Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

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.     

Edge disturbance drives liana abundance increase and alteration of liana–host tree interactions in tropical forest fragments

Closed‐canopy forests are being rapidly fragmented across much of the tropical world. Determining the impacts of fragmentation on ecological processes enables better forest management and improves species‐conservation outcomes. Lianas are an integral part of tropical forests but can have detrimental and potentially complex interactions with their host trees. These effects can include reduced tree growth and fecundity, elevated tree mortality, alterations in tree‐species composition, degradation of forest succession, and a substantial decline in forest carbon storage. We examined the individual impacts of fragmentation and edge effects (0–100‐m transect from edge to forest interior) on the liana community and liana–host tree interactions in rainforests of the Atherton Tableland in north Queensland, Australia. We compared the liana and tree community, the traits of liana‐infested trees, and determinants of the rates of tree infestation within five forest fragments (23–58 ha in area) and five nearby intact‐forest sites. Fragmented forests experienced considerable disturbance‐induced degradation at their edges, resulting in a significant increase in liana abundance. This effect penetrated to significantly greater depths in forest fragments than in intact forests. The composition of the liana community in terms of climbing guilds was significantly different between fragmented and intact forests, likely because forest edges had more small‐sized trees favoring particular liana guilds which preferentially use these for climbing trellises. Sites that had higher liana abundances also exhibited higher infestation rates of trees, as did sites with the largest lianas. However, large lianas were associated with low‐disturbance forest sites. Our study shows that edge disturbance of forest fragments significantly altered the abundance and community composition of lianas and their ecological relationships with trees, with liana impacts on trees being elevated in fragments relative to intact forests. Consequently, effective control of lianas in forest fragments requires management practices which directly focus on minimizing forest edge disturbance.     

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。     

Trees as islands: canopy ant species richness increases with the size of liana‐free trees in a Neotropical forest

The physical characteristics of habitats shape local community structure; a classic example is the positive relationship between the size of insular habitats and species richness. Despite the high density and proximity of tree crowns in forests, trees are insular habitats for some taxa. Specifically, crown isolation (i.e. crown shyness) prevents the movement of small cursorial animals among trees. Here, we tested the hypothesis that the species richness of ants (S_a) in individual, isolated trees embedded within tropical forest canopies increases with tree size. We predicted that this pattern disappears when trees are connected by lianas (woody vines) or when strong interactions among ant species determine tree occupancy. We surveyed the resident ants of 213 tree crowns in lowland tropical forest of Panama. On average, 9.2 (range = 2–20) ant species occupied a single tree crown. Average (± SE) S_a was ca 25% higher in trees with lianas (10.2 ± 0.26) than trees lacking lianas (8.0 ± 0.51). S_a increased with tree size in liana‐free trees (S_a = 10.99A^0.256), but not in trees with lianas. Ant species composition also differed between trees with and without lianas. Specifically, ant species with solitary foragers occurred more frequently in trees with lianas. The mosaic‐like pattern of species co‐occurrence observed in other arboreal ant communities was not found in this forest. Collectively, the results of this study indicate that lianas play an important role in shaping the local community structure of arboreal ants by overcoming the insular nature of tree crowns.     

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。     

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.     

