Boletes of Eastern North America 9780815653943, 9780815634829

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Boletes of Easter n North A mer ica

alan e. beSSette, w illia m c. roody, and ar leen r. beSSette

Sy r acuSe u ni v erSit y pr eSS

NO T IC E : 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.

Copyright © 2016 by Syracuse University Press Syracuse, New York 13244-5290 All Rights Reserved First Edition 2016 16  17  18  19  20  21    6  5  4  3  2  1 ∞ 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. For a listing of books published and distributed by Syracuse University Press, visit www.SyracuseUniversityPress.syr.edu. ISBN: 978-0-8156-3482-9 (h a r d c ov e r) 978-0-8156-1074-8 (pa pe r b ac k) 978-0-8156-5394-3 (e-bo ok) l i br a r y of c ong r e s s c ata l o g i ng -i n-pu bl ic at ion 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

K evin Willia m 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 Stanislaus, 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 University 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 encountered 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 specialists on the systematics, biogeography, and phylogeny of boletes, with particular 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 doctoral candidates in the field of mycology; has held numerous editorial positions, including that of managing editor of Mycologia; and is a member of the Mycological 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 practical 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 Acknowledgments Introduction The Bolete Fruitbody: Macroscopic Features Information about Bolete Identification How to Use This Book Explanation and Use of the Field Keys Field Keys to the Boletes Descriptions and Illustrations of Bolete Genera and Species Alessioporus Aureoboletus Austroboletus Baorangia Boletellus Boletinellus Boletus Bothia Buchwaldoboletus Butyriboletus Caloboletus Chalciporus Cyanoboletus Exsudoporus Frostiella Gyroporus Harrya Heimioporus Hemileccinum Hortiboletus Imleria

xiii xvii 1 3 7 13 15 23 55 57 68 72 76 81 83 171 173 180 185 193 200 202 206 208 215 217 220 221 226

xii  ·  Contents Lanmaoa Leccinellum Leccinum Neoboletus Paragyrodon Phylloporus Pseudoboletus Pulveroboletus Retiboletus Rubroboletus Strobilomyces Suillellus Suillus Sutorius Tylopilus Veloporphyrellus Xanthoconium Xerocomellus Xerocomus Undescribed Bolete Species Pending Publication

228 236 245 275 280 283 287 288 296 301 305 309 316 362 364 405 407 414 420 433

Appendixes A. Chemical Reagents and Bolete Identification B. Microscopic Examination of Boletes C. Collecting, Cooking, and Preserving Boletes D. Common Tree Associates of Eastern Boletes Glossary References and Recommended Resources Index to Common Names Index to Scientific Names Photo Credits

441 443 445 448 449 457 461 463 471

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 mycorrhizal relationships they have with many different tree species. Our goal in writing 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, providing 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 tissues 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, Fuscoboletinus, 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, macrochemical testing reactions, habitat information, edibility, and microscopic features when appropriate.

Preface  ·  xv

More interesting and exciting discoveries will be made in the future, a fact that keeps us happily engaged in the ongoing study of boletes. It is our hope that users of this book will benefit from its content and relaxed presentation, and will share our passion for this beautiful and fascinating group of mushrooms.

Acknowledgments

We could not have written this book without the help of so many friends and colleagues. It is with considerable pleasure that we acknowledge the following individuals who made so many valuable contributions. If we have failed to mention someone, please accept our apologies. Ian Gibson, Roy Halling, Paul Kirk, Shaun Pennycook, Scott Redhead, Tom Volk, and Nathan Wilson provided assistance with Latin nomenclature; Susan Mitchell and Glenn Freeman helped with specimen collecting, testing of the keys, and suggesting improvements to the manuscript. Bill Bakaitis, Charlotte Caplan, James Crane, Jonathan Frank, Roy Halling, David Hibbett, Harry “Whitey” Hitchcock, Mike Hopping, Geoffrey Kibby, Michael Kuo, David Lewis, Owen McConnell, Donna Mitchell, Mitchell Nuhn, Beatriz Ortiz-Santana, Igor Safonov, Noah Siegel, Matthew Smith, Walt Sturgeon, Barbara Thiers, and Bill Yule offered advice, raised questions that made us rethink some of our conclusions, assisted with identification, collected and photographed specimens for study, shared technical knowledge, provided resource information and/or molecular data, and researched unknown or undescribed boletes with us. Each generously shared his or her time and special expertise. For contributing photographs that greatly enhanced the functionality and beauty of this book, we are indebted to Bill Bakaitis, Harley Barnhart, Roy Halling, Mike Hopping, Josh Hutchins, Josh Kielsmeier-Cook, Michael Kuo, Renée LeBeuf, Owen McConnell, John Plischke III, Noah Siegel, Eva Skific, Mary Smiley, Walt Sturgeon, the “3 Foragers,” Greg Thorn, Andrus Voitk, and Bill Yule. Thanks also to David Lewis and Rosemary Williams for permitting us to include the images taken by the late Dan Guravich and Robert Williams. Copy editor Annie Barva’s sharp eyes, keen observations, and clear thinking guided us skillfully through the final manuscript preparation. We would have been lost without her. And to the Syracuse University Press director, Alice Randel Pfeiffer, and all of the staff at Syracuse University Press, a big “thank you.” We couldn’t ask for a better team to work with. If we were to name a “ghost writer,” that honor would go to Donna Mitchell. Not only did Donna collect and identify specimens necessary for this work, but she also continuously tested and pushed the limits of the field keys, read and critiqued the manuscript, sacrificed her time, and allowed her life to be disrupted xvii

xviii  ·  Acknowledgments

by the travel and time away from home that this book demanded of us. Through the entire process, from that first phone call to explore an idea to this tangible book you now hold in your hands, she was quietly and steadfastly involved in every stage and step from start to finish. She held our feet to the fire, demanding the very best of us. Donna, there are not enough words to express our gratitude.

Boletes of Easter n North A mer ica

Introduction

Of all the groups of wild mushrooms, none has engendered more enthusiasm and affection than the boletes. Their inherent beauty, their abundance, and, for many, their culinary appeal have firmly established boletes in the hearts of mushroom hunters around the world. The habitats, geographic distribution, and ecology of boletes—including the intriguing relationships they have with trees and shrubs—only add to their interest. As attractive and recognizable as boletes are as a group, individual species of boletes are not always easy to identify, making them a challenge for amateur and professional mycologists alike. Although some boletes are quite distinctive and easily recognized, others demand careful attention to details of the fruitbody and the habitat in which they appear in order to identify them with confidence. This book is an effort to facilitate the identification of the majority of boletes that occur in the eastern United States and Canada for those without the benefit of formal training in mycology. With the exception of the gilled boletes in the genus Phylloporus, all of the species featured in this guide fit the classic concept of a bolete as a fleshy mushroom having a typically cushion-shaped cap with a fertile layer consisting of vertical tubes beneath it and a stalk. Within this simple design are many variations in color, staining of the tissue when cut or bruised, cap surface, stalk ornamentation, flesh odor and taste, habitat, and growth pattern. The mycelium that is mostly hidden within the soil is another important component of boletes. Whereas the bolete fruitbody is a temporary structure in which the reproductive spores are produced and disseminated, the mycelium is perennial. It is made up of fine filamentous strands, called hyphae, that are typically concealed but may sometimes aggregate into a conspicuous “mycelial mat” at the base of the stalk. When visible, the color of mycelium is occasionally useful in identifying species. The mycelium of most boletes is united with the root system of the trees and shrubs with which it forms an interdependent, beneficial relationship. The fungus mycelium serves the tree or shrub by providing it with otherwise inaccessible moisture and nutrients such as phosphorus, nitrogen, and potassium. In turn, the tree or shrub provides simple sugars produced by photosynthesis that nourishes the fungus. This relationship is called mycorrhiza, an association that may be specific to certain groups of trees and mushrooms. Some 1

2  ·  Boletes of Eastern North America

boletes form mycorrhiza only with conifers, others only with broadleaf trees. In some instances, the mycorrhizal association is host specific, meaning that the mushroom forms a unique union with a single species of tree. For instance, Leccinum luteum (p. 256) will be found only under American hornbeam (Carpinus caroliniana). It is now known that the majority of vascular plants form mycorrhizal relationships with fungal partners. This relationship is so important that some plants cannot survive independent of it. The health of entire ecosystems is determined by this cooperative interaction. For additional information about mycorrhizae, see Kendrick 2000 and Stamets 2005. Although the vast majority of boletes are presumed to form mycorrhiza with trees and shrubs, there are exceptions, such as Pseudoboletus parasiticus (p. 288), which invariably occurs attached to the fruitbody of the earthball, Scleroderma citrinum, on which it is thought to be parasitic. A few bolete species are nonmycorrhizal saprotrophs, growing on tree trunks or stumps or on the ground attached to buried roots. Some species of boletes commonly occur on decaying stumps and logs, suggesting that perhaps they are saprotrophs, but it is more likely that their mycelium has migrated to these substrates and that they are nonetheless mycorrhizal associates of nearby trees. Chemicals applied to the fruitbody sometimes provide color reactions that are valuable for differentiating some boletes. For certain species, we have provided macrochemical test reactions that are reliable, valid, and specifically diagnostic. These tests are mentioned in the descriptions. For additional information, see appendix A, “Chemical Reagents and Bolete Identification” (p. 441). Microscopic features of boletes are discussed in the species descriptions but are not utilized in the field keys for identification. For additional information about microscopic features, see appendix B, “Microscopic Examination of Boletes” (p. 443).

The Bolete Fruitbody Macroscopic Features

Si ze a n d Stat u r e The size and to some degree the shape of a bolete can sometimes be useful diagnostic features, particularly if specimens are unusually large or small. Cap diameters vary from 2 to 20 centimeters or more, though most boletes fall within the 6- to 12-centimeter range. Cap size in relation to stalk length and thickness may result in a stature that is tall and thin or short and stout or one of various other combinations. Regardless of overall size and stature, boletes generally have a characteristic cushion-like appearance that soon enables even beginners to distinguish them from other types of mushrooms even at a distance. Identifying individual species, however, is a much more formidable task, requiring careful scrutiny of each part of the fruitbody. Th e Ca p Features of the cap are of primary importance in identifying boletes. There is a wide range of colors, but most are shades or blends of brown, gray, black, white, yellow, orange, red, or purple. Only occasionally are cap colors bright. More often they are softer earth tones. Although color is an obvious feature, it can be tricky. Not only is there some color variation within species, but cap color can change because of age, exposure to light, or other environmental influences. Surface texture may be glabrous, finely velvety, scaly, fibrillose, or glutinous. When fresh, it can be dry or moist, lubricous or viscid. Some caps may be uneven, pitted, or cracked—especially in age. Viscidity and the degree of cracking of the cap are affected by the weather. Caps that are normally viscid or glutinous when fresh become shiny and dry to the touch in dry weather, usually with particles of soil or other bits of debris adhering to them. Dry conditions also induce or exaggerate cracks and crevices in the cap, often revealing the flesh beneath. On some species, the cuticle extends beyond the diameter of the tube layer to form a narrow sterile band or sometimes flaps of tissue on the margin. Although the color of the flesh is usually white or yellow, it is pinkish or orange buff in some species. Instant staining of exposed flesh can mask the 3

4  ·  Boletes of Eastern North America

original color. Pigments sometimes may leach from the cap surface and stain the flesh, but this discoloration is usually confined to the area just beneath the cap surface. For most boletes, the flesh is firm when young and becomes softer in age. Species in the genus Gyroporus have flesh that remains relatively hard and brittle, a useful trait in recognizing this genus in the field. Th e Tu be L ay er The tube layer of most boletes is easily separable from the cap flesh. This feature is often emphasized in distinguishing boletes from terrestrial polypores. However, there are some boletes, especially some species of Suillus, in which the tubes are not readily removed from the cap. The length of bolete tubes ranges from 2 to 20 millimeters or longer in large specimens. There are differences in how the tube layer is attached to the stalk. In some species, the tube layer may be squarely attached or depressed around the stalk. In others, the tube layer is decurrent and extends down on a portion of the stalk. The tubes and pores can be more or less evenly distributed, or they may be radially aligned. The pores may be circular to angular, irregular, or elongated. In the tube layer’s extreme expression, elongated pores have become gill-like, such as in Phylloporus rhodoxanthus (p. 286).

Phylloporus rhodoxanthus

T he Bolete Fruitbody  ·  5

The size of the pores range from 0.3 to 2 millimeters or larger. Young specimens of certain boletes have pores that are stuffed with mycelium, in which case the pores appear solid until the pores develop as the fruitbody matures. The color of the pores is used to distinguish groups of boletes. It can be white to gray, bright or dull yellow, dull pink to pinkish brown or dark brown, orange or red. The pores and tube walls are often concolorous, but in some specimens with dark-brown, red, or orange pores, the tube walls are typically paler. The color of the pores often changes as the bolete develops from a button to a mature specimen. The color often becomes duller or faded in age. Any staining or color change that is a result of bruising of the pores is also a very important feature. Th e Sta l k The majority of boletes have central, straight stalks. There are exceptions, such as Boletinellus merulioides (p. 82), which typically has an eccentric stalk, and Boletus longicurvipes (p. 112), which has a stalk that is often curved near the base. Many boletes will have “situationally” curved stalks when they originate from somewhat vertical substrates such as a steep bank or tree stump. This curved shape is a geotropic response to properly orient the tube layer. The stalk may be approximately the same diameter throughout, enlarged at the apex and tapered downward, swollen in the midportion, or enlarged downward with a thickened or narrowed base. Most are solid, but a few, such as Suillus cavipes (p. 326) and Gyroporus species, have hollow or pithy centers. In addition to providing definitive characteristics, ornamentation on the stalk contributes greatly to the beauty of boletes. Reticulation found on many species of Boletus and Tylopilus species can be fine and delicate or bold and showy. For example, reticulation on the stalks of Boletus aureissimus var. aureissimus (p. 92) and Boletus nobilis (p. 121) is fine and delicate, whereas reticulation on the stalks of Frostiella russellii (p.  207) and Exsudoporus frostii (p.  205) is coarse, deep, and nearly lacunose. Many species of Suillus have resinous dots and smears, and species of Strobilomyces are woolly or shaggy. Leccinellum and Leccinum species are distinctive because they have stalks with scabers that give them a singed appearance. Some boletes have more subtly patterned or completely smooth and featureless stalks. When some species are young, especially Suillus species, their tube layer is concealed by a membranous, gelatinous, or cottony partial veil that extends from the cap margin to the stalk. As the cap expands, the veil ruptures, usually leaving a ring of material or a clearly defined ring zone on the stalk or sometimes remnants on the cap margin. The pores of some species of Suillus are covered by a false veil because the veil originates from the cap margin but never becomes integrated with the stalk tissue.

6  ·  Boletes of Eastern North America

The base of the stalk is sometimes covered with yellow or red bristle-like hairs, the presence and color of which may be diagnostic, often serving as key identifying features—for example, Suillellus subvelutipes (p. 315). The stalk flesh is usually concolorous with the cap flesh, but the colors are sometimes, especially toward the base, intensified or altogether different.

Information about Bolete Identification

I den t i f ic at ion i n t h e Fi el d If you know the identification of a species and wish to read the description or see a color image of it, simply consult the index. If, however, you wish to identify an unknown bolete, follow the process presented here. Before attempting the identification procedure, try to collect specimens with different stages of growth whenever possible. The process of identifying boletes begins at the time of collection. The importance of recording field observations cannot be overstated. Because nearly all boletes grow in ectomycorrhizal partnership with certain trees, it is particularly important to note which tree species are involved. Some boletes form ectomycorrhiza with a broad range of trees, whereas others may be restricted to a single species. Ectomycorrhizal associations are obvious when boletes are found beneath isolated trees, such as in lawns or park lands, or when they appear among monotypic plantations or groves. In mixed woods, it is more difficult to pinpoint the tree partner, though it is sometimes possible to narrow the choices by knowing which trees are likely candidates. Trees that are prime ectomycorrhizal symbionts include the oaks, birches, beech, aspens and other poplars, hemlocks, pines, firs, and spruces (see appendix D, “Common Tree Associates of Eastern Boletes,” p. 448). Woodlands where these trees are well represented are the most rewarding places to look for boletes. The majority of boletes are terrestrial, but a few will fruit on tree stumps or, more rarely, on the trunks of living trees. Note whether the fruitbodies occur singly, scattered or in groups, or in clusters with their stalk bases united. Keep in mind that the more unusual a feature, either morphological or ecological, the more useful that feature is in making an identification. Other important field observations should include any color changes that occur on the fruitbody from bruising or handling. Bruising or staining of injured flesh, pores, or the cap or stalk surfaces is not characteristic of all boletes, but when it occurs, it is a very important diagnostic feature. Remember that staining reactions can be variable, depending on the species involved, the degree of maturation, weather conditions, and other factors. Staining reactions may also become less pronounced as specimens age and lose moisture after being picked. 7

8  ·  Boletes of Eastern North America

The most commonly observed color change is the development of a blue to grayish-blue or greenish-blue staining reaction when the flesh is exposed to the air or when the pores are scratched or rubbed. Other staining reactions may be yellow, green, olive, red to reddish orange, vinaceous, brown, gray, black, or bluish black. Sometimes there is a succession of color changes on exposed flesh. If so, note the colors and the order in which they appear. To encourage staining of the flesh, cut the bolete in half lengthwise and then rub the exposed surface with the flat of a knife blade. A reaction can be instantaneous, or it may take several minutes or, occasionally, even up to a few hours. With some boletes, such as Suillellus subvelutipes (p. 315), the staining reaction is so rapid and thorough that it is difficult to ascertain the original flesh color. It is important to note that specimens used to determine staining reactions of the flesh should be fresh and unrefrigerated. Placing some bolete species in the refrigerator prior to examining their flesh for color changes may alter the test results. This is especially true for some species of Leccinellum and Leccinum. Staining intensity is also variable. The color change of Gyroporus cyanescens (p. 211) is dramatic. Its entire fruitbody quickly and decisively turns a rich, deep blue whenever it is bruised. In contrast, the flesh of Xerocomellus chrysenteron (p. 416) stains slowly and weakly. With some boletes, especially those that typically stain weakly, the reaction may be erratic, or in some collections the changes are so slight as to be almost imperceptible. Macrochemical testing of the fruitbody is best done on fresh specimens at the time of collection or soon thereafter. For additional information regarding macrochemical testing, see appendix A, “Chemical Reagents and Bolete Identification” (p. 441). Characteristic odors are not common among boletes, but they provide valuable clues when they are present. The strong, spicy odor of Suillus punctipes (p. 346), for instance, will readily and convincingly distinguish it from otherwise similar species. Taste can also be a very useful field character. The flesh of some boletes, especially those in the genus Tylopilus, is exceedingly bitter, whereas the flesh of others may have a distinctly nutty or sweet taste. Chalciporus piperatus (p. 195) has an unforgettable biting, acrid taste. The cuticles of certain species of Suillus have an acidic quality that the tongue is quick to discern. There is no danger in taste testing any bolete as long as you do not swallow your sample. Consumption of alcoholic beverages before taste testing the flesh of a bolete may alter or prevent detection accuracy. Recording an image of a fresh collection either with a camera or a sketchbook can be enormously useful for later referral, particularly if the specimens are likely to deteriorate before you can work on them. Boletes are highly perishable.

