Alan E. Bessette, William C. Roody, and Arleen R

Boletes of Eastern North America

alan e. beSSette, william c. roody, and arleen r. beSSette

SyracuSe univerSity preSS NOTICE: Although this book includes information regarding the edibility of the mushrooms described, it is not intended to function as a manual for the identification and safe consumption of wild mushrooms. Readers interested in consuming wild fungi should consult other sources of information, including experienced mycologists and literary works, before eating any wild mushrooms. The authors and the publisher are not responsible for any undesirable outcomes that may occur for those who fail to read or heed this warning.

First Edition 2016

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∞ The paper used in this publication meets the minimum requirements of the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ANSI Z39.48-1992.

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ISBN: 978-0-8156-3482-9 (hardcover) 978-0-8156-1074-8 (paperback) 978-0-8156-5394-3 (e-book) library of congress cataloging-in-publication data Names: Bessette, Alan, author. | Roody, William C., author. | Bessette, Arleen Rainis, 1951– author. Title: Boletes of eastern North America / Alan E. Bessette, William C. Roody, and Arleen R. Bessette. Description: First edition. | Syracuse, New York : Syracuse University Press, [2016] | Includes bibliographical references and index. Identifiers: LCCN 2016038515 (print) | LCCN 2016039651 (ebook) | ISBN 9780815634829 (hardcover : alk. paper) | ISBN 9780815610748 (pbk. : alk. paper) | ISBN 9780815653943 (e-book) Subjects: LCSH: Boletales—North America—Identification. Classification: LCC QK629.B63 B47 2016 (print) | LCC QK629.B63 (ebook) | DDC 579.6/1632—dc23 LC record available at https://lccn.loc.gov/2016038515

Manufactured in the United States of America pp. ii–iii: Harrya chromapes p. xvi: Frostiella russellii facing page 1: Tylopilus felleus p. 12: Neoboletus luridiformis p. 22: Harrya chromapes p. 432: Hortiboletus rubellus p. 439: Tylopilus plumbeoviolaceus p. 440: Retiboletus ornatipes, gray form p. 456: Baorangia bicolor complex Generously• supported by a gift in honor of

