Phenotypic correlates of the lianescent growth form: a review

Background

As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity.

Scope

In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species.

Conclusions

The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.     

Molecular phylogeny of hyperoliid treefrogs: biogeographic origin of Malagasy and Seychellean taxa and re-analysis of familial paraphyly

Treefrogs of the family Hyperoliidae are distributed in Africa, Madagascar and the Seychelles. In this study, their phylogeny was studied using sequences of fragments of the mitochondrial 16S and 12S rRNA and cytochrome b genes. The molecular data strongly confirmed monophyly of the subfamily Hyperoliinae but indicated that the genus Leptopelis (subfamily Leptopelinae) is more closely related to species of the African family Astylosternidae. The Seychellean genus Tachycnemis was the sister group of the Malagasy Heterixalus in all molecular analyses; this clade was deeply nested within the Hyperoliinae. A re-evaluation of the morphological data did not contradict the sister group relationships of these two genera. The subfamily Tachycneminae is therefore considered as junior synonym of the Hyperoliinae. In addition, the molecular analysis did not reveal justification for a subfamily Kassininae. Biogeographically, the origin of Malagasy hyperoliids may not be well explained by Mesozoic vicariance in the context of Gondwana breakup, as indicated by the low differentiation of Malagasy hyperoliids to their African and Seychellean relatives and by analysis of current distribution patterns.     

Developmental plasticity and biomechanics of treelets and lianas in Manihot aff. quinquepartita (Euphorbiaceae): a branch-angle climber of French Guiana

Background and Aims

Most tropical lianas have specialized organs of attachment such as twining stems, hooks or tendrils but some do not. Many climbers also have an early self-supporting phase of growth and in some species this can produce treelet-sized individuals. This study focuses on how a liana can climb without specialized attachment organs and how biomechanical properties of the stem are modulated between self-supporting treelets and canopy-climbing lianas.

Methods

Biomechanics and stem development were investigated in self-supporting to climbing individuals of Manihot aff. quinquepartita (Euphorbiaceae) from tropical rain forest at Saül, central French Guiana. Bending tests were carried out close to the site of growth. Mechanical properties, including Young''s elastic modulus, were observed with reference to habit type and changes in stem anatomy during development.

Key Results

This liana species can show a remarkably long phase of self-supporting growth as treelets with stiff, juvenile wood characterizing the branches and main stem. During the early phase of climbing, stiff but unstable stem segments are loosely held in a vertical position to host plants via petiole bases. The stiffest stems – those having the highest values of Young''s modulus measured in bending – belonged to young, leaning and climbing stems. Only when climbing stems are securely anchored into the surrounding vegetation by a system of wide-angled branches, does the plant develop highly flexible stem properties. As in many specialized lianas, the change in stiffness is linked to the development of wood with numerous large vessels and thin-walled fibres.

Conclusions

Some angiosperms can develop highly effective climbing behaviour and specialized flexible stems without highly specialized organs of attachment. This is linked to a high degree of developmental plasticity in early stages of growth. Young individuals in either open or closed marginal forest conditions can grow as substantial treelets or as leaning/climbing plants, depending on the availability of host supports. The species of liana studied differs both in terms of development and biomechanics from many other lianas that climb via twining, tendrils or other specialized attachment organs.Key words: Biomechanics, bending, developmental plasticity, French Guiana, liana, Manihot aff. quinquepartita (Euphorbiaceae), treelet, branch angle climber, Young''s modulus     

Leaves of Lianas and Self-Supporting Plants Differ in Mass per Unit Area and in Nitrogen Content

Abstract: The aim of this study was to test the hypothesis that the reduction in supporting tissues in climbers compared to self-supporting plants is also true for their leaves, and that climbers generally require higher leaf nitrogen than self-supporting plants to accomplish fast growth. This hypothesis was tested using paired samples of both growth forms with assessment of leaf area index above the sampled plants (LAI_a) in a tropical rain forest in Gabon. The sampling protocol ensured that within a highly fluctuating low canopy environment, the growth conditions were identical for each pair sampled. The results confirmed the hypothesis. Lianas had significantly lower leaf mass per unit leaf area (LMA) than their supporters. Liana leaves also contained significantly more nitrogen than host tree leaves. The differences in nitrogen concentration between liana and tree leaves reversed for the most shaded sites, when nitrogen was expressed on a leaf area base (N_area). Significant regression between leaf nitrogen and LAI_a was found for the climbers on the shaded sites but not for their supporters. This indicated better acclimation of climbers to prevailing light conditions. Better nitrogen allocation at low LMA, together with lower carbon costs for building supporting tissues, makes lianas highly competitive, especially where high nitrogen availability is assured.     

