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。     

关于对朱华先生"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"一文中有关问题的说明

朱华先生对“西双版纳热带山地雨林的植物多样性研究”（李宗善等,2004）一文中的一些问题提出了质疑,这主要缘于该文着重于群落植物多样性的研究,并未对6块热带山地雨林样地的物种组成、群落结构特征等进行分析,以致朱华先生对“西双版纳热带山地雨林的植物多样性研究”一文中的植被类型的划分及群落特征等方面产生一些不清楚的地方。作为原文的通讯作者,对朱华先生所提出的问题说明如下：     

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。     

中国西南地区热带季节雨林及山地常绿阔叶林热值及能量分配格局

能量分配格局是研究生态系统能量流动的基础,但是由于热带森林结构的高度复杂性和物种多样性,对它的热值和能量分配格局的全面研究还很少。该文研究的热带季节雨林位于西双版纳,是分布于热带亚洲北缘的一种森林类型;山地常绿阔叶林位于云南省中部的哀牢山,属于我国西部亚热带地区的山地常绿阔叶林。该研究的目的是探讨这两种重要森林类型的热值和能量分配格局,验证Golley(1961,1969)提出的世界范围内植被的热值由低纬度向高纬度、由低海拔向高海拔升高的规律。热值的测定采用SDCM-Ⅲa氧弹测量仪。两个森林样地面积都是1 hm²,能量分配格局及年固定量根据生物量和生物量增量计算。研究结果表明,热带季节雨林样地的热值低于山地常绿阔叶林,乔木层的热值>灌木层>草本层,所有器官中叶片的热值较高。由于以前种植砂仁(Amomum villosum)的影响,热带季节雨林样地的能量现存量小于山地常绿阔叶林,但是因为地处高温高湿、光照充足的地区,热带季节雨林的能量年固定量高于山地常绿阔叶林。对于热带季节雨林样地来说,97%的能量储存在乔木层中;山地常绿阔叶林样地的乔木层储存了88%的能量,可见乔木层是维持森林能量结构的关键层。研究结果为Golley的结论提供了更加丰富的实验证据。     

西双版纳20公顷样地热带森林植被数量分类与排序

为了区分热带森林的群落特征并了解其群落特征与地形因子之间的相互关系,以西双版纳20 hm2森林动态监测样地为研究对象,对样地内500个森林群落样方采用Cluster聚类分析方法进行分类和采用除趋势对应分析(DCA)、典范对应分析(CCA)方法进行排序。结果表明:(1)西双版纳热带森林动态监测样地森林群落分为2个类型:热带季节雨林和热带山地常绿阔叶林;(2)植被类型的划分与CCA排序的结果相吻合,CCA排序轴第一、二轴呈现了海拔和凹凸度的梯度变化,并且两者总体上可以对83.9%的植被分布格局进行解释,表明海拔和凹凸度因子对该地区植被类型的分布起着决定性作用;(3)对优势物种的DCA和CCA排序表明不同物种对地形的要求存在差异,其分布格局同样主要受海拔和凹凸度的影响;(4)聚类分析与DCA和CCA排序的结果,同时表明了该地区森林植被的垂直分布格局。此方法可有效解释植被分布格局与地形之间的内在联系,为热带森林的管理和保护提供科学依据。     

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。     

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。     

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。     

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。     

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。     

On the Classification of Forest Vegetation in Xishuangbanna,Southern Yunnnan

Xishuangbanna of southern Yunnan is a region of extremely interest to biologists and also a hotspot for biodiversity conservation . It is located in a transitional zone from tropical Southeast Asia to temperate East Asia biogeographically. The present paper reviewed vegetation types of Xishuangbanna and suggested a revised classification system based on theupdated study results over the last two decades . By combining physiognomic and floristic characteristics with ecological performances and habitats , the primary forest vegetation in Xishuangbanna can be organized into four main vegetation types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad-leaved forest and tropical monsoon forest. The tropical rain forest can be classified into two subtypes , i. e. tropical seasonal rain forest in the lowlands and tropical montane rain forest on higher elevations. The tropical seasonal rain forest in this region shows similar forest profile and physiognomic characteristics to those of equatorial lowland rain forests and is a type of world tropical rain forest. Because of conspicuous similarity on floristic composition , the tropical seasonal rain forest in Xishuangbanna is a type of tropical Asian rain forest . However , since the tropical seasonal rain forest occurs at the northern edge of tropical SE Asia, it differs from typical lowland rain forests in equatorial areas in maintaining some deciduous trees in the canopy layer , fewer megaphanerophytes and epiphytes but more abundant lianas and more plants with microphyll . It is a type of semi-evergreen rain forest at the northern edge of the tropical zone . The tropical montane rain forest occurs in wet montane habitats and is similar to the lower montane rain forests in equatorial Asia in floristic composition and physiognomy . It is a variety of lower montane rain forests at the northern tropical edges of tropical rain forests . The tropical seasonal moist forest occurs on middle and upper limestone mountains and is similar to the tropical montane evergreen broad-leaved forest of the region in physiognomy, but it differs from the latter in floristic composition. The monsoon forest in Xishuangbanna is a tropical deciduous forest under the influence of a strong monsoon climate and is considered to be a transitional vegetation type between tropical rain forest and savanna in physiognomy and distribution. The tropical montane evergreen broad- leaved forest is the main vegetation type in mountain areas . It is dominated by the tree species of Fagaceae , Euphorbiaceae , Theaceae and Lauraceae in majority. It differs from the tropical montane rain forests in lack of epiphytes and having more abundant lianas and plants with compound leaves . It is considered to be a distinct vegetation type in the northern margin of mainland southeastern Asia controlling by a strong monsoon climate, based on its floristic and physiognomic 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.     