Information about Bolete Identification  ·  9

Once mature specimens are picked, they can very quickly turn into an unrecognizable entity. When taking a documentary photo of a bolete, be sure to show the upper cap surface, the pores (including a scrape to show any staining), and the entire stalk. A full-length section of the fruitbody as it appears immediately after cutting is also useful. If there is a tardy staining reaction on the cut flesh, a second photo is desirable to show the change. A sharp color photo or detailed drawing, in combination with good field notes, will enable you to identify a surprising number of boletes long after the actual specimens have been relegated to the compost heap or nearby woodlands. I den t i f ic at ion at Hom e The use of field keys, such as those in this guide, may lead to successful on-thespot identifications of the more distinctive species. Many boletes, however, will require more careful, critical study, best done in the comfort of home or workplace with additional resources on hand. When bringing a collection of boletes in from the field or woodlands for further study, you should include, if possible, several examples to represent all stages of development. Many changes in color and structural appearance, such as a diagnostic cracking of the cap cuticle, can take place during the transition from a button bolete to a mature specimen. Refrigerate any specimens that are not to be worked on right away in order to keep them in favorable condition for a day or two. A mature specimen should be set aside to provide a spore print. Individual spores are too small to be seen without a microscope, but collectively they exhibit a colored deposit that is readily observed with the naked eye. Spore-deposit colors range from yellow to olive, olive brown, pinkish brown to reddish brown, pinkish to reddish pink, vinaceous, yellow brown, chocolate brown, or grayish brown to blackish brown. To make a spore print, cut the stalk squarely off close to the cap when possible, leaving about a one-half-inch stub to serve as a pedestal. To avoid any leaching of pigments from the cap or tubes, make sure the pores do not rest directly on the paper. Place the cap with the pore side facing down on a piece of white paper and cover it with an inverted bowl or similar object to restrict air currents. The spores will drop and form clusters of colored dots on the paper. It may take several hours for a sufficient spore deposit to form for color evaluation. Be sure to maintain a vertical orientation of the tube layer so that the spores can drop freely, especially if the pores are small. Although spore color is a defining characteristic for distinguishing several genera of boletes, it is not always necessary to make a spore print. With a little practice, you can recognize the genus of many boletes on the basis of macroscopic features of the fruitbody. However, with other boletes, spore ornamentation can

Making a spore print

Information about Bolete Identification  ·  11

be critical in determining both genus and species. Observation of this feature requires a microscope. Seen through a microscope, typical bolete spores are long and narrowly elliptical or spindle-shaped in face view and inequilateral in profile. For more discussion of microscopic examination of spores, see appendix B, “Microscopic Examination of Boletes” (p. 443).

How to Use This Book

First and foremost, this book is intended to serve as a guide for the field identification of boletes. The keys and text reflect a minimally technical approach with emphasis on macroscopic features of the fruitbody as well as relevant ecological considerations. The species described and illustrated are organized alphabetically first by genus and then by species for easy reference. They are also organized this way because they share similar macroscopic and microscopic features that define the genus in which they are placed. In some instances, species have been renamed and transferred to a new genus based on their relationships as determined by DNA analysis. Because there are different interpretations of taxonomic status and thus the prioritized species names, it may be easier to locate species descriptions by consulting the index of scientific names, which includes synonyms. The numbers of known species indicated in the introductory matter for each genus may vary from those included in other sources because the numbers depend on which classification scheme one chooses to follow. Each species description begins with the currently accepted Latin name of the genus and species. This Latin name is followed by the name of the author(s) who first described the species and the name(s) of those who subsequently amended the nomenclature. Where there have been revisions, the name of the original author who first described a species is shown in parentheses followed by the name of the person or persons who made a change. For example, you will see Leccinum rubropunctum (Peck) Singer (p. 261). Charles H. Peck was the first to describe this species as Boletus rubropunctus and published it in 1897. Rolf Singer later concluded that this species should be reclassified in the genus Leccinum, so he transferred and published this new combination in 1947. The primary Latin names utilized in this book may vary from those commonly found in other contemporary publications that feature boletes. They reflect the most recent taxonomic trends based on molecular research. Although it is nearly impossible to keep up with the rapidly changing proposed names for mushrooms, we have relied heavily on Index Fungorum (www.indexfungorum.org), which is a perpetually updated source for the most widely accepted current names. Synonyms for species are provided in each of the descriptions. These names have been previously applied to species, many of which appear in recent books and taxonomic literature. 13

14  ·  Boletes of Eastern North America

Each fully described species is accompanied by at least one color illustration. In many cases, more than one depiction is provided to demonstrate the considerable variation exhibited by many boletes. The definitions and terms used in this book are as nontechnical as possible, except where simpler definitions are inadequate or less precise, or when using the technical terms is unavoidable—as in the “Microscopic Features” section of each species description. Such technical terms are defined in the glossary. Dichotomous field keys provide a systematic method for identification of species or closely allied clusters of similar species. Although the keys have been prepared with the nonspecialist in mind, using them requires some familiarity with anatomical terms. Beginners are encouraged to review the section “The Bolete Fruitbody: Macroscopic Features” (p. 3). You should confirm your identifications by careful comparison with the written descriptions. There are many similar species, including some that do not appear in the key. Therefore, it is important to read the information provided in the “Lookalikes” section of each description. It is also a very good practice to “key-out” species whose identity is already known. This will enable you to become comfortable with the key’s language and to sharpen your skills of observation. Do not expect success with every attempt at identifying boletes. You may encounter a species that is undescribed or one that is not included in this work. In addition, characteristics of the fruitbody are often ambiguous, and photographs rarely show the full range of features or possible variations that you may encounter. The field mycologist must develop a measured sense of latitude, which sometimes crosses the line from the science to the art of identifying boletes.

Explanation and Use of the Field Keys

The keys in this guide are based on macroscopic features of the fruitbodies and the ecological situation in which the described species occur. Although field keys are useful tools, they should not be used alone to achieve an identification. Keep in mind that field keys are artificial devices designed to arrange organic entities into tidy concepts with clearly defined margins. However, nature is not tidy, and nearly every feature is expressed within a generous range. Color determination is particularly deceptive. Success in identifying boletes requires a fair measure of flexibility when evaluating features. The art of field identification is recognizing the point at which flexibility exceeds credibility. Only experience and practice will sharpen the skills needed to correctly interpret morphological or ecological ambiguity. Trying to identify boletes can sometimes be difficult and frustrating. Bear in mind the words of the preeminent Swedish mycologist Elias Magnus Fries (1794–1878): “No genus has given me more trouble than that of the Boleti.” The fruitbodies of boletes are relatively simple structures consisting of a cap with a fertile layer of more or less vertical tubes beneath it (in the genus Phylloporus the fertile layer is decurrent and gill-like) and a stalk that elevates the cap and fertile layer in order to facilitate spore dispersal. Within this simple design, there are a limited number of field characters to consider. This limitation makes it challenging, if not impossible, to distinguish every bolete based solely on macroscopic features that may overlap or that exhibit only slight differences. When the field keys are used, too much detail can sometimes become more confusing than convincing in the attempt to separate similar species. In these situations, the couplets (paired numerical choices in a key) will terminate at a small cluster of species that share multiple features. Additional comments that attend the species in a cluster serve as cues or hints that may point to a more plausible determination. However, these observations are not necessarily unique and may also apply to others in the group, whether this is stated or not. At this juncture, turn to the full descriptions and photos to compare similar species in greater detail. Subtle differences that are difficult to convey linguistically are often readily apparent in a photograph. The keys are based on the features exhibited on young, fresh specimens in good condition. Older boletes may differ considerably in stature, cap surface texture, the presence or absence of veils, stalk ornamentation, and especially 15

16  ·  Boletes of Eastern North America

fruitbody colors. Whenever possible, collections for study should include multiple specimens in various stages of development. When fresh young specimens are not available, consider probable characteristics of earlier development, particularly in relation to the color of the cap and pore surface. For example, what is only a trace of a color on a mature cap may have been the dominant color in youth. Pores that start out white often become yellow, pink, or brown as they age. When you are keying out mature boletes with pores in this color range, it may be worthwhile to try an alternative key to species that have white or pallid pores when young. Usi ng t h e K e ys To begin the identification process, review this section’s summary of the five groups of boletes, which are divided according to surface features of the stalk, and select the group that applies. Then proceed to the “List of Field Keys,” which provides page numbers you can use to access the applicable key. The keys are constructed of paired numerical couplets, each of which provides two options. Carefully read each choice and select the one that most accurately fits your specimen. In many cases, ambiguous features are accommodated in either choice of a couplet. Continue the process until you attain a tentative identification. At this point, turn to the species descriptions and photos to confirm that the conclusion reached by using the key is a correct identification. If at some point neither choice in a couplet appears to fit, you may have made an error at a previous couplet, or perhaps the bolete you are attempting to identify is not featured in this book. It is even possible that a collected species may be unknown to science and therefore undescribed. If you are unable to identify a bolete using this guide, try searching other references that describe boletes, including those that encompass regions beyond the boundaries of eastern North America. The distribution ranges for most fungi are imprecise at best. Th e Fi v e Grou ps of Bol et e s Ba sed on Fe at u r e s of t h e Sta l k Ring Stalks

A ring or annular zone that is present on the stalk of some boletes is the remnant from a partial veil that extends from the cap margin to the stalk and encloses the immature pore layer of young fruitbodies. As the cap expands, this veil ruptures and collapses, leaving a membranous or fibrillose ring on the stalk. Some other boletes have a universal veil that encloses the entire young fruitbody. This type of veil leaves a sheathing ring or distinct ring zone on the stalk where the veil ruptures as the fruitbody expands.

Explanation and Use of the Field Keys  ·  17

Ring Stalk

Net Stalks

The net-like pattern (reticulation) is present on many bolete stalks. It can cover the entire stalk or only a fraction of the stalk, and sometimes it is present only at the very apex. Reticulation can be coarse or fine, bold or delicate. A hand lens is helpful to detect particularly fine or delicate reticulation.

Net Stalk

18  ·  Boletes of Eastern North America Resin Stalks

This type of ornamentation exhibits resinous dots and smears on the stalk. It is composed of clusters of cystidia that acquire and often exude pigments as the fruitbody matures. The dots sometimes merge to form irregularly shaped smears or patches. These exudates are more or less tacky to the touch and may stain fingers from handling. They are usually pinkish cinnamon to reddish brown and become darker with age or on drying. In some cases, the dots are whitish at first and do not become pigmented or do so only in age. A hand lens may be required to see them. The scabers and punctations present on Rough Stalks (Key D) are similar in appearance to resin dots but are dry to the touch and nonstaining and tend to be erect. Resinous dots and smears are characteristic of boletes in the genus Suillus.

Resin Stalk

Explanation and Use of the Field Keys  ·  19 Rough Stalks

Rough Stalks include species that have dry scales (scabers) or pointed projections (punctae) on the stalk surface. Scabers are composed of clusters of cystidia that form erect, often pointed scales. Most scabers are pallid at first but soon darken to brown or blackish, giving the stalk a singed or scorched appearance. Unlike the dots on Resin Stalks (Key C), scabers and punctae are dry to the touch and do not stain fingers when handled.

Scabers

20  ·  Boletes of Eastern North America

Punctae

Smooth and Miscellaneous Stalks

This group includes all boletes with stalks that are completely smooth or variously patterned but that lack a ring, reticulation, resinous dots, scales (scabers), or punctae. Because there is considerable variation in stalk appearance within this group, stalk detail illustrations are omitted here.

Explanation and Use of the Field Keys  ·  21

List of Fi el d K e ys Key A Key B-1 Key B-2 Key B-3 Key C Key D Key E-1 Key E-2 Key E-3

Ring Stalks, p. 23 Net Stalks: pores at first white to cream-colored or gray, p. 26 Net Stalks: pores at first some shade of yellow, p. 29 Net Stalks: pores at first orange, tan, buff, red, maroon, brown to dark brown, or nearly black, p. 34 Resin Stalks, p. 35 Rough Stalks, p. 37 Smooth and Miscellaneous Stalks: pores at first white to grayish, buff, olive buff, or pinkish tan, p. 41 Smooth and Miscellaneous Stalks: pores at first some shade of yellow or olive buff, p. 45 Smooth and Miscellaneous Stalks: pores at first orange, red, maroon, cinnamon, buff, or brown, p. 51

Field Keys to the Boletes

Key A Key B-1 Key B-2 Key B-3 Key C Key D Key E-1

Key E-2 Key E-3

Ring Stalks, p. 23 Net Stalks: with white to cream-colored or gray pores at first, p. 26 Net Stalks: with yellow pores, p. 29 Net Stalks: with pores some shade of orange, tan, buff, red, maroon, brown to dark brown or nearly black, p. 34 Resin Stalks, p. 35 Rough Stalks, p. 37 Smooth and Miscellaneous Stalks: with white, whitish, grayish, buff, olive-buff, grayish-olive, or pinkish-tan pores when young, p. 41 Smooth and Miscellaneous Stalks: with yellow to pale-yellow or olive-buff pores at first, p. 45 Smooth and Miscellaneous Stalks: with red to rosy-red or copperyred, orange, maroon, cinnamon-brown, reddish-brown, or coffeebrown pores, p. 51

K ey A Ring Stalks

The ring or ring zone present on the stalk of some boletes is the remnant of a partial veil that extends from the stalk to the cap margin and encloses the pore layer on young specimens, or it may be from a ruptured sheathing universal veil. As the cap develops and expands, the veil breaks and separates from the cap margin. A collapsed partial veil adheres to the stalk as a membranous ring or ring zone. Evidence of a partial or universal veil can sometimes also be seen as hanging fragments of tissue on the cap margin. All of the species keyed here exhibit an obvious ring or distinct annular zone on the stalk. The ring is usually persistent but may become less conspicuous or even disappear on older specimens. Most Ring Stalks are members of the genus Suillus. 1a. Cap tomentose to fibrillose or scaly → 2 1b. Cap glabrous or pulverulent, viscid or glutinous → 8 23

24  ·  Boletes of Eastern North America

2a. Mycorrhizal with larch → 3 2b. Mycorrhizal with pine, beech, or oak → 5 3a. Stalk soon hollow; cap dark brown to yellow brown, dry, suede-like to scaly, ring whitish to ochraceous or inconspicuous or disappearing with age but leaving a distinct annular zone → Suillus cavipes (p. 326) 3b. Stalk solid and remaining so → 4 4a. Stalk ring indistinct or lacking; cap dry, tomentose to fibrillose-scaly, pinkish purple to reddish purple; pores large, angular, and radially arranged when young, becoming gill-like and intervenose at maturity, decurrent, pale yellow becoming golden yellow and finally brownish yellow in age → Boletus paluster (p. 132) 4b. Stalk with a prominent gelatinous reddish to reddish-brown ring on the upper portion, the lower portion with viscid pinkish-red to pinkishgray fibrils over a yellowish ground color; cap with coarse grayish to reddish-brown scales or patches, viscid to glutinous beneath the scales or patches; pores angular, elongated, and somewhat radially arranged, yellow becoming dull yellowish brown, usually staining pinkish when injured → Suillus spectabilis (p. 353) 5a. Cap and stalk densely covered with shaggy, grayish to pinkish-gray or blackish scales; ring zone sometimes obscure; cut flesh soon staining orange red then black; typically associated with oak or beech but sometimes also with pine → 6 5b. Cap and stalk reddish to reddish brown or dull orange → 7 6a. Cap scales purplish gray to blackish; widely distributed → Strobilomyces strobilaceus (p. 309) 6b. Cap scales paler, pinkish gray to pinkish tan or pinkish brown; often in sandy soil, especially in coastal areas; mostly southern in distribution but also reported as far north as Cape Cod, Massachusetts → Strobilomyces dryophilus (p. 307) 7a. Cap red to purplish red or rose red, fading in age to buff or ochraceous, cottony-velvety, soon breaking into fibrils and soft, more or less flattened scales, revealing yellow flesh below; always associated with white pine → Suillus spraguei (p. 356) 7b. Cap dull orange to pale reddish brown, fibrillose-scaly; associated with pines other than white pine in oak–pine woods → Suillus decipiens (p. 330) 8a. Associated with broadleaf trees, especially oak; ring thick, gelatinous; sheathing nearly to the stalk base; cap buff yellow to golden yellow or ocher, becoming dingy yellow brown to reddish brown in age → Paragyrodon sphaerosporus (p. 282) (if cap yellow pulverulent, see Pulveroboletus ravenelii [p. 295]) 8b. Associated with conifer trees → 9 9a. Mycorrhizal with larch → 10