Environmental effects on Neotropical liana species richness

Aim Lianas differ physiologically from trees, and therefore their species‐richness patterns and potential climate‐change responses might also differ. However, multivariate assessments of spatial patterns in liana species richness and their controls are lacking. Our aim in this paper is to identify the environmental factors that best explain the variation in liana species richness within tropical forests. Location Lowland and montane Neotropical forests. Methods We quantified the contributions of environmental variables and liana and tree‐and‐shrub abundance to the species richness of lianas, trees and shrubs ≥ 2.5 cm in diameter using a subset of 65 standardized (0.1 ha) plots from 57 Neotropical sites from a global dataset collected by the late Alwyn Gentry. We used both regression and structural equation modelling to account for the effects of environmental variables (climate, soil and disturbance) and liana density on liana species richness, and we compared the species‐richness patterns of lianas with those of trees and shrubs. Results We found that, after accounting for liana density, dry‐season length was the dominant predictor of liana species richness. In addition, liana species richness was also related to stand‐level wood density (a proxy for disturbance) in lowland forests, a pattern that has not hitherto been shown across such a large study region. Liana species richness had a weak association with soil properties, but the effect of soil may be obscured by the strong correlation between soil properties and climate. The diversity patterns of lianas and of trees and shrubs were congruent: wetter forests had a greater species richness of all woody plants. Main conclusions The primary association of both liana and tree‐and‐shrub species richness with water availability suggests that, if parts of the Neotropics become drier as a result of climate change, substantial declines in the species richness of woody plants at the stand level may be anticipated.     

Effects of Logging on the Diversity of Lianas in a Lowland Tropical Rain Forest in Hainan Island, South China

Lianas are an integral part of tropical forest ecosystems, which usually respond strongly to severe disturbances, such as logging. To compare the effect of different logging systems on the lianas diversity in tropical rain forest, we recorded all lianas and trees ≥1 cm dbh in two 40-year-old forest sites after clear cutting (CC) and selective cutting (SC) as well as in an old-growth (OG) lowland tropical rain forest on Hainan Island in south China. Results showed that OG contained fewer liana stems and lower species richness (stems: 261, richness: 42 in 1 ha) than CC (606, 52) and SC (727, 50). However, OG had the highest Fisher's α diversity index (17.3) and species richness per stem (0.184). Species composition and dbh class distribution of lianas varied significantly with different logging systems. The mean liana dbh in OG (22.1 cm) were higher than those in CC (7.0 cm) and SC (10.4 cm). Stem twining was the most frequent climbing mechanism represented in the forest, as shown by the greatest species richness, abundance, basal area, and host tree number with this mechanism. The percent of host tree stems ≥4 cm dbh hosting at least one liana individual in SC (39%) was higher than CC (23%) and OG (19.5%). Large host trees (dbh≥60 cm) were more likely to be infested by lianas in SC and OG. Our study demonstrated that logging disturbance could significantly change the composition and structure of liana communities in the lowland tropical rain forest of south China.
     

Life History Traits of Lianas During Tropical Forest Succession

Tropical secondary forests form an important part of the landscape. Understanding functional traits of species that colonize at different points in succession can provide insight into community assembly. Although studies on functional traits during forest succession have focused on trees, lianas (woody vines) also contribute strongly to forest biomass, species richness, and dynamics. We examined life history traits of lianas in a forest chronosequence in Costa Rica to determine which traits vary consistently over succession. We conducted 0.1 ha vegetation inventories in 30 sites. To examine the establishment of young individuals, we only included small lianas (0.5–1.5 cm diameter at 1.3 m height). For each species, we identified seed size, dispersal mode, climbing mode, and whether or not the seedling is self‐supporting. We found a strong axis of variation determined by seed size and seedling growth habit, with early successional communities dominated by small‐seeded species with abiotic dispersal and climbing seedlings, while large‐seeded, animal‐dispersed species with free‐standing seedlings increased in abundance with stand age. Contrary to previous research and theory, we found a decrease in the abundance of stem twiners and no decrease in the abundance of tendril‐climbers during succession. Seed size appears to be a better indicator of liana successional stage than climbing mode. Liana life history traits change predictably over succession, particularly traits related to seedling establishment. Identifying whether these trait differences persist into the growth strategies of mature lianas is an important research goal, with potential ramifications for understanding the impact of lianas during tropical forest succession.     

