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.     

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.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Fingerprinting the impacts of global change on tropical forests

Recent observations of widespread changes in mature tropical forests such as increasing tree growth, recruitment and mortality rates and increasing above-ground biomass suggest that ''global change'' agents may be causing predictable changes in tropical forests. However, consensus over both the robustness of these changes and the environmental drivers that may be causing them is yet to emerge. This paper focuses on the second part of this debate. We review (i) the evidence that the physical, chemical and biological environment that tropical trees grow in has been altered over recent decades across large areas of the tropics, and (ii) the theoretical, experimental and observational evidence regarding the most likely effects of each of these changes on tropical forests. Ten potential widespread drivers of environmental change were identified: temperature, precipitation, solar radiation, climatic extremes (including El Niño-Southern Oscillation events), atmospheric CO2 concentrations, nutrient deposition, O3/acid depositions, hunting, land-use change and increasing liana numbers. We note that each of these environmental changes is expected to leave a unique ''fingerprint'' in tropical forests, as drivers directly force different processes, have different distributions in space and time and may affect some forests more than others (e.g. depending on soil fertility). Thus, in the third part of the paper we present testable a priori predictions of forest responses to assist ecologists in attributing particular changes in forests to particular causes across multiple datasets. Finally, we discuss how these drivers may change in the future and the possible consequences for tropical forests.     

Tropical forest carbon balance in a warmer world: a critical review spanning microbial- to ecosystem-scale processes

Tropical forests play a major role in regulating global carbon (C) fluxes and stocks, and even small changes to C cycling in this productive biome could dramatically affect atmospheric carbon dioxide (CO(2) ) concentrations. Temperature is expected to increase over all land surfaces in the future, yet we have a surprisingly poor understanding of how tropical forests will respond to this significant climatic change. Here we present a contemporary synthesis of the existing data and what they suggest about how tropical forests will respond to increasing temperatures. Our goals were to: (i) determine whether there is enough evidence to support the conclusion that increased temperature will affect tropical forest C balance; (ii) if there is sufficient evidence, determine what direction this effect will take; and, (iii) establish what steps should to be taken to resolve the uncertainties surrounding tropical forest responses to increasing temperatures. We approach these questions from a mass-balance perspective and therefore focus primarily on the effects of temperature on inputs and outputs of C, spanning microbial- to ecosystem-scale responses. We found that, while there is the strong potential for temperature to affect processes related to C cycling and storage in tropical forests, a notable lack of data combined with the physical, biological and chemical diversity of the forests themselves make it difficult to resolve this issue with certainty. We suggest a variety of experimental approaches that could help elucidate how tropical forests will respond to warming, including large-scale in situ manipulation experiments, longer term field experiments, the incorporation of a range of scales in the investigation of warming effects (both spatial and temporal), as well as the inclusion of a diversity of tropical forest sites. Finally, we highlight areas of tropical forest research where notably few data are available, including temperature effects on: nutrient cycling, heterotrophic versus autotrophic respiration, thermal acclimation versus substrate limitation of plant and microbial communities, below-ground C allocation, species composition (plant and microbial), and the hydraulic architecture of roots. Whether or not tropical forests will become a source or a sink of C in a warmer world remains highly uncertain. Given the importance of these ecosystems to the global C budget, resolving this uncertainty is a primary research priority.     

Improving predictions of tropical forest response to climate change through integration of field studies and ecosystem modeling

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species‐specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model‐data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate.     

Sources or sinks? The responses of tropical forests to current and future climate and atmospheric composition.

How tropical rainforests are responding to the ongoing global changes in atmospheric composition and climate is little studied and poorly understood. Although rising atmospheric carbon dioxide (CO2) could enhance forest productivity, increased temperatures and drought are likely to diminish it. The limited field data have produced conflicting views of the net impacts of these changes so far. One set of studies has seemed to point to enhanced carbon uptake; however, questions have arisen about these findings, and recent experiments with tropical forest trees indicate carbon saturation of canopy leaves and no biomass increase under enhanced CO2. Other field observations indicate decreased forest productivity and increased tree mortality in recent years of peak temperatures and drought (strong El Niño episodes). To determine current climatic responses of forests around the world tropics will require careful annual monitoring of ecosystem performance in representative forests. To develop the necessary process-level understanding of these responses will require intensified experimentation at the whole-tree and stand levels. Finally, a more complete understanding of tropical rainforest carbon cycling is needed for determining whether these ecosystems are carbon sinks or sources now, and how this status might change during the next century.     

