Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

The negative effect of lianas on tree growth varies with tree species and season

Lianas reduce tree growth, reproduction, and survival in tropical forests. Liana competition can be particularly intense in isolated forest fragments, where liana densities are high, and thus, host tree infestation is common. Furthermore, lianas appear to grow particularly well during seasonal drought, when they may compete particularly intensely with trees. Few studies, however, have experimentally quantified the seasonal effects of liana competition on multiple tree species in tropical forests. We used a liana removal experiment in a forest fragment in southeastern Brazil to test whether the effects of lianas on tree growth vary with season and tree species identity. We conducted monthly diameter measurements using dendrometer bands on 88 individuals of five tree species for 24 months. We found that lianas had a stronger negative effect on some tree species during the wet season compared to the dry season. Furthermore, lianas significantly reduced the diameter growth of two tree species but had no effect on the other three tree species. The strong negative effect of lianas on some trees, particularly during the wet season, indicates that the effect of lianas on trees varies both seasonally and with tree species identity. Abstract in Portuguese is available with online material.     

Functional traits of lianas in an Australian lowland rainforest align with post‐disturbance rather than dry season advantage

Lianas are an important component of tropical forests; they alter tree mortality and recruitment and impact biogeochemical cycling. Recent evidence suggests that the abundance of lianas in tropical forests is increasing. To understand and predict the effect of lianas on ecosystem processes in tropical forests, it is important to understand the mechanisms through which they compete with trees. In this study, we investigated the functional traits of lianas and trees in a lowland tropical forest in northeast Queensland, Australia. The site is located at 16.1° south latitude and experiences significant seasonality in rainfall, with pronounced wet and dry seasons. It is also subject to relatively frequent disturbance by cyclones. We asked the question of whether the canopy liana community at this site would display functional traits consistent with a competitive advantage over trees in response to disturbance, or in response to dry season water stress. We found that traits that we considered indicative of a dry season advantage (xylem water δ¹⁸O as an indicator of rooting depth; leaf and stem tissue δ¹³C and instantaneous gas exchange as measures of water‐use efficiency) did not differ between canopy lianas and canopy trees. On the other hand, lianas differed from trees in traits that should confer an advantage in response to disturbance (low wood density; low leaf dry matter content; high leaf N concentration; high mass‐based photosynthetic rates). We conclude that the liana community at the study site expressed functional traits geared towards rapid resource acquisition and growth in response to disturbance, rather than outcompeting trees during periods of water stress. These results contribute to a body of literature which will be useful for parameterising a liana functional type in ecosystem models.     

No evidence that elevated CO2 gives tropical lianas an advantage over tropical trees

Recent studies indicate that lianas are increasing in size and abundance relative to trees in neotropical forests. As a result, forest dynamics and carbon balance may be altered through liana‐induced suppression of tree growth and increases in tree mortality. Increasing atmospheric CO₂ is hypothesized to be responsible for the increase in neotropical lianas, yet no study has directly compared the relative response of tropical lianas and trees to elevated CO₂. We explicitly tested whether tropical lianas had a larger response to elevated CO₂ than co‐occurring tropical trees and whether seasonal drought alters the response of either growth form. In two experiments conducted in central Panama, one spanning both wet and dry seasons and one restricted to the dry season, we grew liana (n = 12) and tree (n = 10) species in open‐top growth chambers maintained at ambient or twice‐ambient CO₂ levels. Seedlings of eight individuals (four lianas, four trees) were grown in the ground in each chamber for at least 3 months during each season. We found that both liana and tree seedlings had a significant and positive response to elevated CO₂ (in biomass, leaf area, leaf mass per area, and photosynthesis), but that the relative response to elevated CO₂ for all variables was not significantly greater for lianas than trees regardless of the season. The lack of differences in the relative response between growth forms does not support the hypothesis that elevated CO₂ is responsible for increasing liana size and abundance across the neotropics.     