Field Keys to the Boletes  ·  25

9b. Mycorrhizal with pine, spruce, balsam fir, or hemlock → 13 10a. Pores often staining bluish or greenish when bruised; associated with native larch; cap glutinous to viscid when fresh, gluten colorless; ground color variable, from grayish to greenish gray or pale grayish brown to yellow brown or reddish brown → Suillus viscidus (p. 361) 10b. Pores unchanging or staining colors other than bluish or greenish when bruised → 11 11a. Pores white, becoming grayish white and then pale reddish brown, bruising purplish gray and then reddish brown; cap with a dark reddishbrown layer of gluten that covers the whitish ground color, fading to pale reddish brown or yellow brown in age → Suillus serotinus (p. 349) (see also Suillus bresadolae var. flavogriseus [p. 323], Suillus grisellus [p. 338], and Suillus viscidus [p. 361]) 11b. Pores yellow, yellowish, or yellow olive → 12 12a. Pores radially arranged, staining reddish brown; cap viscid to glutinous, dark to pale olive buff, yellowish to tan or olive tan; stalk with a viscid band-like ring; associated with introduced European larch → Suillus flavidus (not illustrated) (p. 336) 12b. Pores not radially arranged, pores staining brownish when bruised; cap glabrous, shiny, viscid to glutinous, color variable, orange yellow, dull red, red brown, reddish brown to dark reddish brown, or dark chestnut brown; occurring only with larch → Suillus grevillei (p. 335) 13a. Pores staining greenish blue when bruised; cap at first covered with a coating of sulfur-yellow powder, later somewhat felt-like, orange red to brownish red → Pulveroboletus ravenelii (p. 295) 13b. Pores unchanging or staining colors other than bluish or greenish blue when bruised → 14 14a. Pores radially arranged → 15 14b. Pores not radially arranged → 16 15a. Stalk viscid from a gelatinous veil; cap glabrous, glutinous when fresh, red to mahogany red, orange chestnut, or reddish brown, often developing black spots and streaks with age → Suillus glandulosus (p. 332) 15b. Stalk dry; cap smooth or somewhat wrinkled, obscurely innately fibrillose to glabrous, viscid, chestnut brown or rusty brown to red brown, fading to orange brown with age → Suillus sinuspaulianus (p. 351) 16a. Cap dark reddish brown to cinnamon brown or sometimes ochraceous; stalk ring large and glutinous with purplish tones on the underside; flesh white to pale yellow → Suillus luteus (p. 342) 16b. Cap yellowish to tan, ocher yellow, orange yellow, olive brown to grayish brown, or pinkish cinnamon, often streaked → 17 17a. Stalk ring thin and band-like; cap cuticle tastes acidic; flesh whitish to pale yellow to orange yellow; pores slowly staining pale reddish brown when

26  ·  Boletes of Eastern North America

bruised or sometimes unchanging, also covered at first with a soft, cottony, yellow partial veil that is coated with gluten → Suillus acidus (p. 318) 17b. Stalk ring baggy, thick, and glutinous; flesh orangish, often marbled → Suillus cothurnatus (p. 328) or Suillus salmonicolor (p. 348) K e y B-1 Net Stalks, with White to Cream-Colored Pores at First

The pores of the boletes keyed here often become yellowish or pinkish as they mature. If this feature is indefinite, refer also to Key B-2. 1a. Pores staining blue to bluish green or bluish gray → 2 1b. Pores unchanging or staining colors other than bluish when bruised → 3 2a. Cap whitish to buff or pale brownish when young, becoming dingy brown with rose tints with age; stalk reticulation slight when present and only at the apex; pore surface bruising greenish blue or bluish gray → Boletus pallidus (p. 131) 2b. Cap rusty brown to pinkish brown, becoming reddish brown to chestnut brown or rusty cinnamon with age, at times mottled dingy yellowish, often wrinkled to shallowly pitted; stalk reticulation prominent, covering at least half of the stalk, stalk whitish near the apex graduating to brownish below; pores bruising bluish gray at first and then ochraceous to brownish → Boletus subcaerulescens (p. 157) 3a. Pores staining variously when bruised (sometimes slowly) → 4 3b. Pores not staining when bruised → 10 4a. Stalk reticulation prominent, coarse or fine (use a hand lens), covering one-third or more of the stalk → 5 4b. Stalk reticulation weakly expressed or limited to near or at the apex only →8 5a. Cap pale to dark gray; stalk coarsely reticulate, whitish with deep yellow or reddish stains at the base → Retiboletus griseus (p. 297) 5b. Cap some shade of brown, reddish brown or maroon or with purplish tones → 6 6a. Flesh taste bitter or astringent; reticulation brown → Tylopilus felleus (p. 375), cap pinkish to reddish purple when young, soon becoming some shade of brown with or without purplish tints and finally brown to tan in age; pores becoming pinkish, vinaceous, or pinkish tan in age, often staining brown when bruised Tylopilus variobrunneus (p. 400), cap dark olive brown to greenish brown or blackish brown when young, becoming nearly glabrous and dull medium brown to chestnut brown; pores becoming yellow brown

Field Keys to the Boletes  ·  27

to brownish pink in age, staining brown to brownish rose or pinkish cinnamon when bruised; stalk reticulation whitish on the upper portion when young Tylopilus tabacinus (p. 398), cap yellow brown to orange brown or tobacco brown, becoming finely cracked in age; pores whitish or sometimes brown at first, becoming brown to yellow brown, with darker-brown patches and stains 6b. Flesh taste mild and pleasant or only slightly bitter → 7 7a. Cap reddish brown to dark rusty brown, typically wrinkled or shallowly pitted → Boletus pseudopinophilus (p. 137) (compare with Boletus chippewaensis (p. 98), Xanthoconium affine (p. 409), and Xanthoconium purpureum (p. 412) 7b. Cap yellow brown to orange brown or tobacco brown, becoming finely cracked in age; pores whitish or sometimes brown at first, becoming brown to yellow brown, with darker-brown patches and stains → Tylopilus tabacinus (p. 398) (see also Tylopilus variobrunneus [p. 400]) 8a. Cap and stalk white to whitish, developing dingy-brownish to yellowishbrown stains when handled; stalk weakly to distinctly reticulate → Tylopilus intermedius (p. 382) 8b. Cap or stalk more highly colored → 9 9a. Pores becoming yellow to rusty ochraceous with age → Xanthoconium affine (p. 409) and Xanthoconium purpureum (p. 412) 9b. Pores becoming pinkish or brownish with age → Tylopilus appalachiensis (p. 367), cap yellow brown over the disc, fading to pale yellow brown to dull tan on the margin or darker brown overall, at times with an orange or grayish tinge Tylopilus balloui (p. 374), cap at first bright orange to bright orange red, fading to dull orange, cinnamon, or tan in age; pores white to dingy white, becoming tan or slightly pinkish in age, staining brown when bruised Tylopilus alboater (p. 366), cap nearly black, velvety, often with a whitish bloom when young; pore surface bruising red and then black (see also Tylopilus atronicotianus [p. 369]) Tylopilus indecisus (p. 380), cap and stalk brown, reddish brown, or maroon or with purplish tones; pores becoming pinkish or brownish with age, staining brown when bruised (see also Tylopilus ferrugineus [p. 377], Tylopilus badiceps [p. 371], Tylopilus plumbeoviolaceus [p. 388], Xanthoconium affine [p. 409], Xanthoconium purpureum [p. 412], Tylopilus variobrunneus [p. 400], and Tylopilus rubrobrunneus [p. 395]) 10a. Stalk reticulation prominent, coarse or fine, covering upper third or more of the stalk → 11

28  ·  Boletes of Eastern North America

10b. Stalk reticulation limited to apex; cap and/or stalk with purple, violet, or lilac tones → 19 11a. Cap and stalk predominantly white, whitish, pale yellow, or grayish white → 12 11b. Cap and/or stalk more highly colored → 14 12a. Stalk reticulation deep with raised ribs, appearing pitted; pores white to grayish and sometimes beaded with clear droplets at first, becoming pinkish at maturity → Austroboletus subflavidus (p. 71) 12b. Stalk reticulation fine or coarse but not with raised ribs or appearing pitted → 13 13a. Flesh taste bitter; cap white to whitish, often with buff, pinkish, or pinkish-tan tinges or sometimes becoming grayish tan, pinkish brown, or golden brown overall with age → Tylopilus rhoadsiae (p. 391) 13b. Flesh taste mild or not distinctive; cap milk white to grayish white, sometimes with yellowish or brownish tinges, especially near the margin → Boletus albisulphureus (p. 86) (see also Retiboletus griseus [p. 297]) 14a. Cap and stalk with purple, violet, or lilac tones; cap color variable from creamy white tinged with lilac to pinkish brown, lilac brown, reddish brown, or sometimes dark purple, often paler near the margin and becoming yellowish brown to almost bronze-colored in age, often pitted, wrinkled, or lumpy and uneven → Boletus separans (p. 153). 14b. Cap and stalk lacking purple, lilac, or violet tones → 15 15a. Stalk with a wide band of yellow or golden yellow near the apex; cap pale brownish yellow to orange yellow, becoming yellow to bright yellow with a paler disc in age → Boletus gertrudiae (p. 108) 15b. Stalk lacking a distinct yellow band near the apex → 16 16a. Associated with conifer trees, especially Norway spruce; cap brown to cinnamon brown or rusty red to tan, often tacky to the touch, smooth or somewhat uneven, and slightly pitted → Boletus edulis (p. 101) (see also Boletus variipes [p. 167] and Tylopilus felleus [p. 375]) 16b. Associated with broadleaf trees, especially oak or beech → 17 17a. Stalk reticulation brown to blackish on mature specimens → Boletus atkinsonii (p. 89), cap grayish brown or yellowish brown, minutely cracked in age; stalk dingy white with fine brown reticulation; flesh taste somewhat nutty-sweet or not distinctive (see also Boletus variipes [p. 167] and Tylopilus tabacinus [p. 398]) Tylopilus minor (p. 384), cap pale brown to brown or dull whitish, often with pinkish tones, typically small, 3–8 cm wide but sometimes up to 15.5 cm wide; stalk whitish at first, becoming brownish to cinnamon with reticulation near the apex or extending to the midportion or sometimes lacking, with white basal mycelium; flesh taste bitter

Field Keys to the Boletes  ·  29

Retiboletus griseus (p. 297), cap pale to dark gray, stalk coarsely reticulate, whitish with deep yellow or reddish stains at the base 17b. Stalk reticulation white → 18 18a. Cap typically pitted to corrugated, yellow brown to deeper vinaceous brown; stalk club-shaped to distinctly bulbous, reticulation consisting of elongated meshes, strongly raised in places; reported from New York and New England, west to Minnesota, but distribution range yet to be determined → Boletus nobilissimus (p. 123) 18b. Cap usually smooth or cracked; stalk reticulation fine to somewhat coarse but not with elongated meshes → Boletus nobilis (p. 121), cap usually smooth but at times somewhat pitted, yellowish brown or reddish brown at first, becoming ochraceous to olive ochraceous or reddish ochraceous with age Boletus variipes (p. 167), cap smooth to somewhat pitted at first, often developing cracks or fissures, color variable, creamy tan to yellowish tan, grayish brown to yellow brown or dark brown (see also Boletus atkinsonii [p. 89]) 19a. Cap brownish to grayish brown or dull cinnamon, at times tinged with purplish areas; stalk purple with white mycelium at the base when young, at times becoming grayish purple or purplish brown → Tylopilus plumbeoviolaceus (p. 388) 19b. Cap grayish violet to bluish violet when young, becoming pale purplish, purplish pink, or tan to dull brown in age, staining rusty violet to dark violet when bruised; stalk concolorous with the cap or paler when young, becoming brown in age, with a white apex and base → Tylopilus violatinctus (p. 402) K e y B-2 Net Stalks, with Yellow Pores

Note: Several boletes have pores that are white when young but then become yellow with age. If this feature is uncertain, refer also to Key B-1. 1a. Pores staining blue, greenish blue, or blackish blue when bruised (sometimes weakly or slowly) → 2 1b. Pores unchanging or staining colors other than blue, greenish blue, or blackish blue when bruised → 16 2a. Cap whitish to dingy white with or without pinkish tones when young, becoming brownish with age; stalk yellow in the upper portion, pinkish to red in the lower; flesh bitter tasting → Caloboletus inedulis (p. 191) 2b. Cap more highly colored at first → 3

30  ·  Boletes of Eastern North America

3a. Cap at first reddish orange, rose red, rose pink, pinkish red, or pinkish purple → 4 (see also Boletus paluster [p. 132]) 3b. Cap at first pinkish cinnamon, dull brick red, brown, olive brown, reddish brown, yellow, or yellow brown → 8 4a. Stalk reticulation prominent, typically covering upper half or more of the stalk → 5 4b. Stalk reticulation or pseudoreticulation delicate to obscure or present only at the stalk apex → 13 5a. Flesh staining quickly and decisively blue when exposed → 6 5b. Flesh unchanging or changing only slightly and slowly blue when exposed →7 6a. Flesh taste bitter to astringent, sour or acidic (like lemon) → Butyriboletus roseopurpureus (p. 184), cap pinkish purple when fresh, becoming darker purplish pink to dark purplish red, at times mottled in these colors, marginal areas becoming grayish with age (compare with Butyriboletus peckii [p. 182]) Alessioporus rubriflavus (p. 56), cap dark wine red at first, soon predominantly a yellow ground color with streaks and splashes of wine red, red brown, or ocher, developing olive tones on the disc in age 6b. Flesh taste not distinctive; cap bright rose red to rose pink, becoming pinkish brown to olive brown with age → Boletus speciosus (p. 155) 7a. Tubes and pores staining bluish green when cut or bruised; cap dull red at first, becoming pinkish red to orange red and fading to reddish orange to dull golden orange in age; distribution in the southeastern United States → Boletus carminiporus (p. 97) 7b. Tubes and pores staining blue and then brownish when cut or bruised; cap deep red to dull red or rose red, fading to brownish rose or pale tan to buffy brown from the disc outward; known distribution eastern Canada south to Georgia, west to Tennessee and Indiana → Butyriboletus peckii (p. 182) 8a. Cap and stalk bright yellow when young, developing reddish-brown tones with age; all parts instantly staining blue when handled or bruised → Neoboletus pseudosulphureus (p. 279) 8b. Cap pinkish cinnamon, dull brick red, brown, olive brown, reddish brown, grayish brown, or yellow brown → 9 9a. Stalk reticulation or pseudoreticulation prominent, extending well below the stalk apex → 10 9b. Stalk reticulation weak to uncertain or limited to the stalk apex only → 14 10a. Stalk reticulation brownish, especially at the central portion of the stalk, possibly yellowish at the apex; cap yellow brown to amber brown, reddish brown, or streaked dull brown over a yellow ground color, becoming ochraceous tawny to hazel with age → Boletus luridellus (p. 114)

Field Keys to the Boletes  ·  31

(compare with Buchwaldoboletus lignicola [p. 177], Boletus ferrugineus [p. 105], and Xerocomus illudens/tenax complex [p. 426]) 10b. Stalk reticulation whitish to pinkish or yellow, sometimes brown → 11 11a. Cap dark chocolate brown to dark purplish brown; stalk entirely red or reddish, reticulation yellow; occurs in sandy soil under bear oak and pitch pine; known only from Cape Cod, Massachusetts, rare → Boletus billieae (p. 95) 11b. Cap dull brick red, pinkish cinnamon, olive brown to gray brown, reddish brown, or dark yellow brown; stalk a mix of red and yellow → 12 12a. Cap light brown to grayish brown or olive brown at first, becoming paler olive brown to dark yellow brown with age; distribution in eastern North America known from the upper Midwest and New England, uncommon → Caloboletus calopus (p. 186) 12b. Cap dull brick red, pinkish cinnamon, reddish brown, or yellow brown to olive brown → Butyriboletus brunneus (p. 181), stalk reticulation fine; cap reddish brown or yellow brown to olive brown; pores instantly bruising blue; flesh pale yellow, quickly staining blue when exposed; distribution from eastern Canada to North Carolina, west to Minnesota; fairly common, especially under hemlock Xerocomus illudens/tenax complex (p. 426), stalk reticulation or pseudoreticulation coarse and wide-meshed; pores usually slowly staining cinnamon red or sometimes slowly staining bluish and then cinnamon red, rarely unchanging when bruised; cap yellow brown to pinkish cinnamon or dull brick red to reddish brown; flesh whitish to pale yellow, unchanging or slowly staining pinkish red when exposed; widely distributed in broadleaf woods with oak, in conifer woods with pine, or in mixed woods 13a. Odor of cut flesh pronounced, often described as similar to beef or chicken bouillon, maple syrup, fenugreek, curry, licorice, or fruit → Boletus pallidoroseus (p. 129), flesh slowly stains blue when exposed; cap at first uniformly pale rose pink, pale purplish pink, or mottled with darker-pink tones, golden yellow or paler yellow along the margin, becoming pale orange brownish in age Boletus sensibilis (p. 150), flesh quickly stains blue when exposed; cap dark to pale brick red, fading to dull rose or sometimes dingy cinnamon in age 13b. Odor of cut flesh not distinctive; reticulation weak or indecisive, usually limited to the very apex of the stalk → Boletus miniatopallescens (p. 119), cap dry, smooth, or somewhat velvety, red to brick red, soon fading to reddish orange or orange yellow, often cracked in age (see also Boletus miniato-olivaceus [p. 117])

32  ·  Boletes of Eastern North America

Boletus purpureorubellus (p. 139), cap viscid when wet, shiny when dry, color variable from dark red or purplish to blood red or reddish orange, with yellow on the margin; currently known from Florida, Georgia, and Cape Cod, Massachusetts Boletus rubricitrinus (p. 145), cap dry, glabrous to slightly velvety, color variable from dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown in age, sometimes with yellow tints, usually bruising bluish black; flesh taste acidic Xerocomus sclerotiorum (p. 430), cap dry, dull or somewhat shiny, pinkish red to rose red or purplish red, sometimes with olive tints, becoming dull rose pink to brownish pink in age, slowly staining blackish blue when bruised Boletus oliveisporus (p. 127), cap dry, slightly velvety, often finely cracked in age, reddish orange overall, sometimes paler yellow orange near the margin when young, becoming dark fulvous tinged with bay, or cinnamon brown to dull brown and somewhat shiny in age, instantly staining blue black when bruised Xerocomus illudens/tenax complex (p. 426), cap yellow brown to pinkish cinnamon or dull brick red to reddish brown; pores usually slowly staining cinnamon red or sometimes slowly staining bluish and then cinnamon red when bruised 14a. Cap deeply corrugated or pitted, tan to reddish tan or ocher brown to reddish brown; reticulation delicate (when present) → Xerocomus hortonii (p. 421) 14b. Cap smooth, wavy, or irregular but not deeply corrugated or pitted → 15 15a. Stalk reticulation restricted (when present) to the very apex of the stalk; cap dull yellow brown, ocher brown, or pale cinnamon brown; flesh pale yellow; occurring with hemlock → Boletus huronensis (p. 110) (see also Caloboletus roseipes [p. 192]) 15b. Stalk reticulation (pseudoreticulation) composed of longitudinal lines that sometimes intersect to form a coarse network; flesh whitish; cap olive brown to yellow brown, often cracking with age → Boletus subtomentosus var. subtomentosus (p. 165) (see also Xerocomus illudens/ tenax complex [p. 426]) 16a. Stalk reticulation or pseudoreticulation fluted or shaggy to laceratereticulate, very slender and tall in proportion to cap diameter → 17 16b. Stalk reticulation or pseudoreticulation coarse or fine but not shaggy or lacerate → 18 (see also Suillus cavipes [p. 326], Leccinum rugosiceps [p. 262] and Boletus chippewaensis [p. 98])