Assessing biome boundary shifts under climate change scenarios in India

Climate change and its cascading impacts are being increasingly recognized as a major challenge across the globe. Climate is one of the most critical factors affecting biomes and their distribution. The present study assessed shifts in biome types of India using the conceptual framework of Holdridge life zone (HLZ) model, minimum distance classifier and climatic datasets to assess the distribution pattern of potential biomes under climate change scenarios in India. Modelling was conducted on the entire region of India using various combinations; (i) current climate scenario, and, (ii) increased temperature and precipitation scenario. The geographical analysis identifies nineteen (19) HLZs in the Indian sub-continent; seven (7) biomes and nineteen (19) sub-biomes. The overall accuracy and kappa coefficient of the biome map prepared for current climate scenario was 82.73% and 0.75, respectively. With the changes in increasing temperature and precipitation scenario, the modelling results predict significant decrease in the area cover for tropical deserts (plains), tropical desert scrubs (lower montane), tropical moist forests (lower montane) and tropical wet forests (lower montane). Along with these changes, there have been substantial increases in the area cover for tropical dry forests (plains) and tropical very dry forests (plains), especially in central and southern India. The results show shifts from very dry tundra (alvar) to dry tundra (alpine) and moist tundra (alpine) and in some places tropical moist forests (sub-alpine) as well. In central India, decrease in tropical moist forests (lower montane) has been observed, while an increase in the area cover of tropical rain forests (plains) in northeastern India has been observed. It is important to understand the impacts and vulnerabilities of projected climate change on forest ecosystems so that better management and conservation strategies can be adopted for biodiversity and forest dependent communities. The knowledge of impact mechanisms will identify adaptation strategies for some conditions which will help in decreasing the susceptibility to anticipated climate change in the forest sector.     

Resilience of tropical dry forests – a meta‐analysis of changes in species diversity and composition during secondary succession

Assessing the recovery of species diversity and composition after major disturbance is key to understanding the resilience of tropical forests through successional processes, and its importance for biodiversity conservation. Despite the specific abiotic environment and ecological processes of tropical dry forests, secondary succession has received less attention in this biome than others and changes in species diversity and composition have never been synthesised in a systematic and quantitative review. This study aims to assess in tropical dry forests 1) the directionality of change in species richness and evenness during secondary succession, 2) the convergence of species composition towards that of old‐growth forest and 3) the importance of the previous land use, precipitation regime and water availability in influencing the direction and rate of change. We conducted meta‐analyses of the rate of change in species richness, evenness and composition indices with succession in 13 tropical dry forest chronosequences. Species richness increased with succession, showing a gradual accumulation of species, as did Shannon evenness index. The similarity in species composition of successional forests with old‐growth forests increased with succession, yet at a low rate. Tropical dry forests therefore do show resilience of species composition but it may never reach that of old‐growth forests. We found no significant differences in rates of change between different previous land uses, precipitation regimes or water availability. Our results show high resilience of tropical dry forests in term of species richness but a slow recovery of species composition. They highlight the need for further research on secondary succession in this biome and better understanding of impacts of previous land‐use and landscape‐scale patterns. Synthesis Secondary forests account for an increasing proportion of remaining tropical forest. Assessing their resilience is key to conservation of their biodiversity. Our study is the first meta‐analysis of species changes during succession focussing on tropical dry forests, a highly threatened yet understudied biome. We show a gradual species accumulation and convergence of composition towards that of old‐growth forests. While secondary tropical dry forests offer good potential for biodiversity conservation, their capacity for recovery at a sufficient rate to match threats is uncertain. Further research on this biome is needed to understand the effect of land use history and landscape processes.     

Differences in epiphyte biomass and community composition along landscape and within-crown spatial scales

Vascular epiphytes contribute to the structural, compositional, and functional complexity of tropical montane cloud forests because of their high biomass, diversity, and ability to intercept and retain water and nutrients from atmospheric sources. However, human-caused climate change and forest-to-pasture conversion are rapidly altering tropical montane cloud forests. Epiphyte communities may be particularly vulnerable to these changes because of their dependence on direct atmospheric inputs and host trees for survival. In Monteverde, Costa Rica, we measured vascular epiphyte biomass, community composition, and richness at two spatial scales: (1) along an elevation gradient spanning premontane forests to montane cloud forests and (2) within trees along branches from inner to outer crown positions. We also compared epiphyte biomass and distribution at these scales between two different land-cover types, comparing trees in closed canopy forest to isolated trees in pastures. An ordination of epiphyte communities at the level of trees grouped forested sites above versus below the cloud base, and separated forest versus pasture trees. Species richness increased with increasing elevation and decreased from inner to outer branch positions. Although richness did not differ between land-cover types, there were significant differences in community composition. The variability in epiphyte community organization between the two spatial scales and between land-cover types underscores the potential complexity of epiphyte responses to climate and land-cover changes.     

