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Climate change and other anthropogenic disturbances are increasing liana abundance and biomass in many tropical and subtropical forests. While the effects of living lianas on species diversity, ecosystem carbon, and nutrient dynamics are receiving increasing attention, the role of dead lianas in forest ecosystems has been little studied and is poorly understood. Trees and lianas coexist as the major woody components of forests worldwide, but they have very different ecological strategies, with lianas relying on trees for mechanical support. Consequently, trees and lianas have evolved highly divergent stem, leaf, and root traits. Here we show that this trait divergence is likely to persist after death, into the afterlives of these organs, leading to divergent effects on forest biogeochemistry. We introduce a conceptual framework combining horizontal, vertical, and time dimensions for the effects of liana proliferation and liana tissue decomposition on ecosystem carbon and nutrient cycling. We propose a series of empirical studies comparing traits between lianas and trees to answer questions concerning the influence of trait afterlives on the decomposability of liana and tree organs. Such studies will increase our understanding of the contribution of lianas to terrestrial biogeochemical cycling, and help predict the effects of their increasing abundance. [ABSTRACT FROM AUTHOR]

Determining population demographic rates is fundamental to understanding differences in species' life‐history strategies and their capacity to coexist. Calculating demographic rates, however, is challenging and requires long‐term, large‐scale censuses. Body size may serve as a simple predictor of demographic rate; can it act as a proxy for demographic rate when those data are unavailable? We tested the hypothesis that maximum body size predicts species' demographic rate using repeated censuses of the 77 most common liana species on the Barro Colorado Island, Panama (BCI) 50‐ha plot. We found that maximum stem diameter does predict species' population turnover and demography. We also found that lianas on BCI can grow to the enormous diameter of 635 mm, indicating that they can store large amounts of carbon and compete intensely with tropical canopy trees. This study is the first to show that maximum stem diameter can predict plant species' demographic rates and that lianas can attain extremely large diameters. Understanding liana demography is particularly timely because lianas are increasing rapidly in many tropical forests, yet their species‐level population dynamics remain chronically understudied. Determining per‐species maximum liana diameters in additional forests will enable systematic comparative analyses of liana demography and potential influence across forest types. [ABSTRACT FROM AUTHOR]

Tree‐tree competition has been widely studied as a mechanism responsible for maintaining forest plant species diversity. Other common plant types, such as lianas, may influence tree species competition. Previous studies reported the negative effects of lianas on trees; however, variation in susceptibility to lianas among tree species and the species‐specificity of liana species for tree hosts remain unclear. If lianas have species‐specific interactions with trees, based either on tree or liana species identity, then lianas may influence tree‐tree interactions by altering tree species‐specific competitiveness.We surveyed 5676 lianas (DBH ≥ 0.5 cm) and 61,538 trees (DBH ≥ 1 cm) in a 12‐ha subtropical secondary forest plot to assess liana‐tree interactions. We tested variation among the 20 most common tree species in their susceptibility to lianas, including whether light‐demanding and shade‐tolerant tree species differed. We quantified liana‐tree network structure and evaluated the specificity between the 20 most common tree species and the 15 most common liana species. We expected that: (1) tree species would vary in their susceptibility to lianas, and the extent of this variation would be predicted by tree shade‐tolerance; and (2) liana species would have distinct species‐specific interactions with host trees.Tree species differed greatly in their susceptibility to lianas, with some tree species accumulating many more lianas than others. Light‐demanding tree species particularly had greater susceptibility to lianas than shade‐tolerant tree species. Both the liana‐tree network structure and the interaction intensity between the 15 liana species and 20 tree species showed distinct specialization of liana‐tree interactions.Synthesis. Variation in trees species' susceptibility to lianas implies that lianas may alter tree species‐specific competitive abilities by disproportionately affecting some tree species more than others. In contrast to studies in neotropical forests, light‐demanding tree species were more susceptible to lianas than shade‐tolerant tree species, suggesting that lianas could reduce light‐demanding tree performance in this forest. The distinct species specialization between liana and tree species suggests that increases in the relative abundance of liana species, which is occurring in many forests, may alter the tree community composition by reducing the relative abundance of preferred host tree species. [ABSTRACT FROM AUTHOR]

Summary: Lightning is an important agent of plant mortality and disturbance in forests. Lightning‐caused disturbance is highly variable in terms of its area of effect and disturbance severity (i.e. tree damage and death), but we do not know how this variation is influenced by forest structure and plant composition.We used a novel lightning detection system to quantify how lianas influenced the severity and spatial extent (i.e. area) of lightning disturbance using 78 lightning strikes in central Panama.The local density of lianas (measured as liana basal area) was positively associated with the number of trees killed and damaged by lightning, and patterns of plant damage indicated that this occurred because lianas facilitated more electrical connections from large to small trees. Liana presence, however, did not increase the area of the disturbance. Thus, lianas increased the severity of lightning disturbance by facilitating damage to additional trees without influencing the footprint of the disturbance.These findings indicate that lianas spread electricity to damage and kill understory trees that otherwise would survive a strike. As liana abundance increases in tropical forests, their negative effects on tree survival with respect to the severity of lightning‐related tree damage and death are likely to increase. [ABSTRACT FROM AUTHOR]

Premise: Determining how xylem vessel diameters vary among plants and across environments gives insights into different water‐use strategies among species and ultimately their distributions. Here, we tested the vessel dimorphism hypothesis that the simultaneous occurrence of many narrow and a few wide vessels gives lianas an advantage over trees in seasonally dry environments. Methods: We measured the diameters of 13,958 vessels from 15 liana species and 10,430 vessels from 16 tree species in a tropical seasonal rainforest, savanna, and subtropical evergreen broadleaved forest. We compared differences in mean and hydraulically weighted vessel diameter (MVD and Dh), vessel density (VD), theoretical hydraulic conductivity (Kt), vessel area fraction (VAF), and wood density (WD) between lianas and trees and among three sites. Results: Nine liana species and four tree species had dimorphic vessels. From the tropical seasonal rainforest to the savanna, liana MVD, Dh and Kt decreased, and VD and WD increased, while only tree WD increased. From the tropical seasonal rainforest to the subtropical forest, six wood traits remained unchanged for lianas, while tree MVD, Dh and Kt decreased and VD increased. Trait space for lianas and trees were more similar in the savanna and more divergent in the subtropical forest compared to the tropical seasonal rainforest. Conclusions: These results suggest that lianas tend to possess greater vessel dimorphism, which may explain how lianas grow well during seasonal drought, influencing their unique distribution across tropical rainfall gradients. [ABSTRACT FROM AUTHOR]

Summary: Among tropical forests, lianas are predicted to have a growth advantage over trees during seasonal drought, with substantial implications for tree and forest dynamics. We tested the hypotheses that lianas maintain higher water status than trees during seasonal drought and that lianas maximize leaf cover to match high, dry‐season light conditions, while trees are more limited by moisture availability during the dry season.We monitored the seasonal dynamics of predawn and midday leaf water potentials and leaf phenology for branches of 16 liana and 16 tree species in the canopies of two lowland tropical forests with contrasting rainfall regimes in Panama.In a wet, weakly seasonal forest, lianas maintained higher water balance than trees and maximized their leaf cover during dry‐season conditions, when light availability was high, while trees experienced drought stress. In a drier, strongly seasonal forest, lianas and trees displayed similar dry season reductions in leaf cover following strong decreases in soil water availability.Greater soil moisture availability and a higher capacity to maintain water status allow lianas to maintain the turgor potentials that are critical for plant growth in a wet and weakly seasonal forest but not in a dry and strongly seasonal forest. [ABSTRACT FROM AUTHOR]

