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Russell Kent Heblack a Thesis Submitted to Sonoma State College smriE SONOMA COUNTY HYPOGEOU s FUNGI by Russell Kent Heblack A thesis submitted to Sonoma State College in partial fulfillment of the requirements for the degree of fr.ASTER OF ARTS in Biology J~~ph (. Powell, Chairman ehrfs K. Xj'e-Idsen I · Harry D'. Thier s I ii SOME SONOMA COUNTY HYPOGEOUS FUNGI A thesis by Russell Kent Heblack ABSTRACT Purpose of the study: None of the early mycologists who explored the Califor­ nia hypogeous fungal flora recorded the occurrence of these fungi in Sonoma County. This study sought to determine which species of hypogeous fungi are present in Sonoma County and with which woody plants they are associated. Procedure: Field research was done during the two years beginning in December 1975. The· method was to locate stands of woody plants and dig under them in search of hypogeous fungi. Once found, the fungi were identified and pre­ served. Notes were made regarding suspected mycorrhizal associates. Findings: Eighteen species of fungi were found. These fungi are representatives of two classes, Ascomycetes: Balsamia magnata, Genabea cerebriformis, Genea arenaria, Genea gardneri, Genea harknessii, Hydnobolites californicus, Hydnotrya ellipsospora, Tuber californicum, Tuber ~­ didum, and Basidiomycetes: Alpova fulvus, Alpova guerc­ icola, Gautieria graveolens, Gymnomyces ferruginascens, Hymenogaster parksii, Hysterangium se~arabile, Leuco­ gaster rubescens, ~elanogaster parksii, Melanogaster tuberiformis. Some collections, representatives of the genera Rhizopogon, Kartellia, llymenogaster, Balsamia, and Genea, were only identifiable to genera. These may represent undescribed species. Two Alpova species, Alpova fulvus Trappe & Heblack sp. nov. ined. and Alpova guercicola Trappe & Heblack sp. nov. ined., were new to science. All of the fungi found were new distribution records for Sonoma County, and some were found with new woody plant associates. Conclusions: This study has shown that the Sonoma County hypogeous fungal flora is diverse. Within this diverse flora, two iii new species were found and described. It is suspected that additional undescribed species will be discovered by fUture researchers. Chairperson: ..,- · ~ . ~ - ,, ,, Dr. tjose''H: · Powe11 M.A. Program: Biology Sonoma State College lV ACKNOWLEDGEMENTS I would like to give sincere thanks to my major professor, Dr. Joseph H. Powell, and my committee members, Drs. Chris K. Kjeldsen and Harry D. Thiers, for their unselfish effort throughout this project. I also wish to thank Mrs. Ellen Thiers for her help with the latin diagnoses, and Dr . .James M. Trappe for his help in identifying collections and the loan of herbarium specimens. R.K.H. v TABLE OF CONTENTS Page ABSTRACT ............................................. ii ACKNOWLEDGEMENTS ..................................... iv LIST OF FIGURES ...................................... vii LIST OF TABLES ....................................... viii INTRODUCTION ......................................... 1 LITERATURE REVIEW .................................... 2 Evolution 2 Basidiomycete Phylogeny ......................... 6 Ascomycete Phylogeny 8 Ascomycete Taxonomy ............................. 9 Basidiomycete Taxonomy 10 Habitat ......................................... 12 Reproduction .................................... 13 Nutrition ....................................... 15 Collection ...................................... 16 Spore Dispersal ................................. 17 Culturing Attempts .............................. 18 Reagents and Stains ............................. 19 Economic Use .................................... 20 METHODS AND MATERIALS ................................ 22 Collection ...................................... 22 Field Notes ............................. 22 Preservation ..................................... 23 vi Page Reagents and Stains ............................. 23 Microtechnique .................................. 25 RESULTS ............................................ 26 Class Basidiomycetes ............................ 26 Class Ascomycetes ............................... 27 Probable Mycorrhizal Associates ................. 27 Keys to the Species of Fungi .................... 29 New Species Descriptions ........................ JJ DISCUSSION ........................................... 37 Taxa Identification ............................. 37 Probable Mycorrhizal Associates 40 CONCLUSIONS .......................................... 4J SUMMARY e I e e e e e e e e e e e e e e e e e e a e e a e e e e a e e e e e e a e e e a e e e e e e 44 APPENDIX ......................................... 45 LITERATURE CITED ..................................... 47 vii LIST OF FIGURES Figure Page 1. Hydnotrya ellipsospora: spores x 14JO .