Terrestrial transects for global change research

The International Geosphere-Biosphere Program has proposed a set of large-scale terrestrial transects to study the effects of changes in climate, land use, and atmospheric composition (global change) on biogeochemistry, surface-atmosphere exchange, and vegetation dynamics of terrestrial ecosystems. The transects ( 1000 km) will be located along existing environmental and land use intensity gradients that span transitions between biomes in regions likely to be widely affected by forcing from components of global change or where the impacts of global change are likely to feed back to affect atmospheric, climatic, or hydrologic systems. Experimental studies on the transects will examine short-term changes in ecosystem function and biosphere-atmosphere interaction in response to variation in primary controlling variables. A hierarchy of modeling approaches will develop predictions of long-term changes in biome boundaries and vegetation distribution. The proposed initial set of IGBP terrestrial transects are located in four key regions: (1) humid tropical forests undergoing land use change, (2) high latitudes including the transition from boreal forest to tundra, (3) semi-arid tropical regions including transitions from dry forest to shrublands and savannas, and (4) mid latitude semi-arid regions encompassing transitions from shrubland or grassland to forests. We discuss here the rationale and general research design of transect studies proposed for each of these priority regions.GCTE Focus 1 Office     

Tree diversity,composition, forest structure and aboveground biomass dynamics after single and repeated fire in a Bornean rain forest

Forest fires remain a devastating phenomenon in the tropics that not only affect forest structure and biodiversity, but also contribute significantly to atmospheric CO2. Fire used to be extremely rare in tropical forests, leaving ample time for forests to regenerate to pre-fire conditions. In recent decades, however, tropical forest fires occur more frequently and at larger spatial scales than they used to. We studied forest structure, tree species diversity, tree species composition, and aboveground biomass during the first 7 years since fire in unburned, once burned and twice burned forest of eastern Borneo to determine the rate of recovery of these forests. We paid special attention to changes in the tree species composition during burned forest regeneration because we expect the long-term recovery of aboveground biomass and ecosystem functions in burned forests to largely depend on the successful regeneration of the pre-fire, heavy-wood, species composition. We found that forest structure (canopy openness, leaf area index, herb cover, and stem density) is strongly affected by fire but shows quick recovery. However, species composition shows no or limited recovery and aboveground biomass, which is greatly reduced by fire, continues to be low or decline up to 7 years after fire. Consequently, large amounts of the C released to the atmosphere by fire will not be recaptured by the burned forest ecosystem in the near future. We also observed that repeated fire, with an inter-fire interval of 15 years, does not necessarily lead to a huge deterioration in the regeneration potential of tropical forest. We conclude that burned forests are valuable and should be conserved and that long-term monitoring programs in secondary forests are necessary to determine their recovery rates, especially in relation to aboveground biomass accumulation.     

Dendrochronological potential of four neotropical dry-forest tree species: Climate-growth correlations in northeast Brazil

Tropical dry forests (TDF) are highly important tropical forest ecosystems. Yet, these forests are highly threatened, usually neglected and only poorly studied. Understanding the long-term influences of environmental conditions on tree growth in these forests is crucial to understand the functioning, carbon dynamics and potential responses to future climate change of these forests. Dendrochronology can be used as a tool to provide these insights but has only scantly been applied in (dry) tropical forests. Here we evaluate the dendrochronological potential of four Caatinga neotropical dry forest tree species – Aspidosperma pyrifolium, Ziziphus joazeiro, Tabebuia aurea, and Libidibia ferrea – collected in two locations in northeastern Brazil (Sergipe state). We provide an anatomical characterization of the ring boundaries for the four species and investigate correlations of their growth with local and regional climatic variables. All four species form annual rings and show high inter-correlation (up to 0.806) and sensitivity (up to 0.565). Growth of all species correlated with local precipitation as well as with sea-surface temperatures in the tropical Atlantic and/or tropical Pacific oceans. We also show teleconnections between growth and the El Niño South Oscillation. The strong dependence of tree on precipitation is worrisome, considering that climate change scenarios forecast increased drought conditions in the Caatinga dry forest. Including more species and expanding dendrochronological studies to more areas would greatly improve our understanding of tree growth and functioning in TDFs. This type of knowledge is essential to assist the conservation, management and restoration of these critical tropical ecosystems.     

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。     

The changing ecology of tropical forests

The threat to tropical forests is often gauged in terms of deforestation rates and the total area remaining. Recently, however, there has been a growing realization that forest can appear intact on a satellite image yet be biologically degraded or vulnerable to degradation. The array of direct threats to humid tropical forest biodiversity, in addition to deforestation, includes: selective extraction of plants; selective extraction of animals; biological invasion; fragmentation; climate change; changing atmospheric composition; and increasing tree turnover rates. The threats are linked to one another by a poorly understood network of causality and feedback effects. Moreover, their potential impacts on forest biodiversity are hard to assess because each threat is as likely to precipitate indirect effects as direct effects, and because several threats are likely to interact synergistically with one another. In spite of the uncertainties, it is clear that the biological health of tropical forests can become seriously degraded as a result of these threats, and it is unlikely that any tropical forest will escape significant ecological changes. Some groups of plants and animals are likely to benefit at the expense of others. Species diversity is expected to decline as a consequence of the changes in forest ecology. In the 21st century scientists and conservationists will be increasingly challenged to monitor, understand, prevent and head off these threats.     