The well‐established pattern of forest thinning during succession predicts an increase in mean tree biomass with decreasing tree density. The forest thinning pattern is commonly assumed to be driven solely by tree‐tree competition. The presence of non‐tree competitors could alter thinning trajectories, thus altering the rate of forest succession and carbon uptake. We used a large‐scale liana removal experiment over 7 years in a 60‐ to 70‐year‐old Panamanian forest to test the hypothesis that lianas reduce the rate of forest thinning during succession. We found that lianas slowed forest thinning by reducing tree growth, not by altering tree recruitment or mortality. Without lianas, trees grew and presumably competed more, ultimately reducing tree density while increasing mean tree biomass. Our findings challenge the assumption that forest thinning is driven solely by tree‐tree interactions; instead, they demonstrate that competition from other growth forms, such as lianas, slow forest thinning and ultimately delay forest succession. [ABSTRACT FROM AUTHOR]

Lianas (woody vines) are abundant and diverse, particularly in tropical ecosystems. Lianas use trees for structural support to reach the forest canopy, often putting leaves above their host tree. Thus they are major parts of many forest canopies. Yet, relatively little is known about distributions of lianas in tropical forest canopies, because studying those canopies is challenging. This knowledge gap is urgent to address because lianas compete strongly with trees, reduce forest carbon uptake and are thought to be increasing, at least in the Neotropics.Lianas can be difficult to study using traditional field methods. Their pliable stems often twist and loop through the understorey, making it difficult to assess their structure and biomass, and the sizes and locations of their crowns. Furthermore, liana stems are commonly omitted from standard field surveys. Remote sensing of lianas can help overcome some of these obstacles and can provide critical insights into liana ecology, but to date there has been no systematic assessment of that contribution.We review progress in studying liana ecology using ground‐based, airborne and space‐borne remote sensing in four key areas: (i) spatial and temporal distributions, (ii) structure and biomass, (iii) responses to environmental conditions and (iv) diversity. This demonstrates the great potential of remote sensing for rapid advances in our knowledge and understanding of liana ecology. We then look ahead, to the possibilities offered by new and future advances. We specifically consider the data requirements, the role of technological advances and the types of methods and experimental designs that should be prioritised.Synthesis. The particular characteristics of the liana growth form make lianas difficult to study by ground‐based field methods. However, remote sensing is well suited to collecting data on lianas. Our review shows that remote sensing is an emerging tool for the study of lianas, and will continue to improve with recent developments in sensor and platform technology. It is surprising, therefore, how little liana ecology research has utilised remote sensing to date—this should rapidly change if urgent knowledge gaps are to be addressed. In short, liana ecology needs remote sensing. [ABSTRACT FROM AUTHOR]

Lianas are a key growth form in tropical forests. Their lack of self‐supporting tissues and their vertical position on top of the canopy make them strong competitors of resources. A few pioneer studies have shown that liana optical traits differ on average from those of colocated trees. Those trait discrepancies were hypothesized to be responsible for the competitive advantage of lianas over trees. Yet, in the absence of reliable modelling tools, it is impossible to unravel their impact on the forest energy balance, light competition, and on the liana success in Neotropical forests. To bridge this gap, we performed a meta‐analysis of the literature to gather all published liana leaf optical spectra, as well as all canopy spectra measured over different levels of liana infestation. We then used a Bayesian data assimilation framework applied to two radiative transfer models (RTMs) covering the leaf and canopy scales to derive tropical tree and liana trait distributions, which finally informed a full dynamic vegetation model. According to the RTMs inversion, lianas grew thinner, more horizontal leaves with lower pigment concentrations. Those traits made the lianas very efficient at light interception and significantly modified the forest energy balance and its carbon cycle. While forest albedo increased by 14% in the shortwave, light availability was reduced in the understorey (−30% of the PAR radiation) and soil temperature decreased by 0.5°C. Those liana‐specific traits were also responsible for a significant reduction of tree (−19%) and ecosystem (−7%) gross primary productivity (GPP) while lianas benefited from them (their GPP increased by +27%). This study provides a novel mechanistic explanation to the increase in liana abundance, new evidence of the impact of lianas on forest functioning, and paves the way for the evaluation of the large‐scale impacts of lianas on forest biogeochemical cycles. [ABSTRACT FROM AUTHOR]

Canopy disturbance explains liana abundance and distribution within tropical forests and thus may also explain the widespread pattern of increasing liana abundance; however, this hypothesis remains untested. We used a 10‐year study (2007–2017) of 117,100 rooted lianas in an old‐growth Panamanian forest to test whether local canopy disturbance explains increasing liana abundance. We found that liana density increased 29.2% and basal area 12.5%. The vast majority of these increases were associated with clonal stem proliferation following canopy disturbance, particularly in liana‐dense, low‐canopy gaps, which had far greater liana increases than did undisturbed forest. Lianas may be ecological niche constructors, arresting tree regeneration in gaps and thus creating a high‐light environment that favours sustained liana proliferation. Our findings demonstrate that liana abundance is increasing rapidly and their ability to proliferate via copious clonal stem production in canopy gaps explains much of their increase in this and possibly other tropical forests. [ABSTRACT FROM AUTHOR]

Lianas are increasing in relative abundance and biomass in many tropical forests. We tested the hypothesis that lianas conform to the fast and acquisitive end of the continuum of plant strategies, allowing lianas to acquire resources faster than trees.We assessed functional traits representative of the leaf (LES) and wood economics spectrum (WES), including plant hydraulics, in 16 liana and 16 tree species in the canopy of two tropical forests at the extremes of the climatic and geological gradient across the Isthmus of Panama.For both forests, we observed a trade‐off between the construction of more productive leaves with rapid turnover and expensive leaves with slower turnover. We also found trait variation associated with wood and hydraulic traits. These two axes were orthogonal, suggesting that trade‐offs at the leaf and stem, including plant hydraulics, operate independently.For the dry forest, lianas had cheaper and more efficient leaves than trees. For the wet forest, lianas and trees overlapped in leaf and stem characteristics. Moreover, the duration of green foliage highly explained the variation between dry forest species, reflecting different adaptations to drought. In the wet forest, fast‐growing species benefited from a higher return on investments of leaf vascular tissues than slow‐growing species and they had a higher capacity to transport water through the leaf.A higher capacity to construct more productive leaves and display leaves with lower costs may favour lianas over trees in light interception, photosynthetic rates, and growth under high light and nutrient availability in dry forests.Synthesis. Lianas in a dry tropical forest had a more acquisitive strategy than trees, characterized by more productive leaves and more efficient display for light interception. In dry environments, lianas appear to benefit from high‐light and nutrient‐rich soils and thus take advantage of higher resource conditions compared to trees. By contrast, in a wet tropical forest, lianas and trees overlapped in leaf and stem characteristics and lianas were not more acquisitive than trees. In wet environments, low light availability and nutrient‐poor soils in a context of low water limitation may constrain variation in resource acquisition strategies between lianas and trees. [ABSTRACT FROM AUTHOR]