Field Keys to the Boletes  ·  33

17a. Cap smooth, glabrous, and viscid, orange to reddish orange, red brown, or bright yellow → Heimioporus betula (p. 219) 17b. Cap velvety, dry, becoming cracked or scaly with age, yellow brown to reddish brown or cinnamon → Frostiella russellii (p. 207) (see also Boletus ferrugineus [p. 105]) 18a. Cap predominantly yellow to orange, including bright yellow, bright orange, golden orange, honey yellow, grayish, or purplish brown → 19 18b. Cap predominantly brick red, pinkish cinnamon, brown or reddish brown, yellow brown, or chestnut brown → 21 19a. Stalk reticulation prominent, typically over most or all of the stalk; handling the cap and stalk often staining fingers yellow → 20 19b. Stalk reticulation obscure or distinct, not prominent; handling the cap and stalk does not stain fingers yellow; distribution from North Carolina south → Boletus aurantiosplendens (p. 90), cap color variable from orange to brownish orange or brownish yellow, sometimes duller yellowish brown or pale brown to rusty brown with a brighter yellowish orange margin Ceriomyces aureissimus var. castaneus (see photo, p. 92), cap velvety, purplish brown, often duller in age 20a. Stalk rich golden orange, reticulation on young specimens indistinct except at the apex; cap bright orange yellow to golden yellow or brownish orange; flesh taste somewhat acidic or not distinctive → Pulveroboletus auriflammeus (p. 291) 20b. Stalk yellow to yellow brown; cap variable from yellow, mustard yellow, olive yellow to yellow brown or gray; flesh bitter tasting → Retiboletus ornatipes (p. 300) 21a. Flesh pale yellow to bright or dark yellow; cap dull brick red, pinkish cinnamon, yellow brown to chestnut brown, or grayish → 22 21b. Flesh whitish; cap reddish to dull brick red, pale to dark cinnamon brown, or reddish brown → 23 22a. Stalk golden yellow with fine yellow reticulation; cap yellowish brown to chestnut brown or grayish brown, becoming paler in age; flesh bright yellow → Boletus auripes (p. 94) (see also Retiboletus ornatipes [p. 300]) 22b. Stalk duller yellow with longitudinal rib-like lines that form a pseudoreticulum; cap pale brownish yellow when young, becoming yellow brown to pinkish cinnamon; flesh pale yellow → Xerocomus illudens/tenax complex (p. 426) 23a. Cap pale to dark cinnamon brown to dull reddish or dark reddish brown, margin with a prominent overlapping sterile band; stalk colored

34  ·  Boletes of Eastern North America

like the cap, with a conspicuous layer of soft white hairs at the base → Aureoboletus projectellus (p. 64) 23b. Cap dull brick red to reddish brown with an olive tint, margin even; stalk woody at the base, reticulation brown, wide-meshed, basal mycelium yellow → Xerocomus illudens/tenax complex (p. 426) K e y B-3 Net Stalks, with Pores Some Shade of Orange, Tan, Buff, Red, Maroon, Brown to Dark Brown, or Nearly Black

1a. Pores radially arranged and more or less decurrent, pinkish brown to golden brown or pale cinnamon brown with pinkish tones; cap brown to reddish brown, burgundy or yellow brown → Bothia castanella (p. 172) 1b. Pores not radially arranged and some shade of orange, red, maroon, or brown to nearly black → 2 2a. Pores brown to maroon or dark brown to nearly black → 3 2b. Pores red to orange → 5 3a. Pores bruising bluish to greenish blue → 4 3b. Pores unchanging or bruising brownish, color grayish orange or darker gray when mature; cap dull reddish brown to brown, dark olive brown, dark gray, brownish gray, or blackish; stalk prominently reticulate nearly overall or at least on the upper one-third → Tylopilus griseocarneus (p. 379) (see also Boletus durhamensis [p. 99], Tylopilus porphyrosporus [p. 389], Tylopilus tabacinus [p. 398], and Bothia castanella [p. 172]) 4a. Pores dark maroon, becoming reddish brown; cap pale brownish olive to yellowish olive; stalk yellow with olivaceous tones at the base, pruinose with reddish pruina that may form a pattern that simulates reticulation on the upper portion → Boletus pseudo-olivaceus (p. 135) 4b. Pores pinkish brown to dark reddish brown to blackish brown; cap dark brown to olive brown or dark vinaceous brown, sometimes staining darker brown when bruised → Tylopilus porphyrosporus (p. 389) 5a. Cap whitish to grayish, grayish brown, or grayish olive; pores pinkish to red or red orange, sometimes yellowish at the margin when young, staining blue when bruised → Caloboletus firmus (p. 189) 5b. Cap red to reddish, brick red, olive yellow, brownish red, brownish orange, or pinkish red → 6 6a. Stalk reticulation coarse and very deep; cap shiny, candy-apple red, often with a narrow, pale margin → Exsudoporus frostii (p. 205) 6b. Stalk reticulation prominent but not coarse or deep, sometimes fine to delicate → 7 7a. Flesh staining bluish when cut → 8

Field Keys to the Boletes  ·  35

7b. Flesh unchanging when cut; cap dull red at first, becoming pinkish red to orange red and fading to reddish orange to dull golden orange with age; known distribution in southeastern United States from North Carolina south to Florida, west to Arkansas and Louisiana → Boletus carminiporus (p. 97) 8a. Associated with oak or other broadleaf trees; stalk reticulation red → 9 8b. Associated with conifers, especially spruce, hemlock, and pine; cap dark red to brick red, rosy red, or brownish red, sometimes with tan areas near the margin in age → Boletus flammans (p. 107) (see also Suillellus luridus [p. 312]) 9a. Cap predominantly red, reddish, pinkish red, brick red, or purplish red → Exsudoporus floridanus (p. 204), occurs on sandy soil under oaks, distribution from Tennessee and North Carolina south; cap pinkish red, rose red, purplish red, or brownish red, typically yellowish or whitish along the margin; flesh pale to bright yellow, quickly staining dark blue when exposed Rubroboletus rhodosanguineus (p. 304), current known distribution from eastern Canada, western New York, Ohio, and West Virginia; cap dark reddish pink to purplish pink or brick red, usually developing brown to olive tones with age 9b. Cap color variable, from yellow to olive yellow, brownish orange, or olive brown, sometimes with pinkish or reddish tints; cap surface dry, dull or shiny, somewhat velvety → Suillellus luridus (p. 312) K ey C Resin Stalks

The boletes keyed here have colored resinous dots or irregularly shaped smears on the stalk that are typically tacky or somewhat sticky to the touch. This feature is characteristic of species in the genus Suillus. 1a. Stalk with a membranous or glutinous ring or distinct ring zone → see Key A, Ring Stalks (p. 23) 1b. Stalk lacking a ring or ring zone → 2 2a. Pores staining blue when bruised; cap fibrillose-scaly when young, becoming nearly glabrous in age, fibers and scales grayish, yellow ocher, dark olive yellow, or reddish brown on a yellow to orange-yellow ground color → Suillus tomentosus (p. 360) 2b. Pores unchanging or staining a color other than blue → 3 3a. Associated with broadleaf trees, especially oak or birch; cap with scattered, appressed, brownish to reddish fibrils over an apricot-orange to

36  ·  Boletes of Eastern North America

yellow ground color, viscid beneath the appressed fibrils, somewhat shiny when dry → Suillus subaureus (p. 358) 3b. Associated with conifer trees, nearly always with pines → 4 4a. Pores at first whitish to very pale yellow, grayish, or pinkish buff → 5 4b. Pores yellow, yellowish orange, ochraceous, brown, or grayish olive → 8 5a. Resinous stalk dots and smears pinkish tan to brownish or reddish brown →6 5b. Resinous stalk dots whitish or yellowish, inconspicuous to lacking in young specimens (use lens) → 7 6a. Cap whitish, becoming yellowish in age; pores whitish at first, soon yellowish to pale ocher yellow in age, often beaded with pinkish droplets when moist; associated with white pine → Suillus placidus (p. 345) 6b. Cap pale yellow, tan, brown, cinnamon, or orangish cinnamon; pore surface often with pinkish-cinnamon moisture beads on young specimens, staining dull cinnamon in age or when bruised or sometimes unchanging → Suillus granulatus (p. 333) 7a. Cap whitish to pale pinkish cinnamon, dull orange yellow, or yellowish tan, becoming pale brown or yellow brown; cap margin with a distinct wide sterile band when young → Suillus neoalbidipes (p. 344) 7b. Cap darker, light to dark brown, vinaceous brown, cinnamon brown, or grayish brown; stalk very short, with inconspicuous resinous dots that become visible only in age → Suillus brevipes (p. 325) (see also Suillus grisellus [p. 338]) 8a. Cap viscid → 9 8b. Cap dry, covered with scattered tufts of reddish, brownish, or grayish fibrils and scales on a yellow ground color, glabrous or nearly so in age; pores pale yellow, sometimes with whitish droplets when young, becoming dull yellow to olive yellow or dingy orange buff in age, staining vinaceous brown when bruised → Suillus hirtellus (p. 340) (see also Suillus subaureus [p. 358]) 9a. Pores bruising vinaceous brown; flesh medium yellow to mustard yellow; cap bright yellow, often with red streaks or patches; associated with white pine → Suillus americanus (p. 319) (see also Suillus hirtellus [p. 340]) 9b. Pores unchanging when bruised; flesh pale yellow; cap pale yellow to ochraceous or rusty orange; odor of flesh fruity or spicy → 10 10a. Pores grayish olive, becoming ochraceous to dull yellow and then olive brown; stalk with minute dots at the apex or glabrous; associated with Scots pine, rare → Suillus bovinus (p. 321) 10b. Pores brown, becoming orange brown to dark dingy yellow; cap viscid, covered with tufts of tiny gray or brown fibrils when young, soon becoming smooth and dull ocher orange, odor spicy; common → Suillus punctipes (p. 346) (see also Suillus subaureus [p. 358])

Field Keys to the Boletes  ·  37

K ey D Rough Stalks, with Scabers or Punctae, or Scurfy

Boletes keyed in this group have stalks that are roughened with dry, scale-like projections (scabers) or points (punctations or punctae). There is sometimes a fine line between stalks that are pruinose and stalks that are punctate. If this character is ambiguous, also consider trying the Key E series. Resin Stalks (Key C) differ in having resinous dots or asymmetrical smears on the stalks that are tacky to the touch. If the stalk is lacerated or shaggy, see Frostiella russellii (p. 207). 1a. Scabers, punctae, or scurfy punctae pinkish, red, or yellowish → 2 1b. Scabers, punctae, or scurfy punctae white, buff, pale brown, brown, reddish brown, reddish orange, or black → 8 2a. Pores soon staining when bruised → 3 2b. Pores unchanging, or very slowly staining, when bruised → 7 3a. Pores bruising blue → 4 3b. Pores bruising yellow to brownish or reddish → 5 4a. Pores red to orange Suillellus hypocarycinus (p. 310), cap dry, somewhat velvety, brown to yellow brown, sometimes with olive or cinnamon tones; distribution south from North Carolina to Florida, west to Mississippi Rubroboletus dupainii (p. 302), cap smooth, slimy-viscid when fresh, becoming shiny when dry, purplish red to pinkish red or bright red, sometimes with yellowish spots; rare, reported only from North Carolina and Iowa 4b. Pores yellow; cap color variable, dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown in age → Boletus rubricitrinus (p. 145) (compare with Xerocomellus intermedius [p. 419], Xerocomus sclerotiorum [p. 430], and Hortiboletus rubellus [p. 225]) 5a. Cap viscid to glutinous when moist, shiny when dry, smooth to slightly wrinkled or pitted, yellow orange to brownish orange or ochraceous, often developing green tones in wet weather or with age; stalk punctae yellow to reddish or reddish brown → Boletus longicurvipes (p. 112) 5b. Cap dry or only slightly viscid when moist → 6 6a. Cap dark brown to chestnut brown, smooth to slightly wrinkled or somewhat shallowly pitted; stalk white on the upper portion, pinkish on the lower portion, covered with coarse rosy-pink scabers that darken to brown with age; known only from Florida, Georgia, and Mississippi → Leccinum roseoscabrum (p. 259)

38  ·  Boletes of Eastern North America

6b. Cap olive brown to reddish brown, often yellow to olive gold on the margin; stalk yellow beneath a covering of reddish or reddish-brown punctae; widely distributed from Georgia northward → Xerocomus morrisii (p. 428) 7a. Pores white at first, becoming pinkish to flesh-colored; cap dry or slightly viscid when moist, pink to rose-colored when young, fading to pinkish tan and then dingy brown with age; flesh white; stalk base deep chrome yellow → Harrya chromapes (p. 217) 7b. Pores yellow at first; flesh yellow; cap furrowed or wrinkled or sometimes smooth, slightly viscid or dry, bay red to chestnut red or reddish brown → Leccinum rubropunctum (p. 261) (see also Hemileccinum subglabripes [p. 220], Leccinum rugosiceps [p. 262], and Xerocomus hortonii [p. 421]) 8a. Pores at first dark chocolate brown to purple brown, becoming reddish brown in age; cap purplish brown to grayish brown; stalk densely covered with darker purple-brown tiny scales → Sutorius eximius (p. 364) 8b. Pores at first white to pale tan, grayish, buff, olive buff, reddish, or yellow →9 9a. Pores at first yellow to pale yellow, dingy yellow, or reddish → 10 9b. Pores at first white to pale tan, grayish, buff, olive buff, or beige → 13 10a. Pores staining brownish or unchanging when bruised → Leccinellum crocipodium (p. 240), cap dry to moist but not viscid, wrinkled, uneven, pitted, often cracked in age, dark brown to blackish brown when young, fading to pale yellow brown in age; associated with broadleaf trees, especially oak Boletus longicurvipes (p. 112), cap shiny, yellow orange to brownish orange or ochraceous, often developing green tones in wet weather or in age Boletus weberi (p. 171), cap dry, cracked, and scaly, with pale-yellow flesh showing in the cracks, brownish olive at first, becoming olive brown, sometimes with reddish tints; pores unchanging or sometimes slowly staining brown when bruised; distribution southern, from Florida west along the Gulf Coast into Texas 10b. Pores bruising blue to greenish blue or grayish blue (sometimes weakly or slowly) → 11 11a. Cap medium to dark brown, blackish brown, chocolate brown, olive brown, grayish brown, or chestnut brown → 12 11b. Cap brick red to rosy red, yellow orange, or mustard yellow, but always with red or rosy areas → Lanmaoa roseocrispans (p. 235) (see also Boletus patrioticus [p. 133], Hortiboletus campestris [p. 223], and Leccinum rugosiceps [p. 262]) 12a. Stalk conspicuously scaly-shaggy; cap medium brown to dark brown, coated with appressed fibers at first, becoming fibrillose-scaly with

Field Keys to the Boletes  ·  39

flattened to erect scales and yellow flesh showing between the scales; known only from Florida → Suillellus pictiformis (not illustrated) (p. 78) 12b. Stalk often scurfy to punctate but not scaly-shaggy → Caloboletus roseipes (p. 192), cap olive brown to grayish brown, becoming pale grayish brown to grayish tan with age; associated with conifers in eastern Canada, New England, and New York Boletellus chrysenteroides (p. 78), cap dark brown and velvety when young, becoming bay brown to chestnut brown and often cracked in age, with whitish to pale-yellow flesh that lacks reddish tints showing in the fissures; often growing on stumps or other woody substrates (see also Boletellus pseudochrysenteroides [p. 80], Xerocomellus chrysenteron [p. 416], and Neoboletus luridiformis [p. 277]) 13a. Pores bruising greenish, blue, brownish, yellowish, yellow brown, pinkish gray, cinnamon, or olivaceous → 14 13b. Pores unchanging when bruised → 20 (if ambiguously bruising slowly and weakly buff, see Leccinum scabrum [p. 265]) 14a. Cap at first some shade of orange to orange brown or yellow → 15 14b. Cap lacking orange, orange brown, or yellow tones → 19 15a. Cap pale yellow at first, darkening to yellow and then olive brown with age, typically wrinkled and shallowly pitted at maturity, becoming cracked in age; mycorrhizal only with American hornbeam (Carpinus caroliniana) → Leccinum luteum (p. 256) 15b. Cap orange, dull orange, orange brown → 16 16a. Pores bruising greenish → Leccinellum quercophilum (p. 244), cap glabrous, wrinkled, and pitted when young, becoming conspicuously cracked with age, whitish to grayish flesh showing in the spaces, dull orangish brown to tawny brown when young, becoming dark brown to grayish brown in age; current known distribution from Illinois and Michigan 16b. Pores bruising brownish, olive, olive fuscous, hazel, or cinnamon → 17 17a. Growing in sandy soil or dunes in northern coastal areas with beach grass, beach heather, and sedges; cap orange to yellowish orange, fading to dull cinnamon or ochraceous, wrinkled, becoming cracked in age → Leccinum arenicola (not illustrated) (p. 274) 17b. Growing with conifer or broadleaf trees → 18 18a. Associated with conifer trees, especially Norway spruce → Leccinum vulpinum (p. 274), cap dull orange to brick red, rusty red, reddish brown, or dark reddish brown, with a sterile margin that splits into hanging flaps at least on young specimens 18b. Associated with broadleaf trees → Leccinum insigne (p. 251), cap orange to rusty cinnamon or brick red, becoming brownish orange in age, with a sterile margin that splits

