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LD5655.V856 1987.C677.Pdf (11.14Mb) , I i.· '.> The systematics and Ecology of Boletes with special reference to the Genus Suillus and its Ectomycorrhizal Relationships in Nepal by Henry Van Tuy! Cotter Dissertation submitted to the Faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Botany APPROVED: Orson K. Mi1ller, Jr., \Chairman l <>< ~--r''=r'< I Khidir W. Hilu John G. Palmer I I I Robert A. Paterson l/' R}Jay StiJes July, 1987 Blacksburg, Virginia The Systematics and Ecology of Boletes with special reference to the Genus Suillus and its Ectomycorrhizal Relationships in Nepal by Henry Van Tuyl Cotter Orson K. Miller, Jr., Chairman Botany (ABSTRACT) The Suillus mycota of Nepal was studied. Nine species are recognized and described; five of the nine appear to be new species. Additional species were collected, but material was inadequate to describe them completely. Cultures of eight and synthesized ectomycorrhizae of six of the Suillus species are described. Synoptic keys to the basidiocarps and to the cultures are presented. Numerical taxonomic analyses of the cultures generated clusters which paralleled the species concepts developed using basidiocarps and ecology. Each species of Suillus from Nepal is host specific based on basidiocarp formation; all hosts are in the Pinaceae. Field associations are Suillus cf. granulatus, S. cf. placidus, S. sibiricus, and greening-foot Suillus with Pinus wallichiana; queen's Suillus with P. roxburghii; waxy Suillus with P. patula; and S. laricinus, himalayan Suillus, and orange-pored Suillus with Larix himalaica. Mycorrhizal syntheses confirmed that the six Suillus-Pinus relationships are ectomycorrhizal. The Suillus mycota of western Virginia has 12 known species. Three, of the five which are ectomycorrhizal with Pinus strobus, have closely related counterparts in Nepal. These counterparts are ectomycorrhizal with P. wallichiana, a five-needled pine closely related to P. strobus. The existence of these three pairs of similar fungi, associated with similar pines, suggests the possibility of cladogenic speciation in parallel by the pine lineage and by its ectomycorrhizal fungal associates. Boletinellus merulioides forms abundant sclerotia in nature throughout its range in eastern North America. Sclerotia collected in the forest germinated to form mycelial colonies that had the same appearance and microscopic characteristics as colonies derived from basidiocarps. Sclerotia which had overwintered in the forest were viable in the spring. The spatial pattern of B. merulioides sclerotia in a forest was compared with basidiocarp frequency recorded over four years. Both estimates of the spatial pattern coincided, but year-to-year basidiocarp frequency varied greatly. Individual B. merulioides dikaryons formed large perennial patches. Basidiocarp and sclerotial densities were centered around and declined outward from Fraxinus americana trees. Boletinellus merulioides and Fraxinus pennsylvanica did not form ectomycorrhizae when grown together in growth pouches. ACKNOWLEDGEMENTS This dissertation is dedicated to my parents and grandparents and to my wife Numerous people have been helpful in a multitude of ways, to all my sincere gratitude; no dissertation is done alone. Dr. 0. K. Miller, Jr., my advisor, and Dr. K. W. Hilu, Dr. J. G. Palmer, Dr. R. A. Paterson, and Dr. R. J. Stipes, my committee members, have been steadfast sources of inspiration, support and advice and have given me the freedom to pursue my ideas and dreams. They are all exemplars in differing ways of what I aspire to become. My heartfelt thanks to to to and , and the other members of the Fulbright community; tc for their confidence; to for valuable discussions of Suillus in the New and Old Worlds; to Tribhuvan Univ. for approval of my research and an affiliation with the Botany Dept.; to the HMG Dept. of National Parks & Wildlife Conservation for permission to collect in Nepal's national parks; to (ACAD), (NY), (Shimla), (TENN) i (NY) and (HKAS) for making specimens available for examination; and to the Fulbright Program, the Graduate School and the Biology Department for support. Acknowledgements iv Cela est bien dit, repondit Candide, mais il faut cultiver notre jardin. Voltaire 1759 v TABLE OF CONTENTS Abstract ii Acknowledgements iv Table of Contents vi List of Figures ix List of Tables xi Chapter 1. Systematics and ectomycorrhizal relationships of the bolete genus Suillus in Nepal . 