Kevin William Pfeiffer •

Dr. Roy E. Halling

We have been fortunate to be able to come to know and collaborate with many outstanding persons while working in the field of mycology. Dr. Roy E. Halling is one of those very special people. We dedicate this book to him with gratitude for his lifelong work and outstanding contributions to the field of mycology, especially in the area of boletology. Roy’s study and passion for mycology had a humble start with trips he took into the mountains of southern California as a Boy Scout. His first love was rock climbing, and through it he developed an interest in the flora of the region. Intending to major in forestry, he enrolled at California State University at Stan- islaus, where he took a mycology course offered by Dr. David Gotelli. Roy began collecting macrofungi, taught himself the basics of mycology, and subsequently developed a specific interest in boletes. He continued his study of the Boletaceae with the eminent mycologist Dr. Harry Thiers at San Francisco State University, viii completing his master’s degree in 1976 with a thesis titled “The Boletaceae of the Sierra Nevada.” Roy received his doctorate degree in 1980 from the Uni- versity of Massachusetts, Amherst, supervised by Dr. Howard Bigelow. He then accepted a two-year postdoctoral position at Harvard University in the Farlow Herbarium. In 1983, he was offered and accepted the position of assistant curator of mycology at the New York Botanical Garden, where he currently holds the position of research mycologist and curator of mycology. While in New York, Roy began to shift his focus to the macrofungi of South America. He obtained a National Science Foundation grant to do a survey of the Collybia in South America. This survey began a fifteen-year collaboration with Dr. Greg Mueller to document the macrofungi of the oak forests in Costa Rica. Also in the 1980s, an opportunity to collect in Australia introduced Roy to the virtually unknown diversity of the boletes that existed there. He encoun- tered boletes with combinations of features that transcended generic concepts as they were known in the Northern Hemisphere and thus began his fascination with the boletes of Australasia that continues today. Since then, Roy has worked intensively on the taxonomy and phylogeny of the fungi of Fraser Island and has collected boletes all over Australia as well as in New Zealand, New Guinea, Indonesia, Malaysia, and Thailand. In addition, Roy serves as adjunct professor of biology at City University of New York Graduate School and as adjunct senior research scientist with the Department of Ecology, Evolution, and Environmental Biology at Columbia University. He is actively involved in international collaboration with other spe- cialists on the systematics, biogeography, and phylogeny of boletes, with partic- ular emphasis in Australia and Southeast Asia. A prolific writer, Roy has published more than one hundred professional papers and more than fifteen abstracts. He has served as adviser for several doc- toral candidates in the field of mycology; has held numerous editorial positions, including that of managing editor of Mycologia; and is a member of the Myco- logical Society of America (MSA) and several other prestigious mycological societies and associations. Roy also contributes his time, energy, and expertise to countless public services, from serving as chairman of the Nomenclature Committee of MSA to being on call with the Poison Control Center & Plant Information Service, appearing on National Public Radio’s program Science Friday, and routinely offering identification assistance and advice to the public at large. Roy is a past president of MSA and in 2006 was bestowed the honor of MSA fellow. There is so much to say about Roy and not enough space to do the telling justice. He is a gentle, kind man with a wry and offbeat sense of humor. When he speaks, there is a smile in his voice. When asked for help, advice, an opinion, or a thought, he unfailingly gives one his full attention and consideration. Roy’s ix character and his love of mycology, the joy he takes in the theoretical and practi- cal understanding of it, and the absolute lack of any personal ownership of it can best be described in his own words: “If there’s anything I can do to help, just let me know. . . . I am happy to share.” Thank you, Roy, for sharing your life’s work with so many.

Contents

Preface xiii Acknowledgments xvii

Introduction 1 The Bolete Fruitbody: Macroscopic Features 3 Information about Bolete Identification 7 How to Use This Book 13 Explanation and Use of the Field Keys 15 Field Keys to the Boletes 23 Descriptions and Illustrations of Bolete Genera and Species Alessioporus 55 Aureoboletus 57 Austroboletus 68 Baorangia 72 Boletellus 76 Boletinellus 81 Boletus 83 Bothia 171 Buchwaldoboletus 173 Butyriboletus 180 Caloboletus 185 Chalciporus 193 Cyanoboletus 200 Exsudoporus 202 Frostiella 206 Gyroporus 208 Harrya 215 Heimioporus 217 Hemileccinum 220 Hortiboletus 221 Imleria 226 xii · Contents

Lanmaoa 228 Leccinellum 236 Leccinum 245 Neoboletus 275 Paragyrodon 280 Phylloporus 283 Pseudoboletus 287 Pulveroboletus 288 Retiboletus 296 Rubroboletus 301 Strobilomyces 305 Suillellus 309 Suillus 316 Sutorius 362 Tylopilus 364 Veloporphyrellus 405 Xanthoconium 407 Xerocomellus 414 Xerocomus 420 Undescribed Bolete Species Pending Publication 433

Appendixes A. Chemical Reagents and Bolete Identification 441 B. Microscopic Examination of Boletes 443 C. Collecting, Cooking, and Preserving Boletes 445 D. Common Tree Associates of Eastern Boletes 448 Glossary 449 References and Recommended Resources 457 Index to Common Names 461 Index to Scientific Names 463 Photo Credits 471 Preface

