Clearance and fragmentation of tropical rain forest in Xishuangbanna, SW, China

Xishuangbanna, situated in the northern margin of the tropical zone in Southeast Asia, maintains large areas of tropical rain forest and contains rich biodiversity. However, tropical rain forests are being rapidly destroyed in this region. This paper analyzed spatial and temporal changes of forest cover and the patterns of forests fragmentation in Xishuangbanna by comparing classified satellite images from 1976, 1988 and 2003 using GIS analyses. The patterns of fragmentation and the effects of edge width were examined using selected landscape indices. The results show that forest cover declined from 69% in 1976 to less than 50% in 2003, the number of forests fragments increased from 6,096 to 8,324, and the mean patch size declined from 217 to 115 ha. It was found that fragment size distribution was strongly skewed towards small values, and fragment size and internal habitat differ strongly among forest types: less fragmented in subtropical evergreen broadleaf forest, but severe in forests that are suitable for agriculture (such as tropical seasonal rain forest and mountain rain forest). Due to fragmentation, the edge width was smaller in 2003 than that in 1976 when the total area of edge habitat exceeded core habitat in different forest types. The core area of tropical seasonal rain forest was smallest among main forest types at any edge width. Fragmentation was severe within 12.5-km buffers around roads. The current forest cover within reserves in Xishuangbanna was comparatively large and less fragmented. However, the tropical rain forest has been degraded inside reserves. For conservation purposes, the approaches to establish forest fragments networks by corridors and stepping stone fragments are proposed. The conservation efforts should be directed first toward the conservation of remaining tropical rain forests.     

Differences in seed rain composition in small and large fragments in the northeast Brazilian Atlantic Forest

Tropical forests are seriously threatened by fragmentation and habitat loss. The impact of fragment size and forest configuration on the composition of seed rain is insufficiently studied. For the present study, seed rain composition of small and large forest fragments (8–388 ha) was assessed in order to identify variations in seed abundance, species richness, seed size and dispersal mode. Seed rain was documented during a 1‐year period in three large and four small Atlantic Forest fragments that are isolated by a sugarcane matrix. Total seed rain included 20,518 seeds of 149 species of trees, shrubs, palms, lianas and herbs. Most species and seeds were animal‐dispersed. A significant difference in the proportion of seeds and species within different categories of seed size was found between small and large fragments. Small fragments received significantly more very small‐sized seeds (<0.3 cm) and less large‐seeded species (>1.5 cm) that were generally very rare, with only one species in small and eight in large fragments. We found a negative correlation between the inflow of small‐sized seeds and the percentage of forest cover. Species richness was lower in small than in large fragments, but the difference was not very pronounced. Given our results, we propose changing plant species pools through logging, tree mortality and a high inflow of pioneer species and lianas, especially in small forest fragments and areas with low forest cover. Connecting forest fragments through corridors and reforestation with local large‐seeded tree species may facilitate the maintenance of species diversity.     

The effect of habitat fragmentation on the bee visitor assemblages of three Australian tropical rainforest tree species

Tropical forest loss and fragmentation can change bee community dynamics and potentially interrupt plant–pollinator relationships. While bee community responses to forest fragmentation have been investigated in a number of tropical regions, no studies have focused on this topic in Australia. In this study, we examine taxonomic and functional diversity of bees visiting flowers of three tree species across small and large rainforest fragments in Australian tropical landscapes. We found lower taxonomic diversity of bees visiting flowers of trees in small rainforest fragments compared with large forest fragments and show that bee species in small fragments were subsets of species in larger fragments. Bees visiting trees in small fragments also had higher mean body sizes than those in larger fragments, suggesting that small‐sized bees may be less likely to persist in small fragments. Lastly, we found reductions in the abundance of eusocial stingless bees visiting flowers in small fragments compared to large fragments. These results suggest that pollinator visits to native trees living in small tropical forest remnants may be reduced, which may in turn impact on a range of processes, potentially including forest regeneration and diversity maintenance in small forest remnants in Australian tropical countryside landscapes.     

西双版纳片断热带雨林的结构、物种组成及其变化的研究

通过对西双版纳地区的６个“龙山”片断热带雨林与同样类型的原始热带雨林的比较研究,探讨了片断热带雨林的群落结构、种类组成、生活型构成、种群结构、植物丰富度及多样性等的变化及其规律。随着热带雨林的片断化,森林边缘效应和人为干扰加剧,热带雨林群落的结构变得不完整,植物丰富度降低。在生活型构成上,附生植物,大、中高位芽植物以及地上芽植物比例减少,藤本植物和小高位芽植物比例明显增高。在种群结构上,受轻、中度干扰的片断雨林,仅含１～２个个体的种类在群落中的相对数量有所增加,乔木种类组成变得不稳定,随着人为干扰的加剧,那些仅有１～２个个体为代表的种类将首先消失。在单位面积种数和物种多样性指数上,情况比较复杂,热带雨林片断化和受到一定干扰可能会增加藤本植物多样性指数,但干扰过度又会减低其多样性指数,附生植物多样性指数则随干扰和隔离时间呈明显下降趋势,但总的说来,片断热带雨林单位面积上的种数远比原始林少得多,随着人为干扰和隔离的加剧,物种多样性指数显著降低。     