Global biodiversity is declining at rates faster than at any other point in human history. Experimental manipulations at small spatial scales have demonstrated that communities with fewer species consistently produce less biomass than higher diversity communities. Understanding the consequences of the global extinction crisis for ecosystem functioning requires understanding how local experimental results are likely to change with increasing spatial and temporal scales and from experiments to naturally assembled systems.Scaling across time and space in a changing world requires baseline predictions. Here, we provide a graphical null model for area scaling of biodiversity–ecosystem functioning relationships using observed macroecological patterns: the species–area curve and the biomass–area curve. We use species–area and biomass–area curves to predict how species richness–biomass relationships are likely to change with increasing sampling extent. We then validate these predictions with data from two naturally assembled ecosystems: a Minnesota savanna and a Panamanian tropical dry forest.Our graphical null model predicts that biodiversity–ecosystem functioning relationships are scale‐dependent. However, we note two important caveats. First, our results indicate an apparent contradiction between predictions based on measurements in biodiversity–ecosystem functioning experiments and from scaling theory. When ecosystem functioning is measured as per unit area (e.g. biomass per m2), as is common in biodiversity–ecosystem functioning experiments, the slope of the biodiversity ecosystem functioning relationship should decrease with increasing scale. Alternatively, when ecosystem functioning is not measured per unit area (e.g. summed total biomass), as is common in scaling studies, the slope of the biodiversity–ecosystem functioning relationship should increase with increasing spatial scale. Second, the underlying macroecological patterns of biodiversity experiments are predictably different from some naturally assembled systems. These differences between the underlying patterns of experiments and naturally assembled systems may enable us to better understand when patterns from biodiversity–ecosystem functioning experiments will be valid in naturally assembled systems.Synthesis. This paper provides a simple graphical null model that can be extended to any relationship between biodiversity and any ecosystem functioning across space or time. Furthermore, these predictions provide crucial insights into how and when we may be able to extend results from small‐scale biodiversity experiments to naturally assembled regional and global ecosystems where biodiversity is changing. [ABSTRACT FROM AUTHOR]

Lianas rely on trees for support and access to high‐light positions in the forest canopy, but the implications for how lianas explore the canopy compared to trees remain understudied. We present an in situ forest canopy study to test the hypotheses that: (1) lianas favour leaf display over stem investment compared to trees and (2) lianas have greater potential to colonize non‐shaded, high‐light areas effectively than trees.We compared branches of 16 liana species with those of 16 sympatric tree species in the canopy of two lowland tropical forests with contrasting rainfall regimes in Panama using 40–50 m tall canopy cranes. One forest was relatively dry and seasonal in rainfall and associated solar radiation. The other forest was relatively wet and had a weaker seasonality.We observed that lianas were more efficient in leaf display over stem investment than trees, particularly in the forest with lower precipitation and stronger seasonality. Specifically, lianas had a lower LMA (leaf mass per unit leaf area), stronger apical dominance, higher stem slenderness and fewer leaf layers than trees. In the forest with higher precipitation and weaker seasonality, lianas also had stronger apical control and fewer leaf layers than trees, but both lianas and trees were relatively similar in LMA and stem slenderness.Our study shows that lianas more effectively explore the canopy than trees under drier conditions, but much less so under wetter conditions. We argue that lianas display a functional strategy that allows them to better intercept light than the tree species in forests with low precipitation and strong seasonality, while they are constrained to display such strategy at high precipitation – light‐limited – sites. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Despite their low contribution to forest carbon stocks, lianas (woody vines) play an important role in the carbon dynamics of tropical forests. As structural parasites, they hinder tree survival, growth and fecundity; hence, they negatively impact net ecosystem productivity and long‐term carbon sequestration.Competition (for water and light) drives various forest processes and depends on the local abundance of resources over time. However, evaluating the relative role of resource availability on the interactions between lianas and trees from empirical observations is particularly challenging. Previous approaches have used labour‐intensive and ecosystem‐scale manipulation experiments, which are infeasible in most situations.We propose to circumvent this challenge by evaluating the uncertainty of water and light capture processes of a process‐based vegetation model (ED2) including the liana growth form. We further developed the liana plant functional type in ED2 to mechanistically simulate water uptake and transport from roots to leaves, and start the model from prescribed initial conditions. We then used the PEcAn bioinformatics platform to constrain liana parameters and run uncertainty analyses.Baseline runs successfully reproduced ecosystem gas exchange fluxes (gross primary productivity and latent heat) and forest structural features (leaf area index, aboveground biomass) in two sites (Barro Colorado Island, Panama and Paracou, French Guiana) characterized by different rainfall regimes and levels of liana abundance.Model uncertainty analyses revealed that water limitation was the factor driving the competition between trees and lianas at the drier site (BCI), and during the relatively short dry season of the wetter site (Paracou). In young patches, light competition dominated in Paracou but alternated with water competition between the wet and the dry season on BCI according to the model simulations.The modelling workflow also identified key liana traits (photosynthetic quantum efficiency, stomatal regulation parameters, allometric relationships) and processes (water use, respiration, climbing) driving the model uncertainty. They should be considered as priorities for future data acquisition and model development to improve predictions of the carbon dynamics of liana‐infested forests.Synthesis. Competition for water plays a larger role in the interaction between lianas and trees than previously hypothesized, as demonstrated by simulations from a process‐based vegetation model. [ABSTRACT FROM AUTHOR]

Over the past two decades, liana density and basal area have been increasing in many tropical forests, which has profound consequences for forest diversity and functioning. One hypothesis to explain increasing lianas is elevated nutrient deposition in tropical forests resulting from fossil fuels, agricultural fertilizer, and biomass burning. We tested this hypothesis by surveying all lianas ≥1 cm in diameter (n = 3,967) in 32 plots in a fully factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment in a mature tropical forest in central Panama. We conducted the nutrient‐addition experiment from 1998 until present and we first censused lianas in 2013 and then again in 2018. After 20 yr of nutrient addition (1998–2018), liana density, basal area, and rarefied species richness did not differ significantly among any of the nutrient‐addition and control treatments. Moreover, nutrient addition in the most recent 5 yr of the experiment did not affect liana relative growth, recruitment, or mortality rates. From 2013 until 2018, liana density, basal area, and species richness increased annually by 1.6%, 1.4%, and 2.4%, respectively. Nutrient addition did not influence these increases. Our findings indicate that nutrient deposition does not explain increasing lianas in this tropical forest. Instead, increases in tree mortality and disturbance, atmospheric carbon dioxide, drought frequency and severity, and hunting pressure may be more likely explanations for the increase in lianas in tropical forests. [ABSTRACT FROM AUTHOR]

