A Monograph of Codonopsis and Allied Genera (Campanulaceae) This page intentionally left blank A Monograph of Codonopsis and Allied Genera (Campanulaceae) De-Yuan HONG State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing In association with Qiang WANG and Kai-Yu PAN AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD PARIS • SAN DIEGO • SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Academic Press is an imprint of Elsevier 125 London Wall, London EC2Y 5AS, UK 525 B Street, Suite 1800, San Diego, CA 92101-4495, USA 225 Wyman Street, Waltham, MA 02451, USA The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK Copyright © 2015 China Science Publishing & Media Ltd. Published by Elsevier Inc. All rights reserved. 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ISBN: 978-0-12-801933-7 British Library Cataloguing-in-Publication Data A catalogue record for this book is available from the British Library Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress For information on all Academic Press publications visit our website at http://store.elsevier.com/ Publisher: Janice Audet Acquisition Editor: Simon Tian Editorial Project Manager: Simon Tian Production Project Manager: Julia Haynes Cover Designer: Victoria Pearson Typeset by TNQ Books and Journals www.tnq.co.in Printed and bound in United States of America Dedication Dedicated to my parents: Brave and diligent father, Guan-Hua HONG (1894–1959); Intelligent and diligent mother, Shi-Ru HU (1901–1971) This page intentionally left blank Contents Codonopsis ovata 74 Synopsis ix Codonopsis clematidea 78 Preface xi Codonopsis obtusa 83 Acknowledgements xiii Codonopsis subsimplex 85 Codonopsis campanulata 87 1. I ntroduction: The Taxonomic Codonopsis alpina 87 History and Questions to be Codonopsis chlorocodon 92 Addressed Codonopsis bhutanica 92 Codonopsis argentea 94 Discoveries in Codonopsis and Allied Genera 1 Codonopsis canescens 96 Circumscription of Codonopsis 4 Codonopsis foetens 99 Systematic Position of Codonopsis and Allied Genera 5 subsp. foetens 99 Taxonomical Revisions of Codonopsis and subsp. nervosa 102 Allied Genera 6 Codonopsis bulleyana 104 Codonopsis thalictrifolia 107 2. B iology subsp. thalictrifolia 108 Gross Morphology 9 subsp. mollis 110 Seed Morphology 19 Codonopsis viridiflora 112 Pollen Morphology 30 Codonopsis tsinlingensis 115 Chromosomes 33 SECTION 2. CODONOPSIS 117 Codonopsis pilosula 117 3. S ystematics and Evolution subsp. pilosula 118 subsp. tangshen 123 Circumscription of Codonopsis and its subsp. handeliana 125 Relationships with Allied Genera 43 Codonopsis hemisphaerica 125 A Three-Tribal Classification System of the Codonopsis kawakamii 126 Campanulaceae S. S. and the Systematic Codonopsis bomiensis 126 Position of Codonopsis and Allied Genera 47 Codonopsis deltoidea 131 Evolution and Biogeography of the Cyanantheae 51 Codonopsis rotundifolia 134 Codonopsis reflexa 136 4. T axonomic Revision of Codonopsis Codonopsis henryi 136 and Allied Genera Codonopsis affinis 138 Codonopsis Wall. 61 Codonopsis viridis 140 SECTION 1. ERECTAE 65 Codonopsis farreri 143 Codonopsis lixianica 65 Codonopsis benthamii 143 Codonopsis elliptica 66 Codonopsis tubulosa 149 Codonopsis chimiliensis 67 subsp. tubulosa 150 Codonopsis meleagris 68 subsp. vadsea 153 Codonopsis gongshanica 69 Codonopsis microtubulosa 153 Codonopsis subscaposa 70 Codonopsis subglobosa 154 Codonopsis cardiophylla 73 Codonopsis bragaensis 156 subsp. cardiophylla 74 Codonopsis cordifolioidea 157 subsp. megaphylla 74 Codonopsis micrantha 158 vii viii Contents SECTION 3. LANCEOLATAE 159 Pseudocodon grey-wilsonii 200 Codonopsis lanceolata 159 Pseudocodon hirsutus 203 Codonopsis ussuriensis 165 Pseudocodon graminifolius 204 SECTION 4. LEPTOCODON 167 Pseudocodon retroserratus 208 Codonopsis gracilis 168 Pseudocodon petiolatus 209 Codonopsis hongii 170 Pseudocodon rosulatus 209 SECTION 5. CAMPANUMOEA 172 Cyclocodon Griff. ex Hook. f. & Thomson 210 Codonopsis javanica 172 Cyclocodon parviflorus 212 subsp. javanica 172 Cyclocodon lancifolius 214 subsp. japonica 177 Cyclocodon axillaris 218 Codonopsis inflata 180 Echinocodon D. Y. Hong 223 Himalacodon D. Y. Hong & Q. Wang 181 Pankycodon D. Y. Hong & X. T. Ma 184 Pseudocodon D. Y. Hong & H. Sun 190 Bibliography 227 Pseudocodon convolvulaceus 191 Index to Specimens Cited 233 subsp. convolvulaceus 192 Index to Botanical Names and Synonyms in subsp. forrestii 194 Codonopsis and Allied Genera 249 Pseudocodon vinciflorus 197 Index 253 subsp. vinciflorus 198 subsp. dianchuanicus 200 Synopsis The author has worked on the Campanulaceae s. s. since Codonopsis and its relationships with allied genera, and 1970’s. Recently after he had partially finished his work on created a new three-tribal classification system of the Paeonia worldwide he has been concentrating his energy on Campanulaceae s. s. The monograph assembles the results studies of Codonopsis and its allies, which is a group very achieved in the author’s work, and discusses evolution of controversial among authors in taxonomy and systematics. He the group, particularly the evolutionary trends of characters and his assistants made expeditions to the Himalaya (S and such as ovary position, fruit morphology, and basic number of SE Tibet and N Pakistan), to the Hengduan Mountains chromosomes. The major part of the monograph is the taxo- (NW Yunnan and SW and W Sichuan), and to E Qinghai, nomic revision of Codonopsis and allied genera: Codonopsis where most members of this group are distributed. During (46 spp.), Cyclocodon (3 spp.), Echinocodon (1 sp.), the field work they conducted population observations, Himalacodon (1 sp.), Pankycodon (1 sp.), and Pseudo- population sampling, and collection of materials for codon (8 spp.). It clarifies the relationships between genera chromosome counting and DNA analyses in addition to and between species, provides an useful key to genera and collection of vouchers and herbarium specimens. The to species for each genus with more than one species, and author examined and identified all specimens available in gives a concise but diagnostic description for each species. 27 herbaria. Based on the material collected in the field and Every species is provided with a fine line illustration, and studied in herbaria they observed gross morphology, seed most species with photos taken in the fields. Specimen morphology, pollen grains, and chromosomes, and made citation and distribution map are presented for each species. intensive DNA sequence analyses. As a result, they have Forty-three collections are designated as types (lectotypes, established three new genera, described seven new species isolectotypes, or neotypes). There is a list of all the herbarium and two new subspecies, clarified the circumscription of specimens examined and their species designations. ix