•.......... 52 2. Genabea cerebriformis: spores x 1110 ............. 52 J. Genea arenaria: spores x 1180 .................... 53 4. Genea sp. (#43): spores x 1420 ................... 53 5. Genea gardneri: spores x 1320 .................... 54 6. Genea harknessii: spores x 1540 .................. 54 7. Balsamia magnata: spores x 1850 .................. 55 8. Balsamia sp. (#47): spores x 1350 . .. 55 9. Hydnobolites californicus: spores x 1550 ......... 56 10. Tuber californicum: spores x 1000 ................ 56 11. Tuber candidium: spores x 1000 ................... 57 12. Hysterangium separabile: spores x 1670 ........... 57 lJ. Gymnomyces ferruginascens: spores x 1570 .......•. 58 14. Martellia sp. (#99): spores x 1500 ............... 58 15. Leucogaster rubescens: spores x 1420 ............. 59 16. Gautieria graveolens: spores x 1680 .............. 59 17. Hymenogaster parksii: spores x 2100 .............. 60 18. Hymenogaster sp. (#90): spores x 1440 ....•....... 60 1?. Rhizopogon sp. (#38): spores x 1300 .............. 61 20. Alpova fulvus: spores x 1560 ......••............. 61 21. Alpova guercicola: spores x 1240 ................. 62 22. l\Ielanogaster tuberiformis: spores x 2000 ......... 62 23. Melanogaster parksii: spores x 3100 .............. 63 viii LIST OF TABLES Table Page 1. Probable mycorrhizal associates ..•..••........... 42 1 INTRODUCTION Truffles and false truffles, hypogeous members of Classes Ascomyceteae and Basidiomyceteae, are relatively little known fungi. Few mycologists collect them and even fewer are well acquainted with their taxonomy(Trappe, 1975). Early in this century there were several workers, centered around the S.F. Bay area, who were quite active in Alameda, Napa, Santa Clara, and Marin Counties. None of these workers (Harkness, 1899; Gilkey, 1916; Parks, 1921) recorded the occurrence of hypogeous fungi in Sonoma County, California. Though these workers reported the occurrence of many hypogeous fungi, they were often vague concerning the woody plant associates of the fungi. It is now known (Trappe, 1974} that most hypogeous rnacromycetes are probably rnycorrhizal. The purpose of this research was to discover which hypogeous fungi occur in Sonoma Co. and with which woody plants they are associated. The research consisted of sampling species of trees and shrubs for hypogeous macro­ mycete associates. This sampling was carried out during the two years beginning in December, 1975. This report demon­ strated the presence of hypogeous fungi in Sonoma Co., and the inventory will undoubtedly expand with further studies. Other workers are urged to continue the search in the county. 2 REVIEW OF THE LITERATURE Evolution There is widespread agreement among mycologists (Thiers, 1971; Heim, 1971) that discussions on the evo­ lution of fungi are speculative, without fossil evidence, and rest on logic, not fact. All of the arguments concerning the origin of hypo­ geous Basidiomycetes are elaborated by workers who are primarily agaricologists like Singer (1962) who are trying to unravel the phylogeny of the Agaricales. These workers often mention the evolution of hypogeous forms because these fungi frequently occupy the spotlight in the argument as to whether evolution has proceeded from agaricoid to gastromycetaceous forms or vice versa. The central argument is agreed upon (Smith, 1971; Singer, 1971; Heim, 1971) and it is best elaborated by Singer (1962): "The three logically possible, and actually inportant theories of today, two of them (2 and J) defended or favored by a group of systematists, are the following: 1. Derivation of the Agaricales from the Gastromycetes and from the Aphyllophorales. 2. Derivation of the Agaricales from the Gastromycetes alone. J. Derivation of the Agaricales from the Aphyllophor­ ales alone." 3 Which group the Agaricales were derived from does not con­ cern this paper; the importance of the above theories is that their elaboration by the three main antagonists, Heim (1948, 1971), Singer (1958, 1962, 1971), Smith (1971, 197J) causes some insight into the evolution of hypogeous forms. Singer and Smith (1960) become the allied proponents of the "Derivation of the Agaricales from the Gastromycetes alone" school, while Heim (1948, 1971) advocates the "Derivation of the Agaricales from the Aphyllophorales alone" theory. The argument centers around whether the Russulaceae evolved via "progressive evolution" (Singer and Smith, 1960) from such hypogeous genera as Gymnomyces and Martellia or vice versa. The arguments become irrational; Singer (1958) states: "Heim and others have drawn rash conclusions from incomplete facts and erroneously believed to have demon­ strated the correctness of their views whereas, in reality, they have only added - valid or questionable: this is a controversial question
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