The spatial habitat heterogeneity hypothesis posits that habitat complexity increases the abundance and diversity of species. In tropical forests, lianas add substantial habitat heterogeneity and complexity throughout the vertical forest profile, which may maintain animal abundance and diversity. The effects of lianas on tropical animal communities, however, remain poorly understood. We propose that lianas have a positive effect on animals by enhancing habitat complexity. Lianas may have a particularly strong influence on the forest bird community, providing nesting substrate, protection from predators, and nutrition (food). Understory insectivorous birds, which forage for insects that specialize on lianas, may particularly benefit. Alternatively, it is possible that lianas have a negative effect on forest birds by increasing predator abundances and providing arboreal predators with travel routes with easy access to bird nests. We tested the spatial habitat heterogeneity hypothesis on bird abundance and diversity by removing lianas, thus reducing forest complexity, using a large‐scale experimental approach in a lowland tropical forest in the Republic of Panama. We found that removing lianas decreased total bird abundance by 78.4% and diversity by 77.4% after 8 months, and by 40.0% and 51.7%, respectively, after 20 months. Insectivorous bird abundance and diversity 8 months after liana removal were 91.8% and 89.5% lower, respectively, indicating that lianas positively influence insectivorous birds. The effects of liana removal persisted longer for insectivorous birds than other birds, with 77.3% lower abundance and 76.2% lower diversity after 20 months. Liana removal also altered bird community composition, creating two distinct communities in the control and removal plots, with disproportionate effects on insectivores. Our findings demonstrate that lianas have a strong positive influence on the bird community, particularly for insectivorous birds in the forest understory. Lianas may maintain bird abundance and diversity by increasing habitat complexity, habitat heterogeneity, and resource availability. [ABSTRACT FROM AUTHOR]

Copyright of Biotropica is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

A central and recurrent discussion in ecology is how deterministic and stochastic processes dictate how communities assemble after severe disturbances. Furthermore, tree biomass converged over time as forests aged, and it was positively affected by edaphic factors, especially soil fertility, implying that fertility can have a determinist effect on tree biomass accumulation. By contrast, liana biomass did not converge, regardless of forest cover or edaphic factors suggesting that liana biomass might be more regulated by stochastic processes than trees. At the moist forest site, rainfall variability in the dry season can change up to 20-fold (17-342 mm between February and April), and it was expected to have important potential consequences in the tree community successional dynamics. [Extracted from the article]

Summary: There are two theories about how allocation of metabolic products occurs. The allometric biomass partitioning theory (APT) suggests that all plants follow common allometric scaling rules. The optimal partitioning theory (OPT) predicts that plants allocate more biomass to the organ capturing the most limiting resource.Whole‐plant harvests of mature and juvenile tropical deciduous trees, evergreen trees, and lianas and model simulations were used to address the following knowledge gaps: (1) Do mature lianas comply with the APT scaling laws or do they invest less biomass in stems compared to trees? (2) Do juveniles follow the same allocation patterns as mature individuals? (3) Is either leaf phenology or life form a predictor of rooting depth?It was found that: (1) mature lianas followed the same allometric scaling laws as trees; (2) juveniles and mature individuals do not follow the same allocation patterns; and (3) mature lianas had shallowest coarse roots and evergreen trees had the deepest.It was demonstrated that: (1) mature lianas invested proportionally similar biomass to stems as trees and not less, as expected; (2) lianas were not deeper‐rooted than trees as had been previously proposed; and (3) evergreen trees had the deepest roots, which is necessary to maintain canopy during simulated dry seasons. See also the Commentary on this article by Lugli, 226: 639–640. [ABSTRACT FROM AUTHOR]

Early successional tropical forests could mitigate climate change via rapid accumulation of atmospheric carbon. However, liana (woody vine) abundance and biomass has been increasing in many tropical forests over the past decades, which may slow the speed at which secondary forests accumulate biomass. Lianas decrease biomass accumulation in tropical forests, and may have a particularly strong effect on young forests by stalling tree growth. As forests mature, trees may outgrow or shed lianas, thus escaping some of the negative effects of lianas. Alternatively, lianas may have the strongest effect in older successional forests if the effect of lianas is commensurate with their density, which increases dramatically in the first decades of forest succession. We tested these two hypotheses using a landscape liana‐removal experiment in 30 forest stands that ranged from 10 to 35 yr old in Central Panama. We measured tree growth and biomass accumulation in the stands every year from 2014 to 2017. We found that the effect of liana removal on large trees (≥20‐cm diameter) decreased with forest age, supporting the hypothesis that lianas have the strongest negative effects on trees, and thus biomass uptake and carbon storage, in very young successional forests. Large trees accumulated more biomass in the absence of lianas in younger forests than in older forests (compared to controls) even after accounting for the effect of canopy completeness and crown illumination, implying that the detrimental effects of lianas go well beyond resource availability and crown health. There was no significant effect of lianas on small trees (1–20‐cm diameter), likely because lianas seek light and thus do not deploy their leaves on small trees that are trapped in the forest understory. Our results show that high liana density early in forest succession reduces forest biomass accumulation by negatively impacting large trees, thus decreasing the capacity of young secondary forests to mitigate climate change. Although the negative effects of lianas on forest biomass diminish as forests age, they do not disappear, and thus lianas are an important component of tropical forest carbon budgets throughout succession. [ABSTRACT FROM AUTHOR]

In tropical regions, rainfall gradients often explain the abundance and distribution of plant species. For example, many tree and liana species adapted to seasonal drought are more abundant and diverse in seasonally‐dry forests, characterized by long periods of seasonal water deficit. Mean annual precipitation (MAP) is commonly used to explain plant distributions across climate gradients. However, the relationship between MAP and plant distribution is often weak, raising the question of whether other seasonal precipitation patterns better explain plant distributions in seasonally‐dry forests. In this study, we examine the relationship between liana abundance and multiple metrics of seasonal and annual rainfall distribution to test the hypothesis that liana density and diversity increase with increasing seasonal drought along a rainfall gradient across the isthmus of Panama. We found that a normalized seasonality index, which combines MAP and the variability of monthly rainfall throughout the year, was a significant predictor of both liana density and species richness, whereas MAP, rainfall seasonality and the mean dry season precipitation (MDP) were far weaker predictors. The strong response of lianas to the normalized seasonality index indicates that, in addition to the total annual amount of rainfall, how rainfall is distributed throughout the year is an important determinant of the hydrological conditions that favor liana proliferation. Our findings imply that changes in annual rainfall and rainfall seasonality will determine the future distribution and abundance of lianas. Models that aim to predict future plant diversity, distribution, and abundance may need to move beyond MAP to a more detailed understanding of rainfall variability at sub‐annual timescales. [ABSTRACT FROM AUTHOR]

Copyright of Journal of Ecology is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

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. [ABSTRACT FROM AUTHOR]

There is mounting empirical evidence that lianas affect the carbon cycle of tropical forests. However, no single vegetation model takes into account this growth form, although such efforts could greatly improve the predictions of carbon dynamics in tropical forests. In this study, we incorporated a novel mechanistic representation of lianas in a dynamic global vegetation model (the Ecosystem Demography Model). We developed a liana‐specific plant functional type and mechanisms representing liana–tree interactions (such as light competition, liana‐specific allometries, and attachment to host trees) and parameterized them according to a comprehensive literature meta‐analysis. We tested the model for an old‐growth forest (Paracou, French Guiana) and a secondary forest (Gigante Peninsula, Panama). The resulting model simulations captured many features of the two forests characterized by different levels of liana infestation as revealed by a systematic comparison of the model outputs with empirical data, including local census data from forest inventories, eddy flux tower data, and terrestrial laser scanner‐derived forest vertical structure. The inclusion of lianas in the simulations reduced the secondary forest net productivity by up to 0.46 tC ha−1 year−1, which corresponds to a limited relative reduction of 2.6% in comparison with a reference simulation without lianas. However, this resulted in significantly reduced accumulated above‐ground biomass after 70 years of regrowth by up to 20 tC/ha (19% of the reference simulation). Ultimately, the simulated negative impact of lianas on the total biomass was almost completely cancelled out when the forest reached an old‐growth successional stage. Our findings suggest that lianas negatively influence the forest potential carbon sink strength, especially for young, disturbed, liana‐rich sites. In light of the critical role that lianas play in the profound changes currently experienced by tropical forests, this new model provides a robust numerical tool to forecast the impact of lianas on tropical forest carbon sinks. [ABSTRACT FROM AUTHOR]

