木质藤本及其在热带森林中的生态学功能

木质藤本是热带森林的一个重要组分,直接或间接地影响着森林中树木的生长和更新,改变森林树木的种类组成,并且可以通过改变森林碳固定量等方式在生态系统水平上发挥作用。全球气候的变化,以及热带森林片断化程度的加剧,将很大程度上影响着木质藤本的多样性和丰富度,其特殊的生物学特性将在森林动态中发挥更加重要的作用。本文结合国内外目前木质藤本研究现状,概述了木质藤本的一般知识（包括木质藤本的定义和生物学特性等）,介绍了木质藤本全球分布格局、其多样性维持机理以及木质藤本在森林生态系统中的功能与作用,并就存在的一些问题以及需进一步开展的工作展开了讨论。     

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...     

元江干热河谷木质藤本的多样性及其与宿主树木的关系

木质藤本是森林生态系统的重要组分。本研究在元江干热河谷地区随机设置了30个20 m×20 m的样方,调查样方中胸径≥0.5 cm的木质藤本多样性及其与宿主树木之间的关系。结果显示:30个样方中记录到胸径≥0.5 cm的木质藤本植物共945株(隶属于22种20属11科),其中,豆科木质藤本的丰富度和多度最高;胸径≤2 cm的木质藤本占个体总数的63.7%;茎缠绕类木质藤本的个体数最多。样方中胸径≥5 cm的树木共有1060株(隶属于38种31属16科),36.0%的树木上至少附藤1株。不同径级和不同树皮粗糙度的树木被木质藤本侵扰的百分比之间存在极显著差异(P0.001)。随着宿主树木平均枝下高的增加,附藤率呈下降趋势。76.5%的木质藤本选择离其根生长点最近的树木进行攀援。表明元江干热河谷中的木质藤本以小径级占优势,树木胸径、枝下高、树皮粗糙度和木质藤本根生长点到树木的距离是影响木质藤本侵扰树木的重要因素,支持木质藤本对宿主树木的侵扰具有选择性的假说。研究结果对中国西南干热河谷退化植被的恢复与物种多样性保护具有重要意义。     

异军突起的藤本植物

对藤本植物(以热带为主)研究的动态与发展进行了综述,其中包括:(1)藤本植物的概念及重要性;(2)藤本植物区系和分布;(3)藤本植物的生物学特性;(4)藤本植物的生态学特性;(5)藤本在森林中的多度(密度)及种的丰富度;(6)藤本在空地上聚生的生态意义以及它们在森林演进过程中的消长;(7)藤本与树木的相互关系;(8)藤本对森林正面和负面的影响;(9)藤本的攀援方式类别及其竞争优势;(10)全球气候变化(CO2浓度增高)促进藤本优势度增长并导致不良的生态后果。     

异军突起的藤本植物

对藤本植物(以热带为主)研究的动态与发展进行了综述,其中包括:(1)藤本植物的概念及重要性;(2)藤本植物区系和分布;(3)藤本植物的生物学特性;(4)藤本植物的生态学特性;(5)藤本在森林中的多度(密度)及种的丰富度;(6)藤本在空地上聚生的生态意义以及它们在森林演进过程中的消长;(7)藤本与树木的相互关系;(8)藤本对森林正面和负面的影响;(9)藤本的攀援方式类别及其竞争优势;(10)全球气候变化(CO₂浓度增高)促进藤本优势度增长并导致不良的生态后果。     

热带森林植被生态恢复研究进展

热带森林是地球上生物多样性最高和生态功能最为强大的植被类型之一,在维护全球生态平衡中起着至关重要的作用,同时也为人类社会提供着多种多样的物质资源和生态系统服务。然而热带森林是目前生物多样性消失最快和生态功能退化最为严重的生态系统之一,如何有效地保护现存的热带森林不再进一步退化,以及如何使已经退化的生态系统尽快得到恢复是生态学工作者面临的重要议题。不同方式、规模和强度的干扰对热带林的破坏程度及其以后的恢复过程产生的影响不同。除少数大型自然干扰事件外,采伐、刀耕火种、农业开发用地等人为干扰是造成当前热带森林植被大面积退化的主要原因。多种干扰交互作用、杂草与外来物种入侵、退化植被和土壤状况、残存植被组分及土壤种子库、退化植被周围的景观格局以及全球气候变化等因素都能够影响热带森林植被恢复的速度和方向。基于功能群的研究思想将可能为物种丰富的热带森林植被恢复的研究提供一个全新途径。     

