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Redacted for Privacy AN ABSTRACT OF THE DISSERTATIONOF Kentaro Hosaka for the degree of Doctor ofPhilosophy in Botany and Plant Pathology presented on October 26, 2005. Title: Systematics, Phylogeny, andBiogeography of the Hysterangiales and Related Taxa (Phallomycetidae, Homobasidiomycetes). Abstract approved: Redacted for Privacy Monophyly of the gomphoid-phalloid dadewas confirmed based on multigene phylogenetic analyses. Four major subclades(Hysterangiales, Geastrales, Gomphales and Phallales) were also demonstratedto be monophyletic. The interrelationships among the subclades were, however, not resolved, andalternative topologies could not be rejected statistically. Nonetheless,most analyses showed that the Hysterangiales and Phallales do not forma monophyletic group, which is in contrast to traditional taxonomy. The higher-level phylogeny of thegomphoid-phalloid fungi tends to suggest that the Gomphales form a sister group with either the Hysterangialesor Phallales. Unweighted parsimonycharacter state reconstruction favorsthe independent gain of the ballistosporic mechanism in the Gomphales, but the alternativescenario of multiple losses of ballistospoiy could not be rejected statistically underlikelihood- based reconstructions. This latterhypothesis is consistent with thewidely accepted hypothesis that the loss of ballistosporyis irreversible. The transformationof fruiting body forms from nongastroid to gastroidwas apparent in the lineage leading to Gautieria (Gomphales), but thetree topology and character statereconstructions supported that truffle-like taxa of thePhallales are ancestral to stinkhorns,which possess more complex, epigeous fruiting bodies. Importantlyall taxa within the Phallales are statismosporic and thusthe derived stinkhorn morphology doesnot require an independent gain of ballistospory. Biogeographical analyses of the Hysterangialesstrongly suggest that the ectomycorrhizal lineages within the Hysterangialesoriginated in the East Gondwana. The synonymous substitutionrate indicated a Paleozoic origin of theHysterangiales although a possibility ofa Cretaceous origin could not be discarded. Becausemodem ectomycorrhizal plantswere absent during the Paleozoic era, a potential existenceof the Hysterangiales during this timemust be explained either by novelectomycorrhizal association of the Hysterangiales withunknown plant lineages,or multiple, independent gains of ectomycorrhizalhabit. The Paleozoic origin of the Hysterangiales also indicates that mycophagousanimals may not be the most important factor forrange expansions of the Hysterangiales. Taxonomic revisionsare made for the gomphoid-phalloid fungi. Onesubclass (Phallomycetidae), two orders (Hysterangialesand Geastrales), four families (Gallaceaceae, Phallogastraceae,Trappeaceae and Sclerogastraceae),7 genera (Austrohysterangium, Cribbangium,Rodwayomyces, Beeveromyces,Cazomyces, Insulomyces and Viridigautieria) and 22new combinations are proposed. ©Copyright by Kentaro Hosaka October 26, 2005 All Rights Reserved Systematics, Phylogeny, and Biogeographyof the Hysterangiales and RelatedTaxa (Phallomycetidae, Homobasidiomycetes) by Kentaro Hosaka A DISSERTATION submitted to Oregon State University in partial fulfillment of the requirement for the degree of Doctor of Philosophy Presented October 26, 2005 Commencement June 2006 Doctor of Philosophy dissertation ofKentaro Hosaka presentedon October 26, 2005. APPROVED: Redacted for Privacy Professor, repesenting Botany and PlantPathology Redacted for Privacy Chair of the Departn*nt of Botanyand Plant Pathology Redacted for Privacy Dean of tM raduate School I understand that my dissertation will become part of the permanentcollection of Oregon State University libraries. Mysignature below authorizes release ofmy dissertation to any readerupon request. Redacted for privacy Kentaro Hosaka, Author ACKNOWLEDGEMENTS Many individuals and organizations have contributedto the completion of this dissertation. This dissertation would not exist without theirhelp. I am grateful tomy committee members; Drs. Joseph Spatafora, MichaelCastellano, Aaron Liston, Jeffrey Stone, Jennifer Parke and James Trappe. I would liketo express my sincere thanks to my major advisor, Joey Spatafora, for welcoming me to his lab and the worldof fungal phylogenetics. His encouraging attitude alwaysmotivated me. Joey also introduced me to the world of Mardi Gras, which I willnever forget. Dr. Mike Castellano has been a great advisor from the beginning ofmy project. His expertise on sequestrate fungi was truly invaluable. Special thanks to the Spatafora lab members, whohave always been supportive mentally, and sometimes financially. The lab hasa great ethnic diversity from which I have learned a lot of things, especially the variety ofcuisine