Lianas are prevalent in Neotropical forests, where liana‐tree competition can be intense, resulting in reduced tree growth and survival. The ability of lianas to grow relative to trees during the dry season suggests that liana‐tree competition is also strongest in the dry season. If correct, the predicted intensification of the drying trend over large areas of the tropics in the future may therefore intensify liana‐tree competition resulting in a reduced carbon sink function of tropical forests. However, no study has established whether the liana effect on tree carbon accumulation is indeed stronger in the dry than in the wet season.Using 6 years of data from a large‐scale liana removal experiment in Panama, we provide the first experimental test of whether liana effects on tree carbon accumulation differ between seasons. We monitored tree and liana diameter increments at the beginning of the dry and wet season each year to assess seasonal differences in forest‐level carbon accumulation between removal and control plots.We found that median liana carbon accumulation was consistently higher in the dry (0.52 Mg C ha−1 year−1) than the wet season (0.36 Mg C ha−1 year−1) and significantly so in three of the years. Lianas reduced forest‐level median tree carbon accumulation more severely in the wet (1.45 Mg C ha−1 year−1) than the dry (1.05 Mg C ha−1 year−1) season in all years. However, the relative effect of lianas was similar between the seasons, with lianas reducing forest‐level tree carbon accumulation by 46.9% in the dry and 48.5% in the wet season.Synthesis. Our results provide the first experimental demonstration that lianas do not have a stronger competitive effect on tree carbon accumulation during the dry season. Instead, lianas compete significantly with trees during both seasons, indicating a large negative effect of lianas on forest‐level tree biomass increment regardless of seasonal water stress. Longer dry seasons are unlikely to impact liana‐tree competition directly; however, the greater liana biomass increment during dry seasons may lead to further proliferation of liana biomass in tropical forests, with consequences for their ability to store and sequester carbon. [ABSTRACT FROM AUTHOR]

Lianas are structural parasites of trees that reduce the growth, survival and reproduction of their hosts. Given that co‐occurring tree species differ strongly in the proportion of individuals that are infested by lianas (liana prevalence), lianas could differentially impact tree species and thereby influence tree community composition. Surprisingly, little is known about what governs variation in liana prevalence.Here, we apply an approach inspired by disease ecology to investigate the dynamics of liana prevalence over 11 years on Barro Colorado Island, Panama. We followed the fate of 1,938 individual trees from 21 tree species, recording deaths and change in liana infestation status. With these data, we fit species‐specific Markov chain models to estimate four rates: colonization by lianas (analogous to disease transmission), shedding or loss of lianas (analogous to host recovery), baseline mortality of uninfested trees (baseline mortality) and additional mortality of infested trees (parasite lethality).Models explained 58% of variation in liana prevalence among tree species, and revealed that host shedding of lianas and parasite lethality were the most important contributors to interspecific variation in liana prevalence at our site. These rates were also strongly related to shade tolerance, with light‐demanding species having greater rates of shedding and lethality, and lower rates of liana prevalence. An indirect path analysis with a structural equation model revealed that both greater rates of liana shedding and liana‐induced lethality contribute to the observed lower rates of liana prevalence for light‐demanding tree species.Synthesis. Our approach revealed that the prevalence of liana infestation among tree species is driven via indirect pathways operating on the rates of shedding and lethality, which relate to the ability (or inability) of trees to shed and/or tolerate lianas. Shade‐tolerant trees have greater proportions of trees infested by lianas because they are both less able to shed lianas and more able to tolerate infestation. Our approach revealed that the prevalence of liana infestation among tree species is driven via indirect pathways operating on the rates of shedding and lethality, which relate to the ability (or inability) of trees to shed and/or tolerate lianas. Shade‐tolerant trees have greater proportions of trees infested by lianas because they are both less able to shed lianas and more able to tolerate infestation. [ABSTRACT FROM AUTHOR]

Contents Summary 366 I. Introduction 366 II. Testing ecological theory: effects of the environment on lianas 369 III. A unified explanation for liana distribution and the maintenance of liana diversity 370 IV. Testing ecological theory: effects of lianas on the environment 373 V. Theoretical effects of lianas on forest diversity 375 VI. Lianas and trophic interactions in forests 375 VII. Unresolved challenges in liana ecology 376 VIII. Conclusions 377 Acknowledgements 377 References 377 Summary: Lianas constitute a diverse polyphyletic plant group that is advancing our understanding of ecological theory. Specifically, lianas are providing new insights into the mechanisms that control plant distribution and diversity maintenance. For example, there is now evidence that a single, scalable mechanism may explain local, regional, and pan‐tropical distribution of lianas, as well as the maintenance of liana species diversity. The ability to outcompete trees under dry, stressful conditions in seasonal forests provides lianas a growth advantage that, over time, results in relatively high abundance in seasonal forests and low abundance in aseasonal forests. Lianas may also gain a similar growth advantage following disturbance, thus explaining why liana density and diversity peak following disturbance at the local, forest scale. The study of ecology, however, is more than the effect of the environment on organisms; it also includes the effects of organisms on the environment. Considerable empirical evidence now indicates that lianas substantially alter their environment by consuming resources, suppressing tree performance, and influencing emergent properties of forests, such as ecosystem functioning, plant and animal diversity, and community composition. These recent studies using lianas are transcending classical tropical ecology research and are now providing novel insights into fundamental ecological theory. [ABSTRACT FROM AUTHOR]

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Abstract: Lianas are structural parasites of trees and reduce individual host tree growth, survival and fecundity. Thus, liana infestation is expected to affect tree population growth rates, with potentially different effects in different species depending on the frequency of liana infestation and the impact of liana infestation on population growth rates. Previous studies have documented the myriad negative effects of lianas on trees and variation in liana infestation among tree species; however, no study has quantified the impact of liana infestation on individual tree species population growth rates. Lianas are increasing in abundance in multiple Neotropical sites, which may have profound consequences for tree species composition if lianas differentially affect host tree species population growth. Here, we use long‐term data to evaluate the effects of liana infestation on the reproduction, growth, survival and ultimately population growth rates of dozens of tree species from Barro Colorado Island, Panama. We then test whether liana infestation affects tree species differentially with respect to two axes of life‐history variation: adult stature and position along the fast–slow axis, a measure of shade tolerance. Liana infestation decreased tree growth, survival and reproduction, with the strongest effects on survival in fast‐growing, light‐demanding species and on reproduction in large‐statured species. In combination, these effects reduced tree population growth rates such that liana‐infested populations declined by an average of 1.4% annually relative to conspecific liana‐free populations. The reduction in population growth rates was greatest among fast‐growing species and smaller in slow‐growing species. Synthesis. Our results demonstrate that liana infestation has strong negative effects on tree population growth rates, which vary systematically among tree species with tree life history. The finding that liana infestation is more harmful to fast‐growing tree species appears to be at odds with the general expectations in the literature. We propose that this is likely due to survivorship bias, as infestation greatly decreases survival in fast‐growing species such that the observable sample is biased towards those that survived and liana‐free. In combination with data on how tree species vary in liana infestation rates, these results provide a basis for predicting the impacts of changes in liana abundance on tree species composition. [ABSTRACT FROM AUTHOR]