藤本植物适应生态学研究进展及存在问题

1　藤本植物的重要性　　藤本植物又称攀缘植物 ,是一类不能自由直立、通过主茎缠绕或攀缘器官攀缘它物升高的植物的总称 ,包括木质藤本和草质藤本两类。藤本植物是热带、亚热带森林重要的外貌和结构特征 ,常把森林冠层打结在一起形成特殊的层片结构[1～4] 。在南美热带森林中 ,在胸径大于 1 5～2 0ｃｍ的树木中有近一半的树冠上攀有藤本[5～ 7] ,森林下层植物中有 1 8%～ 2 2 %是藤本植物的幼藤[6] 。泰国[8] 和Ｇａｂｏｎ[9] 的热带森林的凋落物中藤本植物的叶占 32 %～ 36%。藤本植物也是热带森林种类多样性的重要组成部分。Ｇｅｎｔｒｙ…     

哀牢山中山湿性常绿阔叶林木质藤本植物的多样性与空间分布

木质藤本植物是热带、亚热带山地森林重要的组分之一, 在森林动态、生态系统过程和森林生物多样性形成与维持等方面具有重要作用。本文调查了哀牢山中山湿性常绿阔叶林木质藤本植物的多样性及其在垂直和水平空间上的分布规律。在20个20 m × 50 m的样地中共调查到DBH≥0.2 cm的木质藤本植物1,145株, 隶属于19科25属29种, 其中物种最丰富的科为菝葜科(4种)和蔷薇科(3种), 但多度最高的科为葡萄科(363株, 占总株数的31.7%)。研究发现林下木质藤本(通常DBH < 1 cm)拥有较高的物种丰富度和多度, 对木质藤本植物多样性具有较大的贡献。有55.7%的个体分布在林下层, 林冠层占28.8%, 亚冠层只有15.5%。木质藤本的垂直空间分布在不同径级、不同攀援类型之间具有明显的差异。 从水平空间分布来看, 地形是影响木质藤本的一个重要因素: 沟谷木质藤本的物种丰富度、多度和基面积分别是坡面的171%, 420%和606%; 有12个物种只分布在沟谷生境。这表明哀牢山中山湿性常绿阔叶林木质藤本植物对生境具有偏好性。     

乔木与木质藤本的水力与光合性状的差异:以热带森林崖豆藤属和买麻藤属为例

木质藤本是热带森林的重要组成部分,显著影响森林的结构和功能。已有研究发现木质藤本与乔木的水力结构存在显著差异:木质藤本的缠绕或攀缘茎细小,但其木质部具有粗大的长导管,输水效率高,抗栓塞能力低。为降低基因型差异对比较结果的影响,该研究选取热带崖豆藤属(Millettia)和买麻藤属(Gnetum)的乔木和木质藤本,比较同属内不同生长型植物的水力和光合性状的差异,分析水分传导效率与抗栓塞能力之间以及水力与光合性状之间的相关关系。结果发现:(1)崖豆藤属植物水力性状的种间差异大,与生活型和需光性有关。耐阴的木质藤本反而具有较低的水分传导效率和较高的抗栓塞能力。(2)买麻藤属植物是裸子植物较为进化的类群(具有导管和阔叶),其乔木的水分传导效率很低,但是其木质藤本的水分传导效率高于其他阳生性的被子植物。(3)不论乔木还是木质藤本,水分传导的有效性与安全性在枝条和叶片水平上均没有显著的权衡关系。(4)与同属乔木相比,木质藤本的叶片较枝条的抗栓塞能力更强,在旱季具有更高的最大净光合速率和气孔导度,支持了木质藤本的"旱季生长优势假说"。该研究揭示了热带木质藤本水力性状的多样性和重要性,为阐明环境变化对这一重要植物类群的影响,需要对它们的水力特征进行更广泛的研究。     

Hydraulic and photosynthetic characteristics differ between co-generic tree and liana species: a case study of Millettia and Gnetum in tropical forest

