Genus Vigna Subgenus Ceratotropis Genetic Resources the Asian Vigna: Genus Vigna Subgenus Ceratotropis Genetic Resources

The Asian Vigna: Genus Vigna subgenus Ceratotropis genetic resources The Asian Vigna: Genus Vigna subgenus Ceratotropis genetic resources

Norihiko Tomooka Narional lnstitute of Agrobiofogical Sciences, Tsukuba. lapan

Duncan A. Vaughan NatÎOllallnstitule of Agrobiological Sciences, Tsukuba. Japan.

Helen Moss West Vancouver, Brirish Columbia, Canada

and

Nigel Maxted University of Birmingham. Birmingham, United Kingdolll

....., SPRINGER SCIENCE+BUSINESS MEDIA, B.V. A C.I.P. Catalogue record for this book is available from the Library of Congress.

ISBN 978-94-010-3934-5 ISBN 978-94-010-0314-8 (eBook) DOI 10.1007/978-94-010-0314-8

Printed on acid-free paper

No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording Of otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. National Institute of Agrobiological Sciences (NIAS) The contribution of the National Institute of Agrobiological Resources (NIAS), Japan, towards the costs of publishing this book is acknowledged. NIAS is an independent administrative institution supervised and financially supported by the Ministry of Agriculture, Forestry and Fisheries (MAFF), Japan. NIAS was established in 2001 to conduct basic Life Science research on insects, animals and plants strategically and intensively in order to produce pioneering results. NIAS aims at accelerating genome research and its use for plants (especially rice) and animals (insects and domestic animals) as a leading research institute. Among the several research thrusts of NIAS is to conduct fundamental and technological research on development and use of biological resources in agriculture.

I PGRI

International Plant Genetic Resources Institute (IPGRI) The contribution of the International Plant Genetic Resources Institute (IPGRI) through the subregional network EA-PGR (EXPAND) to some of the studies that contributed to this publication is acknowledged. IPGRI is an autonomous international scientific organization, supported by the Consultative Group on International Agricultural Research (CGIAR). IPGRI's mandate is to advance the conservation and use of genetic diversity for the well-being of present and future generations. IPGRI's headquarters is based in Rome, Italy, with offices in another 19 countries worldwide. It operates through three programmes: (1) the Plant Genetic Resources Programme, (2) the CGIAR Genetic Resources Support Programme, and (3) the International Network for the Improvement of Banana and Plantain (INIBAP). Vll

CONTENTS

1. INTRODUCTION ...... 1 1. VIGNA SUBGENUS CERATOTROPIS IN CONTEXT ...... 1 2. SCOPE AND AIMS OF THE BOOK ...... 6 3. TAXA ABBREVIATIONS ...... 7

2. BIOSYSTEMATIC BACKGROUND ...... 9 1. BASIC STRUCTURE OF SUBGENUS CERATOTROPIS PLANTS ...... 9 2. TAXONOMIC HISTORY OF VIGNA SUBGENUS CERATOTROPIS ...... 9 2.1. Family Leguminosae (Fabaceae) ...... 9 2.2. Genus Vigna Savi ...... 13 2.3. Subgenus Ceratotropis (Piper) Verdcourt: the Asian Vigna ...... 17 2.3.1. Key morphological characteristics of the Asian Vigna ...... 17 2.3.2. Species relationships in the subgenus Ceratotropis ...... 21 3. TAXONOMIC KEY TO TAXA IN SUBGENUS CERATOTROPIS ...... 26

3. GENETIC RESOURCES ...... 29 1. CONSERVED GERMPLASM ...... 29 1.1. Ex situ conservation ...... 29 1.2. In situ conservation ...... 29 1.2.1. Asian Vigna in protected areas ...... 33 1.2.2. Species "hot spots" for in situ conservation ...... 36 1.2.3. Population types for in situ conservation ...... 38 2. CONSERVATION PRACTICES ...... 41 2.1. Collection ...... 41 2.2. Characterization ...... 44 2.2.1. Germination and cultural practices ...... 44 2.2.2. DNA extraction from Vigna for molecular characterization ...... 47 2.3. Evaluation ...... 49 2.3.1. Evaluation for resistance to microorganisms ...... 49 2.3.2. Evaluation for resistance to insects ...... 52 2.4. Use ...... 54