滇南勐宋热带山地雨林木质藤本多样性研究

调查滇南勐宋山地雨林沟谷与坡地两种生境中木质藤本种的丰富度、径级分布、攀缘方式以及样地中被藤本缠绕的树木(dbh≥5cm)的数量和比率。结果表明:沟谷与坡地胸径≥1cm的木质藤本平均密度分别为95.7株/0.1hm2、57株/0.1hm2。调查样地内木质藤本共64种,隶属30个科。茎缠绕是最主要的攀缘方式,占总个体数的57%,卷须缠绕种所占比重最小,仅占3%;沟谷与坡地所调查树木被木质藤本缠绕的比例分别为43.7%和28.6%。与亚洲其它热带地区森林相比,勐宋地区藤本的多样性低,但是木质藤本的密度相当高,并且在一些样地中出现了大型木质藤本,这些可能与该地区森林的演替状态有关。     

Ants inhabiting myrmecophytic ferns regulate the distribution of lianas on emergent trees in a Bornean tropical rainforest

Little is known about the spatial distribution of lianas on emergent trees in tropical rainforests and the factors affecting this distribution. The present study investigated the effects of an arboreal ant species, Crematogaster difformis, which forms myrmecophytic symbioses with two epiphytic ferns, Lecanopteris sp. and Platycerium sp., on the spatial distribution of lianas associated with emergent trees. Living lianas were placed onto trunk surfaces inside and outside the territories of the ants in the canopy, to examine their ability to remove them. The number of leaves pruned by the ants was significantly higher on lianas inside than outside their territories. The spatial overlap of the distributions of lianas and the two ferns on emergent trees were then examined. The frequency of liana colonization of tree crowns was found to be significantly lower on trees with than without ferns. Under the natural conditions, C. difformis workers were observed biting and pruning the lianas. These results suggest that C. difformis regulates the distribution of lianas on emergent trees.     

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.     

Forest Structure and Biomass of a Tropical Seasonal Rain Forest in Xishuangbanna,Southwest China1

Xishuangbanna is a region located at the northern edge of tropical Asia. Biomass estimates of its tropical rain forest have not been published in English literature. We estimated forest biomass and its allocation patterns in five 0.185–1.0 ha plots in tropical seasonal rain forests of Xishuangbanna. Forest biomass ranged from 362.1 to 692.6 Mg/ha. Biomass of trees with diameter at 1.3 m breast height (DBH) ≥ 5 cm accounted for 98.2 percent of the rain forest biomass, followed by shrubs (0.9%), woody lianas (0.8%), and herbs (0.2%). Biomass allocation to different tree components was 68.4–70.0 percent to stems, 19.8–21.8 percent to roots, 7.4–10.6 percent to branches, and 0.7–1.3 percent to leaves. Biomass allocation to the tree sublayers was 55.3–62.2 percent to the A layer (upper layer), 30.6–37.1 percent to the B layer (middle), and 2.7–7.6 percent to the C layer (lower). Biomass of Pometia tomentosa, a dominant species, accounted for 19.7–21.1 percent of the total tree biomass. The average density of large trees (DBH ≥100 cm) was 9.4 stems/ha on two small plots and 3.5 stems/ha on two large plots, illustrating the potential to overestimate biomass on a landscape scale if only small plots are sampled. Biomass estimations are similar to typical tropical rain forests in Southeast Asia and the Neotropics.     

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     

How the diversity,abundance, size and climbing mechanisms of woody lianas are related to biotic and abiotic factors in a subtropical secondary forest,Taiwan

Lianas are woody vines that play an important role in forest dynamics in tropical and subtropical areas. Their relationship to various biotic and abiotic conditions is, however, not yet wholly clear. We explored how the size, climbing mechanisms, diversity and abundance of woody lianas is related to host plant size, environmental factors and topography. Liana assemblages were examined in twenty 20 × 20 m plots in each of three topographic sites (valley, slope and ridge) in a subtropical secondary forest in southeastern Taiwan. The valley site had the highest abundance and species richness of lianas. The abiotic factors, soil pH and rock cover, were related to different topographic sites. Larger lianas were always found on larger host trees, while smaller lianas were found in smaller trees; no lianas with a DBH greater than 10 cm were found. Significantly more adhesive lianas were found on larger trees whereas twining and leaning-hook lianas were found in smaller trees. In conclusion, this study demonstrates that the species of liana is associated with the size and type of tree growing under different topographic conditions.     