Short and Long-Term Soil Moisture Effects of Liana Removal in a Seasonally Moist Tropical Forest

Lianas (woody vines) are particularly abundant in tropical forests, and their abundance is increasing in the neotropics. Lianas can compete intensely with trees for above- and belowground resources, including water. As tropical forests experience longer and more intense dry seasons, competition for water is likely to intensify. However, we lack an understanding of how liana abundance affects soil moisture and hence competition with trees for water in tropical forests. To address this critical knowledge gap, we conducted a large-scale liana removal experiment in a seasonal tropical moist forest in central Panama. We monitored shallow and deep soil moisture over the course of three years to assess the effects of lianas in eight 0.64 ha removal plots and eight control plots. Liana removal caused short-term effects in surface soils. Surface soils (10 cm depth) in removal plots dried more slowly during dry periods and accumulated water more slowly after rainfall events. These effects disappeared within four months of the removal treatment. In deeper soils (40 cm depth), liana removal resulted in a multi-year trend towards 5–25% higher soil moisture during the dry seasons with the largest significant effects occurring in the dry season of the third year following treatment. Liana removal did not affect surface soil temperature. Multiple and mutually occurring mechanisms may be responsible for the effects of liana removal on soil moisture, including competition with trees, and altered microclimate, and soil structure. These results indicate that lianas influence hydrologic processes, which may affect tree community dynamics and forest carbon cycling.     

Why is liana abundance low in semiarid climates?

Lianas are abundant in seasonal tropical forests, where they avoid seasonal water stress presumably by accessing deep‐soil water reserves. Although lianas are favoured in seasonal environments, their occurrence and abundance are low in semiarid environments. We hypothesized that lianas do not tolerate the great water shortage in the soil and air characteristic of semiarid environments, which would increase the risk of embolism. We compared the rooting depth of coarse roots, leaf dynamics, leaf water potential (ψ_leaf), embolism resistance (P₅₀) and lethal levels of embolism (P₈₈) between congeneric lianas that occur with different abundances in two semiarid sites differing in soil characteristics and vapour pressure deficit in the air (VPD_air). Regardless of soil texture and depth, water availability was restricted to the rainy season. All liana species were drought deciduous and had superficial coarse roots (not deeper than 35 cm). P₅₀ varied from ?1.8 to ?2.49 MPa, and all species operated under narrow safety margins against catastrophic (P₅₀) and irreversible hydraulic failure (P₈₈), even during the rainy season. In short, lianas that occur in semiarid environments have lower resistance to cavitation and limit carbon fixation to the rainy season because of leaf fall in the early dry season. We suggest that leaf shedding and shallow roots impairing carbon gain and growth in the dry season may explain why liana abundance is lower in semiarid than in other seasonally dry environments.     

A mechanistic explanation for global patterns of liana abundance and distribution

One of the main goals in ecology is determining the mechanisms that control the abundance and distribution of organisms. Using data from 69 tropical forests worldwide, I demonstrate that liana (woody vine) abundance is correlated negatively with mean annual precipitation and positively with seasonality, a pattern precisely the opposite of most other plant types. I propose a general mechanistic hypothesis integrating both ecological and ecophysiological approaches to explain this pattern. Specifically, the deep root and efficient vascular systems of lianas enable them to suffer less water stress during seasonal droughts while many competitors are dormant, giving lianas a competitive advantage during the dry season. Testing this hypothesis in central Panama, I found that lianas grew approximately seven times more in height than did trees during the dry season but only twice as much during the wet season. Over time, this dry season advantage may allow lianas to increase in abundance in seasonal forests. In aseasonal wet forests, however, lianas gain no such advantage because competing plants are rarely limited by water. I extend this theory to account for the local, within-forest increase in liana abundance in response to disturbance as well as the conspicuous decrease in liana abundance at high latitudes.     