Mechanical,chemical and X-ray analysis of wood in the two tropical lianas<Emphasis Type="Italic"> Bauhinia guianensis</Emphasis> and<Emphasis Type="Italic"> Condylocarpon guianense</Emphasis>: variations during ontogeny

Mechanical and chemical properties as well as microfibril angles of wood tissues from different ontogenetic stages are determined for the neotropical lianas Bauhinia guianensis and Condylocarpon guianense. The mechanical properties include the elastic moduli under bending and under dynamic torsion. The chemical analyses cover (i) the content of cellulose, lignin and hemicelluloses fractions, (ii) the monomeric composition of the uncondensed lignin, and (iii) the composition of the hemicelluloses with respect to neutral monosaccharides. By comparing the wood properties of these lianas with the corresponding properties of wood from self-supporting deciduous trees, common characteristics and differences are revealed. Additionally, the changes in the lignin and polysaccharides fractions as well as the variations in microfibril orientation that occur during ontogeny of the two liana species are discussed with regard to their implications for the mechanical properties of wood.     

Mechanical role of the leaf sheath in rattans

Leaf sheaths of rattans are long, tubular and persistent and unlike many self-supporting palms, extend far from the apex of the plant. The mechanical role of the leaf sheath was investigated in eight rattan species of the subfamily Calamoideae. The main objective was to analyse its influence on the mechanical architecture and contribution to the climbing habit. Bending mechanical properties were measured along climbing axes before and after removal of leaf sheaths. Results were related to stem and leaf sheath geometry and mechanical properties. Contribution of the leaf sheath to axial flexural rigidity was high (c. 90%) in the early stages of growth and towards the apex of older climbing axes for all climbing palms tested. Senescence and loss of the leaf sheath strongly influenced axial stiffness. A nonclimbing species, Calamus erectus, showed a different mechanical architecture. Although lacking secondary growth, palms have been able to develop successful climbers with a mechanical architecture broadly analogous to, although developmentally different from, dicotyledonous lianas. The role of the leaf sheath in modulating mechanical properties during ontogeny ought not to be neglected in studies on monocotyledons, as it possibly contributed significantly to the ways in which different growth forms have evolved in the group.     

Climate niches of milkweeds with plesiomorphic traits (Secamonoideae; Apocynaceae) and the milkweed sister group link ancient African climates and floral evolution

? Premise of the study: Climate change that increases mortality of plants and pollinators can create mate-finding Allee effects and thus act as a strong selective force on floral morphology. Milkweeds (Secamonoideae and Asclepiadoideae; Apocynaceae) are typically small plants of seasonally dry habitats, with pollinia and high pollen-transfer efficiency. Their sister group (tribe Baisseeae and Dewevrella) is mostly comprised of giant lianas of African rainforests, with pollen in monads. Comparison of the two groups motivated a new hypothesis: milkweeds evolved in the context of African aridification and the shifting of rainforest to dry forest. Pollinia and high pollen-transfer efficiency may have been adaptations that alleviated mate-finding Allee effects generated by high mortality during droughts. We formally tested whether milkweeds have a drier climate niche by comparing milkweeds with plesiomorphic traits (Secamonoideae) and the milkweed sister group in continental Africa. ? Methods: We georeferenced specimens of the milkweed sister group and Secamonoideae in continental Africa, extracted 19 climatic variables from the Worldclim model, conducted factor analysis to identify correlated suites of variables, and compared the frequency distributions of the two lineages relative to each factor. ? Key results: The distributions of Secamonoideae and the milkweed sister group differed significantly relative to four factors, each correlated with a distinct suite of climate parameters: (1) air temperature (Secamonoideae: cooler), (2) total and (3) summer precipitation (Secamonoideae: drier), and (4) temperature seasonality and isothermality (Secamonoideae: more seasonal and less isothermal). ? Conclusions: Secamonoideae in continental Africa inhabit drier, cooler sites than do the milkweed sister group, consistent with a shift from rainforests to dry forests in a cooling climate.     

