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。     

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。     

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.     

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。     

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.     

论季雨林的水平地带性

季雨林是受制于湿度因子的经度地带性植被类型,它是随着湿度条件的下降由热带雨林向热带疏林过渡的居间类型,而不是受制于温度因子的纬度地带性植被类型,不是随着温度条件的下降由热带雨林向亚热带常绿阔叶林过渡的居间类型。我国南亚热带的榕树群系、黄桐群系,以及热带北缘的青皮群系均不应是季雨林,前二者是典型的南亚热带低地常绿阔叶林或南亚热带雨林,后者则是热带雨林的一分类群。     

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。     

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。     

Tropical deciduous forest in Yunnan,southwestern China: Implications for geological and climatic histories from a little-known forest formation

In the southern mountain ranges of Yunnan province, China, deep valleys of several large rivers create rain shadows with hot dry summers, and are locally designated tropical; towards the north, notably in the Lancang (Upper Mekong) valley, these regions may experience frost during winter. The woody forest canopy of these valleys is predominantly deciduous, with evergreen elements in the north, where the canopy is open and the forest savanna-like. However, we here present tall forest with a closed deciduous canopy and semi-evergreen subcanopy observed in hot dry valleys of these rivers and their tributaries in the tropical south. The structure and physiognomy of these forests resemble the tall (moist) deciduous forest formation widespread in South Asia and Indo-Burma. Furthermore, these forests are largely composed of tropical elements at both the generic (80%) and the species level (>70%), indicating that these forests are indeed tropical. We originally hypothesized that these isolated forests represent refugia of a pre-Holocene extension of tall (moist) deciduous forest formation of South Asia and Indo-Burma. The sample plot we established to test this hypothesis confirmed that these forests share the structure and physiognomy of the tall (moist) deciduous forest formation; however, the plots also showed that these forests lack the characteristic and dominant species of the formation''s Indo-Burmese range. The tree flora, in particular, indicates that both deciduous and evergreen elements are instead mostly derived from the adjacent tropical semi-evergreen forests of tropical southern China; yet they also include an important endemic element, which implies that these forests have survived as refuges possibly since the Pliocene. The exceptional representation of evergreen elements in these forests indicates that they have rarely been subject to hot fires or domestic cattle browsing, adding to the unique nature of the forests and further justifying their strict conservation.     

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.     

西双版纳自然保护区9种植被亚型象甲科多样性比较

2004年3–4月对西双版纳自然保护区季节性雨林、山地雨林、半常绿季雨林、落叶季雨林、石灰岩山季雨林、季风常绿阔叶林、暖温性落叶阔叶林、暖热性松林、热性竹林等9种植被亚型中象甲科昆虫多样性进行了研究。运用震落法采集标本853号,隶属于18亚科52属96种。跳象亚科个体数量最多,隐喙象亚科种类最丰富。季节性雨林和半常绿季雨林中不仅物种数最丰富,狭适种和稀有种也最多。暖温性落叶阔叶林及季节性雨林个体数量最丰富,落叶季雨林及半常绿季雨林最贫乏。9种植被亚型中象甲群落极不相似。季节性雨林和半常绿季雨林在生物多样性保护上具有重要地位,应予以优先考虑。     

热带季节性湿润林苦楝(Melia azedarach)径向生长季节动态及其对环境因子的响应

气候变化导致的温度升高和降水格局改变可能会影响到树木的生长速率和季节物候。西双版纳热带季节性湿润林分布在石灰岩山中部,属于热带喀斯特生境。由于土层浅薄,土壤保水能力极差,植物生长更容易遭到受到季节性干旱气候的影响。为探究热带季节性湿润林的树木径向生长季节动态及其对环境因子的响应,利用高精度树木生长仪连续两年监测了云南西双版纳热带季节性湿润林中落叶树种苦楝（Melia azedarach）的树干径向变化,并与同步监测的环境因子进行相关分析。结果表明,苦楝径向生长开始、结束以及持续生长的时间在年际间存在差异。与2018年相比,2019年苦楝生长开始和结束的时间较晚,且年生长量较小,这可能是与2019年雨季开始较晚且在生长季早期经历了严重的高温干旱有关。苦楝的径向日生长量与日降水量和相对湿度呈正相关关系,与光合有效辐射、水汽压亏缺和风速呈负相关关系,表明了在苦楝的径向生长主要受水分条件限制。在干旱年份（2019年）,苦楝的日生长量与降水和相对湿度的相关性更强。研究结果有助于进一步了解热带喀斯特生境树木生长对气候变化的敏感性以及树木适应季节性干旱气候的策略。     

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa

BIOME 6000 is an international project to map vegetation globally at mid‐Holocene (6000 ¹⁴C yr bp ) and last glacial maximum (LGM, 18,000 ¹⁴C yr bp ), with a view to evaluating coupled climate‐biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site‐based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method’s skill in reconstructing present‐day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south‐western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 ¹⁴C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial‐interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now‐arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land‐surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere‐biosphere models. The data could also be objectively generalized to yield realistic gridded land‐surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation‐climate feedbacks have focused on the hypothesized positive feedback effects of climate‐induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid‐Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 ¹⁴C yr bp and for the LGM.     