40  ·  Boletes of Eastern North America

into hanging flaps at least on young specimens; flesh stains purplish gray to purplish black when exposed; associated with big-toothed aspen, quaking aspen, or birch Leccinum pseudoinsigne (p. 257), cap bright yellow, orange yellow, or bright orange to orange red, fading to orange buff in age; flesh of young specimens stains reddish cinnamon and then bluish fuscous, but reddish cinnamon stage often lacking on mature specimens; associated with aspen and birch Leccinum versipelle (p. 271), cap dry to slightly viscid, fibrillose, breaking up to form downy patches or small scales with a sterile margin that splits into hanging flaps at least on young specimens, dull orange at first, becoming tan or brownish; flesh staining pinkish and then purple gray to blackish when exposed; associated with birch 19a. Cap brown to dark brown or blackish, often mottled → Leccinum subleucophaeum (p. 269), cap dry or slightly viscid when moist, dark brown with or without paler mottling, often blackish brown over the disc, sometimes fading to yellowish brown in age; flesh slowly staining gray to grayish black; associated with broadleaf trees, especially birch or aspen (if the flesh stains blue, see Tylopilus sordidus [p. 396]) Leccinum snellii (p. 267), cap dark brown to blackish, usually mottled with cream-colored spots; stalk flesh staining orange red at the cap junction, blue green at base; associated with broadleaf trees, especially yellow birch Leccinellum griseum (p. 242), cap wrinkled, uneven, pitted, often cracked in age, dark brown to blackish brown when young, fading to pale yellow brown in age; flesh quickly staining pinkish brown and then darkening to reddish cinnamon; associated with oak Leccinum scabrum (p. 265), cap grayish brown to yellow brown or dark brown, glabrous or matted-fibrillose, breaking up into tiny flattened scales in age; flesh unchanging or slowly staining reddish, especially in the stalk, when exposed; associated with birch 19b. Cap whitish to pale pinkish buff, vinaceous buff, or pale grayish to cinnamon buff → Leccinum holopus (p. 248), cap white or whitish when young, occasionally with gray, buff, tan, or pinkish tints, often darkening and developing a greenish or bluish tinge in age Leccinum insolens (p. 253), cap dry, fibrillose to minutely scaly, sometimes finely cracked, dull white to pale pinkish buff or pale grayish cinnamon buff, typically darkening to pale yellow brown in age; flesh whitish, staining pinkish gray to purple gray and then fuscous when exposed or sometimes lacking the preliminary pinkish coloration before staining purple gray to fuscous

Field Keys to the Boletes  ·  41

Leccinum chalybaeum (p. 245), cap viscid when fresh, somewhat shiny when dry, smooth to slightly subtomentose, buff to pinkish buff when young, becoming dingy yellow brown, sometimes with grayish-green to bluish-gray tints, especially near the margin; distribution from Georgia south to Florida and west to Texas Leccinum glutinopallens (p. 247), cap slimy-viscid when fresh, glabrous, dingy vinaceous buff to pale pinkish gray or pale grayish brown; distribution northern, currently known from eastern Canada west to Michigan 20a. Cap dark brown to blackish, often mottled with cream-colored spots; stalk flesh staining orange red at cap junction, blue green at base → Leccinum snellii (p. 267) 20b. Cap paler → 21 21a. Cap at first dull orange, becoming tan or brownish; associated with birch → Leccinum versipelle (p. 271) 21b. Cap color variable, from whitish to pale tan grayish, brownish, pinkish buff, yellow brown, dark brown, or purplish red, but lacking orange tones → Leccinum scabrum (p. 265), cap grayish brown to yellow brown or dark brown, glabrous or matted-fibrillose, breaking up into tiny flattened scales in age; flesh unchanging or slowly staining reddish when exposed; associated with birch (see also Leccinum holopus [p. 248]) Leccinellum albellum (p. 238), cap color variable, white, whitish, pale tan, pale gray to pinkish gray, or pinkish brown to medium brown, sometimes tinged yellow or with a bluish cast, glabrous or somewhat velvety, often conspicuously pitted at maturity, sometimes cracked in age; associated with broadleaf trees (see also Leccinum holopus [p. 248]) Gyroporus purpurinus (p. 213), cap velvety, purplish red to burgundy; stalk scurfy to somewhat velvety, brittle and hollow, colored like the cap K e y E-1 Smooth and Miscellaneous Stalks, with White, Whitish, Grayish, Buff, Olive-Buff, or Pinkish-Tan Pores When Young

Boletes in this group include those with a smooth or variously textured stalk that lacks reticulation, scabers, scales, rings, or resinous dots. Some may have slightly scurfy stalks or have longitudinal ribs that form a net-like pattern that mimics true reticulation. 1a. Pores staining when bruised, sometimes slowly → 7 1b. Pores unchanging when bruised → 2 2a. Cap viscid to glutinous when fresh → 3

42  ·  Boletes of Eastern North America

2b. Cap dry → 4 3a. Cap bright yellow to orange yellow; stalk viscid to glutinous when fresh, somewhat scurfy near the apex, nearly smooth below, pale yellow to yellow down to the base, which is sheathed with cottony white mycelium → Pulveroboletus curtisii (p. 294) 3b. Cap white to yellowish tan, yellowish brown, cinnamon, vinaceous brown, or dark brown; stalk at first with faint whitish resinous dots (use lens) that darken with age → Suillus brevipes (p. 325), cap light brown to dark brown, vinaceous brown, cinnamon brown, or grayish brown, fading to cinnamon or tan in age; stalk proportionally short, nearly glabrous, white to pale yellow, sometimes with brown stains near the base, with or without inconspicuous resinous dots that become visible only with age Suillus neoalbidipes (p. 344), cap white, pale pinkish cinnamon, dull orange yellow, or yellowish tan with a wide band of sterile tissue on the margin when young, becoming darker cinnamon to pale brown or yellowish brown 4a. Cap shaggy or scaly, often developing a network of ridges and pits, pinkish tan to golden yellow, yellow brown or salmon-tinged; pores white when young, becoming grayish pink to pinkish brown → Veloporphyrellus conicus (p. 407) 4b. Cap glabrous or fibrillose to velvety, smooth or cracking in age → 5 5a. Stalk brittle, soon becoming hollow; flesh firm and brittle → Gyroporus purpurinus (p. 213), cap vinaceous red to burgundy; widely distributed Gyroporus castaneus (p. 209), cap chestnut brown to orange brown or yellow brown; widely distributed Gyroporus subalbellus (p. 215), cap apricot buff to pinkish buff or with cinnamon tones; distribution along the Atlantic Seaboard from Cape Cod, Massachusetts, south to Florida and west to Texas Gyroporus phaeocyanescens (p. 212), cap fulvous to yellow brown or grayish brown; known distribution from Georgia south to Florida, west to Texas 5b. Stalk solid → 6 6a. Cap and stalk white, becoming whitish to pale straw-colored or tinged brownish; distribution southern, from the coastal plain of North Carolina south to Florida and west to Texas → Xanthoconium stramineum (p. 413) (see also Suillus grisellus [p. 338], Tylopilus intermedius [p. 382], and Leccinellum albellum [p. 238]) 6b. Cap and/or stalk with purplish or lilaceous tones, flesh intensely bitter → (if flesh is mild tasting, see Boletus separans [p. 153])

Field Keys to the Boletes  ·  43

Tylopilus plumbeoviolaceus (p. 388), cap brownish to grayish brown, at times tinged with purplish areas; stalk purple with white mycelium at the base when young, at times becoming grayish purple or purplish brown Tylopilus violatinctus (p. 402), cap grayish violet to bluish violet when young, becoming pale purplish, purplish pink, or tan to dull brown in age, staining rusty violet to dark violet when bruised; stalk concolorous with cap or paler (see also Tylopilus ferrugineus [p. 377] and Tylopilus minor [p. 384]) 7a. Cap at first bright orange to bright orange red, fading to dull orange, cinnamon, or tan in age; pores white to dingy white, becoming tan or slightly pinkish in age, staining brown when bruised → Tylopilus balloui (p. 374) 7b. Cap lacking bright-orange to bright orange-red tones → 8 8a. Cap at first white, whitish to straw yellow, buff, apricot buff, pinkish cinnamon, or tinged pinkish or ochraceous → 9 8b. Cap yellow, yellow brown, dark brown, cinnamon brown, chestnut brown, olive brown, grayish brown, maroon, purplish, or black → 12 9a. Pores bruising greenish blue to dark blue → 10 9b. Pores bruising brownish to pinkish cinnamon → 11 10a. Cap, stalk, pores, and flesh instantly staining dark blue to indigo blue when bruised or handled; stalk brittle, stuffed with pith, becoming chambered or hollow with age → Gyroporus cyanescens (p. 211) 10b. Cap and stalk not staining dark blue when bruised or handled; pores bruising greenish blue (sometimes weakly) and then grayish brown; stalk solid → Boletus pallidus (p. 131) (see also Phylloporus boletinoides [p. 285]) 11a. Cap at first apricot buff to pinkish buff; stalk becoming chambered or hollow; pores slowly staining pinkish cinnamon when bruised or in age; flesh unchanging when exposed → Gyroporus subalbellus (p. 215) 11b. Cap at first whitish to pale ochraceous, becoming tan to brownish or orangish brown; stalk solid → Tylopilus rhodoconius (p. 392), cap pale ochraceous to brownish orange at first, becoming pale brown and then darker brown with a paler margin; flesh white to pale creamy white with hyaline marbling, slowly staining pale pinkish and then dingy pinkish red when exposed; odor not distinctive or faintly of chlorine; taste mild at first and then very bitter Tylopilus peralbidus (p. 386), cap white when young, soon becoming ochraceous tan to chamois and finally brownish; flesh white, sometimes staining pale pinkish brown or buff when exposed, often

44  ·  Boletes of Eastern North America

slowly; odor bleach-like or unpleasant; taste bitter (see also Tylopilus intermedius [p. 382] and Boletus pallidus [p. 131]) 12a. Cap yellow, yellow brown, chestnut brown, grayish brown, olive brown, or blackish → 13 12b. Cap maroon to dark brown, cinnamon brown, reddish brown, purple violet, or purplish → 16 13a. Cap black to dark grayish brown, brown, or olive brown → 14 13b. Cap yellow brown, chestnut brown, orange brown, reddish brown, or rusty red → 15 14a. Pores bruising dark blue to dark blue green and then brownish red; cap gray brown to olive brown or dark brown, often with dark greenish or bluish tints along the margin → Tylopilus sordidus (p. 396) 14b. Pores bruising blackish or reddish and then slowly black → Tylopilus alboater (p. 366), cap dry, somewhat velvety, occasionally finely cracked in age, black to dark grayish brown, often covered with a thin whitish bloom when young; stalk sometimes slightly reticulate at the very apex Tylopilus atronicotianus (p. 369), cap dry, glabrous or nearly so, somewhat shiny, olive brown, bronze brown, grayish brown, or pale brownish, at times imbued with yellowish tones, darker in age; stalk occasionally finely reticulate at the very apex 15a. Pores bruising pale cinnamon; cap yellow brown or dark rusty red to reddish brown at first, becoming dull orange to dark orange yellow → Aureoboletus roxanae (p. 67) 15b. Pores bruising dull yellow to brown → Tylopilus appalachiensis (p. 367), cap yellow brown over the disc, fading to pale yellow brown to dull tan on the margin or darker brown overall, at times with an orange or grayish tinge; pores becoming pinkish with age; stalk usually lacking reticulation but sometimes finely reticulate at the apex Xanthoconium affine (p. 409), cap dark brown to chestnut brown or ocher brown, sometimes with white to pale-yellow spots or patches; pores becoming yellowish to dingy yellow brown with age 16a. Flesh staining pinkish to brown when cut (sometimes slowly) → Tylopilus ferrugineus (p. 377), cap somewhat velvety, dark reddish brown; pores whitish at first, becoming pale pinkish buff, bruising dull brown; stalk dull brown to reddish brown, often whitish near the apex, with white basal mycelium; associated with oak Tylopilus badiceps (p. 371), cap maroon and velvety when young and fresh, soon purplish brown to dark reddish brown; pores white for a long time and then dingy white to brownish but not pinkish at maturity;

Field Keys to the Boletes  ·  45

stalk colored like the cap but often with more pronounced violaceous tones or sometimes violaceous nearly overall, usually white at the apex and base 16b. Flesh unchanging when cut → 17 17a. Flesh taste decidedly bitter → Tylopilus williamsii (p. 405), cap dark magenta to purple violet or brownish purple when young, fading to pinkish gray to pale yellow brown; known only from coastal Georgia and Florida west to Mississippi (see also Tylopilus plumbeoviolaceus [p. 388]) Tylopilus rubrobrunneus (p. 395), cap dark to bright purple when young, becoming purple brown, dark reddish brown, dull brown, or cinnamon; widely distributed 17b. Flesh taste mild to slightly tart but not bitter → Austroboletus gracilis var. gracilis (p. 70), cap maroon to reddish brown or cinnamon, at times tawny to yellow brown; pores becoming pinkish to pinkish brown with age Xanthoconium purpureum (p. 412), cap dark red to purple red or maroon, sometimes with pale-brownish tints; pores becoming ochraceous yellowish to rusty ochraceous K e y E-2 Smooth and Miscellaneous Stalks, with Yellow to Pale-Yellow or Olive-Buff Pores at First

Boletes keyed here lack scales, scabers, resin dots, stalk rings, or reticulation on the stalk. Some species may have longitudinal ribs on the stalk that intersect and form a net-like pattern (pseudoreticulation). If this feature is strongly expressed, these species may be placed with Net Stalks (Key B, p. 26). If the stalk is shaggy-lacerate, see Frostiella russellii (p. 207). 1a. Fruitbody attached to the earthball, Scleroderma citrinum; cap ocher brown to tawny olive or like tarnished brass; pores yellow, becoming olivaceous with age, sometimes with reddish or rusty stains, unchanging or very rarely becoming slightly bluish when bruised → Pseudoboletus parasiticus (p. 288) 1b. Fruitbody growing on soil or decaying wood → 2 2a. Pores elongated, decurrent, and radially arranged, at times nearly lamellate and intervenose → 3 2b. Pores not elongated, decurrent, radially arranged or at times nearly lamellate and intervenose → 6

46  ·  Boletes of Eastern North America

3a. Fruitbody occurring strictly with ash trees, stalk usually eccentric; cap slightly viscid and shiny when moist, yellow brown to olive brown or reddish brown → Boletinellus merulioides (p. 82) 3b. Fruitbody occurring with other broadleaf trees or conifers → 4 4a. Pores pale olive buff, becoming dark olive buff with age, gill-like with numerous cross-veins, or poroid; cap dry, somewhat velvety to minutely scaly, becoming nearly smooth with age, cinnamon to dark pinkish brown, fading to dull yellow brown in age → Phylloporus boletinoides (p. 285) 4b. Pores pale to bright yellow → 5 5a. Pores bright yellow, becoming golden yellow with age, strongly decurrent, typically gill-like with cross-veins, often wrinkled and sometimes forked; cap dull red to reddish brown, reddish yellow, or olive brown → Phylloporus rhodoxanthus (p. 286) (see also Boletus subfraternus [p. 159]) 5b. Pores pale yellow, becoming golden yellow and then brownish yellow; cap pinkish purple to reddish purple, covered with minute scales or hairs; associated with larch trees → Boletus paluster (p. 132) 6a. Fruitbody occurring on or around tree stumps and roots or other woody substrates → 7 6b. Fruitbody occurring on soil → 9 7a. Cap blackish brown, becoming chocolate brown to chestnut brown, velvety at first and often cracked in age; pores pale yellow, becoming yellow to greenish yellow, quickly staining blue when bruised; associated with broadleaf trees → Boletellus chrysenteroides (p. 78) (see also Xerocomus hypoxanthus [p. 422]) 7b. Cap bright yellow to golden yellow, orange red, or reddish brown; associated with pines → 8 (see also Boletellus ananas [p. 76] and Imleria badia [p. 227]) 8a. Cap dry, velvety, and suede-like, sometimes finely cracked in age, reddish brown to yellow brown or rusty golden yellow, often darker brown at the margin; pores typically decurrent, yellow to golden yellow at first, becoming brownish yellow in age, bruising dark greenish blue to blackish blue and then fading to reddish brown; flesh pale yellow, staining blue when exposed, sometimes slowly; odor sweet, somewhat minty or citrus-like; associated with and often found with the polypore Phaeolus schweinitzii → Buchwaldoboletus lignicola (p. 177) (see also Boletus oliveisporus [p. 127] and Buchwaldoboletus hemichrysus [p. 174]) 8b. Cap dry to slightly viscid, glabrous, often cracked in age, sulfur yellow when young, becoming pale yellow to whitish with age; flesh sulfur yellow to pale yellow, typically staining blue when exposed but sometimes slowly or not at all; odor not distinctive; pores bruising blue then slowly brownish → Buchwaldoboletus sphaerocephalus (p. 179)

Field Keys to the Boletes  ·  47

9a. Cap covered with coarse, purplish-red to dark-red scales and having whitish veil remnants on the margin; associated with oak or pine; southern in distribution → Boletellus ananas (p. 76) 9b. Cap lacking coarse scales → 10 10a. Cap viscid to glutinous when fresh and moist, shiny when dry → 11 10b. Cap dry, more or less glabrous or velvety, dull or shiny, with or without cracks or fissures → 14 11a. Pores bruising dark bluish gray to blackish blue; cap color variable, dark red or purplish to blood red or reddish orange, yellow on the margin → Boletus purpureorubellus (p. 139) 11b. Pores bruising brownish to dull brick red or unchanging when bruised → 12 12a. Cap bright yellow at first, becoming duller yellow with age; stalk viscid to glutinous when fresh, somewhat scurfy near the apex, smooth below, pale yellow to yellow down to the base, which is sheathed with cottony white mycelium → Pulveroboletus curtisii (p. 294) 12b. Cap light to dark brown, reddish brown, ochraceous to rusty orange or cinnamon → 13 13a. Pores bright golden yellow and remaining so well into age; cap pinkish brown to vinaceous brown when fresh, fading in age or when dry; taste of the cap surface is acidic; stalk viscid when fresh, pale yellow at the apex, streaked and flushed pale pinkish brown downward, with copious white mycelium at the base → Aureoboletus auriporus (p. 60) 13b. Pores pale yellow, becoming olivaceous brown; cap reddish brown to reddish cinnamon, becoming much darker brown in age, often with a pinkish tinge; stalk viscid when wet, glabrous, often longitudinally ridged → Pulveroboletus atkinsonianus (p. 289) (see also Suillus brevipes [p. 325] and Suillus bovinus [p. 321]) 14a. Cap corrugated to wrinkled or with pits or depressions → 15 14b. Cap more or less smooth, with or without cracks or fissures → 16 15a. Pores quickly bruising greenish blue to grayish blue; cap color variable, brick red or rosy red over a yellow ground color when young, becoming yellow orange or mustard yellow with dull pinkish or rosy tones in the folds, burgundy-red or brownish-red splashes on portions of mature specimens, slowly bruising greenish blue to grayish blue especially along the margin; known only from Florida → Lanmaoa roseocrispans (p. 235) 15b. Pores unchanging when bruised, widely distributed; cap tan to reddish tan or ocher brown to reddish brown → Xerocomus hortonii (p. 421) (see also Hemileccinum subglabripes [p. 220]) 16a. Pores bruising blue to greenish blue (sometimes slowly) → 21 16b. Pores unchanging when bruised or changing to colors other than blue or greenish blue → 17

