![Angiosperm phylogeny (from source site Stevens, P.F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 [and more or less continuously updated since].) illustrating the distribution of cambial variants throughout the tree, and the concentration of this feature in the Eudicots, mainly in the Rosidae and Asteridae (inspired in Spicer and Groover 201082.
Spicer , R and
Groover , A. 2010. Evolution of development of vascular cambia and secondary growth. New Phytologist, 186: 577–592. View all references). For references on the distribution of cambial variants by family check Table 1.](https://www.researchgate.net/profile/Lucia-Lohmann/publication/235956623/figure/fig7/AS:316694351368201@1452517225627/Angiosperm-phylogeny-from-source-site-Stevens-PF-2001-onwards-Angiosperm-Phylogeny_Q320.jpg)
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An overview of the anatomy, development and evolution of the vascular system of lianas
Abstract and Figures
Background: Lianas present many interesting structural features that are linked to their climbing habit. Having lost substantial amounts of supporting tissue, these plants depend on external structures for support. Meanwhile, during their evolutionary history, they have gained additional conductive and storage tissues. The wood of lianas generally includes wider vessels, larger amounts of axial parenchyma, larger rays, and longer fibres than those of trees. Cambial variants represent another key anatomical feature of lianas.
Aims: In this paper, we review various aspects of liana biology, including those associated with their vascular system and water conduction, secondary growth and seasonal responses to environmental variability, as well as aspects related to the evolution of their cambial variants. Methods: Examples from the Bignoniaceae and Leguminosae, the two most abundant liana taxa in the Neotropics, are presented in a series of case studies, bringing new data, such as the activity of the cambium during the dry seasons; the radial conducting elements that are associated with the habit; the cambial variant of Bignonieae that has evolved in a recapitulatory fashion; and the increased specialisation for photosynthate conduction by the phloem.
Conclusions: Altogether, lianas represent an excellent model for studies on the convergent evolution of plants.
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... Surprisingly, Carlquist's definition-and those that post-dated him (e.g. Spicer and Groover 2010;Angyalossy et al. 2012Angyalossy et al. , 2015-focusses on cambial activity and modifications during secondary growth, with no reference to procambial patterning (primary growth), despite the multiple reports on modifications in procambial organization in both fossil and extant plant lineages (Worsdell 1906;Esau 1965;Beck 2010). Because alternative vascular ontogenies may originate from modifications in procambium patterning, the term 'cambial variant' is not the most appropriate to broadly describe these phenomena, confusing our scientific understanding of this concept. ...
... Manekia) form a bifacial cambium from the middle ring of medullary bundles producing a continuous cylinder of secondary xylem and secondary phloem (Fig. 3B). Each bundle in the external ring experiences secondary growth, whereby each develops into an 'external vascular cylinder' (Fig. 3B;Angyalossy et al. 2012Angyalossy et al. , 2015. Medullary bundles from the inner ring continue isolated in the pith, which may also undergo secondary growth (Carlquist 2001). ...
... corded stem, Fig. 6). Another example involves 'external vascular cylinders' described for distantly related angiosperms families, including Bignoniaceae, Combretaceae, Euphorbiaceae, Piperaceae, Rubiaceae and Sapindaceae (Angyalossy et al., 2012(Angyalossy et al., , 2015 and in which the 'external vascular cylinders' derived from the activity of both primary and secondary meristems. In Piperaceae, this pattern originates as a procambial variant, in which isolated medullary bundles undergo secondary growth generating the 'external vascular cylinders' (Fig. 3). ...
Over centuries of plant morphological research, biologists have enthusiastically explored how distinct vascular arrangements have diversified. These investigations have focused on the evolution of steles and secondary growth and examined the diversity of vascular tissues (xylem and phloem), including atypical developmental pathways generated through modifications to the typical development of ancestral ontogenies. A shared vernacular has evolved for communicating on the diversity of alternative ontogenies in seed plants. Botanists have traditionally used the term “anomalous secondary growth” which was later renamed to “cambial variants” by late Dr. Sherwin Carlquist (1988). However, the term “cambial variants” can be vague in meaning since it is applied for developmental pathways that do not necessarily originate from cambial activity. Here, we review the “cambial variants” concept and propose the term “vascular variants” as a more inclusive overarching framework to interpret alternative vascular ontogenies in plants. In this framework, vascular variants are defined by their developmental origin (instead of anatomical patterns), allowing the classification of alternative vascular ontogenies into three categories: (1) procambial variants, (2) cambial variants and (3) ectopic cambia. Each category includes several anatomical patterns. Vascular variants, which represent broader developmental-based groups, can be applied to both extant and fossil plants, and thereby offer a more adequate term from an evolutionary perspective. An overview of the developmental diversity and phylogenetic distribution of vascular variants across selected seed plants is provided. Finally, the evolutionary implications of vascular variants are discussed.