Contrasting leaf characteristics of trees and lianas in secondary and mature forests in southwestern China

We compared variation in sun-canopy leaf anatomy, morphology and photosynthetic rates of coexisting woody species (trees and lianas) in an 8-year-old secondary forest (SF) and mature forest (MF) in the wet season in Xishuangbanna, SW China. Variability of leaf traits of 66 species within growth-form groups in each forest was quantified using coefficients of variation (CV). For the mean values, the woody species in the SF had significantly higher leaf thickness and stomatal density, but lower nonmesophyll/mesophyll ratios than those in the MF. The average leaf area and leaf mass area (LMA) in the studied woody species did not change greatly during the successional process, but differed significantly between the growth forms, with trees having higher values than lianas. The light-saturated photosynthetic rate per unit leaf area (A _a) of the woody species in the SF ranged from 11.2 to 34.5 μmol m⁻² s⁻¹, similarly to pioneer tree species from literature data in southeast Asia. The A _a and photosynthetic nitrogen-use efficiency (PNUE) were significantly higher than those in the MF; whereas A _a in the MF ranged between 9 to 21 μmol m⁻² s⁻¹, with similar values between lianas and trees. For all woody species in both SF and MF, there were no significant differences in the average values of the CV of all measured variables for both lianas and trees. However, considerable variation in leaf anatomy, morphology, and photosynthetic rates within both growth forms and forests existed, as well as substantial variation in leaf size and stomatal density. We concluded that the tropical woody species formed a heterogeneous functional group in terms of leaf morphology and physiology in both secondary and mature forests.     

Leaf functional traits of Neotropical savanna trees in relation to seasonal water deficit

The seasonal savannas (cerrados) of Central Brazil are characterized by a large diversity of evergreen and deciduous trees, which do not show a clear differentiation in terms of active rooting depth. Irrespective of the depth of the root system, expansion of new foliage in deciduous species occurs at the end of the dry season. In this study, we examined a suite of leaf traits related to C assimilation, water and nutrients (N, P) in five deciduous and six evergreen trees that were among the dominant families of cerrado vegetation. Maximum CO₂ assimilation on a mass basis (A_mass) was significantly correlated with leaf N and P, and specific leaf area (SLA; leaf area per unit of leaf mass). The highest leaf concentrations of both nutrients were measured in the newly mature leaves of deciduous species at the end of the dry period. The differences in terms of leaf N and P between evergreen and deciduous species decreased during the wet season. Deciduous species also invested less in the production of non-photosynthetic leaf tissues and produced leaves with higher SLA and maintained higher water use efficiency. Thus, deciduous species compensated for their shorter leaf payback period by maintaining higher potential payback capacity (higher values of A_mass) and lower leaf construction costs (higher SLA). Their short leafless period and the capacity to flush by the end of the dry season may also contribute to offset the longer payback period of evergreen species, although it may involve the higher cost of maintaining a deep-root system or a tight control of plant water balance in the shallow-rooted ones.     

Community and ecosystem ramifications of increasing lianas in neotropical forests

Lianas (woody vines) are increasing in neotropical forests, representing one of the first large-scale structural changes documented for these important ecosystems. The potential ramifications of increasing lianas are huge, as lianas alter both tropical forest diversity and ecosystem functioning. At the community level, lianas affect tree species co-existence and diversity by competing more intensely with some tree species than others, and thus will likely alter tree species composition. At the ecosystem level, lianas affect forest carbon and nutrient storage and fluxes. A decrease in forest carbon storage and sequestration may be the most important ramification of liana increases. Lianas reduce tree growth and increase tree mortality—thus reducing forest-level carbon storage. The increase in lianas, which have much less wood than trees, compensates only partially for the amount of carbon lost in the displaced trees. Because tropical forests contribute approximately one-third of global terrestrial carbon stocks and net primary productivity, the effect of increasing lianas for tropical forest carbon cycles may have serious repercussions at the global scale.Key words: carbon cycle, CO₂, disturbance, global change, land use change, liana increases, structural changes, tropical forestsTropical forests contain most of the earth''s plant species and contribute more to carbon storage in the form of above ground biomass than any other terrestrial ecosystem. Temperate and boreal forests are changing rapidly in response to global anthropogenic drivers. Similar large-scale changes are now being detected in tropical forests. One of the largest contemporary changes in tropical forests is an increase in lianas (woody vines),¹ which could have serious consequences for tree species diversity and composition, as well as the reducing capacity of tropical forests to store carbon.^1–3     

What controls liana success in Neotropical forests?