Plant growth forms: an ecological and evolutionary perspective

Trees, shrubs, lianas and herbs have widely different mechanical architectures, which can also vary phenotypically with the environment. This review investigates how environmental effects, particularly mechanical perturbation, can influence biomechanical development in self-supporting and climbing growth forms. The bifacial vascular cambium is discussed in terms of its significance to growth form variation, ecology and evolution among extant plants, and during its appearance and early evolution. A key aspect of this developmental innovation concerned its potential for architectural and mechanical variation in response to environmental effects as well as optimizing hydraulic supply before the appearance of laminate leaves. Growth form diversity and its importance to past and present ecosystems are discussed in relation to both evolutionary constraints and ecological factors such as climatic change and atmospheric CO2 concentrations. We discuss how widely ranging growth forms such as climbers show a large range of developmental and phenotypic variation that has much to offer in understanding how the environment can modify plant development, particularly in terms of the bifacial vascular cambium. The broad approach we propose would benefit a wide range of studies from research into wood development to long-term ecological censuses of today's potentially changing ecosystems.     

Responses of macrophytes to dewatering: effects of phylogeny and phenotypic plasticity on species performance

Temporary dewatering constitutes a drastic change in conditions for aquatic vegetation. Species’ sustained performance under these conditions relies partly on their ability to produce a terrestrial phenotype. Such adaptations may include the development of self-supporting aboveground organs with higher dry matter content enabling plants to withstand gravity and smaller leaves with thicker cuticle to reduce evapotranspiration, leading to lower specific leaf area, higher leaf-construction costs and consequently higher leaf life span. The ability of aquatic plant species to produce a terrestrial-adapted phenotype may differ according to growth form and evolutionary history. The objectives of this study were to (1) measure the effects of dewatering on aquatic plant performance, (2) determine how growth form and phylogenetic position affect performance, and (3) relate plant performance to plasticity. To meet these objectives, we experimentally studied aquatic plant responses to dewatering by measuring survival, growth, and a set of traits describing the morphology and leaf-resource economy of eight aquatic plant species with contrasting phylogeny and growth forms. The ability of aquatic plants to withstand dewatering differed according to phylogeny but not to growth form. The eudicots presented high survival and similar growth rates under terrestrial compared to aquatic conditions, while monocots generally did not survive dewatering. These species produced phenotypic adjustments, such as denser aboveground organs and leaf plasticity, which can explain the maintenance of similar growth rates under terrestrial conditions. The relatively strong plasticity and performance of eudicots in terrestrial habitats suggests that their optimal niche is the interface between aquatic and terrestrial ecosystems.     

Phylogeny of muroid rodents: relationships within and among major lineages as determined by IRBP gene sequences

The rodent family Muridae is the single most diverse family of mammals with over 1300 recognized species. We used DNA sequences from the first exon ( approximately 1200bp) of the IRBP gene to infer phylogenetic relationships within and among the major lineages of muroid rodents. We included sequences from every recognized muroid subfamily except Platacanthomyinae and from all genera within the endemic Malagasy subfamily Nesomyinae, all recognized tribes of Sigmodontinae, and a broad sample of genera in Murinae. Phylogenetic analysis of the IRBP data suggest that muroid rodents can be sorted into five major lineages: (1) a basal clade containing the fossorial rodents in the subfamilies Spalacinae, Myospalacinae, and Rhizomyinae, (2) a clade of African and Malagasy genera comprising the subfamilies Petromyscinae, Mystromyinae, Cricetomyinae, Nesomyinae, and core dendromurines, (3) a clade of Old World taxa belonging to Murinae, Otomyinae, Gerbillinae, Acomyinae, and Lophiomyinae, (4) a clade uniting the subfamilies Sigmodontinae, Arvicolinae, and Cricetinae, and (5) a unique lineage containing the monotypic Calomyscinae. Although relationships among the latter four clades cannot be resolved, several well-supported supergeneric groupings within each are identified. A preliminary examination of molar tooth morphology on the resulting phylogeny suggests the triserial murid molar pattern as conceived by evolved at least three times during the course of muroid evolution.     