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.     

西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成与多样性格局

为探讨西双版纳热带雨林和热带山地常绿阔叶林附生苔藓的组成、多样性格局及其维持机制,该文对两种植被的树干附生苔藓植物进行了调查与分析。所选择的样地位于西双版纳勐腊县,每种植被类型选择10个20 m×20 m的样方,每个样方选择约10棵样树。结果表明:(1)本调查共记录到附生苔藓20科39属60种,其中热带雨林附生苔藓有19科33属48种,热带山地常绿阔叶林附生苔藓有9科14属19种,共有种6种。(2)热带雨林最优势科为平藓科,热带山地常绿阔叶林最优势科为锦藓科。(3)热带雨林比热带山地常绿阔叶林具有更高的物种丰富度、β多样性和γ多样性。(4)不同径级和不同树皮粗糙度宿主上附生苔藓植物的盖度差异显著,而物种丰富度差异不显著。(5)苔藓植物的生活型对植被类型有所偏好,表现为悬垂型、扇型、交织型、粗平铺型集聚于热带雨林,丛集型、垫状型、细平铺型集聚于热带山地常绿阔叶林。(6)直接排序结果表明,宿主特性特别是树皮粗糙度显著影响苔藓植物生活型的组成和分布。热带雨林比热带山地常绿阔叶林能提供更多样的微生境,因而孕育了物种更丰富的附生苔藓植物。考虑到不同的植被类型或宿主特性条件下孕育着不同生活型组成的苔藓植物,因此可将苔藓生活型作为今后森林监测的一项重要指标。     

Combined community ecology and floristics,a synthetic study on the upper montane evergreen broad-leaved forests in Yunnan,southwestern China

The upper montane evergreen broad-leaved forest in Yunnan occurs mainly in the zone of persistent cloud and has a discontinuous, island-like, distribution. It is diverse, rich in endemic species, and likely to be sensitive to climate change. Six 1-ha sampling plots were established across the main distribution area of the upper montane evergreen broad-leaved forest in Yunnan. All trees with d.b.h. > 1 cm in each plot were identified. Patterns of seed plant distributions were quantified at the specific, generic and family levels. The forests are dominated by the families Fagaceae, Lauraceae, Theaceae and Magnoliaceae, but are very diverse with only a few species shared between sites. Floristic similarities at the family and generic level were high, but they were low at the specific level, with species complementarity between plots. Diversity varied greatly among sites, with greater species richness and more rare species in western Yunnan than central Yunnan. The flora is dominated by tropical biogeographical elements, mainly the pantropic and the tropical Asian distributions at the family and genus levels. In contrast, at the species level, the flora is dominated by the southwest or the southeast China distributions, including Yunnan endemics. This suggests that the flora of the upper montane forest in Yunnan could have a tropical floristic origin, and has adapted to cooler temperatures with the uplift of the Himalayas. Due to great sensitivity to climate, high endemism and species complementarity, as well as the discontinuous, island-like, distribution patterns of the upper montane forest in Yunnan, the regional conservation of the forest is especially needed.     

西双版纳地区六种林型地表蜘蛛多样性比较研究

为探索地表蜘蛛多样性及其变化与森林类型和管理方式的关系, 在西双版纳勐仑自然保护区选择热带季节雨林、石灰山季节雨林和山地常绿阔叶林, 在自然保护区附近选择人工纯林、胶茶群落和橡胶林, 共6种林型, 每种林型选择3块样带, 共设置研究样地18块, 分别于2006年12月上旬（雾凉季）、2007年3月下旬（干热季）和2007年7月上旬（雨季）, 以单位地表面积法收集地表蜘蛛的物种组成和数量数据, 并以蜘蛛种类和数量分布为属性进行典范对应分析（CCA）, 探讨不同类型植被与地表蜘蛛多样性的关系。共采集蜘蛛标本9 849头, 用于统计分析的成熟蜘蛛3 119头, 归属于30科, 其中幽灵蛛科、皿蛛科、球蛛科和小密蛛科是地表蜘蛛的优势类群。各林型科的数量为: 热带季节雨林24科, 石灰山季节雨林22科, 山地常绿阔叶林22科, 人工纯林20科, 胶茶群落21科, 橡胶林19科; 各林型特有科数量: 热带季节雨林2科, 山地常绿阔叶林2科, 橡胶林1科; 而仅在雨林中分布的科4个（占全部30个科的13.3%）, 仅在自然林中分布的科6个（20.0%）, 仅在人工林中分布的科1个（3.3%）。从蜘蛛的数量分布看, 个体密度在热带季节雨林显著高于其他5种林型; 橡胶林多样性指数和丰富度指数显著低于3种自然林, 而均匀度指数的最低值也同样在橡胶林出现。CCA分析和聚类分析的结果表明, 6种林型趋于分成2组, 即: 自然林和人工林; 在自然林中两种次生林的相似程度更高; 人工林中人工纯林（非橡胶林）与胶茶群落的相似程度更高。以上结果表明: （1）森林砍伐后种植人工林措施改变了该地区地表蜘蛛群落的物种分布格局; （2）蜘蛛多样性随着人为干扰程度增加有减少的趋势; （3）减少人为干扰和增加植被群落多样性（橡胶林中种植茶树）对保护和恢复物种多样性有重要意义。