Aim   We seek to determine the factors which control the success of lianas across macroecological gradients. Lianas have a strong impact on the growth, mortality and biomass of tropical trees, and are reported to be increasing in dominance, so understanding their behaviour is important from the perspectives of both ecological and global change.
Location   Lowland and montane Neotropical forests.
Methods   Using 65 standardized samples of lianas (≥ 2.5 cm diameter) from across the Neotropics, we attempted to account for characteristics of both the environment and the forest in explaining macroecological variation in liana success in Neotropical forests, using regression analyses and structural equation modelling.
Results   We found that both liana density and basal area were unrelated to mean annual precipitation, dry season length or soil variables, except for a weak effect of mean annual precipitation on liana basal area. Structural characteristics of the forest explained more of the variation in liana density and basal area than the physical environment. More disturbed forests generally tended to have a higher liana density. Liana basal area, however, was highest in undisturbed forests.
Main conclusions   The availability of host trees and their characteristics may be more important than the direct effects of the physical environment in controlling the success of lianas in Neotropical forests. Changes to the tropical climate in the coming century may not strongly affect lianas directly, but could have very substantial indirect effects via changes in tree community structure and dynamics.     

Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

Hydraulic properties and photosynthetic rates in co-occurring lianas and trees in a seasonal tropical rainforest in southwestern China

In this study, we examined wood anatomy, hydraulic properties, photosynthetic rate, and water status and osmotic regulation in three liana species and three tree species co-occurring in a seasonal tropical rain forest. Our results showed that the three liana species had larger vessel diameter, lower sapwood density, and consequently higher branch sapwood specific hydraulic conductivity (K _S) than the three tree species. Across species, K _S was positively correlated with leaf nitrogen concentration and maximum net CO₂ assimilation rate. However, it was also positively correlated with xylem water potential at 50% loss of hydraulic conductivity, indicating a trade-off between hydraulic efficiency and safety. Compared to the tree species, the liana species had higher predawn leaf water potential and lower osmotic adjustment in the dry season. The combination of more efficient water transport, higher photosynthetic rates, and their ability to access to more reliable water source at deeper soil layers in the dry season in the lianas should contribute to their fast growth.     

热带雨林木质藤本植物叶片性状及其关联

热带雨林中木质藤本植物较为丰富。随着全球气候变化加剧,木质藤本植物的丰富度具有不断增加的趋势,有可能对热带森林的结构、功能和动态产生重要影响。然而,目前对木质藤本响应环境变化的机制所知甚少。本研究以13个科20种热带雨林常见木质藤本植物为材料,测定了冠层叶片的17个形态特征及结构性状,并分析了性状间的相互关系。结果表明,叶片相对含水量的种间变异最小（变异系数为5%）,而上表皮厚度的种间变异最大（变异系数为80%）,其它性状的种间变异系数为24%~61%。木质藤本植物的叶脉密度、叶片密度均与气孔密度呈显著正相关,叶片干物质含量与比叶面积呈显著负相关。与相同生境的树木相比,木质藤本的叶面积更小、气孔密度和叶片密度更低、比叶面积更高,但两种植物类群的叶片横切面组织结构厚度无显著差异。研究结果对理解木质藤本植物的生态适应性具有重要意义。     

Effect of liana cutting on water potential and growth of adult<Emphasis Type="Italic"> Senna multijuga</Emphasis> (Caesalpinioideae) trees in a Bolivian tropical forest

