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MANAGING GLOBE GET RESOURCES Fvest frees Committee on Managing Global Genetic Resources: Agricultural Imperatives Board on Agriculture National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1991

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NATIONAL ACADEMY PRESS 2101 Constitution Avenue, NW Washington, DC 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. This material is based on work supported by the U.S. Department of Agriculture, Agricultural Research Service, under Agreement No. 59-32U4-6-75. Additional funding was provided by Calgene, Inc.; Educational Foundation of America; Kellogg Endowment Fund of the National Academy of Sciences and the Institute of Medicine; Monsanto Company; Pioneer Hi-Bred International, Inc.; Rockefeller Foundation; U.S. Agency for International Development; U.S. Forest Service; W. K. Kellogg Foundation; World Bank; and Basic Science Fund of the National Academy of Sciences, contributors to which include the Atlantic Richfield Foundation, AT&T Bell Laboratories, BP America, Inc., Dow Chemical Company, E.I. duPont de Nemours & Company, IBM Corporation, Merck & Co., Inc., Monsanto Company, and Shell Oil Company Foundation. Library of Congress Cataloging-in-Publication Data Forest trees / Committee on Managing Global Genetic Resources: Agricultural Imperatives, Subcommittee on Managing Plant Genetic Resources, Forest Genetic Resources Work Group: Board on Agriculture, National Research Council. p. cm.-(Managing global genetic resources) Includes bibiliographical references and index. ISBN 0-309-04034-5: $19.95 1. Forest genetic resources conservation. 2. Tree~Germplasm resoures. I. National Research Council (U.S.), Forest Genetic Resources Work Group. II. Series. SD399. 7. F67 1991 634. 9-dc20 91-8979 CIP it) 1991 by the National Academy of Sciences. All rights reserved. No part of this book may be reproduced by any mechanical, photographic, or electronic process, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted, or otherwise copied for public or private use without written permission from the publisher, except for the purposes of official use by the U.S. government. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. Printed in the United States of America

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Committee on Managing Global Genetic Resources: Agricultural imperatives PETER R. DAY, Chairman, Rutgers University ROBERT W. ALLARD, University of California, Davis PAULO DE T. ALVIM, Comissao Executiva do Plano da Lavoura Cacaueira, Brasilia JOHN H. BARTON, Stanford University FREDERICK H. BUTTEL, Cornell University TE-TZU CHANG, International Rice Research Institute, The Philippines ROBERT E. EVENSON, Yale University HENRY A. FITZHUGH, International Livestock Center for Africa, Ethiopian MAJOR M. GOODMAN, North Carolina State University JAAP I. HARDON, Center for Genetic Resources, The Netherlands DONALD R. MARSHALL, Waite Agricultural Research Institute, Australia SETIJATI SASTRAPRADJA, National Center for Biotechnology, Indonesia CHARLES SMITH, University of Guelph, Canada JOHN A. SPENCE, University of the West Indies, Trinidad and Tobago Genetic Resources Staff MICHAEL S. STRAUSS, Project Director JOHN A. PING, Project Directort STEVEN KING, Research Associate JOSEPH I. GAGNIER, Senior Project Assistant * Executive Commission of the Program for Strengthening Cacao Production, Brazil. Winrock International, through January 1990. Through June 1990. Through June 1989. . . .

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Subcommittee on Plant Genetic Resources ROBERT W. ALLARD, Chairman, University of California, Davis AMRAM ASHRI, The Hebrew University of Jerusalem, Israel VIRGIL A. JOHNSON, University of Nebraska RAJENDRA S. PARODA, Indian Council of Agricultural Research, New Delhi H. GARRISON WILKES, University of Massachusetts, Boston LYNDSEY A. WITHERS, International Board for Plant Genetic Resources, Italy Forest Genetic Resources Work Group GENE NAMKOONG, Chairman, U.S. Department of Agriculture, Forest Service, Genetics Department, North Carolina State University KAMAL}IT BAWA, Department of Biology, University of Massachusetts, Boston lEFFERY BURLEY, Oxford Forestry Institute, Oxford University, United Kingdom SUSAN S. SHEN, Asia Environment and Social Affairs Division, World Bank, Washington, D.C. IV

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Board on Agriculture THEODORE L. HULLAR, Chairman, University of California, Davis PHILIP H. ABELSON, American Association for the Advancement of Science C. EUGENE ALLEN, University of Minnesota DALE E. BAUMAN, Cornell University R. JAMES COOK, Agricultural Research Service at Washington State University ELLIS B. COWLING, North Carolina State University ROBERT M. GOODMAN, Visiting Professor, University of Wisconsin, and National Research Council Scholar-in-Residence TIMOTHY M. HAMMONDS, Food Marketing Institute PAUL W. JOHNSON, Iowa House of Representatives NEAL A. JORGENSEN, University of Wisconsin ALLEN V. KNEESE, Resources for the Future, Inc. JOHN W. MELLOR, International Food Policy Research Institute ROBERT L. THOMPSON, Purdue University JAN VAN SCHILFGAARDE, U.S. Department of Agriculture, Ft. Collins, Colorado ANNE M. K. VIDAVER, University of Nebraska CONRAD I. WEISER, Oregon State University JAMES E. TAVARES, Acting Executive Director ROBERT M. GOODMAN, NRC Scholar-in-Residence CARLA CARLSON, Director of Communications BARBARA I. RICE, Editor v

