A TAXONOMIC INVESTIGATION OF MYCENA IN CALIFORNIA A thesis submitted to the faculty of San Francisco State University In partial fulfillment of The requirements For the degree Master of Arts In Biology: Ecology and Systematic Biology by Brian Andrew Perry San Francisco, California November, 2002 Copyright by Brian Andrew Perry 2002 A Taxonomic Investigation of Mycena in California Mycena is a very large, cosmopolitan genus with members described from temperate and tropical regions of both the Northern and Southern Hemispheres. Although several monographic treatments of the genus have been published over the past 100 years, the genus remains largely undocumented for many regions worldwide. This study represents the first comprehensive taxonomic investigation of Mycena species found within California. The goal of the present research is to provide a resource for the identification of Mycena species within the state, and thereby serve as a basis for further investigation of taxonomic, evolutionary, and ecological relationships within the genus. Complete macro- and microscopic descriptions of the species occurring in California have been compiled based upon examination of fresh material and preserved herbarium collections. The present work recognizes a total of 61 Mycena species occurring within California, sixteen of which are new reports, and 3 of which represent previously undescribed taxa. I certify that this abstract is a correct representation of the content of this thesis. Dr. Dennis E. Desjardin (Chair, Thesis Committee) Date ACKNOWLEDGEMENTS I am deeply indebted to Dr. Dennis E. Desjardin for the role he has played as a teacher, mentor, advisor, and friend during my time at SFSU and beyond. Dr. Desjardin first introduced me to the amazing world of fungi and instilled in me the desire to become a mycologist, for which I will always be grateful. I would like to thank Dr. Desjardin for encouraging me to tackle the Mycena species of California, as well as the tireless hours he spent with me in the lab and field teaching me about the Agaricales and other fungi. I would also like to thank Dr. Desjardin for the immense amount of patience he displayed during the seemingly endless preparation of this manuscript. I am grateful to Dr. Robert Patterson for his role as a committee member and graduate advisor, as well as teaching me about the vascular plant flora of California. I am also grateful Dr. Greg Spicer for being a committee member, teaching me about phylogenetic systematics, and encouraging me to pursue a PhD. I would also like to thank Dr. Scott Redhead for his correspondence regarding the taxonomy of Mycena species. I would like to thank my labmates and fellow graduate students for making the Biology Department at SFSU such a great place to study biology. In particular I would like to thank Kristin Peterson, Kelly Collins, J.R. Blair, Atik Retnowati, Chuck Bell, Robyn Battaglia, Joshua McDill, and Laura Boykin for their help, encouragement, and friendship. I would also like to thank Michael Wood and Fred Stevens for their continued interest in my thesis project, their companionship on numerous forays, and the use of their beautiful slides of Mycena species. During the course of this study material was borrowed from several institutions, and I would like to thank the following herbaria directors for their assistance: Dr. Robert Fogel of the University of Michigan; Dr. David Largent of Humboldt State University, California; Dr. Donald. H. Pfister of the Farlow Herbarium at Harvard University; and Dr. Scott Redhead of the National Mycology Herbarium, Canada. This project would not have been possible without the collections these herbaria provided. Much of the research presented in this manuscript was made possible by financial assistance from the Mycological Society of San Francisco and the Mycological Society of America. I am very grateful for the assistance these groups provided, and would like to thank them for making such funding opportunities available to students. Finally, I would like thank my wife, Eileen Joy Macapinlac, for all of the support, patience, encouragement, and companionship she continues to provide me with during the course of this and other studies. I would also like to thank my parents, Beverly J. Perry and Richard Perry, for always encouraging me to follow where my interests led me (however strange they may have seemed!). It is to these three people I would like to dedicate the following monograph. iv TABLE OF CONTENTS Introduction..................................................... 1 History and Nomenclature ......................................... 2 History of the Genus in North America ................................ 5 Methods and Materials ............................................ 7 Taxonomically Informative Characters Macromorphological characters................................ 9 Micromorphological characters ............................... 14 Key to California Sections of Mycena ................................ 18 Taxonomic treatment of the sections of Mycena section Oregonenses ....................................... 23 section Fragilipedes ........................................ 24 section Luculentae ......................................... 50 section Rubromarginatae.................................... 56 section Sanguinolentae ..................................... 67 section Galactopoda ....................................... 71 section Longisetae ......................................... 74 section Heimales .......................................... 75 section Aciculae ........................................... 80 section Adonidae .......................................... 82 section Lactipedes ......................................... 87 section Hygrocyboideae..................................... 89 section Fuliginellae ........................................ 96 section Basipedes ........................................ 100 section Amictae .......................................... 103 section Insignes .......................................... 105 section Supinae .......................................... 107 section Polyadelphia ...................................... 109 section Sacchariferae ..................................... 111 section Mycena .......................................... 115 section Calodontes ....................................... 125 section Testudini ......................................... 128 section Intermediae ....................................... 130 section Cinerellae ........................................ 132 section Filipedes ......................................... 141 Literature Cited ................................................ 151 Species Index ................................................. 157 v INTRODUCTION The recent increase in issues of biodiversity, especially those pertaining to the management and conservation of natural resources, underscores the need for baseline knowledge of the organisms functioning in our ecosystems. Unlike mammals, birds, and plants, which appear to be fairly well documented for much of the Northern Hemisphere, groups such as fungi remain relatively unknown for many regions due to their ephemeral nature, and the numerous difficulties that have traditionally been associated with their identification. One such difficulty is the lack of accessible, up-to-date monographs including keys to species identification and detailed species descriptions. Commonly, when monographs do exist for a given group or genus, they tend to cover a very large geographic area and therefore include a high number of species, often making it difficult to identify a given collection to a single species with any degree of certainty. Fortunately, recent investigations have been published which document the diversity of distinct groups and genera of Agaricales within the restricted range of California (Thiers, 1997; Shanks, 1997; Methven, 1997; Desjardin, 1987; Kerrigan, 1986). However, many more genera are relatively poorly known taxonomically, and remain to be investigated. The genus Mycena (Pers. Fr.) Roussel, which is a member of the large family Tricholomataceae, remains largely undocumented in California, as well as many other regions of the United States. The genus is characterized by a white spore print; smooth, hyaline spores; adnate to arcuate or decurrent lamellae; a thin membranous pileus; intricate cystidia; and a hypodermium composed of large, inflated hyphae. Members of the genus are almost exclusively saprotrophic, the exception being a single species [M. citricolor (Ber. & Curt.) Sacc.] reported to be pathogenic on Coffea (Singer, 1986), and two species reported to form mycorrhizal associations with distinct species of the Orchidaceae (Guo et al., 1997). In the field, the basidiomes of Mycena species are commonly encountered growing upon the debris of conifers and woody angiosperms, decaying logs and branches, lignicolous on the bark of living trees, in soil, and more rarely (at least in California) on the debris of ferns, grasses, or other herbaceous plants, or among mosses or grasses. The most recently published material relating to the genus within California is limited to Dr. Alexander Smith’s monograph on North American species of Mycena, published in 1947, and R. A. Maas Geesteranus’ (1992) investigation of Mycena species described from the northern hemisphere. However, both of these works pertain to a very large geographic range, and neither attempt to present a complete survey of the Mycena species found