4. SPECIES CONSPECTUS ...... 57 1. DATA COLLECTION ...... 57 2. CONSPECTUS ...... 59 2.1. Section Angulares N. Tomooka & Maxted ...... 59 2.1.1. Vigna angularis (Willd.) Ohwi & Ohashi ...... 59 Vlll 2.1.2. Vigna dalzelliana (0. Kuntze)Verdcourt ...... 65 2.1.3. Vigna exilis Tateishi & Maxted ...... 68 2.1.4. Vigna hirtella Ridley ...... 72 2.1.5. Vigna minima (Roxb.) Ohwi & Ohashi ...... 76 2.1.6. Vigna nakashimae (Ohwi) Ohwi & Ohashi ...... 80 2.1.7. Vigna nepalensis Tateishi & Maxted ...... 84 2.1.8. Vigna rejlexo-pilosa Hayata ...... 88 2.1.9. Vigna riukiuensis (Ohwi) Ohwi & Ohashi ...... 93 2.1.10. Vigna tenuicaulis N. Tomooka & Maxted ...... 97 2.1.11. Vigna trinervia (Heyne ex Wall.) Tateishi & Maxted ...... 101 2.1.12. Vigna umbellata (Thunb.) Ohwi & Ohashi ...... 108 2.2. Section Ceratotropis N. Tomooka & Maxted ...... 113 2.2.1. Vigna grandiflora (Prain) Tateishi & Maxted ...... 113 2.2.2. Vigna mungo (L.) Hepper ...... 117 2.2.3. Vigna radiata (L.) Wilczek ...... 121 2.2.4. Vigna subramaniana (Babu ex Raizada) M. Sharma ...... 127 2.3. Section Aconitifoliae N. Tomooka & Maxted ...... 131 2.3.1. Vigna aconitifolia (Jacq.) Marechal ...... 131 2.3.2. Vigna aridicola N. Tomooka & Maxted ...... 135 2.3.3. Vigna khandalensis (Santapau) Raghavan & Wadhwa ...... 139 2.3.4. Vigna stipu/acea Kuntze ...... 142 2.3.5. Vigna trilobata (L.) Verdcourt ...... 146

PLATES 1-12 ...... 150

5. ECO-GEOGRAPHIC ANALYSIS ...... 165 I. ECOLOGy ...... 166 1.1. Altitude ...... 166 1.2. Latitude ...... 166 1.3. Phenology ...... 168 1.4. Analysis of population distribution and climate: an example using V. trinervia ... 172 2. GENETIC EROSION ...... 173 3. PHYTOGEOGRAPHY AND CONSERVATION PRIORITIES ...... 174

6. RESEARCH AND FUTURE PERSPECTIVES ...... 179 I. RESEARCH ...... 179 1.1. Cross compatibility studies...... 179 1.2. Gene pools ...... 183 IX

1.3. Crop complexes: V. angularis as a model ...... 184 1.4. Analysis of representative germplasm sets ...... 195 1.4.1. Proteinase inhibitors ...... 195 1.4.2. Insect resistance ...... 199 1.5. Biotechnology ...... 199 1.5.1. Genome maps ...... 201 1.5.2. Comparative genome mapping ...... 201 1.5.3. Gene mapping ...... 203 2. FUTURE PERSPECTIVES ...... 204 2.1. Genetics and cytology ...... 204 2.2. Germplasm ...... 205 2.3. Conservation ...... 205 2.4. Characterization ...... 206 2.5. Evaluation ...... 206 2.6. Last comments ...... 207

APPENDIX I ...... 209 Vigna subgenus Ceratotropis taxa synonymized checklist ...... 209