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The National Academy of Sciences is a private, nonprofit, self- perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal government on scientific and technical matters. Dr. Frank Press is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its members, sharing with the National Academy of Sciences the responsibility for advising the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Robert M. White is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Institute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Samuel O. Thier is president of the Institute of Medicine. The National Research Council was organized by the National Acad- emy of Sciences in 1916 to associate the broad community of science and technology with the Academy's purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in providing services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Frank Press and Dr. Robert M. White are chairman and vice-chairman, respectively, of the National Research Council. . v'

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Preface Fvest trees are integral parts of human society. They provide fuel, fiber, construction and building materials, food, and medicines, among other things. The forest itself is an ecosystem, and as ecosystems, forests stabilize environments and are essential components of the global ecology. Although trees are the dominant vegetation, forests are rich reservoirs of biological diversity. They harbor a major proportion of the world's animal and plant species. Forest trees also enhance and protect our landscapes. They sustain wildlife, industry, and rural economies, and contribute to the quality and richness of our environment. For many years, concern has been expressed about rapid and contin- uing losses of the world's forests. In temperate regions of Europe and North America, the decline in forest health has been attributed to industrial consequences, such as acid rain. In the humid tropics, increased demands on the land resulting from the clearing of forests to accommodate expanding populations and the production of industrial products have been highlighted. The causes notwithstanding, the future for the world's forests, if unprotected, is dim. For many forested areas, efforts to halt or slow losses through the establishment of protected areas will be essential. However, more than protection will be needed. Society will continue to need the services and products derived from the forest. As natural stands of trees are lost, greater efforts to conserve those remaining will continue. There will be more need to select and develop trees and forests that are managed for production purposes to reduce pressure on the remaining . . v''

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viii / Preface natural forests and to provide raw materials in forms more suitable for commercial use and the needs of human society. To develop these future forests, the genetic resources of forest trees must be conserved and developed, whether they exist as trees in planted or protected conservation stands or as seeds or tissue cultures in storage (or one day possibly as DNA libraries). Managing forest genetic resources involves developing overall strategies, applying specific methodologies, developing new techniques, and coordinating local, national, regional, and global efforts. Although tree species are similar in many ways to crop species, managing forest genetic resources is not simply a matter of applying programs for crops to larger plants. Forests generate a wide variety of product values, from the different components of the ecosystems that depend on their structural viability to the industrial and agroforestry crops that can be consumed, and the variety of systems that are used to manage them dictates the variety of ways that the genetic resources are used. Moreover, the genetic architecture of forest trees is poorly known, breeding is slow, and wide variations in ecological and economic environments must be anticipated to use the available genetic variation efficiently. This report, while recognizing serious threats to all the species and ecosystems represented in the world's forests, focuses on managing those forest trees from which harvested materials are currently extracted. Protection of the world's forest ecosystems will require broad efforts by scientists and policymakers. A report prepared by the Commission on Life Sciences in cooperation with the Board on Agriculture, entitled Forestry Research: A Mandatefor Change (National Research Council, 1990), recommends modifications in the way forestry research is conducted to bring about many needed improvements in the forestry research com- munity. Careful, well-coordinated management of those tree species of current or future harvest potential could play an important part in reducing deforestation pressures on the remaining pristine vegetation. Although the total number of known forest tree species exceeds 50,000 and potential extractive use can be made of several thousand species, current efforts to manage trees focus primarily on fewer than 140 species. Clearly, greater efforts and coordination are needed to sustain even the current levels of forest production and to realize the potential produc- tivity of as yet undeveloped species. The Committee on Managing Global Genetic Resources, established by the National Research Council in November 1986, is concerned with genetic resources of identified economic value. These resources are important to agriculture, forestry, fisheries, and industry. The committee