Lianas, or woody climbing plants, are a major constituent of seasonally dry tropical forests, and are thought to impact negatively their host trees. In this study we evaluated whether liana presence was associated with reduced leaf water potentials and growth in adult Senna multijuga trees during the dry season in a lowland Bolivian forest. We used leaf water potentials in trees as a first approach to assess trees’ water status, under the assumption that leaf water potentials become more negative when water losses (via transpiration) exceed gains (by uptake). We measured relative growth in girth at 1.5 m height (gbh) to quantify tree growth. At the beginning of the 1996 dry season (early June), we selected 20 S. multijuga trees 10–20 cm dbh, and measured their gbh. We also recorded pre-dawn and mid-day leaf water potentials in these trees. In ten experimental trees all lianas were then cut, while the remaining trees were used as controls. Pre-dawn and mid-day water potentials were re-measured 1 day after liana-cutting, and then every week in all trees for 1 month and then at 3 and 5 months, until the beginning of the next rainy season (November); gbh was measured again in July 1997 to estimate relative growth rate. Liana removal was associated with less negative pre-dawn (–0.3 vs –0.4 MPa) and mid-day (–0.5 vs –0.7 MPa) water potentials in trees during the dry season. This difference appeared as early as 1 day after cutting, and disappeared once the rainy season began. Liana-cut trees grew more (0.4 mm/mm year) than liana-uncut trees (0.2 mm/mm year). These findings suggest that lianas may interfere with water availability to these trees during the dry season, and may also hinder tree growth. Received: 16 November 1999 / Accepted: 23 March 2000     

Liana Abundance,Diversity, and Distribution on Barro Colorado Island,Panama

Lianas are a key component of tropical forests; however, most surveys are too small to accurately quantify liana community composition, diversity, abundance, and spatial distribution – critical components for measuring the contribution of lianas to forest processes. In 2007, we tagged, mapped, measured the diameter, and identified all lianas ≥1 cm rooted in a 50-ha plot on Barro Colorado Island, Panama (BCI). We calculated liana density, basal area, and species richness for both independently rooted lianas and all rooted liana stems (genets plus clones). We compared spatial aggregation patterns of liana and tree species, and among liana species that varied in the amount of clonal reproduction. We also tested whether liana and tree densities have increased on BCI compared to surveys conducted 30-years earlier. This study represents the most comprehensive spatially contiguous sampling of lianas ever conducted and, over the 50 ha area, we found 67,447 rooted liana stems comprising 162 species. Rooted lianas composed nearly 25% of the woody stems (trees and lianas), 35% of woody species richness, and 3% of woody basal area. Lianas were spatially aggregated within the 50-ha plot and the liana species with the highest proportion of clonal stems more spatially aggregated than the least clonal species, possibly indicating clonal stem recruitment following canopy disturbance. Over the past 30 years, liana density increased by 75% for stems ≥1 cm diameter and nearly 140% for stems ≥5 cm diameter, while tree density on BCI decreased 11.5%; a finding consistent with other neotropical forests. Our data confirm that lianas contribute substantially to tropical forest stem density and diversity, they have highly clumped distributions that appear to be driven by clonal stem recruitment into treefall gaps, and they are increasing relative to trees, thus indicating that lianas will play a greater role in the future dynamics of BCI and other neotropical forests.     

Liana habitat associations and community structure in a Bornean lowland tropical forest

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha⁻¹, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher’s diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial > sandstone hill > kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities.     

广西弄岗五桠果叶木姜子群落木质藤本空间格局及其与树种间的关系

木质藤本是生物多样性的重要组成,木质藤本通过影响支持木进而影响群落的结构和功能,但在生物多样性丰富的北热带喀斯特森林中,木质藤本与支持木的关系鲜为人知。以喀斯特季节性雨林的五桠果叶木姜子（Litsea dilleniifolia）群落为研究对象,对木质藤本的密度、分布格局及其与主要树种的关系进行调查研究,分析木质藤本对树木的影响。结果显示：（1）五桠果叶木姜子群落内木质藤本平均密度为0.0913株/m²,木质藤本在0-20m空间尺度整体表现为聚集分布,且随着尺度增大,聚集强度逐渐减弱;不同径级木质藤本在不同尺度上的分布格局不同。（2）木质藤本对不同径级、不同种类、不同聚集强度的支持木选择表现以下体征：随着支持木径级增加,木质藤本攀附的比例和每木藤本数有增加趋势,且木质藤本胸径与支持木胸径呈极显著正相关;附藤率较高的支持木有紫葳科（Bignoniaceae）种类和东京桐（Deutzianthus tonkinensis）,单木附藤数量多的是南方紫金牛（Ardisia thyrsiflora）;物种的聚集强度与附藤率、附藤数量呈负相关。（3）木质藤本的密度与支持木死亡率关系不显著,而物种的附藤率与死亡率呈极显著负相关。以上结果表明,木质藤本密度在原生性喀斯特季节性雨林中并不高,且木质藤本对支持木具有选择性,但其对五桠果叶木姜子群落的死亡率并未产生显著影响。该研究可为喀斯特原生性季节性雨林的物种共存、极小植物种群保育提供理论依据,也可为石漠化区域的植被修复提供科学参考。     

