Dipterocarpaceae : Mycorrhizae and Regeneration W.T.M. Smits CENTRALE LANDBO UW CATALO GU S 0000 0572 5169 BlöLlOTaiEKM CKNDBOUWUNIVERS TTlCTi , WACiF.NINCEN Promotoren : dr. ir. R.A.A. Oldeman, hoogleraar in de bosteelt en bosoecologie dr. ir. J. Dekker, emeritus hoogleraar in de fytopathologie fJ>J0k'2ö't ,ßrj 2 1 OKT. f994 W.T.M. Smits i .r- » ^ „ UB-CARDEX Dipterocarpaceae : Mycorrhizae and Regeneration Proefschrift ter verkrijging van degraa d van doctor in de landbouw- en milieuwetenschappen op gezag van de rectormagnificus , dr. C.M. Karssen, inhe t openbaar te verdedigen opwoensda g 12 oktober 1994 des namiddags tevie r uur ind eAul a van de Landbouwuniversiteit teWageningen . Uitgegeven als boek door Stichting Tropenbos ,':> n bb'n}û Abstract Smits, W.T.M. (1994). Dipterocarpaceae : Mycorrhizae and Regeneration. PhD Thesis, Wageningen Agricultural University, The Netherlands, 242 pp.,7 6figs., 2 7tables , 9boxes , 8 plates with colour pictures, 242 references, 16 terms in glossary, English, Dutch and Indonesian summaries. Research on mycorrhizae of Dipterocarpaceae is described, involving inventories of both mycorrhizae and sporocarps in natural forest and experimental work in nurseries, green houses, laboratories and gnotobiotic systems. An assessment is made of dipterocarp mycorrhizalspecificit y anda discussio ni spresente do nho wmycorrhiza lspecificit y mayhav e contributed to speciation in Dipterocarpaceae. Other aspects touched upon include work on a non-ectomycorrhizal association of a fungus with dipterocarp roots, proposed to be called amphymycorrhizae. Also discussed are the effects of physical influences upon dipterocarp ectomycorrhizae, demonstrating the negative impacto f hightopsoi ltemperature s and lacko f oxygen upon functioning and survival of dipterocarp ectomycorrhizae. Furthermore how dipterocarp ectomycorrhizae influence regeneration of Dipterocarpaceae through enhanced survival near the mother trees. At the end of the book practical recommendations are given for optimalization of management of mixed dipterocarp forests based upon the conclusions reached in the research, including the use of correct fungus-dipterocarp combinations for different sites. CIP-DATA KONINKLIJKE BIBLIOTHEEK, DENHAA G Smits, W.T.M. Dipterocarpaceae :Mycorrhiza e and Regeneration. Thesis Wageningen, - With réf. - With summaries in Dutch, English and Indonesian. ISBN 90-5485-331-X. Subject headings: Dipterocarpaceae/ tropical forestry/ mycorrhizae/ root research ; forests Tevens uitgegeven alsboek doorStichting Tropenbos I C ;.AJO??0 > )8Z I 1) Direct photosyntheticcarbohydrat e production and translocationfrom leave s to roots is a precondition for formation of ectomycorrhizal sporocarps associated with Dipterocarps. (This book). 2) The so-called delicate balance of speciesric h climax tropical rain forest persists because of the robustness of the forest to overcome disturbances. 3) Conservation of primaryrai n forests through aboycot t of tropical timber cannot bereache d throughindividua lcountr yaction san dwil lhur tth epoo rpeopl elivin g near these forests most. 4) Verdere studie van zwammen kan veel toekomstig gezwam over soortvorming voorkomen. 5) Orangutans are great botanists. 6) Crown shyness,withou t thigmotropy,mus tb e explained through the presence of so far unknown sensitory mechanisms in plants. 7) Specializationi nscienc emostl ylead st oregula rbu tmodes tprogress ,whil ea mor e generalistic approach,combinin gth especialism san d insighti nth efunctionin g of the practical world,wil l bring greatest progress for all. 8) Withincreasin gwelfare ,biodiversit ybecome sa nincreasingl yvaluabl e commodity and the present downward trend innumbe r of speciessurvivin gwil ltur n upward as a result. 9) Therewil lneve rb e an endt o the chain ofsmalle rparticle smakin gu pmatte r nor to the space and time matter occupies. 10) To become a democratic political leader one will always face the dilemma that one is expected to have strong principles but neverthelessb e readyt o abandon ones own principles for the sake of the majority; a good leader must therefore longmor e for power than anything elsean d can neverb e trulytrustfu l tohimsel f or others. W.T.M. Smits, "Dipterocarpaceae :Mycorrhiza e and Regeneration" 12Oktobe r 1994, Wageningen, Nederland. Dipterocarpaceae : Mycorrhizae and Regeneration Table of contents Contents 5 List of figures 3 List of boxes 12 List of tables 13 Preface 15 Acknowledgements 17 Chapter 1 : Introduction 19 1.1 Dipterocarpaceae 19 1.1.1 General 19 1.1.2 History ofutilizatio n 22 1.2 Mycorrhizae 25 1.2.1 General 25 1.2.2 Mycorrhizae associated with Dipterocarps 27 1.3 Purpose and outline of the research 27 1.3.1 General settingo fth e research 28 1.3.2 General description