Liana density tends to increase with decreasing rainfall and increasing seasonality. However, the pattern of liana distribution may be due to differences in soil water retention capacity, not rainfall and seasonality per se. We tested the effect of rainfall and soil substrate with respect to the distribution of liana seedlings in six sites across a rainfall gradient from the wet Atlantic to the dry Pacific in central Panama. Soils were either limestone, with low water-holding capacity, or laterite, with higher water-holding capacity. We sampled liana seedlings at each site using three 1 × 100 m transect. We found that relative liana seedling density was higher on limestone soils compared to laterite soils regardless of the amount of rainfall. Furthermore, liana community composition on limestone soils was more similar to dry forest sites than to adjacent wet and moist forest sites. Liana seedling species diversity relative to trees was significantly higher in a low-fertility dry forest site compared to a high-fertility forest, but did not differ from the other sites. Thus, liana seedling density and community structure may be driven more by soil type and thus by soil moisture availability than strictly by mean annual rainfall and the seasonality of rainfall. [ABSTRACT FROM AUTHOR]

Abstract: Lianas are a key component of tropical forests, where they compete intensely with trees, reducing tree recruitment, growth and survival. One of the most important potential outcomes of liana competition is the reduction of tree reproduction; however, no previous study has experimentally determined the effects of lianas on tree reproduction beyond a single tree species. We used a large‐scale liana removal experiment to quantify the effect of lianas on community‐level canopy and understorey tree and palm reproduction. In 2011, we removed lianas from eight 6,400‐m2 plots (eight plots served as controls) and surveyed understorey tree reproduction in 2012, canopy tree and palm reproduction in 2013, and a second census of all plants in 2016. We found that lianas significantly reduced canopy tree community flowering and fruiting after liana removal. Two years after liana removal, the number of canopy trees with fruits was 173% higher, fruiting individuals had 50% more of their canopy covered by fruits and the number of tree species with fruits was 169% higher than in control plots where lianas were present. Five years after liana removal, the number of canopy trees with fruits was 150% higher, fruiting individuals had 31% more of their canopy covered by fruits and the number of tree species with fruits was 109% higher than in unmanipulated control plots. Liana removal had only a slight positive effect on palms and on understorey tree flower and fruit production, even though understorey light levels had increased 20% following liana cutting. Synthesis. Our findings provide the first experimental demonstration that competition from lianas significantly reduces community‐level canopy tree reproduction. Reduced reproduction increases canopy tree seed and dispersal limitations, and may interfere with deterministic mechanisms thought to maintain tropical canopy tree species diversity, as well as reduce food availability to many animal species. Because lianas are increasing in abundance in many neotropical forests, the effects of lianas on tree reproduction will likely increase, and if the effects of lianas on tree reproduction vary with tree species identity, lianas ultimately could have a destabilizing effect on both tree and animal population dynamics. [ABSTRACT FROM AUTHOR]

Abstract: Dispersal and density dependence are major determinants of spatial structure, population dynamics and coexistence for tropical forest plants. However, because these two processes can jointly influence spatial structure at similar scales, analysing spatial patterns to separate and quantify them is often difficult. Species functional traits can be useful indicators of dispersal and density dependence. However, few methods exist for linking functional traits to quantitative estimates of these processes that can be compared across multiple species. We analysed static spatial patterns of woody plant populations in the 50 ha Forest Dynamics Plot on Barro Colorado Island, Panama with methods that distinguished scale‐specific differences in species aggregation. We then tested how these differences related to seven functional traits: growth form, dispersal syndrome, tree canopy layer, adult stature, seed mass, wood density and shade tolerance. Next, we fit analytically tractable spatial moment models to the observed spatial structure of species characterized by similar trait values, which allowed us to estimate relationships of functional traits with the spatial scale of dispersal, and the spatial scale and intensity of negative density dependence. Our results confirm that lianas are more aggregated than trees, and exhibit increased aggregation within canopy gaps. For trees, increased seed mass, wood density and shade tolerance were associated with less intense negative density dependence, while higher canopy layers and increased stature were associated with decreased aggregation and better dispersal. Spatial structure for trees was also strongly determined by dispersal syndrome. Averaged across all spatial scales, zoochory was more effective than wind dispersal, which was more effective than explosive dispersal. However, at intermediate scales, zoochory was associated with more aggregation than wind dispersal, potentially because of differences in short‐distance dispersal and the intensity of negative density dependence. Synthesis. We develop new tools for identifying significant associations between functional traits and spatial structure, and for linking these associations to quantitative estimates of dispersal scale and the strength and scale of density dependence. Our results help clarify how these processes influence woody plant species on Barro Colorado, and demonstrate how these tools can be applied to other sites and systems. [ABSTRACT FROM AUTHOR]

The effects of lightning on trees range from catastrophic death to the absence of observable damage. Such differences may be predictable among tree species, and more generally among plant life history strategies and growth forms. We used field-collected electrical resistivity data in temperate and tropical forests to model how the distribution of power from a lightning discharge varies with tree size and identity, and with the presence of lianas. Estimated heating density (heat generated per volume of tree tissue) and maximum power (maximum rate of heating) from a standardized lightning discharge differed 300% among tree species. Tree size and morphology also were important; the heating density of a hypothetical 10 m tall Alseis blackiana was 49 times greater than for a 30 m tall conspecific, and 127 times greater than for a 30 m tall Dipteryx panamensis. Lianas may protect trees from lightning by conducting electric current; estimated heating and maximum power were reduced by 60% (±7.1%) for trees with one liana and by 87% (±4.0%) for trees with three lianas. This study provides the first quantitative mechanism describing how differences among trees can influence lightning-tree interactions, and how lianas can serve as natural lightning rods for trees. [ABSTRACT FROM AUTHOR]

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 (Sa) 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) Sa was ca 25% higher in trees with lianas (10.2 ± 0.26) than trees lacking lianas (8.0 ± 0.51). Sa increased with tree size in liana-free trees (Sa = 10.99A0.256), 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. [ABSTRACT FROM AUTHOR]

Accurately describing the ecological relationships between species is more than mere semantics-doing so has profound practical and applied implications, not the least of which is that inaccurate descriptions can lead to fundamentally incorrect predicted outcomes of community composition and functioning. Accurate ecological classifications are particularly important in the context of global change, where species interactions can change rapidly following shifts in species composition. Here, we argue that many common ecological interactions-particularly competition and parasitism-can be easily confused and that we often lack empirical evidence for the full reciprocal interaction among species. To make our case and to propose a theoretical framework for addressing this problem, we use the interactions between lianas and trees, whose outcomes have myriad implications for the ecology and conservation of tropical forests (e.g., Schnitzer et al. 2015). [ABSTRACT FROM AUTHOR]