48  ·  Boletes of Eastern North America

17a. Cap uniformly pinkish red or purplish red to blood red or dark red → Boletus roodyi (p. 141) 17b. Cap tan, yellow, brown, yellow brown, reddish brown, orange brown, grayish brown, or olive brown → 18 18a. Flesh instantly deep indigo blue when cut, brittle; cap fulvous to yellow brown or grayish brown; stalk brittle and soon becoming hollow; current known distribution from Georgia south to Florida and west to Texas → Gyroporus phaeocyanescens (p. 212) 18b. Flesh unchanging or staining weakly when cut → 19 19a. Distribution restricted to the Deep South, currently known only from sandy coastal woodlands in southern Georgia and the Gulf Coast of Florida; cap reddish brown to dark brown, cinnamon brown, or purplish brown; stalk typically abruptly bulbous at the base → Boletus abruptibulbus (p. 84) (see also Xerocomus hypoxanthus [p. 422] and Ceriomyces aureissimus var. castaneus [see photo, p. 92]) 19b. Distribution from South Carolina and northward → 20 20a. Cap glabrous or nearly so → Boletus alutaceus (p. 87), cap pinkish cinnamon to pinkish brown or pinkish tan; pores slightly greenish yellow (see also Pulveroboletus atkinsonianus [p. 289]) Aureoboletus roxanae (p. 67), cap yellow brown to reddish brown, becoming dull orange to dark orange yellow or tawny; pores whitish at first, soon becoming pale yellow and finally pale golden yellow (see also Boletus aureissimus var. aureissimus [p. 92] and Hemileccinum subglabripes [p. 220]) 20b. Cap suede-like to somewhat velvety → Aureoboletus innixus (p. 62), cap dry and somewhat velvety, frequently cracked in age, dull reddish brown to dull cinnamon or yellow brown, sometimes with purplish or reddish tones near the margin; stalk enlarged downward to a pinched base (see also Boletus alutaceus [p. 87] and Aureoboletus roxanae [p. 67]) Boletus subtomentosus var. subtomentosus (p. 165), cap olive brown to yellow brown, often cracked in age Pulveroboletus auriflammeus (p. 291), cap and stalk deep orange yellow when young to golden yellow or brownish orange in age; stalk with longitudinal ribs or reticulation toward the apex 21a. Cap predominantly red, pinkish red, rose red, purplish red, brick red, rusty brown, tawny brown, reddish brown, olive brown, mahogany, or with reddish tones → 25 21b. Cap predominantly whitish to pale buff, bright yellow, ochraceous yellow, orange ocher, yellow brown, ocher brown, or pale cinnamon brown → 22

Field Keys to the Boletes  ·  49

22a. Cap whitish to pale buff or pale brownish; pores whitish to pale yellow when young, becoming yellow to greenish yellow in age, quickly staining greenish blue and then grayish brown when bruised → Boletus pallidus (p. 131) 22b. Cap more highly colored → 23 23a. Cap bright yellow, ochraceous yellow, orange ocher, yellow brown, ocher brown, or pale cinnamon brown → 24 23b. Cap lacking yellow, orange-ocher, yellow-brown, ocher-brown, or cinnamon-brown tones → 25 24a. Distribution southern from Georgia and Florida west to Mississippi; cap ochraceous yellow to orange ocher or pale pinkish cinnamon with yellow on the margin at first, becoming dull brownish yellow in age → Boletus ochraceoluteus (p. 125) 24b. Distribution northern or more widely distributed than just from Georgia and Florida west to Mississippi → Boletus huronensis (p. 110), associated with hemlock trees from eastern Canada south to New York and Connecticut; cap dull yellow brown to pale cinnamon brown (compare with Cyanoboletus pulverulentus [p. 202], Caloboletus roseipes [p. 192], Boletus subtomentosus var. subtomentosus [p. 165], and Hemileccinum subglabripes [p. 220]) Neoboletus pseudosulphureus (p. 279), cap bright yellow at first, becoming reddish brown to brown in age; all parts instantly bruising blue to blackish blue (see also Lanmaoa pseudosensibilis [p. 233]) 25a. Cap olive to olive brown or grayish brown at first, possibly developing pinkish or reddish tones with age; cap surface velvety → 26 25b. Cap lacking olive tones at first → 27 26a. Cap dark olive to olive brown or grayish brown, developing surface cracks, with a red to pinkish color in the cracks, at least on the margin, velvety at first; stalk finely granular to somewhat scurfy, sometimes with longitudinal streaks, often yellow on the upper portion, usually reddish near the base → Xerocomellus chrysenteron (p. 416) 26b. Cap olive when very young, soon pinkish to brick red or dark red, often brownish red toward the margin, usually with olive to tarnished-brass tints, smooth, somewhat velvety; taste of flesh tart to acidic → Boletus patrioticus (p. 133) 27a. Cap at first predominantly brown, dull rusty brown to dull reddish brown, tawny brown, mahogany, cinnamon brown, dark yellow brown, or blackish brown → 28 27b. Cap at first predominantly red, brick red, purplish red, rose red, reddish brown, or pinkish red → 30 28a. Cap mahogany to reddish brown or rose brown, paler toward the margin, margin bruising blue; known only from Mississippi → Boletus mahogonicolor (p. 115)

50  ·  Boletes of Eastern North America

28b. Cap color variable; more widely distributed → 29 29a. All parts of the fruitbody instantly staining dark blue when handled or cut → Cyanoboletus pulverulentus (p. 202), cap dark yellow brown to blackish brown or dark cinnamon brown Boletus oliveisporus (p. 127), cap dark fulvous tinged with bay or cinnamon brown to dull brown 29b. Cap not staining blue when bruised → Lanmaoa pseudosensibilis (p. 233), cap dull rusty brown to dull reddish brown, fading to dull cinnamon or dingy yellow brown Boletus rufomaculatus (p. 148), cap at first dull rusty brown to ocher brown, becoming cracked, mottled with brick red or brownish red in age; usually occurring with beech 30a. Flesh decisively and fairly quickly staining blue when exposed → 31 30b. Flesh slowly and weakly staining blue when exposed or unchanging → 34 31a. Cap small (often 5 cm wide or less), rose red to pinkish red, typically cracking with age → Hortiboletus campestris (p. 223) (see also Boletus subfraternus [p. 159] and Xerocomellus intermedius [p. 419]) 31b. Cap typically broader than 5 cm → 32 32a. Flesh with pronounced odor of curry, fenugreek, maple syrup, or licorice; cap somewhat velvety when young, becoming nearly glabrous in age, dark to pale brick red, fading to dull rose or sometimes dingy cinnamon → Boletus sensibilis (p. 150) (see also Boletus miniatopallescens [p. 119]) 32b. Flesh odor not distinctive → 33 33a. Taste of flesh acidic or tart; occurring in sandy soil under oak or in oak–pine woods; southern in distribution from Florida to Texas but also reported as far north as New Jersey; cap color variable, dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown → Boletus rubricitrinus (p. 145) 33b. Taste of flesh not distinctive; more broadly distributed in eastern North America → Boletus miniatopallescens (p. 119), cap red to brick red, soon fading to reddish orange or orange yellow, often cracked in age Lanmaoa carminipes (p. 231), cap pinkish red to rose red at first, becoming dingy brownish, often yellowish or paler pinkish red along the margin, quickly staining dark blue when bruised or handled Imleria badia (p. 227), cap chestnut brown, reddish brown, or yellow brown, sometimes with olive tones, dry to slightly viscid, smooth or somewhat velvety Xerocomellus intermedius (p. 419), cap red to rose red, often soon developing grayish to olive-brown tones, surface dry, subtomentose, becoming cracked, with pink tints in the cracks (see also Boletellus pseudochrysenteroides [p. 80])

Field Keys to the Boletes  ·  51

34a. Stalk completely glabrous and smooth or in rare instances having slight reticulation at the apex; tube layer shallow → Baorangia bicolor complex (p. 74), cap dark red to rose red, fading to ochraceous or tan, often cracking in dry weather or in age; flesh unchanging or slowly and weakly staining blue when exposed; odor not distinctive or reminiscent of cheese in older specimens Boletus rufomaculatus (p. 148), cap red to dull red, becoming cracked, mottled with brick red or brownish red; flesh slowing staining blue, especially above the tubes, or sometimes unchanging; odor unpleasant or not distinctive Boletus pallidoroseus (p. 129), cap uniformly pale rose pink, pale purplish pink, or mottled with darker-pink tones, golden yellow or paler yellow along the margin, becoming pale orange brownish with age; flesh slowly staining blue when exposed; odor reminiscent of beef or chicken bouillon or of spoiled meat 34b. Stalk somewhat roughened or dotted or with darker flakes → 35 35a. Stalk bruising dark blue or darkening brown when handled → Xerocomus sclerotiorum (p. 430), cap pinkish red to rose red or purplish red, sometimes with olive tints, becoming dull rose pink to brownish pink in age, slowly staining blackish blue when bruised; known from New England south to Florida, west to Tennessee Hortiboletus rubellus (p. 225), cap dark red, fading with age to brick red or olivaceous brown, often becoming finely cracked in age; widely distributed (see also Boletus harrisonii [see photo, p. 222], Hortiboletus campestris [p. 223], and Boletus miniato-olivaceus [p. 117]) 35b. Stalk not bluing or discoloring when handled → Boletus miniato-olivaceous (p. 117), cap red to rosy red when young, becoming pale rose pink and developing olive tones, finally becoming rosy tan to olive yellow with rosy tints, often staining dark blue when bruised; widely distributed (see also Boletus patrioticus [p. 133]) Xerocomus hypoxanthus (p. 422), cap tawny brown to pale red brown or pale yellow brown to orange brown; known only from South Carolina and Florida west to Mississippi K e y E-3 Smooth and Miscellaneous Stalks, with Red to Rosy-Red or Coppery-Red, Orange, Maroon, Cinnamon-Brown, Reddish-Brown, or Coffee-Brown Pores

1a. Pores radially arranged, at times gill-like, buff to olive buff; cap somewhat velvety to minutely scaly, becoming nearly smooth in age, cinnamon

52  ·  Boletes of Eastern North America

1b. 2a. 2b. 3a. 3b. 4a.

4b. 5a.

5b. 6a.

6b. 7a.

7b.

8a.

to dark pinkish brown, fading to dull yellow brown → Phylloporus boletinoides (p. 285) Pores not radially arranged → 2 Pores bruising blue, greenish blue, or blackish blue → 8 Pores not bruising blue, greenish blue, or blackish blue → 3 Flesh taste decidedly acrid-peppery; base of stalk bright yellow → Chalciporus piperatus (p. 195) Flesh lacking acrid-peppery taste → 4 Flesh taste tart and lemony; cap brownish olive at first, becoming olive brown, sometimes with reddish tints, cracked and scaly with pale yellow flesh showing in the cracks; occurring on sandy soil with longleaf pine from Florida west to Texas → Boletus weberi (p. 171) Flesh taste not distinctive or bitter to slightly acidic → 5 Pore surface uniformly dark cinnamon brown to reddish brown; cap medium brown to rusty brown, becoming paler brown to yellowish brown with age, typically yellowish along the margin; known only from the eastern piedmont region of North Carolina → Boletus durhamensis (p. 99) (see also Tylopilus porphyrosporus [p. 389] and Buchwaldoboletus hemichrysus [p. 174]) Pores at first bright rose red to grayish red; cap less than 6 cm wide → 6 Pores at first grayish red to dull red; cap dark orange yellow and staining brown when young, becoming reddish brown to orange brown at maturity; known only from Florida → Chalciporus rubritubifer (p. 199) Pores bright rose red, becoming dull rose, orangish, or yellow brown with age; widely distributed north of North Carolina → 7 Stalk finely pruinose, mostly rose pink, and typically yellow near the base, with yellow basal mycelium; cap yellowish to pinkish red, becoming pinkish cinnamon in age → Chalciporus pseudorubinellus (p. 196) Stalk red or reddish, sometimes mixed with yellow, lacking yellow mycelium at the base; cap red or reddish when young, more yellow and becoming cracked with age → Chalciporus rubinellus (p. 198) Pores dark brown to coffee brown, maroon or brownish yellow; flesh quickly staining blue when cut; odor pungent → Boletus vermiculosoides (p. 169), cap yellow when young, becoming brown with maturity; stalk brownish pruinose over a dull-whitish or pale-yellow ground color (compare with Boletus vermiculosus [see photo p. 169]) Boletus subgraveolens (p. 161), cap dull yellow brown overall or mottled with darker or paler shades of yellow or brown, usually staining dark blue and then darker brown when fresh, becoming cracked in dry weather; stalk dull pale yellow and pruinose on the lower portion with a dark vinaceous-red to maroon base

Field Keys to the Boletes  ·  53

8b. Pores at first red to dark red, pinkish red, orange red, coppery red, or reddish maroon, sometimes with a yellowish margin → 9 (see also comments under Chalciporus piperatus [p. 195] and the photos of Chalciporus piperatoides [p. 195]) 9a. Flesh staining blue when exposed, sometimes slowly → 10 9b. Flesh unchanging or becoming darker yellow when exposed; cap dull red at first, becoming pinkish red to orange red, color fading with age to reddish orange to dull golden orange; current known distribution from North Carolina south to Florida, west to Arkansas and Louisiana → Boletus carminiporus (p. 97) 10a. Stalk glabrous to faintly longitudinally striate; cap dark reddish salmon to bright orange, becoming rosy brick red to reddish brown, fading to brownish orange with yellow tints and a whitish margin; recorded only from McComb, Mississippi → Boletus roseolateritius (p. 143) (see also Boletus fairchildianus [p. 104]) 10b. Stalk more or less glabrous to faintly or at least partially pruinose, scurfy, or punctate → 11 11a. Distribution from North Carolina south to Florida and west to southeast Texas → Suillellus hypocarycinus (p. 310), cap brown to yellow brown, sometimes with olive or cinnamon tones (compare with Boletus subluridellus [p. 163]) Suillellus subluridus (p. 314), cap orange pink to orange yellow with vinaceous tints or red to purplish red, sometimes rusty red to cinnamon red, developing more brownish tones with age 11b. Distributed from South Carolina northward → 12 (see also Suillellus subvelutipes [p. 315]) 12a. Cap brownish to dark olive brown, dark reddish brown, or nearly blackish brown, staining blue to blackish blue when bruised → Neoboletus luridiformis (p. 277), cap velvety when young, dark brown to nearly blackish brown, becoming dark reddish brown to olive brown; known distribution from eastern Canada south to South Carolina and west to Minnesota Boletus pseudo-olivaceus (p. 135), cap pale brownish olive to yellowish olive; stalk with reddish pruina that may form a pattern that simulates reticulation on the upper portion; known distribution from New England south to North Carolina and west into the Great Lakes region 12b. Cap bright red to pinkish red, orange red, brick red, purplish red, reddish brown, or cinnamon brown, all parts staining blue to blackish blue when cut or bruised → 13

54  ·  Boletes of Eastern North America

13a. Stalk yellow, pruinose; cap purplish red, brownish red, pinkish red, or orange red; distribution from New England and New York south to Georgia and west to Minnesota → Boletus subluridellus (p. 163) 13b. Stalk reddish to orange cinnamon (may be yellow on the upper portion) → Suillellus subvelutipes (p. 315), cap color variable, cinnamon brown to yellow brown, reddish brown, or reddish orange to orange yellow; stalk base often with stiff yellow or reddish hairs; all parts instantly bruising blackish blue Boletus rufocinnamomeus (p. 146), cap dull brick red with an ochraceous overtone from an evanescent bloom when young, becoming reddish brown to dull cinnamon brown with age Neoboletus luridiformis (p. 277), cap velvety when young, dark brown to nearly blackish brown, becoming dark reddish brown to olive brown; known distribution from eastern Canada south to South Carolina and west to Minnesota Lanmaoa borealis (p. 229), cap dark red to bright apple red, becoming dull brick red to rusty rose in age; yellow flesh that slowly stains blue at the stalk apex; orange red to red pores that stain greenish blue; and a red stalk that may be reticulate at the apex

Descriptions and Illustrations of Bolete Genera and Species

Gen us A l e s siop oru s Gel a r di, Vi zzi n i & Si mon i n i The genus Alessioporus was erected in 2014 to accommodate species formerly classified in several genera, including Boletellus and Boletus. Alessioporus was named in honor of the late Carlo Luciano Alessio (1919–2006), who dedicated most of his mycological studies to the investigation of Italian boletes. Species in this genus have dry caps, reticulate stalks, and yellow flesh that quickly stains dark blue when exposed. At present, it is a small genus consisting of two European and one American species that have been shown to be closely related when multigene molecular analysis was performed.

