City University of New York (CUNY) CUNY Academic Works All Dissertations, Theses, and Capstone Projects Dissertations, Theses, and Capstone Projects 5-2018 A Contribution Toward a Global Monograph of Gyroporus: Taxonomy, Phylogeny, Biogeography Naveed Davoodian The Graduate Center, City University of New York How does access to this work benefit ou?y Let us know! More information about this work at: https://academicworks.cuny.edu/gc_etds/2597 Discover additional works at: https://academicworks.cuny.edu This work is made publicly available by the City University of New York (CUNY). Contact: [email protected] A Contribution Toward a Global Monograph of Gyroporus: Taxonomy, Phylogeny, Biogeography by Naveed Davoodian A dissertation submitted to the Graduate Faculty in Biology in partial fulfillment of the requirements for the degree of Doctor of Philosophy, The City University of New York 2018 © 2018 NAVEED DAVOODIAN All Rights Reserved ii A Contribution Toward a Global Monograph of Gyroporus: Taxonomy, Phylogeny, Biogeography by Naveed Davoodian This manuscript has been read and accepted for the Graduate Faculty in Biology in satisfaction of the dissertation requirement for the degree of Doctor of Philosophy. _________________ ____________________________________ Date Roy E. Halling Chair of Examining Committee _________________ ____________________________________ Date Cathy Savage-Dunn Executive Officer Supervisory Committee: Joseph W. Rachlin James C. Lendemer Amy C. Berkov Edward J. Kennelly Sarah E. Bergemann THE CITY UNIVERSITY OF NEW YORK iii ABSTRACT A Contribution Toward a Global Monograph of Gyroporus: Taxonomy, Phylogeny, Biogeography by Naveed Davoodian Advisor: Roy E. Halling Gyroporus (Sclerodermatineae, Boletales, Agaricomycetes, Basidiomycota, Fungi) is a genus of ectomycorrhizal mushroom-forming fungi distributed throughout the world in suitable habitats. Previous attempts to untangle the diversity of this genus proved difficult due to the presence of semi-cryptic species and equivocal results from phylogenetic analysis of ribosomal RNA markers. To overcome these obstacles, a combined taxonomic and phylogenetic (emphasizing protein-coding genes) approach is used here to delimit species and elucidate geographic and evolutionary patterns of Gyroporus. Careful study of relevant literature and herbarium specimens was augmented by field work in North America, Australia, and East Asia for observation and collection of fresh material. For phylogenetic analyses, the mitochondrial gene atp6 and the nuclear gene rpb2 were utilized. The nuclear gene tef1 was also utilized for select exemplars. Several distinct clades are inferred, most notably a clade corresponding to Gyroporus castaneus as a speciose northern hemisphere group (the castaneus clade), a clade unifying Gyroporus cyanescens and similar entities (the cyanescens clade), and a clade unifying Gyroporus longicystidiatus and a New World sister species (the longicystidiatus clade). Also highly notable iv is the recovery of a sister relationship between the cyanescens and longicystidiatus clades. Abundant sister relationships between Eurasian and North American (including Central American) Gyropori are apparent and suggestive of boreotropical biogeographic scenarios; the Australian Gyropori display various biogeographic affinities. Eighteen new Gyroporus taxa are outlined and several biogeographic hypotheses are discussed. This study provides key groundwork for future efforts on this well-known but poorly-understood group of fungi. v For Casie Keyon Mom & Dad and Mark Reed (1962–2017) vi ACKNOWLEDGEMENTS I am deeply indebted to my advisor, Roy Halling, for his guidance and support over the years, and to Barbara Thiers, whose perspectives and assistance greatly enhanced my graduate school experience. I sincerely thank Larry Kelly for his tireless advocacy on behalf of the NYBG graduate students, and Joan Reid for her assistance to the CUNY Biology PhD students. Joe Rachlin and Dwight Kincaid are thanked for the support and exchange of ideas they facilitated since the start of my graduate education. Tim Baroni is greatly thanked for his committee service and specimen loans in support of my dissertation research. I truly appreciate the participation of Amy Berkov and Ed Kennelly in this effort. I am thankful for the participation of James Lendemer at various stages of my graduate education, especially for providing important fieldwork opportunities. I am extremely grateful for the training and guidance I received from Sarah Bergemann, especially in the earliest stages of my research. The Department of Biology at Middle Tennessee State University was extraordinarily accommodating, and I am grateful to the faculty, staff, and students at MTSU. Kentaro