Diversity and community composition of ectomycorrhizal fungi in a dry deciduous dipterocarp forest in Thailand

Large forest areas of South-East Asia, are dominated by the Dipterocarpaceae tree family, which contains many important timber species. Unlike many other tropical trees, Dipterocarpaceae rely on ectomycorrhizal (ECM) root symbiosis for their mineral nutrition. This study aims to document the richness and community composition of ECM fungi in a dry deciduous forest in Thailand. Combining morphological and molecular identification methods revealed 69 species of ECM fungi that belong to 17 phylogenetic lineages. The /russula-lactarius, /tomentella-thelephora, /sordariales, /sebacina and /cantharellus lineages were the most species-rich. The fungal richness is comparable to other tropical rain forest sites, but the phylogenetic community structure has elements of both tropical and temperate ecosystems. Unlike tropical rain forests, the Cenococcum geophilum complex was one of the most frequent fungal taxa that had a relatively high ITS genetic diversity over the small sampling area. This study provides the first snapshot insight into the fungal community of dry dipterocarp forests. However, it is necessary to broaden the spatial and temporal scales of sampling to improve our understanding of the below-ground relations of dry and humid tropical forests.     

Tropical rain forest fragmentation and wild populations of primates at Los Tuxtlas,Mexico

In view of the extensive destruction, fragmentation, and conversion of primate habitats to anthropogenic vegetation, information on Neotropical primate ability to use a landscape consisting of forest fragments and agricultural habitats is necessary to understand the ecological flexibility of the species involved and it is of relevance to the design of conservation scenarios at the landscape level. We censused howlers and spider monkeys in 126 forest fragments and in 44 agricultural sites at Los Tuxtlas, southern Veracruz, Mexico, and used the IDRISI Geographic Information System to assess the extent of primate habitat remaining. We conducted economic surveys to assess the productivity of several systems of land management, including cattle ranching. Seventy- five percent of native habitat has been lost at Los Tuxtlas, 20% remains in the form of isolated fragments of vegetation, and only 5% consists of contiguous rain forest at high elevations (> 800 m). Howlers and spider monkeys were present in only 60 and 8% of the forest sites investigated, respectively, attesting to extensive local extinction. Populations of both species are small and found in a fragmented and isolated condition throughout the landscape. A large number of howlers were detected in artifactual habitats such as cacao, coffee and mixed (cacao and coffee) plantations shaded by rain forest trees. Residual rain forest vegetation along streams, rivers, and lakes facilitated the interfragment and interhabitat movement of howlers. Economic surveys showed that some of the agricultural habitats were more productive than cattle ranching. Conservation of spider monkeys requires large areas of contiguous forest, but only small areas are present at high elevations. Howlers are restricted to elevations < 800 m, where most of the forest has disappeared, but can apparently exist in a matrix of forest fragments, arboreal agricultural habitats, and pasture lands. Structural aspects of the vegetation and connectivity among habitat islands may be indispensable components of both landscape scenarios.     

The Importance of Tropical Rain Forest Fragments to the Conservation of Plant Species Diversity in Los Tuxtlas, Mexico

Of what value are forest fragments to the conservation of the tropical rain forest diversity for a landscape? We compared the changes in composition, diversity, and plant structure of 15 small (1–76 ha) relatively unprotected forest fragments with those of a large (700 ha) well protected fragment (LWPF) in Los Tuxtlas, Mexico. The trees, shrubs, lianas, palms, and herbs with dbh?≥?2.5 cm were sampled in 1000 m² at each site. For each ecological group (based on light requirements for germination: primary, secondary, and non-secondary light demanding, NSLD, species) and life form, estimates of species diversity, density, and basal area were analyzed using a stepwise multiple regression to determine whether there were any relationships between these variables and fragment characteristics (size, shape index, and isolation). The composition and plant structure of LWPF were different from those of the small fragments and large trees are absent from the canopy of the latter. Fragment size best explained the differences in composition and plant structure. Species composition in the largest fragments was similar to that of LWPF, but there was no significant difference in total richness between LWPF and the fragments, even though both the richness and abundance of secondary and NSLD species did differ. LWPF had more large primary trees in the canopy, and a greater abundance and basal area of palms and herbs. For tropical rain forest conservation, it is important to maintain the greatest possible number of large fragments and establish policies that prevent forest remnants from being further reduced in size and increasingly isolated from each other.     