Because boletes are fairly large and colorful, and especially because many of them are excellent edibles, they are one of the most popular groups sought after by mushroom hunters. They are also important ecologically because of the mycor- rhizal relationships they have with many different tree species. Our goal in writ- ing this book was to provide a means to identify the majority of boletes found in eastern North America within the current context of accepted taxonomy. This book is not just a “redo” or update of North American Boletes (Bessette, Roody, and Bessette 2000). In the sixteen years since that book was published, many new American bolete species have been discovered, and major changes in bolete taxonomy have taken place. In our ongoing study and appreciation of boletes, we have learned much in recent years. This book includes these changes, additions, and corrections and presents them in a less-formal manner. We have restricted the species featured to those that are known to occur throughout eastern North America, extending west to Louisiana and north to Manitoba, Canada, provid- ing more-focused and less-burdensome geographic coverage. A large number of new photographs, new information, and restructured field keys contribute to the ease of use for enthusiasts at all levels of expertise. With the ongoing process of molecular studies, construction of phylogenetic trees, and analysis of new data, many of the species currently placed in the genera we have listed will be transferred in the future to new or different genera. This is especially true for species currently in the genus Boletus. The process of erecting new genera to accommodate species in this very large genus and to some extent in other bolete genera has created a difficult issue for taxonomists, who must decide which genus name should be accepted. Some of the recently proposed genera are based only on macroscopic features, whereas others are based on microscopic characteristics or molecular analysis and the position of species in phylogenetic trees. As more and more DNA analysis of bolete species becomes available, it is uncertain which of the newly proposed genera will be changed and which will be retained. It is, however, quite likely that recently proposed new genera based only on macroscopic features will most likely be reassigned following DNA analysis. For example, Alfredo Vizzini, Giampaolo Simonini, and Matteo Gelardi (2014) erected the genus Exsudoporus to accommodate Boletus floridanus and Boletus frostii, two species that exude golden-yellow or amber-yellow droplets xiii xiv · Preface when young. The resulting new combinations are Exudoporus floridanus and Exudoporus frostii. A second example further describes the issue of retaining newly proposed genera based on macroscopic features. Alan Bessette and Orson K. Miller (2000) published Boletus rainisii. Gelardi, Vizzini, and Simonini (2014) erected the genus Cyanoboletus to accommodate several boletes with tis- sues that instantly discolor dark indigo blue when handled or bruised. One of the new combinations proposed was Cyanoboletus rainisii. In a subsequent publication, Noah Siegel, Christian F. Schwarz, and Jonathan L. Frank (2014) using DNA analysis, placed Boletus rainisii in the genus Xerocomellus and published the new combination Xerocomellus rainisiae. Recently proposed name changes based only on macroscopic features are briefly mentioned in the overview section of each description when necessary. By the time this book is published, it is quite likely that many more of these changes will have occurred. Although many scientific names have changed and more will be changed, the mushrooms themselves remain the same. No matter what name is used, the reader can always refer to the synonyms provided with each description or check other resources, such as Index Fungorum, to search for additional synonyms or to check for more recent name changes. To access Index Fungorum, use the link www.indexfungorum.org. Select “Search Index Fungorum” in the upper-right portion of the page. Make certain “name” is selected under “Search By” on the top left side of the new page. Enter the name of the genus and species you wish to research in the field provided under “Enter a search term.” Hitting “Search” will bring you to a screen that displays the genus and species name in blue type, the authority(ies) and the year it was published in black and in parentheses, respectively, and the currently accepted name in green. Clicking on the current name provided allows you to search Species Fungorum, which provides the current name, synonyms, authorities, and the year and publication site(s). Some previously accepted genera have been eliminated, and all of the species in those genera have been reassigned. Examples include Boletinus, Fuscoboleti- nus, and Gyrodon. In this work, species descriptions are preceded by genus descriptions. In most instances, the genus descriptions list the key macroscopic features on which the genus is based. Examples include Suillus and Tylopilus. In other instances, the genus is based on molecular or chemical analysis, and the number of macroscopic features listed may be significantly reduced. Examples include Imleria, Lanmaoa, and Retiboletus. Whenever possible, previously unpublished color images of the described species have been included. The known geographical distribution ranges for many of the species described have been expanded. We have modified species descriptions to include our personal observations of macroscopic features, mac- rochemical testing reactions, habitat information, edibility, and microscopic features when appropriate.