Alessioporus rubriflavus (A)

55

56  ·  Boletes of Eastern North America

Alessioporus rubriflavus (B)

Alessioporus rubriflavus J. L. Frank, A. R. Bessette & A. E. Bessette, nom. prov. common name(s): none. overview: The term rubriflavus means “red and yellow,” a reference to the colors of the fruitbody. Red portions of the cap cuticle immediately stain amber with KOH, and the flesh immediately stains orange with KOH. Cap: 5–14(21) cm wide, convex, becoming broadly convex to nearly plane at maturity, margin with a narrow band of sterile tissue when young, becoming even; surface slightly viscid or dry, slightly velvety, sometimes finely cracked in age, dark wine red on very young buttons, soon developing a yellow ground color covered with streaks and splashes of various shades of wine red, red brown, and ocher, sometimes retaining wine-red coloration well into maturity, becoming olive to brownish over the disc in age, staining greenish blue to bluish black when bruised; flesh firm, bright yellow, quickly staining dark blue when exposed; odor not distinctive or mildly unpleasant; taste unpleasant, sour, or sometimes astringent. pores and tubes: yellow at first, soon becoming pale orange yellow and then olive brown, deeply depressed at maturity, quickly bruising blue then slowly reddish brown; tubes 8–25 mm deep. stalk: 4.5–11 cm long, enlarged downward, ventricose to clavate, typically with a pinched base, solid; surface dry, conspicuously reticulate over at

Aureoboletus   ·  57

least the upper half, finely velvety over the lower half, bright yellow near the apex or nearly overall, with dark wine-red to red-brown streaks and splashes below, especially near the base, quickly staining blue and then slowly brownish when bruised; reticulation yellow toward the apex, yellow brown below, darkening when handled, with a whitish mycelium; flesh bright yellow, quickly staining blue when exposed, usually dark wine red near the base. habitat and season: solitary, scattered, or in groups on the ground in mixed pine and oak woods; summer and fall. distribution and frequency: Georgia and South Carolina northward to southern New York, distribution limits yet to be determined; occasional to locally common. spore print: olive brown. microscopic features: spores 13–19 × 4–6 µm, subellipsoid to subfusiform, smooth, pale brownish yellow. edibility: unknown. lookalikes: Boletus flavissimus (Murrill) Murrill (not illustrated), originally described from Florida, has a bright-yellow cap, lacks red coloration in all stages of development, does not stain when bruised, and its stalk lacks reticulation. Butyriboletus brunneus (p. 181) has a reddish-brown or yellow-brown to olive-brown cap; pale-yellow flesh that quickly stains blue; a yellow, finely reticulate stalk with red on the lower portion; and smaller spores, 10–15 × 3–5.5 µm. Neoboletus pseudosulphureus (p. 279) has a bright-yellow cap that becomes duller yellow to tawny at maturity and typically develops brown to brownish-red tints, especially over the disc, in age. It has a yellow stalk that sometimes develops reddish tints, especially near the base, and usually lacks reticulation or is sometimes reticulate on the upper portion. It has smaller spores that measure 10–16 × 4–6 µm. Gen us A u r eob ol e t u s Pouz a r The genus Aureoboletus was erected in 1957 to accommodate species formerly classified in the genus Boletus. The name Aureoboletus means “brilliant golden yellow,” a reference to the color of the pores of the species in this genus. It is a small genus of medium-size terrestrial boletes, with only five species occurring in eastern North America. They have viscid or dry caps that sometimes have a projecting sterile margin. A white mycelium is often present at the base of the stalk. The pores are brilliant golden yellow at first and become dull golden yellow to greenish yellow or brownish yellow in age. The cap and stalk flesh is white and does not stain when cut or bruised. These species have olive-brown spore prints.

Aureoboletus auriporus (A)

Aureoboletus auriporus (B)

Aureoboletus auriporus (C)

Aureoboletus gentilis

60  ·  Boletes of Eastern North America

Aureoboletus auriporus (Peck) Pouzar = Boletus auriporus Peck common name(s): Acidic Golden-pored Bolete, Golden-pored Bolete overview: The combination of the pinkish-brown to vinaceous-brown cap and brilliant golden-yellow pores is very striking. Both the cap cuticle, which is distinctly acidic tasting, and the tube layer of this bolete are easily removed. The term auriporus means “golden pores,” in reference to the color of this beautiful bolete’s pores. cap: 2–8 cm wide, convex, becoming broadly convex to nearly plane, margin with a narrow band of sterile tissue; surface coated with tiny appressed fibrils, smooth or sometimes bumpy and appearing blistered, moist and viscid when fresh, becoming dull when dry, pinkish brown to vinaceous brown when fresh, fading in age or when dry; taste of the cap surface is acidic; flesh white to pale yellow, except vinaceous under the cap surface, typically not staining blue when cut or bruised, sometimes staining weakly blue or pinkish; odor somewhat pungent or not distinctive; taste acidic or not distinctive. Pores and tubes: brilliant golden yellow when young and fresh and remaining so well into maturity, becoming dull golden yellow to greenish yellow and then brownish yellow in age, usually slowly staining dull brick red when bruised or sometimes unchanging; tubes 6–15 mm deep. Stalk: slightly enlarged downward or nearly equal, typically narrowed abruptly at the base; surface viscid to tacky when fresh, pale yellow at the apex, streaked and flushed pale pinkish brown downward, with copious white mycelium at the base; flesh concolorous with the cap flesh. Habitat and season: solitary to scattered or in groups under oak; late spring, summer, and fall. Distribution and frequency: New Jersey to Florida, west to Mississippi and Texas; fairly common. Spore print: olive brown. Microscopic features: spores 11–16 × 4–6 µm, fusiform-ellipsoidal, smooth, pale brown. Edibility: edible and somewhat acidic tasting. Lookalikes: Aureoboletus gentilis (Quél.) Pouzar (see photo, p. 59) has a pale gray-pink to pinkish-brown streaked cap and grows under broadleaf trees, especially oak and beech. Aureoboletus innixus (p. 62) has a darkerbrown dry cap, a cap surface lacking an acidic taste, and flesh that has an odor reminiscent of witch hazel. It often grows in clusters fused at the stalk bases.

Aureoboletus innixus (A)

Aureoboletus innixus (B)

62  ·  Boletes of Eastern North America

Aureoboletus innixus (Frost) Halling, A. R. Bessette & Bessette = Boletus innixus Frost = Boletus caespitosus Peck = Pulveroboletus innixus (Frost) Singer Common name(s): Clustered Brown Bolete Overview: This common small- to medium-size brown bolete, which lacks any prominent distinguishing feature, is often present during dry periods when other boletes are scarce. The term innixus means “reclining,” alluding to the fruitbodies, which are often leaning, especially when tightly clustered. The cap surface stains mahogany red to reddish brown with KOH, develops a green flash and then stains dull orange red with NH4OH, and is pale olive with FeSO4. The flesh stains pale dull pinkish orange with KOH or NH4OH and pale gray with FeSO4. Cap: 3–7.5 cm wide, convex, becoming broadly convex to nearly plane at maturity, margin with a narrow band of sterile tissue; surface dry and somewhat velvety, frequently cracked in age, dull reddish brown to dull cinnamon, or yellow brown, sometimes with purplish or reddish tones near the margin; flesh white to pale yellow or tinged vinaceous beneath the cuticle; odor pungent, reminiscent of witch hazel; taste slightly lemony or not distinctive. Pores and tubes: bright yellow when young, becoming dull yellow, unchanging when bruised; tubes 3–10 mm deep. Stalk: short and stout, typically enlarged downward, club-shaped and distinctly swollen above a tapered base, solid; surface moist or dry on the upper portion, viscid near the base, yellowish and streaked with darkbrown tones, with yellow basal mycelium sometimes visible; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or often in caespitose clusters in broadleaf woods, especially with oak; late spring through fall. Distribution and frequency: eastern Canada south to Georgia, west to Michigan; occasional to locally common. Spore print: olive brown. Microscopic features: spores 8–11 × 3–5 µm, ellipsoidal, smooth, pale brown. Edibility: edible but not highly regarded. Lookalikes: Aureoboletus auriporus (p. 60) has a viscid cap with a peelable cuticle that tastes acidic and flesh that lacks a distinctive odor.

Aureoboletus projectellus (A)

64  ·  Boletes of Eastern North America

Aureoboletus projectellus (B)

Aureoboletus projectellus (Murrill) Halling = Boletellus projectellus (Murrill) Singer = Boletus projectellus (Murrill) Murrill = Ceriomyces projectellus Murrill Common name(s): none

Aureoboletus   ·  65

Overview: Although not common for much of its range, this tall, slender-stalked bolete is not difficult to recognize. The dull-pinkish to cinnamon-brown cap, concolorous coarsely ribbed stalk, and association with pines are distinctive features. The term projectellus means “extending or projecting,” a reference to the cap cuticle, which conspicuously projects beyond the tube layer. The identification features include a large pinkish to reddish-brown cap with an overlapping cap cuticle, a long reddish-brown reticulate stalk with a copious white basal mycelium, growth under pines in late summer and fall, and very large spores. Cap: 4–20 cm wide, convex, becoming broadly convex to nearly plane in age, margin with a distinct band of sterile tissue; surface dry, somewhat velvety when young, often cracked in age, pale to dark cinnamon brown or dark reddish brown, occasionally with gray or olive shades, especially when young; flesh whitish, often with a rosy tinge, not staining blue when exposed but slowly changing to yellow brown; odor not distinctive; taste somewhat acidic. Pores and tubes: pale yellow to olive yellow when fresh, becoming brownish olive in age, staining lemon yellow when bruised, depressed at the stalk in age; tubes 1–2.5 cm deep. Stalk: unusually long, up to 24 cm, and up to 5 cm thick, equal or enlarging downward, solid; concolorous with the cap or somewhat paler, often with a conspicuous covering of soft white mycelium at the base and with prominent coarse reticulation overall or at least on the upper two-thirds, distinctly viscid at the base in wet weather; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered under pines, especially two-needle Scots pine; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally common. Spore print: olive brown. Microscopic features: spores 18–33 × 7.5–12 µm, oval to ventricose, smooth, pale brown. Edibility: edible and good when fresh, flavorless when dried and reconstituted. Lookalikes: Perhaps the bolete that is most likely to be confused with Aureoboletus projectellus is Frostiella russellii (p. 207), which has a scaly cap and a more lacerated stalk. Imleria badia (p. 227) is similarly colored but smaller and has a nonreticulate stalk and pale-yellow pores that stain blue when bruised.

Aureoboletus roxanae (A)

Aureoboletus roxanae (B)

Aureoboletus   ·  67

Aureoboletus roxanae (Frost) Klofac = Boletus roxanae Frost = Boletus roxanae var. auricolor Peck = Ceriomyces roxanae (Frost) Murrill = Xerocomus roxanae (Frost) Snell Common name(s): Roxanna’s Bolete Overview: Although this small and relatively nondescript bolete lacks dramatic features, it is easily recognized once encountered a time or two. Its distinguishing features include a granulose yellow-brown cap and a club- or bowling-pin-shaped yellow to pale orange-yellow stalk, typically with a distinct dull-orange zone at the apex. The term roxanae means “Roxanna’s”: this bolete is named after the wife of American mycologist Charles C. Frost (1805–1880). Cap: 3–9 cm wide, convex to broadly convex, becoming nearly plane in age, margin even; surface dull, dry, granular-scaly when young, becoming nearly glabrous, sometimes cracking at maturity, yellow brown or dark rusty red to reddish brown at first, becoming dull orange to dark orange yellow, fading somewhat in age, margin often paler than the disc; flesh whitish to pale yellow, not staining when exposed; odor weakly pungent or not distinctive; taste unpleasant. Pores and tubes: whitish at first, soon becoming pale yellow and finally pale golden yellow in age, darkening somewhat or developing pale cinnamon stains when injured; tubes 6–10 mm deep. Stalk: enlarged downward, solid; surface dry, faintly pruinose and longitudinally striate, at least on the upper portion, yellow to pale orange yellow or tawny, typically with a distinct dull-orange zone at the apex; flesh concolorous with the cap flesh or at times having pinkish tones. Habitat and season: solitary or scattered under oaks, usually in mixed oak and pine woods; summer and fall. Distribution and frequency: eastern Canada south to South Carolina, west to Minnesota; occasional. Spore print: dark olive brown. Microscopic features: spores 8–13 × 3–5 µm, fusiform-ellipsoid, smooth, yellowish. Edibility: edible but usually having an unpleasant taste. Lookalikes: Boletus alutaceus (p. 87) has been reported from New England south to Kentucky and west to Ohio and Wisconsin, and it grows in broadleaf and mixed woods, especially with oak and beech. It initially has a yellowish-brown cap that becomes pale tan with a pinkish tint, pale greenish-yellow pores that do not stain when bruised, and whitish to pale-tan flesh that does not stain when exposed. Aureoboletus innixus (p. 62) has

68  ·  Boletes of Eastern North America

much brighter-yellow pores and a brownish stalk, and it usually grows in clusters fused at the stalk bases. Gen us A u s t rob ol e t u s (Cor n er) Wol f e The genus Austroboletus was erected in 1980 to accommodate species formerly classified in the genus Tylopilus. The name Austroboletus means “southern bolete.” These boletes are medium-size terrestrial boletes with viscid or dry caps that sometimes have a projecting sterile margin and often have unusually long stalks in relation to their cap diameters. A conspicuous white mycelium is usually evident at the base of the stalks. The cap and stalk flesh is white to pale vinaceous and does not change color when cut or bruised. These mushrooms’ pores are white at first, darken to pinkish brown at maturity, and do not stain blue; they may darken or stain brownish when bruised. The colors of their spore prints vary from vinaceous pink to pinkish brown or reddish brown. Microscopically, their spores are obscurely to distinctly pitted or sinuously pitted. Two species of Austroboletus currently occur in eastern North America.

Austroboletus gracilis var. gracilis

Austroboletus gracilis var. flavipes

Austroboletus gracilis var. pulcherripes

70  ·  Boletes of Eastern North America

Austroboletus gracilis var. gracilis (Peck) Wolfe = Tylopilus gracilis (Peck) Henn. = Porphyrellus gracilis (Peck) Singer Common name(s): Graceful Bolete Overview: The Graceful Bolete is not uncommon, though it seldom occurs in large numbers. It is typically solitary or scattered on the ground but occasionally will fruit on decaying logs and stumps. The term gracilis means “slender,” a reference to the stalk. Cap: 3–10 cm wide, convex to broadly convex, margin even; surface dry, finely velvety when young, sometimes cracked in age, maroon to reddish brown or cinnamon, at times tawny to yellow brown; flesh white or tinged pink, not staining when exposed; odor not distinctive; taste slightly tart or not distinctive. Pores and tubes: white when young, becoming pinkish to pinkish brown or burgundy tinged in age, darkening or staining brownish when bruised; tubes 1–2 cm deep. Stalk: long and slender in relation to the cap diameter, solid, enlarged downward or nearly equal, often curved; surface dry, concolorous with the cap or paler, with elevated and anastomosing rib-like lines that sometimes form an obscure, narrow reticulation overall or at least on the upper half, basal mycelium white; flesh concolorous with the cap flesh or sometimes yellow toward the base. Habitat and season: solitary, scattered, or in groups on the ground or sometimes on decaying wood in conifer and broadleaf forests; late spring, summer, and fall. Distribution and frequency: eastern Canada south to Florida, west to Tennessee and Michigan; occasional. Spore print: pinkish brown to reddish brown. Microscopic features: spores 10–17 × 5–8 µm, narrowly ovoid to subellipsoidal, pitted, pale brown. Edibility: edible. Lookalikes: Some species of Tylopilus are similar and have the same sporeprint colors. However, they have smooth spores. Gyroporus castaneus (p. 209) and Gyroporus purpurinus (p. 213) have white pores that do not become pinkish brown at maturity, hollow stalks, yellow spore prints, and smooth spores. Austroboletus gracilis var. flavipes T. J. Baroni, Halling & Both (see photo, p. 69), differs from Austroboletus gracilis var. gracilis by having more yellowish colors overall and by lacking both anastomosing riblike lines on the stalk and reticulation. Austroboletus gracilis var. pulcherripes Both & Bessette (see photo, p. 69), differs from var. gracilis by having darker colors overall and a stouter stalk that is coarsely reticulate over the upper two-thirds.

Austroboletus   ·  71

Austroboletus subflavidus

Austroboletus subflavidus (Murrill) Wolfe = Tylopilus subflavidus Murrill = Porphyrellus subflavidus (Murrill) Singer Common name(s): White Coarsely-ribbed Bolete Overview: The term subflavidus means “nearly yellow,” a reference to the color of this bolete’s cap and stalk. Specimens of this beautiful bolete are often stunted because they dry out quickly in their sandy habitat. The white to pale-yellow cap and stalk, coarsely ribbed reticulation, and bitter-tasting flesh are the key identification features. Cap: 3–10 cm wide, convex, becoming nearly plane and usually slightly depressed in age, margin even; surface dry, finely velvety at first soon becoming cracked, white at first, becoming buff or yellowish to grayish, often with a pale pinkish-orange tinge in age; flesh white, unchanging when exposed or bruised; odor somewhat fruity; taste bitter. Pores and tubes: white to grayish, sometimes beaded with clear droplets at first, becoming pinkish at maturity, not staining when bruised; tubes 1–2 cm deep. Stalk: nearly equal or tapered downward, rarely enlarged at the base, solid; surface dry, concolorous with the cap, with thick, raised, and coarsely ribbed reticulation that has a pitted appearance; flesh yellow in the base and white above. Habitat and season: scattered or in groups under oak and pine; late spring, summer, and fall.

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Distribution and frequency: New Jersey Pine Barrens south to Florida, west to Mississippi along the Gulf Coast; infrequent or sometimes fairly common locally. Spore print: reddish brown. Microscopic features: spores 15–20 × 6–9 µm, fusoid, minutely pitted, pale brown. Edibility: inedible because of the bitter taste. Lookalikes: Tylopilus rhoadsiae (p. 391) is similarly colored and has bitter-tasting flesh. Its stalk is typically clavate with a pinched base and prominent brown reticulation that is not coarsely ribbed, thick, raised, or appearing pitted, and its spores are smaller and smooth. Gen us B aor a ngi a G. Wu & Zh u L . Ya ng The genus Baorangia was erected in 2015 to accommodate species formerly classified in the genus Boletus. The name Baorangia is a combination of the Chinese words bao, meaning “thin,” and rang, meaning “hymenium,” a reference to short tubes. Species in this genus have tubes that are one-fifth to one-third the thickness of the cap flesh at a position halfway to the cap center. They have yellow tubes and pores that stain blue when bruised and yellow flesh that is unchanging or that slowly and weakly stains blue when exposed. At this time, there are four known species of Baorangia worldwide, with only one occurring in eastern North America.

