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.     

Prediction of neotropical tree and liana species richness from soil and climatic data

We present an analysis of local species richness in neotropical forests, based on a number of 0.1 ha samples of woody plants collected by the late Alwyn Gentry. For each of 69 forests, soils were analysed and climatic data were collated. Using transformed independent variables and interaction terms, multiple regression equations were developed that explained the greatest possible amount of variation in species richness, and the best equations were selected on the basis of regression diagnostics. The best models are presented for (a) all neotropical forests, (b) forests west of the Andes (transandean) and (c) east of the Andes (cisandean), and for various subsets based on elevation and annual rainfall. For the whole dataset, and for most subsets, annual rainfall and rainfall seasonality were the most important variables for explaining species richness. Soil variables were correlated with precipitation — drier forests have more nutrient-rich soils. After the inclusion of rainfall variables, available soil nutrient concentrations contributed little to explaining or accounting for additional variation in species numbers, indicating that tropical forest species richness is surprisingly independent of soil quality. The results are consistent with the hypothesis that plants in mature tropical forests may obtain nutrients through the process of direct cycling, in which mineral nutrients are extracted from litterfall before they enter the soil. The strong relationship between community species richness and rainfall patterns has implications for biodiversity conservation. Wet forests with an ample year-round moisture supply harbour the greatest number of woody plant species and should be a focus of conservation efforts.Died 3 August 1993.     

Identification of characteristic plant co-occurrences in neotropical secondary montane forests

Aims Inferring environmental conditions from characteristic patterns of plant co-occurrences can be crucial for the development of conservation strategies concerning secondary neotropical forests. However, no methodological agreement has been achieved so far regarding the identification and classification of characteristic groups of vascular plant species in the tropics. This study examines botanical and, in particular, statistical aspects to be considered in such analyses. Based on these, we propose a novel data-driven approach for the identification of characteristic plant co-occurrences in neotropical secondary mountain forests.Methods Floristic inventory data were gathered in secondary tropical mountain forests in Ecuador. Vegetation classification was performed by coupling locally adaptive isometric feature mapping, a non-linear ordination method and fuzzy- c -means clustering. This approach was designed for dealing with underlying non-linearities and uncertainties in the inventory data.Important findings The results indicate that the applied non-linear mapping in combination with fuzzy classification of species occurrence allows an effective identification of characteristic groups of co-occurring species as fuzzy-defined clusters. The selected species indicated groups representing characteristic life-form distributions, as they correspond to various stages of forest regeneration. Combining the identified 'characteristic species groups' with meta-information derived from accompanying studies indicated that the clusters can also be related to habitat conditions. In conclusion, we identified species groups either characteristic of different stages of forest succession after clear-cutting or of impact by fire or a landslide. We expect that the proposed data-mining method will be useful for vegetation classification where no a priori knowledge is available.     

Soil and Geographic Distance as Determinants of Floristic Composition in the Azuero Peninsula (Panama)

Many studies analyzing the relative contribution of soil properties versus distance‐related processes on plant species composition have focused on lowland tropical forests. Very few have investigated two forest types simultaneously, to contrast ecological processes that assemble the communities. This study analyses—at the landscape scale—the relative contribution of soil and distance on lowland and submontane tropical forests, which co‐occur in two reserves of the Azuero peninsula (Panama). Floristic inventories and soil sampling were conducted in 81 0.1‐ha plots clustered in 27 sites, and data were analyzed using Mantel tests, variance partitioning and non‐metric multidimensional scaling. The largest differences in floristic composition occurred between reserves in both forest types. Soil variation and geographic distance were important determinants of floristic composition, but their effects were highly correlated; together they explained 7–25 percent and 46–50 percent of the variation in lowland and submontane forests, respectively. Soil variables that had the best correlations with floristic composition were iron, zinc, and silt content in lowland, and calcium, copper, iron, potassium, magnesium, phosphorus, zinc, and sand content in submontane forests. The studied forests showed a high beta diversity that seems to be related primarily with soils and, secondarily, with dispersal limitation and stochastic events. The results reveal a response of tree assemblages to environmental gradients, which are particularly conspicuous in Panama. The effects of limited dispersal seem to be more important in submontane than in lowland forests, probably as a result of higher isolation.     