Hosaka, Olivier Raspé, and Matt Smith are thanked for their major supportive roles in this project. Visits by myself and my advisor to the National Museum of Nature and Science (Japan), Botanic Garden Meise, and the University of Florida, and the generous sharing of specimens and data were instrumental to this project. This project would not have been possible without help from Hosaka, Raspé, Smith, their institutions, and their staff and students. I am indebted to Nishi Rajakaruna, whose guidance and mentorship during college brought me to this point. I am also indebted to Tom Volk, who encouraged my interest in fungi and facilitated a summer research experience for me in Wisconsin when I was in college. I am vii very grateful to David Porter and Marc Cubeta, who provided feedback and advice at various stages of my education. Ellen Bloch and Laura Briscoe are thanked for their invaluable assistance in the herbarium. Bart Buyck, Terry Henkel, Beatriz Ortiz-Santana, Terrence Delaney, Joe Ammirati, Barrie Overton, Alan Franck, Igor Safonov, Rod Tulloss, Walt Sturgeon, Christian Schwarz, Richard Robinson, and Neale Bougher are greatly thanked for providing specimens, especially specimens that would otherwise have not been possible to obtain. Don Pfister and the staff at the Farlow Herbarium are sincerely thanked for facilitating the study of critical specimens housed at FH. The staff members at the University of Michigan Herbarium are thanked for providing access to key specimens housed at MICH. Zev Reuter, Annie Christian, and Caltha are thanked for sharing space, food, and collecting spots in the Hudson Valley. T. Triggs and M. Aristova are thanked for sharing space and resources in New England and the Midwest; M. Aristova is greatly thanked for providing translations from Russian. My sincere gratitude to Ben Stewart for providing lodging during a visit to Michigan. Karim Ahmed and Pete Pavicevic are thanked for their guidance in navigating NYC and environs. Many thanks to T. McLeod and C. Montgomery for their generosity and kindness when I was in North Carolina. There are countless people who contributed positively to this project and my intellectual development over the years. They cannot all be enumerated here; they nonetheless were crucial to reaching this point. viii Table of Contents Chapter 1: Background and Methodology…………………………………………………...…...1 Introduction…………………………………………………………………………...…..1 Methodological Approach……………………………………………………………...…8 Specimens Examined…………………………………………………………………….13 References………………………………………………………………………………..29 Chapter 2: Taxonomic Treatment………....……………………………………………………..33 Taxonomic Treatment………....………………………………………………………....33 A Selection of New Species (including Segregates)…………………………………….59 Excluded Taxa and Names………………………………...…………………………….77 References……………………………………………………………………………….83 Chapter 3: Phylogenetics and Biogeography…………………………………………………….85 Analysis of Atp6 and Rpb2 Sequences in MrBayes……………………………………...85 Discussion………………………………………………………………………………..85 Parsimony Analysis of Rpb2 and Atp6 Data in NONA, with Additional Discussion…...92 Biogeography and Further Discussion…………………………………………………..93 Conclusions and Future Directions……………………………………………………..111 References………………………………………………………………………………113 Addendum………………………………………………….………………………..………….118 ix Figure 1-1. Gyroporus castaneus iconotype…………………………….…………………...…...4 Figure 1-2. Gyroporus cyanescens iconotype…………………………...…………………...…...5 Figure 1-3. 28s phylogram…………………………...……………………………………….......7 Table 1-1. Gyroporus specimens………………………………………………………………...13 Figure 2-1. Specimen ND 12…………………………………………………………………….41 Figure 2-2. Specimen KH-JPN15-733………………………………………………………...…41 Figure 2-3. Cheilocystidia, G. longicystidiatus holotype………………………………………..45 Figure 2-4. G. longicystidiatus holotype pileus…………………………………………...……..46 Figure 2-5. G. longicystidiatus holotype hymenophore……………………….………...…….....47 Figure 2-6. KH-JPN15-569………………………………………………………………….…...48 Figure 2-7. KH-JPN15-569 close...……………………………………………………....……...49 Figure 2-8. Gyroporus australiensis……………………………………………………………..61 Figure 2-9. Gyroporus mcnabbii…………………………………………………………………66 Figure 2-10. REH 8821……………………………………………………………………….….72 Figure 2-11. REH 8821 (2nd photo).………………...……………………………………….….72 Figure 2-12. Specimen ND 13 view of hymenophore…………………………………………...74 Figure 2-13. Specimen ND 13 cap in cross-section…………………………………..………….75 Figure 2-14. ND 57……………………………………………………………………………....75 Figure 2-15. KH-TBG12-712…………………………………………………………………....76 Figure 3-1. Atp6 phylogram……………………………………………………………………...86 x Figure 3-2. Rpb2 phylogram……………………………………………………………………..88 Figure 3-3. Pileipellis types……………………………………………………………………...91 Figure 3-4. Rpb2 parsimonious dendrogram…..………………………………………………...94