1 The geology, climate and vegetation of Nepal 3 History of mycology in Nepal 7 Boletes of Nepal 13 Comments on the genus Suillus 16 Materials and Methods 22 Field work 22 Descriptions of basidiocarps 25 Cultures and cultural descriptions 25 Numerical taxonomic analysis of cultural data 29 Vegetative compatibility experiment 31 Ectomycorrhizal syntheses 32 Results 34 Field work and summary of collections 34 Definition of Suillus species from Nepal 35 Numerical taxonomic analysis of cultural data 36 Vegetative compatibility experiment 41 Table of Contents vi Ectomycorrhizal syntheses 42 Descriptions of Nepali Suillus species & their ectomycorrhizae 44 Su illus cf. granulatus 45 Su illus cf. placidus 48 Suillus sibiricus 51 Su illus II greening-foot" 54 Su illus II queen I s II 57 Su illus II waxy II 60 Su illus laricinus 63 Su illus 11 himalayan" 66 Su illus II orange-pored" 68 Synoptic keys to Suillus species of Nepal 70 Synoptic key to basidiocarps of Suillus from Nepal 72 Synoptic key to cultures of Suillus from Nepal 75 Discussion 77 Figures and Tables for Chapter 1 91 Chapter 2. The genus Suillus in western Virginia: Checklist and host relationships 113 Comments on the genus Suillus 113 Materials and Methods 113 Results 114 Host relationships 115 Checklist of western Virginia species of Suillus 116 Discussion 118 Table for Chapter 2 120 Chapter 3. Sclerotial production and ecology of Boletinellus merulioides ....... 121 3.1. Sclerotia of Boletinellus merulioides in nature 121 Materials and Methods 122 Field collections 122 Laboratory cultures 123 Herbarium collections 123 Table of Contents vii Results ....... 124 Description and identification of the naturally occurring sclerotia 124 Occurrence and distribution 125 Germination and overwintering 127 Discussion 127 3.2. Comparison of spatial patterns of sexual and vegetative states of Boletinellus merulioides 129 Materials and Methods 130 Results 134 Discussion 138 Figures and Table for Chapter 3 142 Literature Cited 148 Appendix A. List of fungal collections from Nepal 158 Appendix B. Forms used in describing cultures, basidiocarps and ectomycorrhizae . 167 Appendix C. Data set of 82 characters for 28 Nepali bolete cultures 174 Vita 176 Table of Contents viii .!.!ll OF FIGURES Figure 1.1. Van and Irene Cotter's collecting routes in Nepal 91 Figure 1. 2. Dendrogram of Nepali Suillus cultures, host data not included . 92 Figure 1. 3. Dendrogram of Nepali Suillus cultures, host data included . 93 Figure 1. 4. PCA three-dimensional graph of Nepali Su illus cultures 94 Figure 1.5. PCA three-dimensional graph of Nepali Su illus cultures, different orientation of axes . 95 Figure 1.6. PCA three-dimensional graph of Nepali Suillus cultures, cultures interconnected by minimum spanning tree linkages 96 Figure 1.7. a. Suillus cf. granulatus. b. Synthesized ecto- mycorrhizae between Pinus wallichiana and Suillus cf. granulatus 97 Figure 1.8. a. Suillus cf. placidus. b. Synthesized ectomycorrhizae between Pinus wallichiana and Suillus cf. placidus . 98 Figure 1.9. a. Suillus sibiricus. b. Synthesized ectomycorrhizae between Pinus wallichiana and Suillus sibiricus . 99 Figure 1.10. a. Suillus "greening-foot". b. Synthesized ecto- mycorrhizae between Pinus wallichiana and greening-foot Suillus 100 Figure 1.11. a. Suillus "queen's". b. Synthesized ectomycorrhizae between Pinus roxburghii and queen's Suillus 101 Figure 1.12. Su illus "waxy" 102 Figure 1.13. Suillus laricinus 102 Figure 1.14. Su illus "himalayan" 103 Figure 1.15. Su illus "orange-pored" 103 Figure 3.1. Sclerotia of Boletinellus merulioides 142 Figure 3.2. Basidiocarps and sclerotia of Boletinellus merulioides 142 Figure 3.3. Cultures of Boletinellus merulioides 142 Figure 3.4. Range of Boletinellus merulioides basidiocarps and sclerotia . 143 List of Figures ix Figure 3.5. Vertical profile of sclerotia of Boletinellus merulioides . 144 Figure 3.6. Spatial pattern of Boletinellus merulioides in plot 6 145 Figure 3.7. Boletinellus merulioides basidiocarp frequency year by year . 146 List of Figures x LIST OF TABLES Table 1.1. Summary of the members of the Pinaceae which occur in Nepal . 104 Table 1.2. Outline of Van and Irene Cotter's collecting activity in Nepal 106 Table 1.3. Summary by genus of the fungal collections from Nepal 108 Table 1.4. Species of Suillus in Nepal and their ectomycorrhizal partners ..... 111 Table 1.5. Comparison of Suillus species associated with Pinus strobus of North America and P. wallichiana of the Himalaya 111 Table 1.6. Suillus cultures from Nepal 112 Table 2.1. Species of Suillus in western Virginia 120 Table 3.1. Vegetative compatibility among isolates of Boletinellus merulioides . 147 List of Tables xi CHAPTER .L.. SYSTEMATICS AND ECTOMVCORRHIZAL RELATIONSHIPS OF THE BOLETE GENUS SUILLUS IN NEPAL The bolete genus Suillus is ectomycorrhizal almost exclusively with members of the Pinaceae, especially the genera Pinus and Larix (Miller, 1982; Singer, 1986). Species of Suillus have been reported from around the world wherever Pinus and Larix grow either as native or introduced trees. Therefore, although Suillus had not been reported from Nepal, the genus was expected to occur there because of the presence of both Pinus and Larix (Table 1.1). Nepal has two native pines; Pinus wallichiana, a 5-needled pine, and P. roxburghii, a 3-needled pine; and two Himalayan endemic native larches; Larix himalaica and L. griffithiana. Exotic pines, especially P. patula (3(-5)-needled), are used in reforestation efforts in Nepal. The objectives of my research in Nepal were to: 1) Collect and identify the Suillus species associated with the conifer genera Abies, Larix, Picea, Pinus and Tsuga.
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