... These descriptions greatly contrast with Valerianoideae species in the Northern Hemisphere, which are characterized by the regular eustelic organization with a single cambium producing inner secondary xylem and outer secondary phloem, with xylem parenchyma scant or absent (Solereder 1908, Metcalfe andChalk 1957). A detailed study on the anatomy of South American Valerianoideae is necessary since wood anatomy has been proven to be an important source of data for studying evolution (Carlquist 1980, Onyenedum andPace 2021), including its relatedness with specific growth forms (Angyalossy et al. 2012, Pace andAngyalossy 2013). ...
... Because all these species have a rosulate leaf arrangement, these anatomical variants may complement the protection provided by their external morphology. These cambial variants do not represent disordered growth and have a polyphyletic origin in the angiosperms (Carlquist 2001), and are particularly common in lianas (Angyalossy et al. 2012). Surprisingly, no cambial variants have been described for Valeriana lianas in this or previous studies. ...
... The presence of many parts of unlignified parenchyma may be justified because this habit does not require supportive tissue since it grows on a phorophyte or other support. The presence of unlignified, soft elements blending with rigid elements is quite common in lianas (Angyalossy et al. 2012, Bastos et al. 2016 and is related to the flexibility and torsion resistance of these plants. However, this may be a secondary function, as the non-lianescent ancestor of this group already had these parts of unlignified wood. ...
Andean species of Valeriana are frequently pointed to as an example of island woodiness, i.e. plants with herbaceous ancestors that usually evolve woodier forms on islands. We investigated this phenomenon through morphoanatomical and phylogenetic analyses. Plants were collected in the Páramos of Ecuador and had their vegetative morphology described. Stems were sectioned for histological analyses. We mapped the morphoanatomical data plus the maximum reported size for these and other species of the genus, on phylogenetic trees reconstructed on the basis of previously published sequences. Bigger than their Holarctic counterparts, the ancestor of the South American Valeriana was likely to have had a maximum size of 132 cm, and then after evolved to bigger and smaller sizes in a pattern similar to Brownian motion, as supported by phylogenetic signal values. We classified the collected plants into six growth forms (rosette herbs, semirosette herbs, elevated rosettes, elevated semirosettes, shrubs, and lianas), that are not directly related to variable levels of woodiness, as pointed out by our histological analyses. However, the production of unlignified parenchyma in the wood is very frequent, except in older regions of shrubs. The ancestor of the South American Valeriana is also very likely to have unlignified parenchyma in its wood.
... The ability of lianas to expand their stem systems in conditions of low water availability is due to their ability to store water and nutrients in the non-lignified parenchyma of their stems (Mooney and Gartner, 1991;Angyalossy et al., 2012). After flowering and fruiting events, lianas lose their aerial biomass, remaining alive through underground systems where reserves are allocated during the leafless phase (Mooney and Gartner, 1991;Angyalossy et al., 2012). ...
... The ability of lianas to expand their stem systems in conditions of low water availability is due to their ability to store water and nutrients in the non-lignified parenchyma of their stems (Mooney and Gartner, 1991;Angyalossy et al., 2012). After flowering and fruiting events, lianas lose their aerial biomass, remaining alive through underground systems where reserves are allocated during the leafless phase (Mooney and Gartner, 1991;Angyalossy et al., 2012). In our study, we observed that carbohydrate reserves in the form of amyloplasts were located only in the endodermis of the petiole of the stem in primary and secondary growth. ...