Pattern of changes in species diversity,structure and dynamics of forest ecosystems along latitudinal gradients in East Asia

We examined effects of seasonality of climate and dominant life form (evergreen/deciduous, broad-leaf/coniferous) together with energy condition on species diversity, forest structure, forest dynamics, and productivity of forest ecosystems by comparing the patterns of changes in these ecosystem attributes along altitudinal gradients in tropical regions without seasonality and along a latitudinal gradient from tropical to temperate regions in humid East Asia. We used warmth index (temperature sum during growing season, WI) as an index of energy condition common to both altitudinal and latitudinal gradients. There were apparent differences in patterns of changes in the ecosystem attributes in relation to WI among four forest formations that were classified according to dominant life form and climatic zone (tropical/temperate). Many of the ecosystem attributes—Fishers alpha of species-diversity indices, maximum tree height and stem density, productivity [increment rate of aboveground biomass (AGB)], and population and biomass turnover rates—changed sharply with WI in tropical and temperate evergreen broad-leaved forests, but did not change linearly or changed only loosely with WI in temperate deciduous broad-leaved and evergreen coniferous forests. Values of these ecosystem attributes in temperate deciduous broad-leaved and evergreen coniferous forests were higher (stem density was lower) than those in tropical and temperate evergreen broad-leaved forests under colder conditions (WI below 100°C). Present results indicate that seasonality of climate and resultant change in dominant life form work to buffer the effects of energy reduction on ecosystem attributes along latitudinal gradients.     

Drought and presence of ants can influence hemiptera in tropical leaf litter

Climate change is predicted to impact tropical rain forests, with droughts becoming more frequent and more severe in some regions. We currently have a poor understanding of how increased drought will change the functioning of tropical rain forest. In particular, tropical rain forest invertebrates, which are numerous and biologically important, may respond to drought in different ways across trophic levels. Ants are a diverse group that carry out important ecosystem processes, shaping ecosystem structure and function through predation and competition, which can influence multiple trophic levels. Hemiptera are a mega-diverse order, abundant in tropical rain forests and are ecologically important. To understand the roles of ants in exerting predation and competition pressure on invertebrates in tropical rain forests during drought and a post-drought period, we established a large-scale ecosystem manipulation experiment in Maliau Basin Conservation Area in Malaysian Borneo, suppressing the activity of ants on four 0.25 ha plots over a two-year period. We sampled hemipterans found in the leaf litter during a drought (July 2015) and a post-drought period (September 2016) period. We found significant shifts in the assemblage of hemipterans sampled from the leaf litter following ant suppression. Specifically, for ant-suppression plots, the species richness and abundance of herbivorous hemipterans increased only during the post-drought period. For predatory hemipterans, abundance increased with ant-suppression regardless of drought conditions, and we found marginal evidence for a species richness increase during the post-drought period with little or no change in the drought period. These results illustrate how ants in tropical forests structure invertebrate communities and how these effects may vary with climatic variation.     

Tropical rainforest canopies and climate change

Abstract There is less certainty about the impact of climate change on tropical rainforests than on temperate forests because of the comparative lack of background data and because few large scale experiments have been, and are being, carried out in tropical rainforests. Many of the factors critical to the future of tropical rainforests concern canopies, the key processes that take place there, and the roles and interactions of canopy biodiversity. In particular there are almost no data on how forest canopies and processes are changing with increased carbon dioxide levels. The implications of elevated carbon dioxide, climatic stress and related changes in water‐use efficiency, nutrient availability and other such changes are discussed particularly with references to Australia's tropical rainforests.     

Tropical forest composition and function across space and time: Insights from diverse gradients in Área de Conservación Guanacaste

Environmental gradients have played a pivotal role in the history and development of plant ecology and are useful for testing ecological and evolutionary theory. Área de Conservación Guanacaste is a spatio-temporal mosaic of forests that have evolved continuously across elevation, topography, soil types, succession, and annual and inter-annual climatic change. Studies of plant ecology across diverse gradients of ACG have shaped functional ecology, successional theory, community assembly, plant–herbivore interactions, among many other fields. In this review, we synthesize the, perhaps overlooked, role plant ecological studies of ACG have had on our understanding of tropical forest dynamics. We outline present-day processes that will have major impacts on forest dynamics of ACG in the future and highlight how ACG will continue to shape future research priorities in plant ecology.     

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。