西双版纳热带森林地区不同生境蜜蜂的物种多样性研究

本文研究了西双版纳热带森林地区不同生境蜜蜂的物种多样性。共鉴定蜜蜂总科5科、22属、77种。8个生境的物种丰富度指数D_MA的取值范围在1.914～7.877之间,多样性指数H′的范围是0.608～1.398,均匀度指数J_sw为0.716～0.956,其中保护区、雨林和灌丛拥有较高的物种丰富度指数和多样性指数,橡胶林的最小,而均匀度指数以曼养广龙山最大,农田最小。根据不同生境物种的相似性系数C进行系统聚类,8个生境在相似性系数0.169的水平时全部聚合在一起。傣族龙山在保持雨林的蜜蜂物种多样性中可以起一定的作用,但很有限,且当龙山面积越小,受人为干扰越严重时,物种多样性的损失就越大。雨林一旦被开垦为橡胶林或农田,雨林中原有的许多蜜蜂将会消失,物种多样性显著减少。     

Gap-crossing movements predict species occupancy in Amazonian forest fragments

In fragmented landscapes, species persistence within isolated habitat patches is governed by a myriad of species life‐history, habitat patch and landscape characteristics. We investigated the inter‐specific variation in non‐forest gap‐crossing abilities of an entire tropical forest‐dependent avifauna. We then related this measure of dispersal ability to species life‐history characteristics and occupancy data from 31 variable‐sized forest patches sampled within the same fragmented forest landscape. A total of 5436 gap‐crossing movements of 231 forest‐dependent bird species were observed across ten linear forest gaps of varying widths, adjacent to large areas of undisturbed forest. Species persistence in isolated fragments was strongly linked to gap‐crossing ability. The most capable gap‐crossers were medium to large‐bodied species in the large insectivore, frugivore and granivore guilds, matching the most prevalent subset of species in small forest patches. However, some competent gap‐crossing species failed to occur in small patches, and minimum forest‐patch area requirements were more important in determining patch occupancy for these species. Narrow forest gaps (4–70 m) created by roads and power‐lines may become territory boundaries, thereby eliminating home‐range gap‐crossing movements for many forest species, but permit rarer dispersal events. Wider gaps (>70 m) may inhibit gap‐crossing behaviour for all but the most vagile species. Although patch size and quality may be the most important factors in structuring species assemblages in forest fragments, our results show that the degree of patch isolation and permeability of the surrounding matrix also explain which species can persist in forest isolates. Reducing the number and width of forest‐dividing gaps; maintaining and/or creating forest corridors and increasing matrix permeability through the creation and maintenance of ‘stepping‐stone’ structures will maximise the species retention in fragmented tropical forest landscapes.     

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。     

Plant Community Structure in Tropical Rain Forest Fragments of the Western Ghats,India1

Changes in tree, liana, and understory plant diversity and community composition in five tropical rain forest fragments varying in area (18–2600 ha) and disturbance levels were studied on the Valparai plateau, Western Ghats. Systematic sampling using small quadrats (totaling 4 ha for trees and lianas, 0.16 ha for understory plants) enumerated 312 species in 103 families: 1968 trees (144 species), 2250 lianas (60 species), and 6123 understory plants (108 species). Tree species density, stem density, and basal area were higher in the three larger (> 100 ha) rain forest fragments but were negatively correlated with disturbance scores rather than area per se. Liana species density, stem density, and basal area were higher in moderately disturbed and lower in heavily disturbed fragments than in the three larger fragments. Understory species density was highest in the highly disturbed 18‐ha fragment, due to weedy invasive species occurring with rain forest plants. Nonmetric multidimensional scaling and Mantel tests revealed significant and similar patterns of floristic variation suggesting similar effects of disturbance on community compositional change for the three life‐forms. The five fragments encompassed substantial plant diversity in the regional landscape, harbored at least 70 endemic species (3.21% of the endemic flora of the Western Ghats–Sri Lanka biodiversity hotspot), and supported many endemic and threatened animals. The study indicates the significant conservation value of rain forest fragments in the Western Ghats, signals the need to protect them from further disturbances, and provides useful benchmarks for restoration and monitoring efforts.     