... For example, in M. cordifolia, the vascular cambium has a variant activity; the interfascicular vascular cambium only has a radial initial, forming only broad rays facing the secondary xylem and phloem. Some authors call this "xylem in plates" (Carlquist, 2001) and others "axial elements separated by rays" (Angyalossy et al., 2012). This cambium variation can also be observed in Aristolochia L. (Trueba et al., 2015). ...
... 400 samples of lianas, affirms that a simple cut on the transverse plane of their stem "provides a highly efficient descriptive tool for the identification of taxa (families, genus and species)"; in the absence of fertile material, this is a valuable tool that can be used by specialists. In addition, liana featuressuch as cambial variants -are commonly taxonspecific (Angyalossy et al. 2015). For instance, Amorim et al. (2017) mention that parenchyma type and crystal abundance support the distinction between Heteropterys Kunth species, a Malpighiaceae genus that is similar to Banisteriopsis. ...
... Wood anatomical features were analyzed using the lianescent xylem. According to Angyalossy et al. (2015), it has vessels dimorphism, cambial variants, fewer fibers (mostly gelatinous), greater quantity of parenchyma cells, and area of conduction (large vessels). The caupuri ethnotaxon samples were analyzed in the internode region only, since the node area usually has no defined planes. ...
... The terminology used in the anatomical descriptions was based on Angyalossy et al. (2015), IAWA committee (1989), and Pace et al. (2018), and the measurements were based on IAWA committee (1989) recommendations. In addition, for species that have vessel dimorphism, such as lianas, IAWA committee (1989) recommends recording the larger size class (mean tangential diameters 100-200 μm large; > 200 very large). ...
The worldwide use of ayahuasca, an entheogenic tea originally used in ceremonial context by Amazonian peoples, has increased in the last decade. In Brazil, “seringueiros” or rubber tappers have incorporated its use into urban settings, creating three main religions that continue to drink the sacred tea: Santo Daime, Barquinha, and União do Vegetal. A neo-ayahuascan network has arisen along with the expansion of ayahuasca, with distinct meanings associated with the tea, including using it for therapeutic, artistic, religious, and playful purposes. Brazilian ayahuascan groups (BAGs) commonly prepare the tea using the stems of Banisteriopsis caapi (Malpighiaceae) along with the leaves of Psychotrya viridis (Rubiaceae) or sometimes with those of Diplopterys cabrerana (Malpighiaceae). There are citations of traditional Amazonian peoples using the stem of other Malpighiaceae species to prepare ayahuasca in the literature, also several papers refer to “ethnotaxa” of B. caapi recognition. Fertile samples must be collected in order to identify genera and species of Malpighiaceae, which is not always possible due to the short flowering period. There is also a difficulty in collecting sacred plants due to access limitation inside the traditional communities, and due to the natural obstacles in liana sampling in the Amazon Forest. Data in the literature shows that wood anatomy is a good tool to identify plants. Our study synthesizes ethnobotanical material and wood anatomy data in order to understand: (1) whether there is consensus between the respondents of the BAGs of the identity of ethnotaxa; (2) whether there is a hierarchical classification of the ethnotaxa, and whether it can be described and reproduced, illustrating the variation of B. caapi in more detail; (3) whether stem anatomy can be used to distinguish the genera and ethnotaxa used in the production of ayahuasca; and (4) whether there is homogeneity in the use of ethnotaxa or if any of them is more commonly used. Thirty-eight people belonging to BAGs were interviewed. Eighteen ethnotaxa and 30 names for B. caapi were documented. Among the respondents, there are disagreements about the ethnotaxa identities. There is an ethnotaxa hierarchical classification based on stem morphology, which groups them as lianas with swollen nodes (caupuri) and without swollen nodes. However, the tea chemical effects are equally important, as there are lianas called caupuri which do not have swollen nodes, but have the same effect. Wood anatomy and morphology can help in understanding the categorization of ethnotaxa. Stem wood anatomy was also used to verify the identity of Diplopterys cf. pubipetala, which was cited by one respondent. Some ethnotaxa are more used for their chemical effects or their ease of cultivation. Our data suggest that B. caapi has some degree of domestication and that BAGs help to maintain a significant portion of B. caapi diversity. The role of these groups as plant “guardians” used to make ayahuasca is unique at this historical moment, in which the Amazon rainforest and its great diversity is being cruelly and irresponsibly decimated.