The derivation, utility and implications of a divergence index for the fynbos genus Leucadendron (Proteaceae)

The derivation, utility, and implications of a divergence index for the fynbos genus Leucadendron (Proteaceae). The genus Leucadendron is useful for a phylogenetic study because it contains many species (79), is morphologically very diverse and appears to have diverged early on in the history of the family in Africa. Polarity of many characters was determined by outgroup analysis and the ontogenetic method. Exceptionally high levels of convergence made the determination of the phylogeny of the subsections problematic. An index, which is the sum of weight of all derived characters present in a species, was determined.
The genus should be removed from subfamily Aulacinae and placed closer to the Leucospermum 'alliance'. I suggest an arid temperate, shrub-like origin for the African Proteaceae, which is in contrast to some published viewS. Modifications to accepted concepts of fruit homology and evolutionary trends such as pollination and dispersal are presented for the family. The divergence index, if used as a 'morphological clock', appears to be useful for further biogeographic and ecological analyses.     

Nuclear gene phylogeny of narrow-mouthed toads (Family: Microhylidae) and a discussion of competing hypotheses concerning their biogeographical origins

The family Microhylidae has a large circumtropic distribution and contains about 400 species in a highly subdivided taxonomy. Relationships among its constituent taxa remained controversial due to homoplasy in morphological characters, resulting in conflicting phylogenetic hypotheses. A phylogeny based on four nuclear genes (rag-1, rag-2, tyrosinase, BDNF) and one mitochondrial gene (CO1) of representatives of all currently recognized subfamilies uncovers a basal polytomy between several subfamilial clades. A sister group relationship between the cophylines and scaphiophrynines is resolved with moderate support, which unites these endemic Malagasy taxa for the first time. The American members of the subfamily Microhylinae are resolved to form a clade entirely separate from the Asian members of that subfamily. Otophryne is excluded from the subfamily Microhylinae, and resolved as a basal taxon. The placement of the Asian dyscophine Calluella nested within the Asian Microhyline clade rather than with the genus Dyscophus is corroborated by our data. Bayesian estimates of the divergence time of extant Microhylidae (47-90 Mya) and among the subclades within the family are discussed in frameworks of alternative possible biogeographic scenarios.     

Evolutionary history and climate conditions constrain the flower colours of woody plants in China

进化历史和气候条件共同影响中国木本植物花色的分布 本研究以中国木本植物为研究对象,主要探讨两个问题：(1)不同生活型物种花色组成的差异;(2)生物地理区、进化年龄和气候条件对不同花色地理分布格局的影响。研究使用7673种木本植物的物种分布数据和花色信息(分为白色、红色、黄色、黄绿色、绿色和蓝紫色),并结合属级系统进化树来比较不同生活型(包括灌木、乔木和藤本)物种花色组成的差异,分析不同生物地理区、进化年龄和现代气候对花色地理格局的影响。研究结果表明,与乔木和藤本植物相比,灌木具有更高比例 的由花青素着色的红色花和蓝紫色花物种。中国木本植物的花色地理格局受到区域效应和现代气候(尤其是降水和UVB辐射)的共同影响。倾向于蜂媒传粉的黄色花和蓝紫色花物种和由花青素着色、耐环境胁迫的红色花和蓝紫色花物种比例在中国西北部地区更高。绿色花物种的进化起源更早,但进化时间对花色地理格局的解释力很弱。这些结果说明中国木本植物花色的地理格局受到进化历史和现代环境的共同影响。     

Do lianas really have wide vessels? Vessel diameter–stem length scaling in non-self-supporting plants