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。     

中国热带森林植被类型研究历史和划分探讨

热带森林是我国森林植被的重要组成部分,明确其森林植被类型分类对于生物多样性维持机制研究和保护管理等都具有重要意义。该文以中国热带森林植被分类研究中存在的问题为出发点,通过阐述我国各省区植被类型分类的研究历史,在综合考虑多种生物和非生物影响因素的基础上,提出一个新的热带森林植被类型分类框架以供探讨。结果表明:(1)尽管针对我国热带森林的分布范围和群落特征等都已开展了诸多研究,但对我国热带森林植被类型的划分依据和分类体系仍存在争议。(2)尽管我国的热带森林都处于季风气候区带内,但许多热带地区的森林植被类型并不只是受季风影响,而是气候带、关键气候因子、地形、土壤反馈和物种适应等多种因素共同作用的结果。(3)我国的热带森林植被包括5个植被型,即非典型性热带雨林、热带季雨林、热带山地雨林、热带山顶苔藓矮林(热带云雾林)和热带针叶林,其中热带季雨林植被型包含4种植被亚型[热带落叶季雨林、热带半落叶(半常绿)季雨林、热带常绿季雨林和热带石灰岩(石山)季雨林]。(4)阐明了上述热带森林植被型和植被亚型在我国各省区的分布情况,并提出未来有必要对人工恢复后的热带森林进行评估和植被类型划分。综上所述,该文提出一个新的热带森林植被类型分类框架,以期为今后基于不同地区开展热带森林比较研究提供参考。     

广东季雨林的几个问题

 季雨林是分布在具有明显干湿季节变化热带地区,在干季或多或少,甚至全部落叶的森林植被;是介于热带雨林向热带稀疏林过渡的居间类型,而不是由热带雨林向亚热带常绿阔叶林过渡的植被类型。应归属于经向地带性植被,而非纬度地带性植被。并且,在广东南部沿海地区的气候条件下,也不会有季雨林发育。广东南部沿海地区的榕树+香蒲桃+野苹婆(Ficus microcarpa+Syzygium odoratum+Sterculia lanceolata)群落等类型不是季雨林,而是热带季节雨林,属广义的热带雨林范畴。     

Tree species diversity of tropical forest vegetation in Xishuangbanna, SW China

Tree species diversity of four tropical forest vegetation types was investigated in Xishuangbanna, southwestern China. These are: tropical seasonal rain forest, tropical montane rain forest, evergreen broad-leaved forest and monsoon forest over limestone. A total of 17 samples were taken and four species diversity indices were calculated: Shannon-Wiener's H, the complement of Simpson's index, d, Fisher's and evenness index E. The results reveal the long-tailed rank/abundance diagrams of these forests. However, this feature is greatly reduced in the samples of monsoon forest over limestone. Tropical seasonal rain forest shows the highest tree species diversity of all four vegetation types. Owing to the variation of microenvironment, diversity values within the same vegetation type vary between the samples from different patches. The tree species diversity of single-dominant rain forest is not significantly lower than that of mixed rain forest, because the dominant species of some single-dominant rain forests are principally in the emergent layer. This is composed of sparse and huge trees of one species and, consequently, creates a unique canopy architecture and more heterogeneous microenvironments for the more diversified species composition under the emergent layer. The occurrence of tree species with small population sizes, particularly of species represented by only one individual, is highly correlated with the tree species diversity of the local forest vegetation. They are crucial elements in the richness of local biodiversity.     

Leaf flushing during the dry season: the paradox of Asian monsoon forests

Aim Most deciduous species of dry monsoon forests in Thailand and India form new leaves 1–2 months before the first monsoon rains, during the hottest and driest part of the year around the spring equinox. Here we identify the proximate causes of this characteristic and counterintuitive ‘spring‐flushing’ of monsoon forest trees. Location Trees of 20 species were observed in semi‐deciduous dry monsoon forests of northern Thailand with a 5–6‐month‐long severe dry season and annual rainfall of 800–1500 mm. They were growing on dry ridges (dipterocarp–oak forest) or in moist gullies (mixed deciduous–evergreen forest) at 680–750 m altitude near Chiang Mai and in a dry lowland stand of Shorea siamensis in Uthai Thani province. Methods Two novel methods were developed to analyse temporal and spatial variation in vegetative dry‐season phenology indicative of differences in root access to subsoil water reserves. Results Evergreen and leaf exchanging species at cool, moist sites leafed soon after partial leaf shedding in January–February. Drought‐resistant dipterocarp species were evergreen at moist sites, deciduous at dry sites, and trees leafed soon after leaf shedding whenever subsoil water was available. Synchronous spring flushing of deciduous species around the spring equinox, as induced by increasing daylength, was common in Thailand's dipterocarp–oak forest and appears to be prevalent in Indian dry monsoon forests of the Deccan peninsula with its deep, water‐storing soils. Main conclusions In all observed species leafing during the dry season relied on subsoil water reserves, which buffer trees against prolonged climatic drought. Implicitly, rainfall periodicity, i.e. climate, is not the principal determinant of vegetative tree phenology. The establishment of new foliage before the summer rains is likely to optimize photosynthetic gain in dry monsoon forests with a relatively short, wet growing season.