Leaf attributes and tree growth in a tropical dry forest

Questions: How are leaf attributes and relative growth rate (RGR) of the dominant tree species of tropical deciduous forest (TDF) affected by seasonal changes in soil moisture content (SMC)? What is the relationship of functional attributes with each other? Can leaf attributes singly or in combination predict the growth rate of tree species of TDF? Location: Sonebhadra district of Uttar Pradesh, India. Methods: Eight leaf attributes, specific leaf area (SLA); leaf carbon concentration (LCC); leaf nitrogen concentration (LNC); leaf phosphorus concentration (LPC); chlorophyll concentration (Chl), mass‐based stomatal conductance (Gs_mass); mass based photosynthetic rate (A_mass); intrinsic water use efficiency (WUEi); and relative growth rate (RGR), of six dominant tree species of a dry tropical forest on four sites were analysed for species, site and season effects over a 2‐year period. Step‐wise multiple regression was performed for predicting RGR from mean values of SMC and leaf attributes. Path analysis was used to determine which leaf attributes influence RGR directly and which indirectly. Results: Species differed significantly in terms of all leaf attributes and RGR. The response of species varied across sites and seasons. The attributes were positively interrelated, except for WUEi, which was negatively related to all other attributes. The positive correlation was strongest between Gs_mass and A_mass and the negative correlation was strongest between Gs_mass and WUEi. Differences in RGR due to site were not significant when soil moisture was controlled, but differences due to season remained significant. The attributes showed plasticity across moisture gradients, which differed among attributes and species. Gs_mass was the most plastic attribute. Among the six species, Terminalia tomentosa exhibited the greatest plasticity in six functional attributes. In the step‐wise multiple regression, A_mass, SLA and Chl among leaf attributes and SMC among environmental factors influenced the RGR of tree species. Path analysis indicated the importance of SLA, LNC, Chl and A_mass in determining RGR. Conclusion: A _mass, SMC, SLA and Chl in combination can be used to predict RGR but could explain only three‐quarters of the variability in RGR, indicating that other traits/factors, not studied here, are also important in modulating growth of tropical trees. RGR of tree species in the dry tropical environment is determined by soil moisture, whereas the response of mature trees of different species is modulated by alterations in key functional attributes such as SLA, LNC and Chl.     

森林木质藤本数量过度增长的机制与生态效应

木质藤本是维持森林物种多样性的组成部分。近年来，热带、亚热带森林尤其是次生林内的木质藤本数量的过度增长给森林的恢复和健康发展造成了威胁。至今，国内尚少有研究对森林木质藤本过多的现象、增长机制与生态效应进行综合认识。该文梳理国内外相关文献，从木质藤本数量增长的机制与生态效应进行分析和总结，综合相关研究认为：(1)木质藤本数量的增长与干旱化加剧、大气CO₂浓度上升、自然干扰和森林破碎化有关，在环境变化的情况下，木质藤本在形态、行为、生理等方面比树木更具优势，表现为更快的生长速率、更强的繁殖力和可塑性以及资源利用效率。(2)木质藤本主要通过遮荫胁迫、资源竞争和机械压力与损伤等方式对树木造成影响。(3)木质藤本过度增长在个体水平上会阻碍树木生长、生殖并引起树木死亡，在群落水平上会改变物种组成、降低多样性，在生态系统水平上会降低森林碳储量，改变碳、矿质养分和水分循环过程等。因此，建议结合野外长期监测与控制实验开展木质藤本数量动态与环境变化关系、森林干扰对木质藤本生长的影响、木质藤本对环境变化的响应及适应机理、木质藤本数量过度增长的生态效应评价研究。同时，应积极探索合理的森林...