we shared at potlucks. Thanks to Andrea Humpert, Eric Muench, Drs. AnnetteKretzer, Jamie Platt and Rekha Meyer, for a warm welcome when I first arrivedin the U.S. to join the lab. Special thanks to (soon to be Dr.) Gi-Ho Sung fora great friendship even though all Beaver's football games we watched together resultedin a huge disappointment. Also thanks to Gi-Ho's wife, Chang-Ae Song, for providingme authentic kimchee numerous times. Thanks to Drs. Conrad Schoch and Barbara Robbertse for introducing me to the world of "Settlers". Thanksto Drs. Admir Giachini and Eduardo Nouhra for our friendship and discussionon the gomphoid-phalloid fungi. Kris Whitbeck was a great teacher and taughtme how to identify trees in Oregon. Thanks to Dr. Susie Dunham for sharing my interest in fungal ecology, and arranginga vacation house in Hawaii during the annual meeting of MycologicalSociety of America in 2005. Thanks to Desiree Johnson and Mike Crockettfor sharing the excitement for the white Suillus. Jason Barker and Tn Tran helpedme with data collection. Special thanks to the Forest Mycology including Dan Luoma,Joyce Eberhart, Tom Horton, Randy Molina, Efren Cázares, Jane Smith, Nancy Weber,Doni McKay, Bruce Caidwell, Charles Lefevre, David Pilz, Matt Trappe, and KellyCollins, who supported and encouraged me in one wayor another. Thanks to Dr. Wes Colgan, Ray Kenny and Nich Simpson for sharingsome data from the Mesophelliaceae and stinkhoms. Numerous important specimens have been provided frommany parts of the world. Hiromi Sasaki, Youko Andoh, Tetsuo Muroi, Drs.Kazumasa Yokoyama and Akiyoshi Yamada kindly provided me several undescribed truffle-likefungi from Japan. Thanks to Dr. Steve Miller for Guyanan Hysterangium, Dr. LisaGrubisha for Hysterangium from the Channel Islands, Dr. Jean-Mark Moncalvo forThailand Protubera, and Dr. Manfred Binder for Chinese stinkhonns. Specialthanks to Dr. David Hibbett for providing me important literatureon Geastrum ectomycorrhizae. Thanks to Dr. Wilford Hess for providingme a reprint on the SEM study of the gasteromycetes spores. Thanks to Dr. Hanns Kreisel for givingme insightful ideas on the Geastrales taxonomy. Dr. Michael Weiss kindly providedme a Sebacinales dataset. Thanks to Mark Dasenko, the Center for Gene Research and Biotechnology, for processing numerous sequencing plates ina timely manner. Thanks to Chris Sullivan for providing me very helpful computer assistance and answeringmany ridiculous questions. Special thanks to Dr. Daniel Barker for developinga newer version of the Bayesian-Multistate script. One of the biggest events during my Ph.Dprogram was a research trip to the Southern Hemisphere. Financial support for this tripcame from the National Science Foundation, Doctoral Dissertation Improvement Grant (DEB0309412).Many individuals made this trip very successful. Special thanksgo to Edith Birky for processing the travel budget before the trip, and infinite numbers of receiptsafter the trip. Also thanks to Wendy Woolsey, Teel's Travel Planners, for herattempt to get the cheapest options for my travel itinerary. Drs. Greg Mueller, RytasVilgalys, Orson Miller and Darlene Judd provided me useful tips in Papua New Guinea. I deeply thank to Dr. Roland Treu, University of Goroka, for everythingthat happened in Papua New Guinea. From the first day in PapuaNew Guinea, I could not have done anything without him. Several people at the PapuaNew Guinea government including Barbara Roy, Roy Banka, James Robins and CepasKayo went through my research visa application. Thanks to Yoshika Sawairi, PapuaNew Guinea Embassy in Japan, for dealing with the visa application in the finalmoment. Thanks to Dr. Debra Wright for arranging the local travel. Special thanksto the local people in the village of Maimafu for their warm welcome. Charles Pepe guidedand body guarded me through the woods with a huge bush knife. Thanks to Dr. Tanguy Jaffré, Institute de Reserchepour le Devéloppement, for providing me the exact location on Nothofagus stands inNew Caledonia. He also showed me many Nothofagus specimensso that I could identify Nothofagus species more easily in the field. Joseph Manuete and David Kurpisz at Direction des Ressources Naturelles de la Province Sud issuedme a collecting permit in a timely manner. Special thanks to the park rangers at Parc Provincial de la Riviere Bleuefor allowing me to excavate truffles in the park,even when the park was closed for holidays. In New Zealand, Drs. Ross and Jessica Beevergave me a warm welcome and provided me a room to stay. Ross kindly guidedme to virtually every place in New Zealand, including the Nothofagus stands and hot springs.Thanks to Nick Singer, Department of Conservation, for issuinga collecting pennit. Thanks to Paula Wilkie for arranging my registration
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