热带森林的低密度种及其形成及维持机制

低密度种指在热带森林中存在的成年个体密度很低的物种 ,是热带森林中树种的主要存在形式。对低密度种的认识和保护是热带森林生物多样性保育的关键之一。与稀有种相比 ,低密度种的含义更为广阔 ,它还包括那些密度低、分布范围广和绝对个体数量大的物种。低密度种形成的主要原因是母树附近的幼苗和种子的存活率较低 ,密度制约和有限更新也是低密度种形成的重要原因。维持低密度种最重要的外部原因是昆虫较强的飞行能力和传粉能力 ,而其最重要的内部原因是低密度种的生殖特性。     

Drier tropical forests are susceptible to functional changes in response to a long‐term drought

Climatic changes have profound effects on the distribution of biodiversity, but untangling the links between climatic change and ecosystem functioning is challenging, particularly in high diversity systems such as tropical forests. Tropical forests may also show different responses to a changing climate, with baseline climatic conditions potentially inducing differences in the strength and timing of responses to droughts. Trait‐based approaches provide an opportunity to link functional composition, ecosystem function and environmental changes. We demonstrate the power of such approaches by presenting a novel analysis of long‐term responses of different tropical forest to climatic changes along a rainfall gradient. We explore how key ecosystem's biogeochemical properties have shifted over time as a consequence of multi‐decadal drying. Notably, we find that drier tropical forests have increased their deciduous species abundance and generally changed more functionally than forests growing in wetter conditions, suggesting an enhanced ability to adapt ecologically to a drying environment.     

热带森林植物功能群及其动态研究进展

热带森林极高的物种多样性和结构复杂性给生态学研究带来了很多挑战。植物功能群是对特定环境响应相似或对主要的生态过程具有相似作用的物种组合。应用植物功能群的方法是有效减少热带森林群落复杂性,并揭示其格局和过程的良好途径。在介绍植物功能群概念和划分途径的基础上,分析了热带森林植物功能群的时空动态规律。一般来讲,划分植物功能群通常有3种途径,并可通过5个步骤来完成。热带森林植物功能群的空间分布常受景观格局的制约,而环境异质性往往是影响不同植物功能群组配比例变化的直接原因。不同类型的植物功能群随演替过程发生显著的替代,而干扰体系和全球气候变化对功能群的动态过程具有重要的驱动作用。以功能群为基础的动态模型在模拟热带林群落动态和预测植被潜在分布等方面具有广阔的发展前景。探索有效的植物功能分类方法、构建完善的植物功能性状数据库、开发更为精确的功能群动态模型以及加强基于景观水平的植物功能群动态机制的认识等是未来热带森林植物功能群及其动态研究的重要方向。     

Valuing environmental functions in developing countries

The interface between ecology and economics is the valuation of environmental goods, services and attributes. The total economic value of an ecosystem is derived by estimating monetary values for it's direct use, indirect use, option and non-use values. Recent advances have been made in the economic valuation of direct, option and non-use values. Much less attention has been paid to measuring the indirect use value provided by environmental functions. These values may be particularly significant in developing countries. This paper details the challenge presented by valuing environmental functions to ecologists and economists and synthesizes the methodological advances that have occurred. Using tropical forests, wetlands and biodiversity as illustrations, the application of this methodology to valuing the functions of complex natural systems is investigated and existing studies reviewed. Conclusions on further research are presented.     

Research Priorities for Neotropical Dry Forests1

Our understanding of the human and biophysical dimensions of tropical dry forest change and its cumulative effects is still in the early stages of academic discovery. The papers in this special section on Neotropical dry forests cover a wide range of sites and problems ranging from the use of multispectral and hyperspectral remote sensing platforms to the impact of hurricanes on tropical dry forest regeneration. Here, we present to the scientific community the results of a workshop on which research priorities for tropical dry forests were discussed. This discussion focuses on the need to develop linkages between remote sensing, ecological, and social science research. The incorporation of social sciences into ecological research could contribute dramatically to our understandings of tropical dry forests by providing important contextual information to ecologists, and by helping to develop an important science–policy–public nexus on which environmental management can succeed.     

Rapid Liana Colonization along a Secondary Forest Chronosequence

Lianas (woody vines) can have profound effects on tree recruitment, growth, survival, and diversity in tropical forests. However, the dynamics of liana colonization soon after land abandonment are poorly understood, and thus it is unknown whether lianas alter tree regeneration early in succession. We examined the liana community in 43 forests that ranged from 1 to 31 yr old in central Panama to determine how fast lianas colonize young forests and how the liana community changes with forest succession. We found that lianas reached high densities early in succession, commonly exceeding 1000 stems/ha within the first 5 yr of forest regeneration. Lianas also increased rapidly during early succession in terms of basal area but did not show evidence of saturation within the 30 yr of our chronosequence. The relative contribution of lianas to total woody plant community in terms of basal area and density increased rapidly and reached a saturation point within 5 yr (basal area) to 15 yr (density) after land abandonment. Our data demonstrate that lianas recruit early and in high density in tropical forest regeneration, and thus lianas may have a large effect on the way in which secondary forests develop both early and throughout succession.     