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Preface I ix has been assisted by two subcommittees and several work groups that gathered information or prepared specific reports. One of the work groups, chaired by Gene Namkoong, examined the management of forest genetic resources and drafted this report. It is one of five reports to be published in a series entitled, Managing Global Genetic Resources. The other reports prepared by the committee, its subcommittees, and work groups address issues related to the management of plant genetic resources by the U.S. National Plant Germplasm System and the global management of livestock, fish and shellfish, and crop plants. The examination of crop plants will be included in the committee's main report, which will address the legal, political, economic, and social issues surrounding global genetic resources management as they relate to agricultural imperatives. In addition, a work group was appointed to provide information that would aid in planning and designing a new storage facility for the U.S. National Seed Storage Laboratory. The committee released the report, Expansion of the U.S. National Seed Storage Laboratory: Program and Design Considerations, in April 1988. Copies of this report are available from the Board on Agriculture. The Forest Genetic Resources Work Group was asked to do the following: Examine the uses and status of forest tree genetic resources globally. Examine in situ and ex situ methods for the conservation of forest genetic resources. Assess current germplasm conservation activity for forest tree genetic resources by national, regional, and international organizations. Identify the major problems in implementing action, including coordination, information dissemination, and training. Recommend future actions to solve or alleviate technical and financial problems. Present a global strate~v for conserving and managing forest tree ~ i, genetic resources. (JJ (J {J OCR for page R1
x I Preface that determine the structure of genetic variation of managed and unmanaged forest tree populations and points out how this knowledge is important to efforts to create, monitor, conserve, and manage reserves for tree species. Chapter 4 defines the methods and technology available for the management of trees through complementary in situ and ex situ conservation activities, and outlines the importance of greater emphasis on developing long-term in situ conservation programs. Chapter 5 describes the activities of the national, regional, and international organizations that are involved with the management and conservation of tree genetic resources. Chapter 6 presents the committee's recom- mendation on how to implement rapidly a much needed global strategy for conserving and managing forest tree genetic resources. The sciences related to genetics, plant breeding, and resource man- agement are advancing rapidly. The commitment of increasing numbers of nations to genetic resources conservation is growing. The committee believes that the conclusions and recommendations presented in this report can contribute to improved conservation and management of a major resource the world's forest trees. PETER R. DAY, Chairman Committee on Managing Global Genetic Resources: Agricultural Imperatives

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Acknowledgments Many people and agencies have contributed their support, time, and creative analysis to this report. Because of the global nature of the study, it was critical that scientists from numerous countries took the time to respond to a variety of questions posed by the committee. To all of those researchers, special thanks are offered. The committee would like to thank Robert Kellison, Frank Santamour, Karl Keipi, John Spears, and Carl Gallegos for providing initial advice and direction for this report. In particular, the help of John C. Gordon, Raymond Curies, Colin Hughes, R. Sniezko, Milton Kaneshiro, Lilia Barrientos, and Lucy Nunnally is gratefully acknowledged. Special thanks are offered to Christel Palmberg of the Food and Agriculture Organization of the United Nations and to Stanley Krugman of the U.S. Forest Service, who contributed their knowledge and time from the beginning of this study. In addition the committee acknowledges Grace Jones Robbins, Nicole L. Kelsey, Philomina Mammen, Sherry Showell, Carole Spalding, and Maryann Tully of the Board on Agriculture staff for their assistance throughout various drafts of the report. x'

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Contents EXECUTIVE SUMMARY The Crisis of World Forest Decline, 3 Benefits, Uses, and Development of Forest Trees, 4 Structure of Genetic Variation in Trees, 7 Recommendations, 8 1 WORLD FORESTS Losses in Forests, 21 Basic Inventory Data, 25 Causes of Forest Tree Loss, 26 What Can Be Done?, 29 Recommendations, 32 MULTIPLE USES OF FOREST TREES Benefits of Trees and Forests, 37 Land Use Systems Involving Trees, 41 Increasing the Usefulness of Tree Genetic Resources, 45 Recommendations, 47 STRUCTURE OF GENETIC VARIATION Mating Systems and Gene Flow, 51 Estimating Genetic Variation, 59 Conclusions, 71 Recommendations, 71 . . . Xlll 21 .37

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xivlContents 4 CONSERVATION AND MANAGEMENT OF TREE GENETIC RESOURCES ...................... Conservation and Management Strategies, 73 In Situ Conservation, 75 Ex Situ Conservation, 85 Choice of Method, 89 Management Status of Tree Species, 93 Conclusions, 95 Recommendations, 96 5 INSTITUTIONS INVOLVED IN MANAGING TREE GENETIC RESOURCES ................ In Situ Conservation Activities, 102 Ex Situ Conservation Activities, 103 International Institutions, 105 National and Regional Institutions, 116 Conclusions, 124 Recommendations, 127 6 ORGANIZING A GLOBAL SYSTEM OF COOPERATION ..................... Coordinating and Expanding Existing Programs, 1ZY Developing Regional and National Programs, 132 Developing an International Forest Tree Genetic Resources Program, 134 Recommendation, 137 REFERENCES .................... APPENDIXES A Forest Tree Species Used in Breeding or Testing Activities, 147 B Literature Survey of Threatened Provenances or Species, 161 C Sources of Seed for Research, 183 GLOSSARY ABBREVIATIONS AUTHORS INDEX ..73 .99 ....... 129 .139 - 201 ..207 ..209 213

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Forest Trees

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