Tropical lianas deploy most of their leaves towards the top of the forest canopy, whereas trees exhibit a more stratified crown. Forest canopies are often exposed to hot and windy conditions, and how lianas cope with the extremely high transpirational demands under these environments remains unknown., We investigated stem hydraulic properties, leaf drought tolerance, diurnal changes in leaf and stem water potentials (Ψleaf and Ψstem), stomatal conductance ( gs), photosynthetic rate, sap flow and stem native percentage loss of conductivity ( PLC) for four liana species in a tropical forest in southwest China. Five co-occurring tree species were also selected for comparison., Lianas reached maximal transpiration at a relatively lower vapour pressure deficit (<1 kPa) than did co-occurring trees, suggesting vigorous photosynthesis during the morning. However, liana gs declined markedly over the day, with low gs at midday and afternoon. Lianas generally had higher stem sapwood-specific conductivity and maximum sap flux density but were less tolerant to drought-induced cavitation than were evergreen trees. Both lianas and trees lost leaf turgor in the top canopy at midday, but lianas lost leaf turgor earlier (∼2 h) than trees., Seven of eight species exhibited midday increases in PLC when xylem tensions were released to −0·3 to −0·5 MPa for PLC measurements. On average, lianas experienced high PLC (35·9%), along with a greater degree of disequilibrium between leaf and stem water potentials than trees (ΔΨstem-leaf: 1·37 MPa vs. 0·75 MPa) during the day. Earlier stomatal closure and efficient water transport may help lianas maintain higher Ψstem than trees despite having similar Ψleaf., Our results provide evidence that physiological regulation and efficient water transport mediate daily water relations in tropical lianas and may explain how lianas operate efficiently in tropical seasonal forests. Further studies involving a broader range of species are needed to confirm our findings. [ABSTRACT FROM AUTHOR]

Lianas are an important component of tropical forests, where they reduce tree growth, fecundity, and survival. Competition for light from lianas may be intense; however, the amount of light that lianas intercept is poorly understood. We used a large-scale liana-removal experiment to quantify light interception by lianas in a Panamanian secondary forest. We measured the change in plant area index ( PAI) and forest structure before and after cutting lianas (for 4 yr) in eight 80 m × 80 m plots and eight control plots (16 plots total). We used ground-based Li DAR to measure the 3-dimensional canopy structure before cutting lianas, and then annually for 2 yr afterwards. Six weeks after cutting lianas, mean plot PAI was 20% higher in control vs. liana removal plots. One yr after cutting lianas, mean plot PAI was ~17% higher in control plots. The differences between treatments diminished significantly 2 yr after liana cutting and, after 4 yr, trees had fully compensated for liana removal. Ground-based Li DAR revealed that lianas attenuated light in the upper- and middle-forest canopy layers, and not only in the upper canopy as was previously suspected. Thus, lianas compete with trees by intercepting light in the upper- and mid-canopy of this forest. [ABSTRACT FROM AUTHOR]

Prediction of carbon dynamics in response to global climate change requires an understanding of the processes that govern the distribution of carbon stocks. Above-ground biomass ( AGB) in tropical forests is regulated by variation in soil fertility, climate, species composition and topography at regional scales, but the drivers of fine-scale variation in tropical forest AGB are poorly understood. The factors that control the growth and mortality of individual trees may be obscured by the low resolution of studies at regional scales., In this paper, we evaluated the effects of soil nutrients, topography and liana abundance on the fine-scale spatial distribution of AGB and density of trees for a lowland tropical moist forest in Panama using additive regression models., Areas with larger values of AGB were negatively associated with the presence of lianas, which may reflect competition with lianas and/or the association of lianas with disturbed or open-canopy patches within forests. AGB was positively associated with soils possessing higher pH and K concentrations, reflecting the importance of below-ground resource availability on AGB independently of stem density., Synthesis. Our results shed new light on the factors that influence fine-scale tree AGB and carbon stocks in tropical forests: liana abundance is the strongest predictor, having a negative impact on tree AGB. The availability of soil nutrients was also revealed as an important driver of fine-scale spatial variation in tree AGB. [ABSTRACT FROM AUTHOR]

Lianas are a common plant growth form in tropical forests, where they compete intensely with trees, decreasing tree recruitment, growth, and survival. If the detrimental effects of lianas vary significantly with tree species identity, as is often assumed, then lianas may influence tree species diversity and community composition. Furthermore, recent studies have shown that liana abundance and biomass are increasing relative to trees in neotropical forests, which will likely magnify the detrimental effects of lianas and may ultimately alter tree species diversity, relative abundances, and community composition. Few studies, however, have tested the responses of multiple tree species to the presence of lianas in robust, well-replicated experiments. We tested the hypotheses that lianas reduce tree seedling growth and survival, and that the effect of lianas varies with tree species identity. We used a large-scale liana removal experiment in Central Panama in which we planted 14 replicate seedlings of 14 different tree species that varied in shade tolerance in each of 16 80 × 80 m plots (eight liana-removal and eight unmanipulated controls; 3136 total seedlings). Over a nearly two-yr period, we found that tree seedlings survived 75% more, grew 300% taller, and had twice the aboveground biomass in liana-removal plots than seedlings in control plots, consistent with strong competition between lianas and tree seedlings. There were no significant differences in the response of tree species to liana competition (i.e., there was no species by treatment interaction), indicating that lianas had a similar negative effect on all 14 tree species. Furthermore, the effect of lianas did not vary with tree species shade tolerance classification, suggesting that the liana effect was not solely based on light. Based on these findings, recently observed increases in liana abundance in neotropical forests will substantially reduce tree regeneration, but will not significantly alter tropical tree species diversity, relative abundance, or community composition. [ABSTRACT FROM AUTHOR]

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Plants compete with their neighbours for a finite set of limiting resources, and this decreases individual plant performance, growth and survival. However, neighbouring plants also affect each other in positive ways., Positive facilitative effects can occur when neighbouring plants ameliorate harsh abiotic conditions (temperature, wind and high irradiation). Thus, when environmental conditions are severe, the importance of facilitation may increase. The co-occurrence and masking effects of competition and facilitation among neighbouring plants have made it difficult to tease them apart in the past., We planted bur oak acorns ( Quercus macrocarpa) into an experimental diversity gradient in a central MN grassland that provided a gradient in plant biomass. We predicted that greater biomass of neighbours would increase both competition and facilitation as measured by impacts on the minimum leaf water potential reached on any given day. Under moderate conditions, competition should predominate, but under hot/dry conditions, facilitation should become more important. We measured temperature, humidity and soil moisture in these plots for two growing seasons, as well as oak seedling leaf water potential across a range of daily conditions., On cool/humid days, plant interactions were dominated by competition for soil water: leaf water potentials of juvenile oaks were lower in plots with greater herbaceous biomass (and higher diversity). Conversely, on hot/dry days, facilitation of the microclimate determined the net effect of plants on their neighbours: leaf water potentials of juvenile oaks were higher in plots with higher herbaceous diversity and biomass., Synthesis. In terms of plant water status, plant interactions among neighbours can flip from net negative (competition) to net positive (facilitation) depending on daily abiotic conditions. The relative importance of both positive and negative interactions for plant water status may affect the overall performance of plants over time. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Lianas in tropical forests compete intensely with trees for above- and belowground resources and limit tree growth and regeneration. Liana competition with adult canopy trees may be particularly strong, and, if lianas compete more intensely with some tree species than others, they may influence tree species composition. We performed the first systematic, large-scale liana removal experiment to assess the competitive effects of lianas on multiple tropical tree species by measuring sap velocity and growth in a lowland tropical forest in Panama. Tree sap velocity increased 60% soon after liana removal compared to control trees, and tree diameter growth increased 25% after one year. Although tree species varied in their response to lianas, this variation was not significant, suggesting that lianas competed similarly with all tree species examined. The effect of lianas on tree sap velocity was particularly strong during the dry season, when soil moisture was low, suggesting that lianas compete intensely with trees for water. Under the predicted global change scenario of increased temperature and drought intensity, competition from lianas may become more prevalent in seasonal tropical forests, which, according to our data, should have a negative effect on most tropical tree species. [ABSTRACT FROM AUTHOR]