... Some have suggested to us that it might be that narrow-stemmed plants do not produce wide vessels because there is some developmental linkage between vessel diameter and stem diameter. This possibility is rejected by lianas, which routinely produce very wide vessels in slender stems (Carlquist 1985a;Angyalossy et al. 2012Angyalossy et al. , 2015. Similarly, distal roots far from the plant base also have very wide vessels for their diameter . ...
... Lianas have wider vessel diameter variances than self-supporting plants (Carlquist 1985a;Ewers et al. 1997;Angyalossy et al. 2012Angyalossy et al. , 2015, a pattern consistent with the vulnerability-diameter link. Lianas are well known for supposedly having very wide vessels. ...
The best explanations of the relationship between organismal form and function-those regarded by scientists as the most solid — always account for both comparative, across-species, patterns, as well as experimental results. This is true in all of biology, as it is for the study of xylem structure-function relations, where there is still a need for xylem physiology and functional comparative wood anatomy to mutually complement each other. To illustrate the magnitude and urgency of this need, we discuss Sherwin Carlquist’s postulate of a link between vulnerability to drought-induced embolism and conduit diameter, summarizing some of the major global patterns of xylem trait variation that are currently explained by postulating a vulnerability–diameter link. These include wider community mean and maximum vessel diameters in wetter vs drier vegetation types; that vessels can be >700 μ m in diameter but plants virtually always produce much narrower ones; that dryland plants with wider vessels drop their leaves earlier; wide-to-narrow vessels across growth rings; and the wide vessels of lianas surrounded by narrow vessels. These patterns are global, and we are aware of no anatomical evidence contradicting a vulnerability–diameter link. Despite the pervasiveness of these patterns, many xylem biologists do not regard the patterns as providing guidance for research in functional xylem biology. Instead, proposing and testing hypotheses to account for all of the data — xylem physiology experiments and comparative anatomical patterns in all their complexity and with all of their contradictions — provides the best way forward for the field. This effort requires proposing and testing hypotheses that are consistent with both experimental as well as comparative data. Crucially, it also requires not rejecting the vulnerability–diameter link without providing an alternative explanation that better explains the patterns currently explained by appeal to the link.
... Only uniseriate and biseriate rays as seen in M. mirabilis have also been found in Creochiton and Pseudodissochaeta (Van Vliet 1981). In general, lianas tend to have large vessel elements for efficient water and nutrient transport (Carlquist 1985, Ewers and Fisher 1991, Angyalossy et al. 2012. The widest vessel element observed in M. mirabilis was only 76.9 µm, which is narrower than in some genera of Dissochaeteae [e.g. ...
The increasing availability of DNA sequence data, in particular target enrichment data based on the universal Angiosperms353 probe set, but also accumulated Sanger data from previous phylogenetic studies, is facilitating the placement of taxa that are difficult to place with certainty based on morphological evidence alone. Here, we investigate phylogenetic relationships of Medinilla mirabilis (Melastomataceae), a species distributed in central Africa and currently classified in the mega-diverse genus Medinilla of tribe Sonerileae. Medinilla mirabilis is a twining liana with verticillate leaves when young, spherical inflorescences, 4-merous flowers, dimorphic stamens, and baccate fruits. Our results revealed that M. mirabilis is sister to tribe Dissochaeteae and only distantly related to Medinilla. We also provide new data on wood anatomical and seed morphological characters of M. mirabilis. The alternate inter-vessel pits in M. mirabilis and Dissochaeteae are consistent with the phylogenetic placement. Seeds of M. mirabilis are similar to those of Dissochaeteae and of Medinilla. Due to its unique morphology and phylogenetic position, we propose to reinstate the monospecific genus Myrianthemum with Myrianthemum mirabile. This necessitates expansion of the Southeast Asian tribe Dissochaeteae to include Myrianthemum as its only African member. Our study of M. mirabile demonstrates that the combined application of Angiosperms353 and Sanger data is a cost-effective approach to phylogenetically place enigmatic taxa.