Lianas and other climbing plants are known for their extraordinarily wide vessels. Wide vessels are thought to contribute to the extreme hydraulic efficiency of lianas and to play a part in their ability to dominate many tropical habitats, and even their globally increasing abundance with anthropic disturbance. However, recent hydraulic optimality models suggest that the average vessel diameter of plants generally is the result of tip-to-base vessel widening reflecting the effects of selection buffering conductive path length-imposed hydraulic resistance. These models state that mean vessel diameter should be predicted by stem length, by implication even in lianas. We explore vessel–stem relations with 1409 samples from 424 species in 159 families of both self- and non-self-supporting plants. We show that, far from being exceptional in their vessel diameter, lianas have average natural (not hydraulically weighted) vessel diameters that are indistinguishable for a given stem length from those in self-supporting plants. Lianas do, however, have wider variance in vessel diameter. They have a small number of vessels that are wider than those in self-supporting plants of similar stem lengths, and also narrower vessels. This slightly greater variance is sufficient to make hydraulically weighted vessel diameters in lianas higher than those of self-supporting counterparts of similar stem lengths. Moreover, lianas have significantly more vessels per unit of wood transection than self-supporting plants do. This subtle combination of slightly higher vessel diameter variance and higher vessel density for a given stem length is likely what makes lianas hydraulically distinctive, rather than their having vessels that are truly exceptionally wide.     

Opening the black box: phylogenetics and morphological evolution of the Malagasy fossorial lizards of the subfamily "Scincinae"

The island of Madagascar harbors a highly endemic vertebrate fauna including a high diversity of lizards of the subfamily "Scincinae," with about 57 species in eight genera. Since limb reduction seems to have been a common phenomenon during the evolution of Malagasy "scincines," diagnosing evolutionary relationships based on morphology has been difficult. Phylogenetic analyses of multiple mitochondrial DNA sequences including the entire ND1, tRNA(LEU), tRNA(ILE), tRNA(GLN) genes, and fragments of the 12S and 16S rRNA and tRNA(MET) genes were conducted to test the monophyly of the largest genus Amphiglossus, and to evaluate the various formal and informal species groupings previously proposed for this species-rich group. A further objective was to determine the phylogenetic placements of the several greatly limb-reduced and limbless Malagasy "scincines" and ascertain whether any of these are derived from within the morphologically plesiomorphic Amphiglossus. As limb reduction in skinks is mostly associated with body elongation via an increase in the number of presacral vertebrae, we evaluate the pattern of evolution of the numbers of presacral vertebrae in the context of our phylogeny. We demonstrate that Amphiglossus as currently diagnosed is non-monophyletic, and the species fall into two major groups. One of these groups is a clade that contains the included species of the subgenus Amphiglossus (Madascincus) among other species and is a member of a larger clade containing Paracontias and Pseudoacontias. In the second group, the nominate subgenus Amphiglossus (Amphiglossus) forms several subclades within a larger clade that also contains Androngo crenni and Pygomeles braconnieri, and is sister to Voeltzkowia. All analyses provide strong support for the monophyly of Paracontias and Voeltzkowia. Based on the preferred phylogenetic hypothesis and weighted squared-change parsimony we show that the ancestor of the Malagasy clade was already elongated and had a moderately high number of presacral vertebrae (46-48), which is hypothesized to be the ancestral condition for the whole Malagasy "scincine" clade. We further demonstrate that both multiple increases and reductions of presacral vertebrae evolved in many clades of Malagasy "scincines" and that the use of presacral vertebrae as a major character to diagnose supraspecific units is dubious. Based on our results and published morphological evidence we consider Scelotes waterloti Angel, 1930 to be a junior synonym of Amphiglossus reticulatus (Kaudern, 1922).     

Phylogeny and biogeography of cichlid fishes (Teleostei: Perciformes: Cichlidae)

Family level molecular phylogenetic analyses of cichlid fishes have generally suffered from a limited number of characters and/or poor taxonomic sampling across one or more major geographic assemblage, and therefore have not provided a robust test of early intrafamilial diversification. Herein we use both nuclear and mitochondrial nucleotide characters and direct optimization to reconstruct a phylogeny for cichlid fishes. Representatives of major cichlid lineages across all geographic assemblages are included, as well as nearly twice the number of characters as any prior family‐level study. In a strict consensus of 81 equally most‐parsimonious hypotheses, based on the simultaneous analysis of 2222 aligned nucleotide characters from two mitochondrial and two nuclear genes, four major subfamilial lineages are recovered with strong support. Etroplinae, endemic to Madagascar (Paretroplus) and southern Asia (Etroplus), is recovered as the sister taxon to the remainder of Cichlidae. Although the South Asian cichlids are monophyletic, the Malagasy plus South Asian lineages are not. The remaining Malagasy lineage, Ptychochrominae, is monophyletic and is recovered as the sister group to a clade comprising the African and Neotropical cichlids. The African (Pseudocrenilabrinae) and Neotropical (Cichlinae) lineages are each monophyletic in this reconstruction. The use of multiple molecular markers, from both mitochondrial and nuclear genes, results in a phylogeny that in general exhibits strong support, notably for early diversification events within Cichlidae. Results further indicate that Labroidei is not monophyletic, and that the sister group to Cichlidae may comprise a large and diverse assemblage of percomorph lineages. This hypothesis may at least partly explain why morphological studies that have attempted to place Cichlidae within Percomorpha, or that have tested cichlid monophyly using only “labroid” lineages, have met with only limited success. © The Willi Hennig Society 2004.     