Let's Not Forget the Biodiversity of the Cities

Cities contain over half of the world's population and are the source of many anthropogenic effects on the world's biota. Close examination of San Juan, Puerto Rico, a tropical city, reveals a significant green infrastructure. The city contains natural and human-constructed forests, urban aquatic systems with native and introduced species, and a thriving and diverse biota adapted to urban conditions. It behooves tropical ecologists to pay attention to the biodiversity of urban environments to learn lessons about ecological persistence and adaptation to novel anthropogenic environments.     

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。     

Birds,Cattle, and Bracken Ferns: Bird Community Responses to a Neotropical Landscape Shaped by Cattle Grazing Activities

Large areas of tropical moist forests have been converted to cattle pastures, generating complex landscapes where different habitats are represented by small patches with an uneven spatial distribution. Here, we describe how bird communities respond to the different elements present in a livestock landscape that was originally dominated by tropical moist forest. We surveyed six habitats: open pastures, pastures with shrubs, early‐ and middle‐secondary forests, mature forest, and pastures invaded by bracken ferns (Pteridium aquilinum). Bird diversity was high in secondary and mature forests, and low in fern‐invaded sites and open pastures. Fern‐dominated sites had the lowest bird species richness, and trophic guild diversity of all habitats. Habitat structure affected both bird species richness and densities in similar ways. Tree species richness was the habitat attribute that had a bigger positive effect on bird species richness. Bird community structure varied among sampled habitats, separating habitats in two major groups (forests and pastures). Our data indicate that bracken fern‐invaded pastures were the worst habitat condition for avian communities. To increase bird diversity, we recommend to eliminate or manage bracken fern and to increase shrub and tree cover in open pastures to provide food resources and shelter for birds. Finally, we encourage the maintenance of secondary and mature forest remnants as a strategy to conserve resident birds within a landscape dominated by livestock activities.     

Species Richness,Forest Structure,and Functional Diversity During Succession in the New Guinea Lowlands

Much of the world's tropical forests have been affected by anthropogenic disturbance. These forests are important biodiversity reservoirs whose diversity, structure and function must be characterized across the successional sequence. We examined changes in structure and diversity along a successional gradient in the lowlands of New Guinea. To do this, we measured and identified all stems ≥5 cm diameter in 19 0.25 ha plots ranging in age from 3 to >50 yr since disturbance. We also measured plant functional traits related to establishment, performance, and competitive ability. In addition, we examined change in forest structure, composition, species diversity, and functional diversity through succession. By using rarefaction to estimate functional diversity, we compared changes in functional diversity while controlling for associated differences in stem and species density. Basal area and species density increased with stand age while stem density was highest in intermediate secondary forests. Species composition differed strongly between mature and secondary forests. As forests increased in basal area, community‐weighted mean wood density and foliar carbon increased, whereas specific leaf area and proportion of stems with exudate decreased. Foliar nitrogen peaked in medium‐aged forests. Functional diversity was highest in mature forests, even after accounting for differences in stem and species diversity. Our study represents one of the first attempts to document successional changes in New Guinea's lowland forest. We found robust evidence that as succession proceeds, communities occupy a greater range of functional trait space even after controlling for stem and species density. High functional diversity is important for ecological resiliency in the face of global change.     

Principles of Natural Regeneration of Tropical Dry Forests for Restoration

Tropical dry forests are the most threatened tropical terrestrial ecosystem. However, few studies have been conducted on the natural regeneration necessary to restore these forests. We reviewed the ecology of regeneration of tropical dry forests as a tool to restore disturbed lands. Dry forests are characterized by a relatively high number of tree species with small, dry, wind‐dispersed seeds. Over small scales, wind‐dispersed seeds are better able to colonize degraded areas than vertebrate‐dispersed plants. Small seeds and those with low water content are less susceptible to desiccation, which is a major barrier for establishment in open areas. Seeds are available in the soil in the early rainy season to maximize the time to grow. However, highly variable precipitation and frequent dry spells are important sources of mortality in seeds and seedlings. Collecting seeds at the end of the dry season and planting them when soil has sufficient moisture may increase seedling establishment and reduce the time they are exposed to seed predators. Germination and early establishment in the field are favored in shaded sites, which have milder environment and moister soil than open sites during low rainfall periods. Growth of established seedlings, however, is favored in open areas. Therefore, clipping plants around established seedlings may be a good management option to improve growth and survival. Although dry forests have species either resistant to fire or that benefit from it, frequent fires simplify community species composition. Resprouting ability is a noticeable mechanism of regeneration in dry forests and must be considered for restoration. The approach to dry‐forest restoration should be tailored to this ecosystem instead of merely following approaches developed for moister forests.     