... Whatever the case, these variants generate complex morphologies given the unusual arrangement of vascular tissues, which have been interpreted as ecological adaptations to many different conditions, such as underground reservoir organs, cushion plants, halophytes and especially the climbing habit, as they generate increased hydraulic conduction, flexibility and mechanical resistance (Carlquist, 1991(Carlquist, , 2013Fisher and Ewers, 1991;Rowe et al., 2004). With multiple independent evolutions across gymnosperms and especially angiosperms (Angyalossy et al., 2012(Angyalossy et al., , 2015, vascular variants are a compelling system to explore how development is modified to shape plant form during evolutionary time. ...
Background and aims:
The tribe Paullinieae have the highest diversity of vascular variants among the seed plants. The developmental diversity is better understood in the species rich genera Paullinia and Serjania, however, the phylogeny and diversity of vascular variants in the smaller genera of Paullinieae remain understudied. Here we investigate the evolution of development of stem vasculatures in the small genus Urvillea.
Methods:
We generate the first molecular phylogeny of Urvillea derived from 11 markers using a maximum likelihood and Bayesian approach. In combination with phylogenetic reconstruction, stochastic character mapping is used to assess evolutionary changes in stem ontogenies, determined from developmental anatomy of stems collected in the field or from herbarium and wood collections.
Key results:
Urvillea is supported as a monophyletic group and sister to Serjania. There are five stem ontogenies in Urvillea, including a typical growth and four vascular variants. Most stem ontogenies initiate with lobed stems. Lobed adult stems are conserved in Urvillea, but this ontogeny was lost multiple times. A reversal to typical growth occurred in non-climbing species. Phloem wedges, fissured stems and ectopic cambia evolved independently once. Phloem wedges is an intermediate developmental stage in the formation of fissured stems, which is characterized by a continuous fragmentation of vascular tissues. Lobed stems may generate constriction zones and lobes may split or not.
Conclusions:
Urvillea stands out as the third most diverse genus in number of vascular variants within Paullinieae, but only one ontogeny (fissured stems) is exclusive for the genus. Differential cambial activity and ectopic cambia are the main ontogenetic processes generating stem diversity. The evolutionary history of vascular variants demonstrates the large developmental plasticity of the cambium in such a small genus and corroborates a scenario of repeated evolution of complex anatomies within Paullinieae lianas.
... Para caracterizar la corteza se utilizaron la clasificación de cortezas de Bohren et al. (2003), el atlas de Agueda Castro (2009), el manual de León Gómez et al. (2001 y los glosarios de Junikka (1994) y IAWA (Angyalossy et al., 2016). En la descripción de la organización vascular, se emplearon los criterios de clasificación de variantes cambiales de Metcalfe (1989), Carlquist (2001), Larson (1994) y Angyalossy et al. (2012). Para la caracterización anatómica de leño se siguió al glosario de terminología de maderas de IAWA (1989). ...
Debido a su forma de vida, las lianas presentan una gran diversidad de adaptaciones en sus tallos, y, como consecuencia, son fácilmente distinguibles a partir de su morfología caulinar. Generalmente, las hojas y órganos reproductivos de las lianas se localizan por encima de las copas de los árboles, por lo que muchas veces resulta difícil reconocer estas plantas dentro de la selva y por ello los caracteres de los tallos son muy útiles para su identificación taxonómica. Este trabajo tiene como objetivo caracterizar la morfología y anatomía caulinar de las lianas del Parque Provincial Moconá (Misiones, Argentina) para su reconocimiento macroscópico. Se relevaron los senderos de fácil acceso e importancia turística del parque: Sendero de la Gruta, Sendero de los Chachíes, Humedal y Camino al embarcadero "Piedra Bugre". Se determinaron y caracterizaron macroscópicamente 16 especies de lianas, pertenecientes a 8 familias de angiospermas: Apocynaceae (3 spp.), Aristolochiaceae (1 sp.), Bignoniaceae (4 spp.), Cactaceae (1 sp.), Fabaceae (2 spp.), Nyctaginaceae (1 sp.), Sapindaceae (3 spp.), Vitaceae (1 sp.). En las lianas estudiadas se reconocieron diferentes tipos de características adaptativas al hábito trepador, las que pueden agruparse de la siguiente manera: forma de crecimiento del tallo, morfología externa del tallo, superficie del tallo, sección del tallo, estructuras de anclaje, variantes cambiales y características anatómicas del leño. Se elaboraron dos claves dicotómicas para su reconocimiento, una utilizando sólo caracteres externos de los tallos y otra con caracteres externos y anatómicos.