APPENDIX II ...... 213 Specimen citation - herbarium specimens ...... 214

REFERENCES ...... 247

INDEX TO SCIENTIFIC NAMES ...... 259

INDEX ...... 265 xi

AUTHORS

Norihiko Tomooka, obtained his PhD from the University of Kyoto, Japan. The title of his thesis was "Genetic diversity and landrace differentiation of mungbean, Vigna radiata (L.) Wilczek, and evaluation of its wild relatives (the subgenus Ceratotropis) as breeding materials". Much of the research for his PhD was conducted during 5 years that he spent based at the Chai Nat Field Crops Research Center, Department of Agriculture, Thailand, as a visiting scientist from the Ministry of Agriculture, Forestry and Fisheries (MAFF), Japan. From 1992 to the present he has been responsible for the Vigna grain legumes collection in the MAFF, Japan, Genebank system based in the National Institute of Agrobiological Sciences (NIAS). He has participated in 17 germplasm collecting missions for grain legumes in Japan and 12 abroad. In 1998, he visited Europe as an OECD fellow and visited major herbaria, while based at Kew, U.K., to gather data on Vigna subgenus Ceratotropis germplasm.

Duncan Vaughan obtained his BSc in Agricultural Botany from the University of Reading, UK, and MSc and PhD from the University of Illinois, Champaign-Urbana, USA. His graduate research was on the wild relatives of soybean in the subgenus Glycine and genetics of soybean seed quality. He spent in total 8 years at the International Rice Research Institute, the Philippines during which he wrote a genetic resources handbook for the genus Oryza. In 1993, he joined the MAFF, Japan, where he has been working together with Dr. Tomooka in the Crop Evolutionary Dynamics Team ofNIAS. This team has focused much of its research effort on the wild relatives of crops including the Asian Vigna, particularly the Vigna angularis complex, molecular analysis and evaluation of gemplasm for various traits.

Helen Moss has an MSc degree in the Conservation and Use of Plant Genetic Resources from the University of Birmingham, UK. During the 1980's she worked as a consultant for the International Board for Plant Genetic Resources and undertook an extensive field collection and ecogeographic studies of both African and Asian Vigna.

Nigel Maxted is Senior Lecturer in Biodiversity Conservation at the University of Birmingham. He has three degrees in biology, taxonomy and conservation, his PhD research focused on the application of database technology to taxonomy and conservation. He has twenty years experience in research, consultancy and teaching biodiversity conservation. He has primarily focused on the taxonomy and ecogeography oflegume diversity, as well as the in situ and ex situ conservation of plant genetic resources. He regularly works as a consultant for the leading international conservation agencies. He has published over 100 research papers and in the last 3 years has written two books on in situ and ex situ conservation and one on legume genetic diversity in the Mediterranean. He is Chair of the ECP/GR Wild Species Conservation in Genetic Reserves Task Force. Xlll ACKNOWLEDGEMENTS

