Kun-fang Cao FAGUSDOMINANC E IN CHINESE MONTANE FORESTS: natural regeneration of Fagus lucida and Fagushayatae var .pashanica Ontvangers 16 JON, 1995 UB-CARDEX CENTRALE.LANDBOUWCATALOGU S 0000 0546 8513 *i°3- Promotoren: Dr. Ir. R.A.A. Oldeman hoogleraar in de bosteelt en bosoecologie Prof. X.P. Wang hoogleraar in de botanie (Beijing), Botanisch Instituut, Chinese Academie van Wetenschappen Co-promotor: Dr. Ir. R. Peters Gast onderzoeker in bosecologie /ûvKtèTOl , W*. Kun-fang Cao F^GTO DOMINANCE IN CHINESE MONTANE FORESTS: natural regeneration of Fagus lucida and Fagushayatae var .pashanica Proefschrift ter verkrijging van de graad van doctor in de landbouw- en milieuwetenschappen op gezag van de rector magnificus, dr. C.M. Karssen, in het openbaar te verdedigen op maandag 9 januari 1995 des namiddags te vier uur in de Aula van de Landbouwuniversiteit te Wageningen l/;M ~C]G>[ DG O CIP-DATA KONINKLIJKE BIBLIOTHEEK, DEN HAAG Cao, Kun-Fang FAGUS dominance in Chinese montane forests: natural regeneration of Faguslucida and Fagushayatae var . pashanica I Kun-fang Cao. -[S.L.: s.n.].-Ill. Thesis Wageningen. - With ref. - With summary in English, Dutch and Chinese. ISBN 90-5485-330-1 Subject headings: beech forests, regeneration, China, forest ecology. Cover design by: Dr. R. Peters Printed by: Grafisch Service Centrum Van Gils B.V. fjpofzo fn PROPOSITIONS 1. Notwithstanding the actual scarcity ofjuvenil e beeches, Fagus dominance in the mixed montane beech forests of southern China will be maintained as long as the actual climate of this beech region persists. (This thesis) 2. In shaded environments within forests, minimizing self-shading is a common and general plant strategy which is followed, however, by means of numerous different modes of plant development. (This thesis) 3. Reiteration strategies in the form of basal sprouting and as mechanism of crown building define an important, frequently occurring but still under-documented developmental pattern allowing numerous tree species to survive in shade. (This thesis) 4. Modelling tree species' or forest ecosystem's responses to global climatic change should consider not only temperature and precipitation but also natural hazards. 5. Because natural events repeatedly open up the forest canopy and so stimulate forest regeneration and interfere with the competitive exclusion of species and also exert selective pressure on plant populations and therefore increase genetic fitness of successive generations for survival, such events should not be considered as "disasters" or 'disturbances' in natural forests. 6. Because chance plays an important role in ecology (cf. Hubbell &Foste r 1986), many natural phenomena are unpredictable. 7. In recent Chinese policy, there have been three major steps towards deforestation: "steel production" in late 1950s, "grain production" between 1960s and 1970san d privatization of forests in 1980s. 8. Today, the Chinese government gives priority to economical development, whereas it can be demonstrated that education, scientific research, conservation of the environment and political reform are equally important. 9. To save natural resources and protect the environment, people in developed countries should limit their living requirements, people in developing countries should control their population growth. 10. Socialist and capitalist systems will have much in common in the next century. To my parents BIBLIOTHEEK LANDBOUWUNIVERSITETr WAGENTNf—ï VII PREFACE The work described in this thesis was initiated in 1987whe n the author and Dr. R. Peters worked together in Chinese beech forests. This joint work has been extended into a joint research project between the Institute of Botany in Beijing, Chinese Academy of Sciences, Department of Forestry, Wageningen Agricultural University, and Department of Plant Ecology and Evolutionary Biology, Utrecht University, which focuses on the ecology of Chinese beech forests. The project has being carried out under the Scientific Cooperation Agreement between the Chinese Academy of Sciences and the Royal Dutch Academy of Sciences. The work described in this thesis waspar t of this joint research project. First of all, I thank you, Prof. R.A.A. Oldeman and Prof. Xian-pu Wang for your enthusiasm, supervision and support, for valuable discussions and for your friendship. I am indebted to you, Dr. R. Peters for your co-supervising this study, for your valuable advice and discussions, for your cooperation, andfo r your encouragement and kindness. I thank you, Dr. P. Schmidt and Dr. F. J.J.M. Bongers for your advice on the research plan and critical comments on parts of the thesis. I am grateful to you, Prof. F. Halle, Dr. C. Edelin for your advice and discussions on tree architecture, and to you, Dr. B. Thiébaut