Alternative pathways of liana communities in the forests of northwestern Argentina

Liana dynamics in secondary and mature forests are well known in tropical areas dominated by native tree species. Outside the tropics and in secondary forests invaded by exotic species, knowledge is scarce. In this study, we compare liana communities between secondary and mature forests dominated by native species in a subtropical montane area of Sierra de San Javier, Tucuman, Argentina. Additionally, we evaluate changes of liana communities in secondary forests with increasing densities of Ligustrum lucidum and Morus alba, two of the most invasive exotic trees of the area. We surveyed liana species richness and density in three 30-year secondary patches, four 60-year secondary patches, and four mature patches dominated by native tree species, to analyze changes in liana communities with forest age. Within each patch, we sampled 10–25 20 × 20 m quadrats. Additionally, we surveyed liana density and species richness in secondary forest patches with different densities of L. lucidum and M. alba. In native-dominated forests, liana species richness increased and showed a tendency of increasing basal area from 30-year secondary forests to mature forests. Liana density was highly variable, and most of the species were shared between native-dominated secondary and mature forests. Liana density and species richness decreased with L. lucidum density, whereas in secondary forests highly dominated by M. alba, lianas increased in density. Overall, lianas followed different pathways influenced by native forest succession and exotic tree invasions.     

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.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Opposing mechanisms affect taxonomic convergence between tree assemblages during tropical forest succession

Whether successional forests converge towards an equilibrium in species composition remains an elusive question, hampered by high idiosyncrasy in successional dynamics. Based on long‐term tree monitoring in second‐growth (SG) and old‐growth (OG) forests in Costa Rica, we show that patterns of convergence between pairs of forest stands depend upon the relative abundance of species exhibiting distinct responses to the successional gradient. For instance, forest generalists contributed to convergence between SG and OG forests, whereas rare species and old‐growth specialists were a source of divergence. Overall, opposing trends in taxonomic similarity among different subsets of species nullified each other, producing a net outcome of stasis over time. Our results offer an explanation for the limited convergence observed between pairwise communities and suggest that rare species and old‐growth specialists may be prone to dispersal limitation, while the dynamics of generalists and second‐growth specialists are more predictable, enhancing resilience in tropical secondary forests.     

Climate-growth relations of congeneric tree species vary across a tropical vegetation gradient in Brazil

Seasonally dry tropical forests are an important global climatic regulator, a main driver of the global carbon sink dynamics and are predicted to suffer future reductions in their productivity due to climate change. Yet, little is known about how interannual climate variability affects tree growth and how climate-growth responses vary across rainfall gradients in these forests. Here we evaluate changes in climate sensitivity of tree growth along an environmental gradient of seasonally dry tropical vegetation types (evergreen forest – savannah – dry forest) in Northeastern Brazil, using congeneric species of two common neotropical genera: Aspidosperma and Handroanthus. We built tree-ring width chronologies for each species × forest type combinations and explored how growth variability correlated with local (precipitation, temperature) and global (the El Niño Southern Oscillation - ENSO) climatic factors. We also assessed how growth sensitivity to climate and the presence of growth deviations varied along the gradient. Precipitation stimulates tree growth and was the main growth-influencing factor across vegetation types. Trees in the dry forest site showed highest growth sensitivity to interannual variation in precipitation. Temperature and ENSO phenomena correlated negatively with growth and sensitivity to both climatic factors were similar across sites. Negative growth deviations were present and found mostly in the dry-forest species. Our results reveal a dominant effect of precipitation on tree growth in seasonally dry tropical forests and suggest that along the gradient, dry forests are the most sensitivity to drought. These forests may therefore be the most vulnerable to the deleterious effects of future climatic changes. These results highlight the importance of understanding the climatic sensitivity of different tropical forests. This understanding is key to predict the carbon dynamics in tropical regions, and sensitivity differences should be considered when prioritizing conservation measures of seasonally dry topical forests.