Lianas exhibit peak abundance in tropical forests with strong seasonal droughts, the eco-physiological mechanisms associated with lianas coping with water deficits are poorly understood., We examined soil water partitioning, sap flow, and canopy eco-physiological properties for 99 individuals of 15 liana and 34 co-occurring tree species in three tropical forests that differed in soil water availability., In the dry season, lianas used a higher proportion of deep soil water in the karst forest ( KF; an area with severe seasonal soil water deficit ( SSWD)) and in the tropical seasonal forest ( TSF, moderate SSWD), permitting them to maintain a comparable leaf water status than trees in the TSF or a better status than trees in the KF. Lianas exhibited strong stomatal control to maximize carbon fixation while minimizing dry season water loss. During the dry period, lianas significantly decreased water consumption in the TSF and the KF. Additionally, lianas had a much higher maximum photosynthetic rates and sap flux density in the wet season and a lower proportional decline in photosynthesis in the dry season compared with those of trees. , Our results indicated that access to deep soil water and strong physiological adjustments in the dry season together with active wet-season photosynthesis may explain the high abundance of lianas in seasonally dry forests. [ABSTRACT FROM AUTHOR]

Treefall gaps are the "engines of regeneration" in tropical forests and are loci of high tree recruitment, growth, and carbon accumulation. Gaps, however, are also sites of intense competition between lianas and trees, whereby lianas can dramatically reduce tree carbon uptake and accumulation. Because lianas have relatively low biomass, they may displace far more biomass than they contribute, a hypothesis that has never been tested with the appropriate experiments. We tested this hypothesis with an 8-yr liana removal experiment in central Panama. After 8 years, mean tree biomass accumulation was 180% greater in liana-free treefall gaps compared to control gaps. Lianas themselves contributed only 24% of the tree biomass accumulation they displaced. Scaling to the forest level revealed that lianas in gaps reduced net forest woody biomass accumulation by 8.9% to nearly 18%. Consequently, lianas reduce whole-forest carbon uptake despite their relatively low biomass. This is the first study to demonstrate experimentally that plant-plant competition can result in ecosystem-wide losses in forest carbon, and it has critical implications for recently observed increases in liana density and biomass on tropical forest carbon dynamics. [ABSTRACT FROM AUTHOR]

Recent work has demonstrated that competition and facilitation likely operate jointly in plant communities, but teasing out the relative role of each has proven difficult. Here we address how competition and facilitation vary with seasonal fluctuations in environmental conditions, and how the effects of these fluctuations change with plant ontogeny. We planted three sizes of pine seedlings (Pinus strobus) into an herbaceous diversity experiment and measured pine growth every two weeks for two growing seasons. Both competition and facilitation occurred at different times of year between pines and their neighbors. Facilitation was important for the smallest pines when environmental conditions were severe. This effect decreased as pines got larger. Competition was stronger than facilitation overall and outweighed facilitative effects at annual time scales. Our data suggest that both competition and the counter-directional effects of facilitation may be more common and more intense than previously considered [ABSTRACT FROM AUTHOR]

Negative density dependence (NDD) and habitat specialization have received strong empirical support as mechanisms that explain tree species diversity maintenance and distribution in tropical forests. In contrast, disturbance appears to play only a minor role. Previous studies have rarely examined the relative strengths of these diversity maintenance mechanisms concurrently, and few studies have included plant groups other than trees. Here we used a large, spatially explicit data set from Barro Colorado Island, Panama (BCI) to test whether liana and tree species distribution patterns are most consistent with NDD, habitat specialization, or disturbance. We found compelling evidence that trees responded to habitat specialization and NDD; however, only disturbance explained the distribution of the majority of liana species and maintained liana diversity. Lianas appear to respond to disturbance with high vegetative (clonal) reproduction, and liana species' ability to produce clonal stems following disturbance results in a clumped spatial distribution. Thus, clonal reproduction following disturbance explains local liana spatial distribution and diversity maintenance on BCI, whereas negative density dependence and habitat specialization, two prominent mechanisms contributing to tree species diversity and distribution, do not. [ABSTRACT FROM AUTHOR]

Copyright of Biotropica is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

Copyright of Biotropica is the property of Wiley-Blackwell and its content may not be copied or emailed to multiple sites or posted to a listserv without the copyright holder's express written permission. However, users may print, download, or email articles for individual use. This abstract may be abridged. No warranty is given about the accuracy of the copy. Users should refer to the original published version of the material for the full abstract. (Copyright applies to all Abstracts.)

1. Lianas are a conspicuous element of many tropical forests, accounting for up to 40% of woody stem density and 20% of species richness in seasonal forests. However, lianas have seldom been surveyed at sufficiently large spatial scales to allow an assessment of the importance of habitat variables in structuring liana communities. 2. We compare the association patterns of 82 liana species and an equivalent sample of tree species on the 50 ha Forest Dynamics Project plot on Barro Colorado Island, Panama, with topographic habitat variables (high and low plateau, slope, swamp and streamside), and thirteen mapped soil chemical variables. In addition, we test for liana species associations with canopy disturbance using a canopy height map of the plot generated using light detection and ranging. 3. For all liana species combined, densities differed among topographic habitat types in the plot, with significantly higher densities on the seasonally drier lower plateau habitat (1044 individuals ha−1) than the moister slope habitat (729 individuals ha−1). Lianas were also significantly more abundant than expected in areas with low canopy height. 4. The proportion of liana species associated with one or more topographic habitat variables (44%) was significantly lower than that for trees (66%). Similarly, liana species were significantly less frequently associated with PC axes derived from soil chemical variables (21%) than trees (52%). The majority of liana species (63%) were significantly associated with areas of the plot with low canopy height reflecting an affinity for treefall gaps. 5. Synthesis. The habitat associations detected here suggest that liana density is associated primarily with canopy disturbance, and to a lesser extent with topography and soil chemistry. Relative to trees, few liana species were associated with local variation in topography and soil chemistry, suggesting that nutrient availability exerts only weak effects on liana community composition compared to trees. Results from this study support the contention that increases in forest disturbance rates are a driver of recently observed increases in liana abundance and biomass in neotropical forests. [ABSTRACT FROM AUTHOR]