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Taxonomy of Hawaiʻi Island's Lepiotaceous (Agaricaceae

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Taxonomy of Hawaiʻi Island's Lepiotaceous (Agaricaceae TAXONOMY OF HAWAIʻI ISLAND’S LEPIOTACEOUS (AGARICACEAE) FUNGI: CHLOROPHYLLUM, CYSTOLEPIOTA, LEPIOTA, LEUCOAGARICUS, LEUCOCOPRINUS A THESIS SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAIʻI AT HILO IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN TROPICAL CONSERVATION BIOLOGY AND ENVIRONMENTAL SCIENCE MAY 2019 By Jeffery K. Stallman Thesis Committee: Michael Shintaku, Chairperson Don Hemmes Nicole Hynson Keywords: Agaricaceae, Fungi, Hawaiʻi, Lepiota, Taxonomy Acknowledgements I would like to thank Brian Bushe, Dr. Dennis Desjardin, Dr. Timothy Duerler, Dr. Jesse Eiben, Lynx Gallagher, Dr. Pat Hart, Lukas Kambic, Dr. Matthew Knope, Dr. Devin Leopold, Dr. Rebecca Ostertag, Dr. Brian Perry, Melora Purell, Steve Starnes, and Anne Veillet, for procuring supplies, space, general microscopic and molecular laboratory assistance, and advice before and throughout this project. I would like to acknowledge CREST, the National Geographic Society, Puget Sound Mycological Society, Sonoma County Mycological Society, and the University of Hawaiʻi Hilo for funding. I would like to thank Roberto Abreu, Vincent Berrafato, Melissa Chaudoin, Topaz Collins, Kevin Dang, Erin Datlof, Sarah Ford, Qiyah Ghani, Sean Janeway, Justin Massingill, Joshua Lessard-Chaudoin, Kyle Odland, Donnie Stallman, Eunice Stallman, Sean Swift, Dawn Weigum, and Jeff Wood for helping collect specimens in the field. Thanks to Colleen Pryor and Julian Pryor for German language assistance, and Geneviève Blanchet for French language assistance. Thank you to Jon Rathbun for sharing photographs of lepiotaceous fungi from Hawaiʻi Island and reviewing the manuscript. Thank you to Dr. Claudio Angelini for sharing data on fungi from the Dominican Republic, Dr. Ulrich Mueller for sharing information on a fungus collected in Panama, Dr. Matthew Smith for sharing information on a fungus collected from Florida, and Dr. Teresa Lebel for sharing information on a mushroom from Australia. Special thanks to Dr. Else Vellinga on a variety of topics related to Lepiota taxonomy and reviewing species descriptions, and to my committee members Michael Shintaku and Nicole Hynson, and in particular Dr. Don Hemmes, whose knowledge of macrofungi on Hawaiʻi Island and choice to serve on my committee was necessary to begin, and complete this project. ii Abstract The fungi are a hyperdiverse group of important organisms, many of which remain undescribed and unknown, particularly from tropical habitats. This project documenting the lepiotaceous fungi (Agaricaceae: Lepiotae, Leucocoprinae) of Hawaiʻi Island resulted in 148 collections from September 2016–January 2019. Based on these collections and prior records, 38 species of lepiotaceous fungi from Hawaiʻi Island are reported: two in Chlorophyllum, one in Cystolepiota, seven in Lepiota, 15 in Leucoagaricus, and 13 Leucocoprinus. Two new records of previously undocumented species from Hawaiʻi, Leucocoprinus straminellus and Leucoagaricus cf. americanus, are reported, along with 21 additional species that could not be assigned to a known taxon. Two potentially undescribed Lepiota spp. likely contain poisonous amatoxins. General descriptions and a dichotomous key are provided for all taxa with sufficient information, and a phylogenetic analysis of the genus Lepiota, including the two potentially- deadly species based on nrITS sequences, is presented. iii Table of Contents Acknowledgements .................................................................................................................... ii Abstract ...................................................................................................................................... iii List of Tables .............................................................................................................................. vi List of Figures ........................................................................................................................... vii Introduction ................................................................................................................................. 1 Kingdom Fungi’s Importance and Diversity .............................................................................................. 1 Fungal Taxonomy and Systematics .......................................................................................................... 2 Taxonomy and Systematics of the Genus Lepiota and Relatives ............................................................ 3 Hawaiian Fungi, Including Lepiota and Relatives in Hawaiʻi ..................................................................... 5 Questions and Importance ......................................................................................................... 7 Methods ....................................................................................................................................... 8 Collection and Field Site Information ........................................................................................................ 8 Specimen Descriptions ............................................................................................................................. 9 DNA Extraction, Amplification, and Sequencing ..................................................................................... 11 Sequence Editing and Phylogenetic Analysis ......................................................................................... 12 Additional Considerations ....................................................................................................................... 13 Reagents and Chemicals ........................................................................................................................ 13 Results and Discussion ........................................................................................................... 15 Molecular and Phylogenetic Analysis...................................................................................................... 15 Meixner Test ............................................................................................................................................ 17 Research Questions, Species Totals, and Seasonality .......................................................................... 19 Key to the Lepiotaceous Fungi of Hawaiʻi Island .................................................................. 22 Key 1: Chlorophyllum, Cystolepiota, and lepiotaceous fungi outside the Agaricaceae .......................... 22 Key 2: Lepiota ......................................................................................................................................... 23 Key 3: Leucocoprinus and Leucoagaricus .............................................................................................. 24 iv Species Descriptions ................................................................................................................ 29 Chlorophyllum ......................................................................................................................................... 29 Cystolepiota ............................................................................................................................................. 37 Lepiota ..................................................................................................................................................... 41 Leucoagaricus ......................................................................................................................................... 71 Leucocoprinus ....................................................................................................................................... 122 Additional Taxa ....................................................................................................................... 168 Lepiotaceous taxa known from Hawaiʻi Island not collected during project .......................................... 168 Lepiotaceous taxa not occurring in Hawaiʻi, on Hawaiʻi Island, or lacking information ........................ 169 Lepiotaceous look-alikes outside the Agaricaceae ............................................................................... 174 Tables ....................................................................................................................................... 177 References ............................................................................................................................... 183 v List of Tables 1. Genera of lepiotaceous fungi reported from the Hawaiian Islands ....................................................... 177 2. Morphological overview of subsections of Lepiota sensu stricto .......................................................... 177 3. Lepiota sensu stricto clades by nrITS and nrLSU DNA analysis .......................................................... 178 4. Species of lepiotaceous fungi reported from the Hawaiian archipelago ............................................... 179 5. Sequences downloaded from GenBank for phylogenetic tree construction ......................................... 180 6. nrITS sequences generated for this project with GenBank accession numbers .................................. 181 7. nrITS pairwise genetic distance comparisons between Lepiota cf. elaiophylla and close relatives.. ... 182 vi List of Figures 1. Map of commonly visited field sites .........................................................................................................
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