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      Convergent Evolution and the Diverse Ontogenetic Origins of Tendrils in Angiosperms

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          Abstract

          Climbers are abundant in tropical forests, where they constitute a major functional plant type. The acquisition of the climbing habit in angiosperms constitutes a key innovation. Successful speciation in climbers is correlated with the development of specialized climbing strategies such as tendrils, i.e., filiform organs with the ability to twine around other structures through helical growth. Tendrils are derived from a variety of morphological structures, e.g., stems, leaves, and inflorescences, and are found in various plant families. In fact, tendrils are distributed throughout the angiosperm phylogeny, from magnoliids to asterids II, making these structures a great model to study convergent evolution. In this study, we performed a thorough survey of tendrils within angiosperms, focusing on their origin and development. We identified 17 tendril types and analyzed their distribution through the angiosperm phylogeny. Some interesting patterns emerged. For instance, tendrils derived from reproductive structures are exclusively found in the Core Eudicots, except from one monocot species. Fabales and Asterales are the orders with the highest numbers of tendrilling strategies. Tendrils derived from modified leaflets are particularly common among asterids, occurring in Polemoniaceae, Bignoniaceae, and Asteraceae. Although angiosperms have a large number of tendrilled representatives, little is known about their origin and development. This work points out research gaps that should help guide future research on the biology of tendrilled species. Additional research on climbers is particularly important given their increasing abundance resulting from environmental disturbance in the tropics.

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          Absolute diversification rates in angiosperm clades.

          The extraordinary contemporary species richness and ecological predominance of flowering plants (angiosperms) are even more remarkable when considering the relatively recent onset of their evolutionary diversification. We examine the evolutionary diversification of angiosperms and the observed differential distribution of species in angiosperm clades by estimating the rate of diversification for angiosperms as a whole and for a large set of angiosperm clades. We also identify angiosperm clades with a standing diversity that is either much higher or lower than expected, given the estimated background diversification rate. Recognition of angiosperm clades, the phylogenetic relationships among them, and their taxonomic composition are based on an empirical compilation of primary phylogenetic studies. By making an integrative and critical use of the paleobotanical record, we obtain reasonably secure approximations for the age of a large set of angiosperm clades. Diversification was modeled as a stochastic, time-homogeneous birth-and-death process that depends on the diversification rate (r) and the relative extinction rate (epsilon). A statistical analysis of the birth and death process was then used to obtain 95% confidence intervals for the expected number of species through time in a clade that diversifies at a rate equal to that of angiosperms as a whole. Confidence intervals were obtained for stem group and for crown group ages in the absence of extinction (e = 0.0) and under a high relative extinction rate (epsilon = 0.9). The standing diversity of angiosperm clades was then compared to expected species diversity according to the background rate of diversification, and, depending on their placement with respect to the calculated confidence intervals, exceedingly species-rich or exceedingly species-poor clades were identified. The rate of diversification for angiosperms as a whole ranges from 0.077 (epsilon = 0.9) to 0.089 (epsilon = 0.0) net speciation events per million years. Ten clades fall above the confidence intervals of expected species diversity, and 13 clades were found to be unexpectedly species poor. The phylogenetic distribution of clades with an exceedingly high number of species suggests that traits that confer high rates of diversification evolved independently in different instances and do not characterize the angiosperms as a whole.
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            Tempo and mode in evolution: phylogenetic inertia, adaptation and comparative methods

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              A mechanistic explanation for global patterns of liana abundance and distribution.

              One of the main goals in ecology is determining the mechanisms that control the abundance and distribution of organisms. Using data from 69 tropical forests worldwide, I demonstrate that liana (woody vine) abundance is correlated negatively with mean annual precipitation and positively with seasonality, a pattern precisely the opposite of most other plant types. I propose a general mechanistic hypothesis integrating both ecological and ecophysiological approaches to explain this pattern. Specifically, the deep root and efficient vascular systems of lianas enable them to suffer less water stress during seasonal droughts while many competitors are dormant, giving lianas a competitive advantage during the dry season. Testing this hypothesis in central Panama, I found that lianas grew approximately seven times more in height than did trees during the dry season but only twice as much during the wet season. Over time, this dry season advantage may allow lianas to increase in abundance in seasonal forests. In aseasonal wet forests, however, lianas gain no such advantage because competing plants are rarely limited by water. I extend this theory to account for the local, within-forest increase in liana abundance in response to disturbance as well as the conspicuous decrease in liana abundance at high latitudes.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                03 April 2018
                2018
                : 9
                : 403
                Affiliations
                [1] 1Laboratório de Sistemática, Evolução e Biogeografia de Plantas Vasculares, Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo , São Paulo, Brazil
                [2] 2Department of Plant Biology, University of California, Davis , Davis, CA, United States
                [3] 3Genomics and Transposable Elements Laboratory (GaTE-Lab), Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo , São Paulo, Brazil
                Author notes

                Edited by: Elena M. Kramer, Harvard University, United States

                Reviewed by: Barbara Ambrose, New York Botanical Garden, United States; Stefanie M. Ickert-Bond, University of Alaska Fairbanks, United States

                *Correspondence: Mariane S. Sousa-Baena m.sousabaena@ 123456yahoo.com

                This article was submitted to Plant Evolution and Development, a section of the journal Frontiers in Plant Science

                †Present Address: Mariane S. Sousa-Baena, Genomics and Transposable Elements Laboratory (GaTE-Lab), Departamento de Botânica, Instituto de Biociências, Universidade de São Paulo, São Paulo, Brazil

                Article
                10.3389/fpls.2018.00403
                5891604
                29666627
                1a5b3774-5c01-41f4-bca2-83ae08949b4b
                Copyright © 2018 Sousa-Baena, Sinha, Hernandes-Lopes and Lohmann.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 17 November 2017
                : 13 March 2018
                Page count
                Figures: 5, Tables: 1, Equations: 0, References: 139, Pages: 19, Words: 13909
                Categories
                Plant Science
                Review

                Plant science & Botany
                climbing habit,helical growth,lianas,ontogenetic origin,primary homology,recurrent evolution,tendrils,vines

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