This book could not have been prepared without the help of many people. We would like to acknowledge the support and encouragement of the present and former management of the National Institute of Agrobiological Sciences (NIAS) particularly: M. Nakagahra, N. Katsura, S. Miyazaki, K. Okuno, and J. Kurisaki. Legume scientists that have had a major influence on our thinking and have cooperated with us in our studies, Y. Egawa, Japan International Research Center for Agricultural Sciences (JIRCAS), Y. Tateishi, University of the Ryukyus and K. Murata, Hokkaido Prefectural Agricultural Experiment Stations. We acknowledge the great contribution of present and former staff and visitors within the Crop Evolutionary Dynamics Team that have undertaken much of the research reported in this book and collected germplasm with us: A. Kaga, K. Doi, K. Kashiwaba, S. Tsukamoto, K. Motoyoshi, M. Inoue, R. Q. Xu, X. W. Wang, M. S.Yoon, O. K. Han, B. Chaitieng, A. Konarev, E. Potokina, M. T. Federici Rodriguez, 1. Kalubowila, L. E. Nsapato, S. S. Rao, Samsudin, P. Dominguez de Sanctis, Sayafrudin, D. Weerasekera, and X. X. Zong. Present or former staff of the National Institute of Agrobiological Sciences that have helped us in collecting germplasm in various parts of Japan and our Vigna field operations: H. Okano, T. Yoshida, T. Nobori, S. Hirashima, H. Kuwahara, T. Komatsuzaki, M. Akiba, T. Oomizu, S. Yanagiya, T. Taguchi, Y. Tsubokura, H. Tomiyama, T. Chibana, S. Hattori, and T. Yokoyama. Other visitors that have collected with us and supplied ideas for our work that include K. Hammer, University of Gesamthochschule Kassel, Germany, D. Jarvis, IPGRI, Italy, B. Pickersgill, University of Reading, U. K. The staff of the Japan International Cooperation Agency (JICA) have helped greatly with our field work, particularly we would like to thank M. Kawase, K. Egara, K. lrie and J. Takahashi. We would like to thank Imetyzez Khodabux a former student at Birmingham University for help with geographic information system analysis particularly in relation to figures 5.7 and 5.9. Our collaboration in various countries of Asia would not have been possible without the kind help and support of many people. We would like to acknowledge from Thailand: C. Thavarasook, C. Lairungreang, S. Pichitporn, S. Chotechuen, P. Nakeeraks, P. Poonsavasde and S. Nuplean, Chai Nat Field Crops Research Center N. Boonkerd, Suranaree Industrial University, D. Boonmalison, National Genebank, P. Ornnaichart and B. Taengsan, JIRCAS office, P. Srinives and S. Siripin, Kasetsart University; in Sri Lanka Department ofAgriculture, S. D. G. Jayawardena, M. Jayasuriya, A. S. U. Liyanage M. Samarasinghe, S. K. Senevirathne, D. K. Edirisinghe, H. M. Tilakaratne Banda, H. J. Warshakoon, J. Samaranayake, J. Simon, Raja and M. D. Dassanayake; in Malaysia: S. Anthonysamy and Ithin bin Binjang, Universiti Pertanian Malaysia; in Myanmar: U Tin Soe, U Kyaw Soe, U Tan Sein and Daw 00 Mar Awn. We are grateful for information on germplasm and genebank collections supplied by the following people: L. Engle and S. Shanmugasundaram, AVRDC, Wang Shurnin, Institute for Crop Germplasm Resources, China, and S. Dillon, Australian Tropical Crops Genetic Resource XIV

Centre, Australia, N. Q. Ng, IITA, T. Vanderborght, National Botanic Garden of Belgium, C. L. Guevara and D. Debouck, CIAT, and the curator of the seed bank, USDA. We are grateful to the staff of IPGRI in Asia for their cooperation particularly P. Sajise, V. Ramantha Rao, Z.W. Zhang and M. D. Zhou We appreciate assistance of taxonomists at the herbarium the Royal Botanic Gardens, Kew, particularly B. Verdcourt, G. P. Lewis, B. Schrire and Rico-Arce M. de Lourdes, during the first authors stay as an OECD fellow in 1998. We thank curators of the following herbaria for permission to consult specimens: R. Vickery, British Museum (Natural History), UK; D. Reue, Paris, Natural History Museum, France; A. Robyns, National Botanic Garden of Belgium; G. Thijsse, Rijksherbarium, Leiden, The Netherlands; H. Ohashi, Tohoku University, Japan; N. Murakami, Kyoto University, Japan. We thank to the following for permission to use copyright material in this book: The Botanical Society of Japan (Journal of Plant Research) for Figs. 3.1, 3.2; American Society of Agronomy (Crop Science) for Figs. 6.7, 6.8; Springer-Verlag (Theoretical and Applied Genetics) and N. Young for Fig. 6. lOa; Springer-Verlag (Molecular and General Genetics) and A. Kaga for Fig. 6 lOb; Kluwer Academic publishers (Euphytica) for Figs. 6.9, Table 6.5, (Genetica) for Figs. 2.8.b, (Genetic Resources and Crop Evolution) for Figs. 2.7a and b; Finally we would like to especially acknowledge the inputs of Kyoko Motoyoshi who not only prepared the figures that appear in chapter 4 but also help very much with preparation of the book in camera ready format.

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