The Contribution of Tropical Secondary Forest Fragments to the Conservation of Fruit-feeding Butterflies: Effects of Isolation and Age

Concomitant with the rapid loss of tropical mature forests, the relative abundance of secondary forests is increasing steadily and the latter are therefore of growing interest for conservation. We analysed species richness of fruit-feeding nymphalid butterflies in secondary forest fragments of different age and isolation and in mature forest at the eastern margin of the Lore Lindu National Park in Central Sulawesi, Indonesia. From April to August 2001 we collected 2322 individuals of fruit-feeding butterflies, belonging to 33 species. Butterfly species richness increased with succession, but was significantly higher in mature forests than in all types of secondary forest. Isolation of the forest fragments did not have a significant effect on butterfly species richness in the range of distances (up to 1700 m) studied. Rather it appeared to affect only a few species. Species richness of endemic species was higher than of non-endemic species. Although endemic species were most diverse in mature forests, many species captured were restricted to secondary forests. Our results show that mature forest is essential for the conservation of nymphalid butterflies and for the endemic species in this area. However, considering the relatively large number of species found in these rather small habitat islands, secondary forest fragments, especially older successional stages, can be taken into account in conservation efforts and thus contribute to the preservation of tropical biodiversity on a landscape scale.     

西双版纳片断热带雨林植物区系成分及变化趋势

本文以西双版纳地区的6个“龙山”片断热带雨林的植物物种多样性编目为基础,通过与同样类型的原始热带雨林的比较,探讨了片断热带雨林植物区系成分的变化。随着热带雨林的片断化,森林边缘效应和人为干扰加剧,阳性先锋植物和藤本植物等的侵入,导致这些先锋植物和藤本植物所属的科属在群落中的种数增多,优势度增大。而一些含喜荫湿的荫生植物、群落顶极树种及附生植物较多的科属则在群落中的种数减小了,优势度下降。在植物属的地理成分构成上,片断热带雨林中全热带分布和热带亚洲—热带非洲分布成分比例相对增高,而热带亚洲成分相对减少。在种的地理成分构成上,片断热带雨林中广域分布种比例相对增高,显示了与属的地理成分类似的变化。这也是森林片断化后,由于边缘效应和林内生境在一定程度上干暖化,喜光的先锋成分增多,而先锋成分大多为一些广域分布属种所致。     

Edge effects on palm diversity in rain forest fragments in western Ecuador

At the edges of tropical rain forest fragments, altered abiotic and biotic conditions influence the structure and dynamics of plant communities. In Neotropical rain forests, palms (Arecaceae) are important floristic and ecological elements. Palms’ responses to edge effects appear to be idiosyncratic and to depend on the level of disturbance at edges. This paper explores how variation in forest structure at the edges of two old-growth forest fragments in a tropical rain forest in western Ecuador affects palms of different species, life-forms, and size classes. We investigate (1) how edge effects influence the relative proportion of palm adults and juveniles, (2) how distance from the forest edge affects palm density and species richness, (3) how altered forest structure along edges affects palm density. We found that at edges (1) palm communities had a lower proportion of adults relative to juvenile individuals compared to continuous forests, (2) the density of two species of palms and the overall species richness of the palm community tended to decrease toward the edges within forest fragments, and, (3) altered forest structure decreased the density of adult palms. Hence, edge effects on palms were controlled by the degree of modification of the forest structure, and by species responses to edge-related disturbance.     

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。     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

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。     

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.     

Continental rain forest fragments in Singapore resist invasion by exotic plants

Abstract Aim In general, the plant communities of oceanic islands suffer more from exotic plant invasions than their continental equivalents. At least part of this difference may be contributed by differences in non‐biological factors, such as the antiquity and intensity of human impacts and the absence of internal barriers to dispersal, rather than differences in inherent invasibility. We tested the resistance of species‐rich continental rain forests to plant invasion on a small, continental island that has been subject to prolonged and intensive human impact. Location Singapore is a 683‐km² equatorial island <1 km from the Asian mainland and with a population of 4 million people. It has a continental biota but has been subject to human impacts as intense as on any oceanic island. Methods We sampled twenty‐nine sites in seven vegetation types, ranging from urban wasteland to fragments of primary lowland rain forest. In each sample plot, all plant species were identified, exotic cover was estimated, and a range of environmental variables measured. Additional qualitative surveys for exotic invasion were made in other forest areas in Singapore. The data were analysed by Spearman's rank correlation coefficient. Results The number of exotic species recorded at a site was unrelated to the number of native species. Across all sites, percentage canopy opening had the highest correlation with the number of exotic species, while soil pH (which largely reflects the incorporation of calcareous construction wastes) had the highest correlation if the mangrove sites were excluded. There were no exotics in mangrove forest and only a tropical American, bird‐dispersed shrub, Clidemia hirta (L.) D. Don (Melastomataceae: Koster's Curse), in primary and tall secondary forest patches. The species‐poor early stages of woody plant succession on highly degraded soils were also very resistant to exotic plant invasion. Main conclusions Long‐isolated rain forest fragments in an exotic‐dominated continental island landscape resist invasion by exotic plants, suggesting that the problems on oceanic islands may reflect an inherently greater invasibility. This study also adds to the increasing evidence that the floras of tropical rain forest fragments in South‐east Asia are remarkably resilient on a time‐scale of decades to a century or more.     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.