for your introduction to architectural plasticity of Fagussylvatica. To you, Prof. T. Poulson, I am obliged for your instruction in using Ozalid light-sensitive paper to assess light intensity. Prof. E. Box, Prof. Hsin-shih Chang and two anonymous referees, I amgratefu l to you for your comments and advice on Chapter 2. For your reviews, comments and discussions on parts of this thesis, I thank all ofyou : Ir. H.H. Bartelink, Ir. J. den Ouden, Ir. P. van der Meer, Ir. M.E. van der Zee, Dr. R. van Rompaey, Ir. R.B. Shrestha, Ir. F. Sterk, Prof. Hui-ying Zhen. Several persons also provided me with discussions and advice. Thankyou : Prof. Ling-zhi Chen, Prof. Wei-lie Chen, Prof. J. Fanta, Prof. Miao-guang Hé, the late Prof. Hsioh-yu Hou, Prof. Shun-shi Hu, Dr. W. B.J. Jonkers, Asso. Prof. Shi-zhou Sun, Prof. Zhao-chen Kong, Ir. K.F. Wiersum, Mr. Yi-qin Yang, Mr. Wei-cao Yu, Prof. Tsun-shen Ying. In China, several people helped met o carry outth e fieldwork. Thankyou : Mr. Hé-ping Chen, Mr. Ke Guo,Mr . Shi-qiang Hé and Mrs. Xia Li for Daba; Mr.Jin - sheng Hé, Mr. Xiong Lin and Asso. Prof. Yuan-guang Wen for Miao'ershan and Lilio; Mr. Chuan-dong Yang, 'Lao' Yang and 'Xiao' Yang for Fanjingshan; Asso. Prof. De-fa Shi for four sites in Zhejiang Province; Mr. Fan-zhang Du and Mr. Fong-bing Yan for Hupingshan. Mr. M.H.J.S.M. Hegeman, thank you for your help with the computer network. Mrs. Chun-ying Liu, for your assistance in running the Thornth.For program, Prof. Dian-'an Yang, for your assistance in using the EIS (a modified GIS) software, and Ir. P. vande r Meer, for your assistance in the light measurement in Wageningen; please accept my warm acknowledgements. Ir. G. Gort and Ir. J.J. Jansen, I am grateful to you for your advice on statistics. Mrs. J. Burrough-Boenish, thank youfo r editing the English VIII of this thesis. Mrs. A.D. Bosch-Jonkers, thank you for your translation of the summary into Dutch. Mrs. Fan-zhi Kong and Mrs. Yi-chuan Yao, I am obliged to you for analysing silica content of bamboo leaves. For your relevant information, I thank all ofyou : Dr. F.J.J.M. Bongers, Prof. Hsin-shih Chang, Dr. Cornelissen, Dr. C. Edelin, Dr. J.Y. Fang, Dr. N.R. De Graaf, the late Prof. Hsioh-yu Hou, Dr. Mo-yi Kang; Dr. T. Kohyama, Asso. Prof. De-zhen Kong, Prof. Zhao-chen Kong, Mr. Ren-ling Liu, Dr. T. Nakashizuka, Dr. T. Ohkubo, Prof. M. Ohsawa, Dr. R. Peters, Prof. R.A.A. Oldeman, Dr. P. Schmidt, Asso. Prof. De-fa Shi, Dr. B. Thiébaut, Prof. Xian-pu Wang, Prof. M. Werger, Asso. Prof. Yuan-guangWen . I appreciate thepermissio n to carry out the fieldwork, given by: Guiling Department of Forestry for Miao'ershan Nature Reserve; Guizhou Department of Forestry for Fanjingshan MABNatur e Reserve; Huangshan National Park for Huangshan; Hunan Department of Forestry for Hupingshan Nature Reserve; Longsheng Department of Forestry for Liluo Forest Station; Nanjiang Forest Service for Daba Forest Station; Yongjia Department of Forestry for Shihaishan Forest Station; Zhejiang Department of Forestry for Tianmushan National Park, for Longwangshan Nature Reserve and for Baishanzu Nature Reserve. I sincerely thank you, my colleagues in the Laboratory of plant Ecology and the Laboratory of Quantitative Vegetation Ecology, Institute of Botany in Beijing, Chinese Academy of Sciences, and you, the colleagues in the Departments of Forestry and Ecological Agriculture, Wageningen Agricultural University, for your assistance of all kinds you have offered to me from time to time, your hospitality and friendship. This study was co-funded by the National Science Foundation of P.R. China (project no. 39070188) and a doctoral fellowship of Wageningen Agricultural University, by the Chinese Academy of Sciences, the Netherlands Ministry of Education and Sciences &th e Netherlands Academy of Sciences (KNAW; Scientific Cooperation with China). The Institute of Botany in Beijing provided the regular salary and research facilities. The Department of Forestry, Wageningen Agricultural University, provided working facilities during my three periods of stay in Wageningen. I always felt welcome for the discussions with my promotor at the Department of Ecological Agriculture of Wageningen Agricultural University. My wife Ling Wang and my daughter Dongna Cao gave me support, freedom and cheerfulness; thank you for all of these. Wageningen, The Netherlands September 1994 ft-# ^Tf 4 Kun-Fang Cao IX CONTENTS Preface VII Chapter 1 Introduction 1 1.1 Importance of beeches and beech forests 1 1.2 Juvenile beeches are rare in southern Chinese beech forests 3 1.3 Abiotic factors that may affect beech regeneration 3 1.4 Biotic factors that affect beech regeneration 4 1.5 Objectives of the study 5 1.6 Time of field work 5 Chapter 2 Climatic range and distribution