of flow,rs and fruits

Transcription

1

2 Roland Keller Identifi~cation of tropical woodyi plants in the absence of flow,rs and fruits I I A field guide Springer Basel AG

3 Author: Dr. Roland Keller Institut de Botanique Systematique et de Geobotanique Batiment de Biologie CH-1015 lausanne A CIP catalogue record for this book is available from the library of Congress, Washington, D.C., USA Deutsche Bibliothek Cataloging-in-Publication Data Keller, Roland: Identification of tropical woody plants in the absence of flowers and fruits: a field guide / Roland Keller. ISBN ISBN (ebook) DOI / The publisher and author cannot assume any legal responsibility for information on drug dosage and administration contained in this publication. The respective user must check its accuracy by consulting other sources of reference in each individual case. The use of registered names, trademarks, etc. in this publication, even if not identified as such, does not imply that they are exempt from the relevant protective laws and regulations or free for general use. This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. For any kind of use, the permission of the copyright holder must be obtained Springer Basel AG Originally published by Birkhauser Verlag, P.O. Box 133, CH-4010 Basel, Switzerland 1996 Printed on acid-free paper produced from chlorine-free pulp. TCF ~ Cover design: Markus Etterich, Basel ISBN

4 Acknowledgements I wish to thank the following for their assistance and their friendship, Daniel Atuany (guide PHPA at the Manusela, National Park, Moluccas Islands), Claude Edelin (CNRS research scientist, at the University Montpellier II), Peter Endress (professor at the University of Zurich), Jacques Fournet (INRA research scientist Guadeloupe), Francis Halle (professor at the University Montpellier II), Rene Hebding (botanist at the Villa Les Cedres, Saint-Jean-Cap-Ferrat), Pak Ijun (botanist, the BIOTROP Institute, Bogor, Java), Anton Leeuwenberg (taxonomist at the Agricultural University ofwageningen, The Netherlands), Pierre Lombion (botanist in Guadeloupe), Jean-Franc;ois and Caroline Molino (scientists and hosts at Bukittinggi, Sumatra), Jeanine Raharilala (taxonomist at the Parc Botanique et Zoologique de Tsimbazaza, Madagascar), Mamisoa Rapanoelina (Montagne d' Ambre Project manager at Diego Suarez, Madagascar), Elio Sanoja (professor at the Universidad Nacional Experimental de Guayana, Venezuela), without whom this identification key would simply have been mere speculation. The author is indebted to Claire Muller, to the Societe Academique Vaudoise and to the Fondation du 450e me Anniversaire de l'universite de Lausanne who generously helped finance the edition. The drawing up of this document was carried out under the auspices of the Institut de Botanique Systematique et de Geobotanique at Lausanne University. Adrian Bell contributed to the translation of the French version into English and made numerous technical and scientific suggestions. Angelika Collis and Colin Preston also helped to correct and translate.

5 A Zoe et Lucien

6 CONTENTS Foreword... XI Introductory remarks Part I: Keys and glossary General Key... 9 KeysA-Z Glossary, notes and illustrations Geographical distribution Outer bark and lenticels Macroanatomy I: Inner bark, rays and exudates Macroanatomy II: Internal phloem, wood and pith Climbing systems Unit of extension, monopodium and sympodium Ramification: Rhythm and position Architectural models Phyllotaxy and torsion Heterophylly Shape of stems Axillary buds and prophylls Stipules and interpetiolar ridge Leaf: Blade and rachis Petioles and petiolules Leaf folding and aestivation Venation I Venation II Glands and translucent dots Teeth and indumenta Part II: The principal families of tropical woody Dicotyledons illustrated by means of their vegetative characters Families identified by the key and the orders or classes to which they belong Piperaceae, Chloranthaceae, Meliosmaceae, Araliaceae, and 2. Menispermaceae, Aristolochiaceae, Hernandiaceae, Illiciaceae, Canellaceae Lauraceae, Monimiaceae, and 4. Annonaceae, Magnoliaceae, Myristicaceae