ofth e research area 30 Chapter 2 : Inventory of mushrooms found near Dipterocarps 33 2.1 Introduction 33 2.2 Methods 33 2.2.1 Layout ofth e plot 33 2.2.2 Collection and treatment ofsporocarp s 37 2.2.3 Assessment ofth e mycorrhizal status and/or specificity ofth e associations 42 2.3 Results 45 2.3.1 Tree species composition ofth eplo t 45 2.3.2 The mushrooms: keys and descriptions 60 2.3.3 Combinations ofDipterocarp s and ectomycorrhizal sporocarps 83 2.3.4 Factors affecting mushroom development 86 Contents - 5 - a. Influence ofrainfall upo n sporocarp appearance 86 b. Substrate preferences of several suspected ectomycorrhizal fungi 87 C. The influence of light and temperature upon sporocarp formation 88 d. Relation between physiological condition of phytobionts and sporocarp formation 88 e. Rate of appearance and deterioration of sporocarps 92 2.4 Discussion 92 2.4.1 Reliability of thesurve y data with respect to assessing host-specificity 92 a. Omission of small and hypogeous sporocarps 93 b. The limitation to the crown projection areas 93 C. Thepossibilit y of other tree species being associated with the encountered sporocarps 94 d. The limitation to dipterocarp canopy trees 96 e. The differences in vitality of the sampled trees 97 f. The differences innumber s of sampled trees per species 97 g. The possible correlation between topography, site of the sampled trees and differences in encountered sporocarps 98 h. The possibility of incomplete sampling 98 i. Possible interactions between physical influences and appearance of sporocarps 101 j. The limitation to one sample plot 101 2.4.2 Specificity 103 Chapter 3 : Inventories of ectomycorrhizae 107 3.1 Introduction 107 3.2 Methods (description andevaluation) 107 3.2.1 Method 1,direc t systematic root sampling 107 3.2.2 Method 2, direct root sampling of individual trees 109 3.2.3 Method 3,roo t sampling onspot s of mycobiont sporocarp appearances Ill 3.2.4 Method 4,to p soil washing and plotting of ectomycorrhizal types Ill 3.2.5 Method 5,collectin g of roots of dipterocarp seedlings resulting from natural regeneration 113 3.2.6 Method 6, sampling of seedlings resulting from natural regeneration and transferred to perforons 113 3.2.7 Method 7,plantin g of non-mycorrhizal dipterocarp seedlings under seed trees followed by evaluation of mycorrhizaltype s established 114 3.3 Results 114 3.3.1 Results of direct systematic root sampling 114 3.3.2 Results of topsoi l washing and plotting of ectomycorrhizal types 117 3.3.3 Resultsfrom collectin g roots of natural regeneration of Dipterocarps 118 3.4 Discussion 120 Chapter 4 :Inoculatio n experiments 123 4.1 Introduction 123 4.2 Methods 123 4.2.1 Infection of non-mycorrhizal dipterocarp seedlings related todistanc e from large dipterocarp trees 124 4.2.2 Influence of seed collecting method upon mycorrhizal infection after transplanting to medium without inoculum 124 4.2.3 Inoculation of non-mycorrhizal dipterocarp seedlings with chopped sporocarps of potential ectomycorrhizal fungi 125 4.3 Results 125 4.3.1 Correlation between mycorrhizal infection and distance from large dipterocarp trees. 125 4.3.2 Occurrence of first ectomycorrhizal infection 126 4.3.3 Compatibility ofphytobiont-mycobion t combinations under greenhouse conditions . 127 4.4 Discussion 128 Chapter 5 : Perforon studies 131 5.1 Introduction 131 5.2 Methods 138 5.2.1 Ectomycorrhizal development in soil and in perforations 138 5.2.2 Process of mycorrhizal infection inperforon s and amount of inoculum needed for growth enhancement of previously non-mycorrhizal plants 138 5.2.3 Rate ofroo t growthan d spread of ectomycorrhizae studied in perforons with Shorea stenoptera andAnisoptera marginata 139 5.2.4 Inter-species exchange of ectomycorrhizal fungi amongst dipterocarp seedlings . 139 5.3 Results 140 5.3.1 Influence of perforations upon roots and ectomycorrhizae 140 5.3.2 Infection process, start of growth enhancement and amount of inoculum needed . 141 5.3.3 Growth of roots and ectomycorrhizalhypha e 143 5.3.4 Selective mycorrhizal establishment inpresenc eo f various ectomycorrhizal fungi .. 143 5.4 Discussion and conclusions 144 5.4.1 Methodology 144 5.4.2 The obligate nature of dipterocarp mycorrhizae 146 5.4.3 Specificity of dipterocarp ectomycorrhizae 147 Chapter 6 : Physical influences upon ectomycorrhizae 149 6.1 Introduction 149 6.2 Methods 149 6.2.1 Experiment 1 (controlled heating of mycorrhizal root systems) 149 6.2.2 Experiment 2 (planting under three different field conditions with varying light intensities andsoi ltemperatures ) 151 6.2.3 Experiment 3 (separation of light effects upon the parts above ground and soil temperature upon ectomycorrhizae) 152 6.2.4 Experiment 4 (effect of soil compaction) 152 6.2.5 Experiment 5 (effect of soil