Major trends in stem anatomy and growth forms in the perianth-bearing Piperales,with special focus on Aristolochia

Background and Aims

The order Piperales has the highest diversity of growth forms among the earliest angiosperm lineages, including trees, shrubs, climbers and herbs. However, within the perianth-bearing Piperales (Asarum, Saruma, Lactoris, Hydnora, Prosopanche, Thottea and Aristolochia), climbing species only occur in the most species-rich genus Aristolochia. This study traces anatomical and morphological traits among these lineages, to detect trends in growth form evolution and developmental processes.

Methods

Transverse stem sections of different developmental stages of representatives of Asarum, Saruma, Lactoris, Hydnora, Thottea and Aristolochia were compared and anatomical traits were linked to growth form evolution. Biomechanical properties of representative climbers were determined in three-point bending tests and are discussed based on the anatomical observations. Growth form evolution of the perianth-bearing Piperales was reconstructed by ancestral character state reconstruction using Mesquite.

Key Results

While species of Asarum and Saruma are exclusively herbaceous, species of the remaining genera show a higher diversity of growth habit and anatomy. This growth form diversity is accompanied by a more complex stem anatomy and appropriate biomechanical properties. The ancestral growth form of the perianth-bearing Piperales is reconstructed with either a shrub-like or herbaceous character state, while the following three backbone nodes in the reconstruction show a shrub-like character state. Accordingly, the climbing habit most probably evolved in the ancestor of Aristolochia.

Conclusions

Since the ancestor of the perianth-bearing Piperales has been reconstructed with a herb- or shrub-like habit, it is proposed that the climbing habit is a derived growth form, which evolved with the diversification of Aristolochia, and might have been a key feature for its diversification. Observed anatomical synapomorphies, such as the perivascular fibres in Lactoris, Thottea and Aristolochia, support the phylogenetic relationship of several lineages within the perianth-bearing Piperales. In addition, the hypothesis that the vegetative organs of the holoparasitic Hydnoraceae are most probably rhizomes is confirmed.     

Optimisation of spatial allocation patterns in lianas compared to trees used for support

There are only limited possibilities to study the competition between trees and lianas in the top canopy of tropical rain forests. Furthermore, the important question how the leaf traits are related to twig support is rarely studied, especially regarding growing space partitioning between the self-supporting and the climbing growth form. Our study used the hot-air balloon within the “Operation Canopee” in the Masoala National Park, Madagascar, to test the differences in spatial allocation patterns of leaves and twigs in lianas and tree parts used for support. The sampling design emphasised to obtain a common assembly of twigs and leaves from both, trees and lianas. The results from the top canopy were compared to the data from the understorey regarding biomass and nutrients in leaves and distal twigs. In the understorey the reduction in structural investment was much stronger in lianas than in trees. The results showed that lianas reduced carbon investment per volume, but increased leaf nitrogen concentration and leaf area ratio (LAR), the latter driven by a reduction in leaf mass per area (LMA). In the top canopy, lianas contributed about one third of the leaf area density of 3 m² m⁻³. For distal twigs, no relationship was found between twig biomass per volume and leaf area density for trees, but lianas balanced both structural parameters closely. The climbers benefit from the external support provided by the trees and optimise the area of assimilation tissue at low per volume investment for mechanical stability. Several traits such as low LMA and high leaf nitrogen concentrations together with higher LAR and optimised spatial investment advantage the climbing growth form and enable a fast acquisition of growing space. The results emphasize the necessity to consider spatial and structural features of growing space acquisition when dealing with plant competition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.