7 5. Hamamelidaceae, Fagaceae, Polygonaceae, and 6. Ulmaceae, Urticaceae, Moraceae Dilleniaceae, Quiinaceae, Ochnaceae, and 8. Clusiaceae, Theaceae, Actinidiaceae, Marcgraviaceae, Pellicieraceae, Bonnetiaceae Lecythidaceae, Myrsinaceae, Theophrastaceae, and 10. Ebenaceae, Sapotaceae, Thymeleaceae, Proteaceae Aquifoliaceae, Olacaceae, Icacinaceae, and 12. Combretaceae, Myrtaceae, Melastomataceae Capparidaceae, Violaceae, Flacourtiaceae, PassifIoraceae, and 14. Dipterocarpaceae, Elaeocarpaceae, Tiliaceae, Sterculiaceae, Bombacaceae Malvaceae Euphorbiaceae Leguminosae Chrysobalanaceae, Rosaceae, Zygophyllaceae, Oxalidaceae, Connaraceae, and 18. Sapindaceae, Meliaceae, Rutaceae, Anacardiaceae, Burseraceae Rhamnaceae, Vitaceae, Leeaceae, and 20. Cunoniaceae, Rhizophoraceae, Cornaceae, Alangiaceae, Anisophylleaceae Dichapetalaceae, Malpighiaceae, Polygalaceae, Xanthophyllaceae, Vochysiaceae, and 22. Celastraceae, Irvingiaceae, Humiriaceae, Linaceae, Ixonanthaceae, Erythroxylaceae Convolvulaceae, Boraginaceae, Solanaceae, Bignoniaceae, Oleaceae, Verbenaceae, and 24. Loganiaceae, Apocynaceae Rubiaceae Bibliography Indexes Index of the families mentioned in the glossary Index of the genera x

8 Foreword While studies of forest vegetation may differ in their underlying objective, be it physiology, ecology or biodiversity, common to all these is the fact that all require taxonomic knowledge. The process of taxonomy or of forest ecology begins in principle with an inventory of the flora, the evaluation of this inventory still being based to a large extent of reproduction-related organs. In a tropical forest, the majority of flowers or fruits are most often found in the canopy. The canopy, however, is difficult to reach and to do so necessitates heavy, expensive or sophisticated equipment such as a tower, a crane, or a hanging platform suspended from a dirigible balloon. Thus, most of the time, botany is practised near the ground with the aid of light equipment such as ladders, branchloppers, and climbing irons. Furthermore, a large proportion of the trees and lianas of the understorey bear neither flowers nor fruits at certain times of the year. Despite this absence of seasonal characters it should at least be possible to recognize the families of plants by means of easily observable and permanent characters. To meet this need, an identification system has been designed in the form of a dichotomous key. In addition to permitting recognition ofplantfamities in the field in all seasons, this system can serve as a starting-point for a more detailed knowledge of the forest taxa. Many plant families with a pantropical distribution are recognized by the botanist, forestry engineer, or knowledgeable native, thanks to certain non-reproductive morphological characters. Taxonomic features of this kind, such as the fibrous bark of Annonaceae, the disposition in tiers of the branches of Diospyros, the translucent dots of the Rutaceae, etc., are often mentioned in descriptions of families or species. However, they seldom if ever appear in the identification keys of regional flora. Vegetative organs can present morphological or macroanatomical features that are perfectly appropriate for identification in the field. It is therefore worth verifying on a pantropical scale the taxonomic value of traits that are little known or,jorgotten". This may also lead to the discovery of new taxonomic characters. The diversity of herbaceous species in a tropical forest is less than that of woody species (although it can attain that of ligneous plants in a cloud forest housing numerous epiphytes). And even though the physiognomy of tropical forests is marked by the abundance of large woody plants, the efforts of taxonomists have been mainly directed towards plants that are relatively small in size. This is clearly shown by the importance given to these groups in the floras now being published; the volumes on Bromeliaceae, Melastomataceae, Ericaceae, have already appeared, while those on Anacardiaceae and Leguminosae are in preparation. On the other hand, certain floras treat trees as their priority but often exclude woody lianas. A key for tropical forest identification must apply primarily to trees, shrubs and woody lianas. Woody plants possess long-lived inner bark and dead outer bark. These tissues exhibit easily recognizable traits which are almost always absent in herbs. The architecture of woody plants is in general more

9 complex than that of herbs, with stems differentiated into trunks, branches and twigs. Through their structure, bark, rhytidome and the morphological characters of branches and leaves, woody plants provide a great deal of taxonomic information. This underlines the potential utility of a vegetative key for ligneous plants. Plant communities living in temperate or cold climates consist mostly of herbs or small woody plants. The long-lived vegetative organs of these plants are often reduced or considerably modified; tubers, short-shoots, c1adodes, phyllodes, fleshy scales, areoles, etc. allow the plant to stand up to harsh climatic conditions. The morphological classification of these organs is not always easy, whereas the short-lived organs of the same plants are readily identifiable (the leaves of caducifolious, short-shooted trees, the flowers of aphyllous, phyllode or c1adode bearing shrubs and the aerial leaves of tubers or bulbs). The long-lived vegetative organs of plants adapted to contrasted climates are often difficult to identify or describe, whereas their flowers pose relatively few problems of interpretation. These difficulties have probably inhibited attempts at vegetative identification that could have been (or should have been) undertaken by European botanists in previous centuries. If we consider tropical trees, shrubs and lianas on the other end, it is evident that numerous attributes of their leaves, branches and trunks are easy to observe owing to their relative size and distinctive traits. From the vegetative point of view, the trees and vines of tropical forests permit a new view of the plant world. This presents an opportunity to create,for woody plants, a new key: vegetative and pantropical. XII