Species with lianescent habit account for half of the diversity of Bignoniaceae. Recent molecular phylogenetic studies have provided the basis for new circumscriptions of entire liana lineages within tribes Bignonieae and Tecomeae s.s., where only monophyletic taxa are recognized. However, some clades remain without good morphological synapomorphies. In search of features of taxonomic potential, we collected, sectioned, and analyzed the bark of 83 lianescent species of the Bignoniaceae, covering all 20 genera from tribe Bignonieae currently recognized, plus three of the most widely cultivated lianas of Tecomeae s.s. Detailed bark descriptions are given to major lineages within both tribes, following their most recent phylogenetic hypotheses and classifications. Our anatomical studies allowed us to identify 19 potential synapomorphies for large clades or specific genera of lianas, such as the fibrous phloem found in members of the Fridericia Mart. emend L.G.Lohmann and allies clade, the exclusive presence of sclereids in the regular phloem of Pleonotoma Miers, and the presence of radially elongated fibers in Manaosella L.C.Gomes, among others. Using a combination of features, we were able to produce the first bark key to identify genera of lianescent Bignoniaceae. Our work reinforces the importance of bark features for a deeper understanding of taxonomic and phylogenetic relationships among taxa. © Publications scientifiques du Muséum national d’Histoire naturelle, Paris.
Climbers were among the least studied plant growth forms until recently for a variety of reasons. However, research on climbers, particularly the ecology of woody climbers, has flourished over the past four decades. This exponential rise is linked to a growing recognition of climbers’ importance to the dynamics, structure, and function of tropical forests. Since then several studies have been conducted on various research themes covering the broad spectrum of climber biology. In this chapter, I examine the trends in climber research from the past to the present and provide a thorough analysis of climber studies that have advanced our understanding of their biology.
RESUMO O corpus lignosum compositum, típico para as lianas da família Sapindaceae, é designado neste trabalho como "cilindro vascular composto". No caule de Serjania caracasana (Jacq.) Willd. essa variação cambial está representada por um cilindro vascular central circundado por oito cilindros vasculares periféricos. Não existe consenso quanto à terminologia que envolve essa estrutura, o que torna difícil uma abordagem anatômica desagregada de uma adequação terminológica. Nesse estudo, por meio da análise anatômica do caule, verificou-se que mesmo antes da vascularização há indícios do aspecto composto, com a formação de oito lobos que circundam a região central. Com o início da vascularização, cada lobo e a região central são denominados "cilindro vascular". O termo aqui adotado "cilindro vascular composto" é adequado, pois reflete a homologia entre os cilindros vasculares em S. caracasana através da origem procambial. Esse termo exibe um caráter descritivo que facilita a compreensão do conceito e mantém a relação de equivalência lingüística com o termo original - corpus lignosum compositum. Rejeita-se o termo "caule poliestélico" ou "caule multiestelar", pois os resultados aqui apresentados indicam a presença de um único estelo no caule.
The climbing habit in plants has apparently evolved numerous times. Species that climb are well represented in habitats ranging from tropical rain forests through temperate forests to semi-deserts. The Biology of Vines, first published in 1992, is a treatment of what is known about climbing plants, written by a group of experts and covering topics ranging from the biomechanics of twining to silvicultural methods for controlling vine infestations. Also included are detailed accounts of climbing plant evolution, stem anatomy and function, climbing mechanics, carbon and water relations, reproductive ecology, the role of vines in forest communities and their economic importance. The chapters are based on research on herbaceous vines and woody climbers (lianas) in both temperate and tropical zones, deserts and rain-forests and Old and New World areas. Much remains to be learned about the biology of these plants, but this volume provides a substantial foundation upon which further research can be based.
The accessory tissues of the hydrosystem consist of the parenchymatic contact tissue of the xylem rays, as weIl as the paratracheidal parenchyma in gymnosperms, and the paratracheal contact-parenchyma in angiosperms.