西双版纳热带次生林生物量的初步研究

 本文采用“空间代替时间”和维量分析的方法研究了西双版纳热带次生林4块不同年龄林分的生物量,并详细分析了热带次生林在演替初期阶段生物量的变化趋势。结果表明：林分总生物量随林龄而增加,5年生林分的总生物量为41.932t·hm-2,10年生林分的总生物量为52.116t·hm-2,14年生林分的总生物量为88.284t·hm-2,22年生林分的总生物量为113.743t·hm-2。林分生物量的层次分配比例以乔木层所占的比例最大,占4/5以上,随林龄而增加;灌木层增长到14年生林分后又下降,草本层随林龄而递减,层间植物则上升。生物量的器官分配比例以干材所占的比例最大,占1/2以上,随林龄而递增;而枝、根和叶的生物量分配比例则随林龄而下降。并建立了4个林分主要优势种及乔木层器官生物量的回归模型。     

A comprehensive synthesis of liana removal experiments in tropical forests

Lianas are a quintessential feature of tropical forests and are often perceived as being poorly studied. However, liana removal studies may be one of the most common experimental manipulations in tropical forest ecology. In this review, we synthesize data from 64 tropical liana removal experiments conducted over the past 90 yr. We explore the direction and magnitude of the effects of lianas on tree establishment, growth, survival, reproduction, biomass accretion, and plant and animal diversity in ecological and forestry studies. We discuss the geographical biases of liana removal studies and compare the various methods used to manipulate lianas. Overall, we found that lianas have a clear negative effect on trees, and trees benefitted from removing lianas in nearly every study across all forest types. Liana cutting significantly increased light and water availability, and trees responded with vastly greater reproduction, growth, survival, and biomass accumulation compared to controls where lianas were present. Removing lianas during logging significantly reduced damage of future merchantable trees and improved timber production. Our review demonstrates that lianas have an unequivocally detrimental effect on every metric of tree performance measured, regardless of forest type, forest age, or geographic location. However, lianas also appear to have a positive contribution to overall forest plant diversity and to different animal groups. Therefore, managing lianas reduces logging damage and improves timber production; however, the removal lianas may also have a negative effect on the faunal community, which could ultimately harm the plant community.     

A survey of vessel dimensions in stems of tropical lianas and other growth forms

Summary Vessel dimensions (total diameter and length) were determined in tropical and subtropical plants of different growth forms with an emphasis upon lianas (woody vines). The paint infusion and compressed air methods were used on 38 species from 26 genera and 16 families in the most extensive survey of vessel length made to date. Within most stems there was a skewed frequency distribution of vessel lengths and diameter, with many short and narrow vessels and few long and wide ones. The longest vessel found (7.73 m) was in a stem of the liana (woody vine) Pithecoctenium crucigerum. Mean vessel length for 33 species of lianas was 0.38 m, average maximum length was 1.45 m. There was a statistically significant inter-species correlation between maximum vessel length and maximum vessel diameter. Among liana stems and among tree+shrub stems there were statistically significant correlations between stem xylem diameter and vessel dimensions. Lianas with different adaptations for climbing (tendril climbers, twiners, scramblers) were similar in their vessel dimensions except that scramblers tended to have shorter (but not narrower) vessels. Within one genus, Bauhinia, tendril climbing species had greater maximum vessel lengths and diameters than tree and shrub species. The few long and wide vessels of lianas are thought to hydraulically compensate for their narrow stem diameters. The many narrow and short vessels, which are present in the same liana stems, may provide a high resistance auxiliary transport system.