木质藤本是热带森林的重要组成部分, 显著影响森林的结构和功能。已有研究发现木质藤本与乔木的水力结构存在显著差异: 木质藤本的缠绕或攀缘茎细小, 但其木质部具有粗大的长导管, 输水效率高, 抗栓塞能力低。为降低基因型差异对比较结果的影响, 该研究选取热带崖豆藤属(Millettia)和买麻藤属(Gnetum)的乔木和木质藤本, 比较同属内不同生长型植物的水力和光合性状的差异, 分析水分传导效率与抗栓塞能力之间以及水力与光合性状之间的相关关系。结果发现: (1)崖豆藤属植物水力性状的种间差异大, 与生活型和需光性有关。耐阴的木质藤本反而具有较低的水分传导效率和较高的抗栓塞能力。(2)买麻藤属植物是裸子植物较为进化的类群(具有导管和阔叶), 其乔木的水分传导效率很低, 但是其木质藤本的水分传导效率高于其他阳生性的被子植物。(3)不论乔木还是木质藤本, 水分传导的有效性与安全性在枝条和叶片水平上均没有显著的权衡关系。(4)与同属乔木相比, 木质藤本的叶片较枝条的抗栓塞能力更强, 在旱季具有更高的最大净光合速率和气孔导度, 支持了木质藤本的“旱季生长优势假说”。该研究揭示了热带木质藤本水力性状的多样性和重要性, 为阐明环境变化对这一重要植物类群的影响, 需要对它们的水力特征进行更广泛的研究。     

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     

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.     

Variation in liana abundance and biomass along an elevational gradient in the tropical Atlantic Forest (Brazil)

Lianas play a key role in forest structure, species diversity, as well as functional aspects of tropical forests. Although the study of lianas in the tropics has increased dramatically in recent years, basic information on liana communities for the Brazilian Atlantic Forest is still scarce. To understand general patterns of liana abundance and biomass along an elevational gradient (0–1,100 m asl) of coastal Atlantic Forest, we carried out a standard census for lianas ≥1 cm in five 1-ha plots distributed across different forest sites. On average, we found a twofold variation in liana abundance and biomass between lowland and other forest types. Large lianas (≥10 cm) accounted for 26–35% of total liana biomass at lower elevations, but they were not recorded in montane forests. Although the abundance of lianas displayed strong spatial structure at short distances, the present local forest structure played a minor role structuring liana communities at the scale of 0.01 ha. Compared to similar moist and wet Neotropical forests, lianas are slightly less abundant in the Atlantic Forest, but the total biomass is similar. Our study highlights two important points: (1) despite some studies have shown the importance of small-scale canopy disturbance and support availability, the spatial scale of the relationships between lianas and forest structure can vary greatly among tropical forests; (2) our results add to the evidence that past canopy disturbance levels and minimum temperature variation exert influence on the structure of liana communities in tropical moist forests, particularly along short and steep elevational gradients.     

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.     

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.     

The relative importance of trees versus lianas as hosts for phytophagous beetles (Coleoptera) in tropical forests

Aim Insect assemblages associated with lianas in tropical forests are poorly studied compared with those associated with trees. The importance of lianas for the maintenance of local species richness of insect herbivores in tropical forests is therefore poorly understood. With this in mind, a comparative study of the relative importance of trees and lianas as hosts for phytophagous beetles was carried out. Location The study area was located in the canopy of a dry tropical forest in Parque Natural Metropolitano, Panama province, Republic of Panama. Methods A crane system was utilized to access the canopy. The number of species and host specialization of adult phytophagous beetles associated with twenty‐six liana species of ten different families, and twenty‐four tree species of twelve different families were compared. Results A total of 2561 host associations of 697 species of beetles were determined (1339 for trees and 1222 for lianas). On average 55.8 ± 6.8 beetle species were found to be associated with each tree species while the comparable number for lianas was 47.0 ± 6.1. The pooled numbers of phytophagous beetle species associated with trees and lianas, respectively, were not significantly different. However, there were significantly more species feeding on green plant parts on lianas than on trees, and there were significantly more wood eaters on trees than on lianas. Phytophagous beetles associated with lianas were significantly more specialized than the tree associates due to a higher degree of specialization among the species feeding on green plant parts of lianas. Wood eaters and flower visitors showed no differences in host specialization on different growth forms. Main conclusion The present study shows that lianas are at least as important as trees for the maintenance of local species diversity of phytophagous beetles at this site. The mechanisms that drive the patterns can only be hypothesized. Plant architecture, size, and length of growing season are probably involved. Further studies, should include measurements of plant traits to elucidate experimentally what mechanisms that drive the patterns. Additional insight would come from similar studies in other forest types, and also studies of other major taxonomic groups of arthropod herbivores.     

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.     

Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO₂ assimilation per unit mass (A _mass), nitrogen concentration (N _mass), and δ¹³C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO₂ assimilation per unit area (A _area), phosphorus concentration per unit mass (P _mass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree A _area decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ¹³C increased four times more than tree δ¹³C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher A _mass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.