阔河段两岸和河谷盆地, 其群落结构相对简单, 乔木一般仅有1至2层, 上层树种在干季落叶或上层及下层树种在干季都落叶; 在生活型组成上, 季雨林的木质藤本相对较少, 大高位芽植物及地上芽植物很少, 但地面芽植物很丰富, 地下芽植物和一年生植物也相对丰富; 在叶级和叶型特征上, 季雨林植物的小叶和复叶比例相对较高, 分别占到24%和44%; 在植物区系地理成分构成上, 季雨林的热带分布属合计也占绝对优势, 但以泛热带分布属的比例相对较高, 约占到总属数的30%, 热带亚洲至热带非洲分布属的比例也较高, 约占总属数的12%。季雨林的地理成分更为多样性, 起源与发展历史也更复杂和古老。     

云南常绿阔叶林的植被地理研究

云南具有极其丰富的生物多样性和以常绿阔叶林为优势的植被类型。该研究利用6个基于样方层面的1 hm²样地资料, 以及通过对整个植被类型的植物区系的调查, 对云南常绿阔叶林植被型的3个植被亚型(季风常绿阔叶林、半湿润常绿阔叶林和中山湿性常绿阔叶林)的生态外貌特征、植物区系组成及其生物地理演化进行了研究。在样方层面, 尽管这3个常绿阔叶林在树种组成上优势种均为壳斗科、樟科和山茶科植物, 但它们在种类组成、多样性、生态外貌和生物地理特征上呈现多样化。分布在南部及西南部的季风常绿阔叶林物种组成极其丰富, 具有热带森林的生态外貌, 并以热带亚洲分布种为优势种。主要分布在云南高原的半湿润常绿阔叶林和云南中部和北部山地的中山湿性常绿阔叶林具有亚热带常绿阔叶林的生态外貌特征和以中国-喜马拉雅及中国特有种占优势, 是中国西南独特的植被类型。在植被亚型层面, 这3个常绿阔叶林的植物区系(包括所有生活型的种子植物)中种数最多的科, 按地理成分均为世界分布型的科, 含种数较少的科则为其他各种分布型的科。半湿润常绿阔叶林和中山湿性常绿阔叶林的植物区系, 热带分布属分别占总属数的44.91%和44.04%, 温带分布属占46.29%和48.19%, 其中北温带分布属比例最高, 分别为18.36%和19.95%。季风常绿阔叶林植物区系则显示了不同的地理成分格局: 热带分布属占总属数的78.05%, 并以热带亚洲分布属占最高比例。通过对这3个常绿阔叶林的比较发现, 半湿润常绿阔叶林和中山湿性常绿阔叶林除生态外貌特征有一定区别外, 在植物区系组成和地理成分上很接近, 它们在种的组成上, 与季风常绿阔叶林的类似性仅为17.1%和15.4%。季风常绿阔叶林因其在植物区系和生态外貌上与后二者区别明显, 建议在云南植被分类上划分一个独立的植被型, 它是东南亚低山常绿阔叶林分布在中国西南部热带北缘山地的一个植被类型。结合云南的地质历史和古植物学资料, 认为云南的常绿阔叶林及其植物区系受晚中新世以来的地质历史事件深刻影响。半湿润常绿阔叶林是中国西南独特而特有种丰富的植被类型, 由于严重的人为干扰破坏, 现已片段化或成为萌生灌丛状, 应给予优先保护。     

西双版纳热带季节雨林的研究

本文以多个样方的资料分析,从群落综合特征上研究了西双版纳热带季节雨林的特点和类型、物种组成、生活型谱、叶级谱等。根据上层标志树种、群落生态外貌特征和生境把西双版纳的热带季节雨林区分为低丘雨林和沟谷雨林二个群系组,各包括若干群系。低丘雨林中面积最大和最有代表性的群系是大药树、龙果林,而沟谷雨林中最有代表性的群系是番龙眼、千果榄仁林。二者相比,低丘雨林的物种多样性要小一些,附生植物相对少一些,小、矮高位芽植物和小叶、落叶树种比例稍高,在生态上向季雨林和热带山地的常绿阔叶林过渡,有更强的地方代表性。沟谷雨林则更接近典型的湿热带雨林。     

Patterns of variation among forests of tropical Asian mountains,with some explanatory hypotheses

Decline in mean temperature with elevation correlates with a more or less gradual change in forest floristics mediated by minimum temperature tolerances and competition, and in its diversity, well documented on wet equatorial mountains. On the mountains of monsoon tropical Asia, the predominance of a single wet and dry season offers opportunity to relate increasing cloud and rainfall seasonality with a departure in a zonation most clearly recognised on equatorial mountains. Observations indicate that the lowland to lower montane forest transition is gradual and constant in elevation throughout the monsoon tropical Asia region, and contrasts with the variable elevation at which forest structure and physiognomy, alternatively floristics alone, changes between lower and upper equatorial montane forests. Both ecotones are observed to correlate with the elevation and seasonality at which decline in temperature daily results in condensation of water vapour into cloud, resulting in cloud shadow, and fog therefore water supply where cloud penetrates the canopy. Seasonal variation in these climatic interactions influence forest structure and physiognomy and, by mediating the interaction between vegetation and substrate, also leads to zonation in soils and consequently floristics. Historical biogeographical factors also influence zonal floristics.