Baorangia bicolor complex (A)

Baorangia bicolor complex (B)

Baorangia bicolor complex (C)

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Baorangia bicolor complex (D)

Baorangia bicolor complex (Kuntze) G. Wu, Halling & Zhu L. Yang = Baorangia rubelloides G. Wu, Halling & Zhu L. Yang = Boletus bicolor Peck = Boletus rubellus subsp. bicolor (Kuntze) Singer = Boletus bicolor var. subreticulatus A. H. Sm. & Thiers Boletus bicolor Massee and Boletus bicolor Raddi are names assigned to different species. Common name(s): Red and Yellow Bolete, Two-toned Bolete Overview: The term bicolor means “two colors.” The Red and Yellow Bolete is generally considered to be a good edible but is easily confused with similar species that are suspected to be poisonous or of uncertain edibility. Its distinguishing features are a red cap and stalk, bright-yellow pores that quickly stain blue, a tube layer that is shallow in proportion to the thickness of the flesh, and yellow flesh that is unchanging or slowly and weakly stains blue when exposed. Although most collections exhibit the macroscopic and microscopic features that closely agree with those provided in the original description, some collections display a wide variation in color and staining of the bruised cap surface, the degree of blue staining of the flesh, and the amount of the red color present on the stalk. Odor of the flesh is also inconsistently interpreted. These differences strongly suggest a group of closely allied species that ultimately will require molecular analysis to confirm their relationship. Until more information is available, we have

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decided to treat Baorangia bicolor as a complex and have included photos of several examples that illustrate this variability. Cap: 5–12.5 cm wide, convex to broadly convex, becoming nearly plane in age, margin incurved at first; surface dry, somewhat velvety when young, often cracking in dry weather or with age, dark red to rose red or brick red, rarely yellow, fading to ochraceous or tan with age, sometimes staining blue when bruised; flesh yellow, unchanging or slowly and weakly staining blue when exposed; odor not distinctive or reminiscent of cheese in older specimens; taste not distinctive. Pores and tubes: bright yellow, becoming olive yellow in age, quickly staining blue when bruised; tubes 3–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, smooth, yellow at the apex, red or rosy red on the lower two-thirds or more, unchanging or sometimes slowly staining blue when bruised, typically lacking reticulation but sometimes reticulate on the upper portion; flesh usually bright golden yellow, deeper yellow than the cap, unchanging or erratically staining blue. Habitat and season: solitary, scattered, or in groups under oaks; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: olive brown. Microscopic features: spores 8–12 × 3.5–5 µm, oblong to slightly ventricose, smooth, pale brown. Edibility: edible with caution and very good. Lookalikes: Boletus bicoloroides A. H. Sm. & Thiers (not illustrated), reported from Michigan, has a deep vinaceous-red cap that slowly fades to dull ochraceous with red splashes and does not become cracked in age. It has yellow flesh that stains blue when exposed, yellow pores that stain blue when bruised, tubes up to 15 mm deep, and a dark-red stalk from the base to the apex. Microscopically, it has large strongly amyloid spores that measure 14–17 × 4–5.5 µm and strongly dextrinoid pileus trama. The combination of the strongly amyloid spores with the strongly dextrinoid pileus trama is most unusual and clearly separates this species from other similar boletes. Boletus sensibilis (p. 150) quickly bruises blue on all parts and has a mostly yellow stalk and longer tubes, and its flesh has a sweet odor of fenugreek or curry. Boletus pallidoroseus (p. 129) is somewhat similar. It has a pale rose-pink to pale purplish-pink cap typically with a yellow margin and flesh with an odor of beef or chicken bouillon. Also compare with Boletus rufomaculatus (p. 148), which has a mottled brick-red to brownish-red cap with a narrow band of sterile tissue and paler-yellow stalk flesh and grows under beech.

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Gen us B ol e t e l lu s Mu r r i ll The genus Boletellus was erected in 1909 to accommodate species formerly classified in the genera Boletus and Suillus. The name Boletellus means “small Boletus.” It is a fairly small genus with less than a dozen species known from North America. They are small- to medium-size terrestrial or lignicolous boletes that closely resemble species of the genus Boletus. Their caps are typically dry, somewhat velvety or scaly, and often cracked in age, sometimes with a sterile projecting margin. Stalks are typically solid at maturity and lack scabers and resinous dots. They have white to yellow flesh that often stains blue when cut. Spore-print colors may be olive brown or dark rusty brown to dark brown. Microscopically, their spores are ornamented with longitudinal ridges, striations, or wings.

Boletellus ananas (A)

Boletellus ananas (M. A. Curtis) Murrill = Boletus ananas M. A. Curtis Common name(s): Pineapple Bolete Overview: The Pineapple Bolete is a distinct species unlikely to be confused with any other southern bolete. The term ananas refers to the genus of pineapple because of the scales on the cap that extend beyond the cap margin, which give this bolete its unique appearance.

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Boletellus ananas (B)

Cap: 3–10 cm wide, obtuse at first, becoming convex to broadly convex; surface dry, coated with coarse and overlapping purplish-red to dark-red scales that become dull pinkish tan to dingy yellow in age and extend beyond the margin; flesh whitish on exposure, quickly becoming yellowish and then blue and finally bluish gray; odor and taste not distinctive. Pores and tubes: yellow when fresh, sometimes tinged reddish brown in age, staining blue when bruised, covered when young by a whitish partial veil that usually leaves remnants on the cap margin and normally does not form a ring on the stalk; tubes 9–16 mm deep. Stalk: enlarged downward or nearly equal; surface dry, glabrous to slightly fibrillose, white to pale tan, sometimes with a reddish zone near the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups, often on the bases of trees or on the ground, under oaks and pines; year-round. Distribution and frequency: North Carolina south to Florida, west to Texas; occasional. Spore print: dark rusty brown to dark brown. Microscopic features: spores 15–24 × 7–11 µm, fusoid, with spirally arranged, conspicuous, longitudinally ridged, thin wings and often with an indistinct apical pore, pale brown.

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Edibility: edible. Lookalikes: Suillellus pictiformis Murrill = Boletellus pictiformis (Murrill) Singer (not illustrated) is a similar and rare species known only from Florida. It has a dry cap coated with appressed to erect, shaggy, medium-brown to darkbrown scales. Its cap becomes cracked in age, with yellow flesh showing in the fissures. Its stalk is sheathed with shaggy to fibrillose, medium-brown to dark-brown scales, and its greenish-yellow to olive-yellow pores stain blue when bruised. Boletellus flocculosipes (Murrill) Perr.-Bertr. (not illustrated), known only from Gainesville, Florida, has a dry, coarsely shaggy, dark grayish-brown to grayish-black cap, pale-yellow flesh with a pinkish zone near the cuticle that stains blue when exposed; sulfur-yellow pores that become dingy yellow to brownish at maturity and rapidly stain dark blue green when bruised; and a dry, shaggy, reddish-brown stalk with a glabrous yellow apex and a dark grayish-brown to grayish-black base. It has spores that measure 12–14 × 6–7 µm, and it grows in broadleaf woods near hickory.

Boletellus chrysenteroides

Boletellus chrysenteroides (Snell) Snell = Boletus chrysenteroides Snell Common name(s): none. Overview: This small, dark bolete is unusual in that it most often grows on decaying wood rather than in soil. The epithet chrysenteroides means

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“resembling chrysenteron,” a reference to its similarity with Xerocomellus chrysenteron (p. 416). Cap: 3–6 cm wide, convex, becoming broadly convex, margin even; surface dry, dark brown and velvety when young, becoming bay brown to chestnut brown and often cracked in age, with whitish to pale-yellow flesh that lacks reddish tints showing in the fissures; flesh whitish to pale yellow, staining blue, often slowly, when exposed; odor and taste not distinctive. Pores and tubes: pale yellow, becoming yellow to greenish yellow, quickly staining blue when cut or bruised; tubes 1.2–1.8 cm deep. Stalk: equal or enlarged slightly downward, solid; surface dry, reddish brown to blackish brown, often yellowish at the apex, often decorated with punctae that may align to simulate a net-like pattern, staining blue then slowly reddish when bruised or handled; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or in groups on decaying logs, stumps, at the base of standing trunks, or sometimes on the ground, in broadleaf or mixed oak and pine woods, especially with oak; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to fairly common. Spore print: olive brown to dark brown. Microscopic features: spores 10–16 × 5–8 µm, narrowly ovate to nearly oblong, longitudinally striate, yellowish to brownish. Edibility: edible. Lookalikes: Xerocomellus chrysenteron (p. 416) grows on the ground and has a dark-olive to olive-brown or grayish-brown cap, often with a reddish zone at the margin. Its cap soon becomes cracked and shows a red to pinkish color in the fissures. Xerocomellus intermedius (p. 419) has a cap that is grayish olive to brownish olive or red to rose red with olive tints and becomes cracked in age, with pinkish tints showing in the fissures. It grows on the ground. Boletellus pseudochrysenteroides (p. 80) grows on the ground or on decaying wood in broadleaf woods, especially with beech or oak. It has a dark rose-red to red-brown cap when young that typically becomes olivaceous to ochraceous brown and develops prominent cracks as it ages. It has whitish to pale-yellow flesh that quickly stains blue when exposed, and its stalk base is sometimes coated with yellowish mycelium.

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Boletellus pseudochrysenteroides

Boletellus pseudochrysenteroides A. H. Sm. & Thiers Common name(s): none. Overview: The term pseudochrysenteroides means “resembling Boletellus chrysenteroides” (p. 78). The key identification features include the dark rosered to red-brown cap when young, blue-staining pores, and large spores with sublongitudinal ridges. NH4OH stains the cap yellowish. Cap: 5–10 cm wide, broadly convex, expanding to plane or nearly so, margin with a narrow band of sterile tissue when young, even at maturity; surface dry, velvety, dark rose red to red brown at first, typically developing olivaceous to ochraceous-brown tones as it ages, soon becoming cracked and showing pale-yellow to pinkish flesh in the spaces, not staining blue when bruised; flesh pale yellow, instantly staining pink under the cuticle and pale blue above the tubes when exposed; odor not distinctive; taste acidic or not distinctive. Pores and tubes: pale yellow at first, becoming olive brown at maturity, staining blue and then slowly brown when bruised, strongly depressed at the stalk in age; tubes 8–15 mm deep, staining blue when bruised. Stalk: nearly equal or slightly enlarged downward, solid; surface scurfy, dull yellow brown or colored like the cap, sometimes with a yellow apex; covered on the upper half or more by reddish to brownish punctae; not staining when bruised; base sometimes coated with yellowish mycelium; flesh pale yellow to ochraceous, quickly staining blue when exposed.

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Habitat and season: solitary or scattered under broadleaf trees, especially beech or oak, sometimes on decaying logs or stumps; summer and fall. Distribution and frequency: Reported from Michigan and North Carolina, distribution limits yet to be established; rare. Spore print: dark olive brown to dark olive fuscous. Microscopic features: spores 11–18 × 5–8 µm, subellipsoidal to fusoid, smooth, thick-walled, ornamented with nine to twelve sublongitudinal ridges, with an apical break in the outer wall in the form of a slit, brown to ochraceous. Edibility: unknown and too rare to gather for food. Lookalikes: Boletellus chrysenteroides (p. 78) has a dark-brown cap that is velvety when young and becomes bay brown to chestnut brown and often cracked in age. It has whitish to pale-yellow flesh that lacks reddish tints showing in the fissures, and it grows on decaying logs and stumps, at the base of standing trunks, or sometimes on the ground in broadleaf or mixed oak and pine woods, especially with oak. Xerocomellus intermedius (p. 419) has weakly striate spores that measure 9–12 × 4–5 µm. Gen us B ol e t i n e l lu s Mu r r i ll The genus Boletinellus was erected in 1909 to accommodate some species formerly classified in several genera, including Daedalea, Boletus, and Suillus. Boletinellus is a small genus of medium-size boletes found in Asia and eastern North America. They have a dry to slightly viscid, typically smooth, and soft cap; yellow flesh that sometimes stains blue or greenish blue; radially arranged and elongated pores; and an eccentric to lateral stalk. The spore-print color is olive brown.

Boletinellus merulioides

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Boletinellus merulioides (Schwein.) Murrill = Gyrodon merulioides (Schwein.) Singer Common name(s): Ash Tree Bolete Overview: The term merulioides means “resembling Merulius,” referring to a genus of resupinate crust fungi with a wrinkled and ridged fertile surface. This unique and fascinating bolete is easily recognized by the dark reddish-brown cap, eccentric stalk, yellow radiating pores, and association with ash trees. Although the Ash Tree Bolete occurs with ash trees, it is not mycorrhizal with them. The Ash Tree Bolete has struck up a relationship with a parasitic aphid that is sheltered in sclerotia produced by the bolete. The sclerotia surround and protect the aphid, which provides honeydew nutrients to the bolete. For additional information on this relationship, see Brundrett and Kendrick 1987. Cap: 4–12.5 cm wide, often kidney-shaped in outline, convex with an incurved margin at first, becoming depressed in the center to nearly funnel-shaped in age; surface slightly velvety at first, smooth, slightly viscid and shiny when moist, yellow brown to olive brown or reddish brown; flesh thick in the center, thin at the margin, yellow, unchanging or sometimes staining bluish when exposed, reddish brown around larval tunnels; odor musty or unpleasant or not distinctive; taste variously described as pungent, resembling raw potatoes, or not distinctive. Pores and tubes: decurrent, not easily separated from the cap flesh, with elongated and radially arranged pores and gill-like ridges and crosswalls, yellow to dull golden yellow or olive, usually slowly bruising greenish blue; tubes 3–6 mm deep. Stalk: short, solid, eccentric to nearly lateral or central, often curved and pinched at the base; surface dry, concolorous with the pores on the upper portion, brownish to blackish at the base, sometimes mottled with reddish brown, staining reddish brown when bruised; flesh concolorous with and staining like the cap flesh, usually reddish brown near the base. Habitat and season: solitary or more often in groups on the ground or infrequently on wood, near or under ash trees; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Alabama and Wisconsin; fairly common. Spore print: olive brown. Microscopic features: spores 7–11 × 6–7.5 µm, broadly ellipsoidal to nearly globose, smooth, pale yellow in water mounts, ochraceous in KOH. Edibility: edible but usually rated as mediocre. Lookalikes: Boletinellus proximus (Singer) Murrill (not illustrated), known only from Florida, has a dark-brown to purplish-brown cap that fades to brown or cinnamon buff in age; pale-brown flesh that does not stain when exposed; decurrent, yellow to yellow-brown or olive-brown, angular to

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irregular, often elongated and radially arranged pores; and an eccentric to central stalk that is yellow or concolorous with the cap. Its spores measure 7–10 × 5–7 µm, and it grows on the ground or on decaying wood in swampy woods. Gen us B ol e t u s L . The genus Boletus was erected in 1753 to accommodate species formerly classified in several genera, including Agaricus, Suillus, and Polyporus. The name Boletus means “fleshy-pored fungus.” It is the largest genus of boletes, with more than one hundred species known from North America. Recent molecular research involving DNA analysis has resulted in the placement of several species formally classified as Boletus into other preexisting genera or into newly erected genera. It can be expected that this process will continue into the foreseeable future. Members of the genus Boletus are small to large terrestrial or sometimes lignicolous boletes with dry or viscid caps and typically solid stalks. Their flesh is white or yellow and stains blue or brown in some species when cut. Pores may be white, yellow, orange, red, brown, or grayish and may stain blue, brown, blackish, or some other color when bruised. Their stalks can be smooth and reticulate or have raised, intersecting, longitudinal ribs that mimic reticulation.

Boletus abruptibulbus (A)

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Boletus abruptibulbus (B)

Boletus abruptibulbus Roody, Both & B. Ortiz Common name(s): Abruptly Bulbous Sand Bolete Overview: The term abruptibulbus means “having an abruptly bulbous base,” a reference to this unusual bolete’s stalk base. The strongly bulbous to abruptly bulbous base, the prominent pseudorhiza, the fairly large spores, and growth in coastal woodlands are the diagnostic features that set this species apart from other boletes. The cap surface and bulbous base of dried material immediately stain dark red and then slowly fade with NH4OH. Cap: 3–8 cm wide, convex, becoming broadly convex, with a narrow sterile margin; surface dry to subviscid when wet, shiny, glabrous, occasionally appressed-fibrillose, often developing tiny cracks in age, reddish brown or dark brown, becoming cinnamon brown; flesh white to very pale yellow, unchanging when exposed; odor pleasant; taste not distinctive. Pores and tubes: narrowly depressed at the stalk, with short decurrent teeth, angular to nearly gill-like pores near the stalk, pale yellow at first, becoming pale golden yellow and then yellowish olive to greenish olive in age; tubes 3–8 mm deep. Stalk: nearly equal on the upper portion, strongly bulbous to abruptly bulbous on the lower portion, solid, with a prominent pseudorhiza covered in sand; surface dry, pallid or yellowish at the apex, gradually darker and becoming concolorous with the cap toward the base, with white basal mycelium; flesh yellow, unchanging when cut. Habitat and season: solitary or in scattered groups in sandy soil in coastal oak and pine woods; winter and spring. Distribution and frequency: at present known only from the Gulf Coast of the Florida Panhandle and southeastern Georgia; occasional.

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Spore print: olive brown. Microscopic features: spores 13.5–19.8(22.5) × 5–7.2 µm, smooth fusoid, grayish yellow in KOH, dextrinoid in Melzer’s. Edibility: edible but usually permeated with sand when collected. Lookalikes: Aureoboletus auriporus (p. 60) has similar colors and is shiny and viscid. Its pores are brilliant yellow; it has smaller spores; and it lacks a strongly developed pseudorhiza.

Boletus albisulphureus (A)

Boletus albisulphureus (B)

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Boletus albisulphureus (Murrill) Murrill = Gyroporus albisulphureus Murrill Common name(s): Chalk-white Bolete Overview: The term albisulphureus means “white and yellow,” referring to this handsome bolete’s overall colors. The key identification features include the white cap; white, unchanging flesh; yellow pores; and a white, reticulated stalk. Cap: 4–12 cm wide, obtuse to convex, becoming broadly convex at maturity, margin even; surface dry, glabrous, sometimes finely cracked in age, milk white to grayish white, sometimes with yellowish or brownish tinges, especially near the margin; flesh white, unchanging when exposed; odor somewhat pungent and medicinal or not distinctive; taste not distinctive. Pores and tubes: white to buff at first, becoming yellow at maturity and then dingy olive yellow in age, not staining blue when bruised, attached to subdecurrent; tubes 5–15 mm deep. Stalk: nearly equal or enlarged downward, sometimes abruptly narrowed at the base, solid; surface dry, milk white to whitish, yellowish at the apex, with white to yellowish reticulation on the upper portion; flesh concolorous with the cap flesh, typically with yellow tints at the base. Habitat and season: solitary, scattered, or in groups in sandy soil under oaks; summer, fall, and early winter in the southern part of its range. Distribution and frequency: Maine south to Florida, west to Texas; infrequent. Spore print: olive brown. Microscopic features: spores 11–15 × 3.5–4.5 µm, subfusiform, smooth, hyaline to pale yellowish, some dextrinoid in Melzer’s. Edibility: edible. Lookalikes: This species is often confused with Xanthoconium stramineum (p. 413), which is similar but has brighter-yellow pores and lacks reticulation on its stalk. Tylopilus rhoadsiae (p. 391) has a white cap and stalk that often have pinkish or pinkish-tan tinges, prominent white to brown reticulation, white pores that becomes dull pinkish at maturity, and bitter-tasting flesh.

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Boletus alutaceus

Boletus alutaceus Morgan = Xerocomus alutaceus (Morgan) E. A. Dick & Snell Common name(s): none. Overview: The term alutaceus means “leather-colored.” The key identification features include the brown to tan cap; whitish flesh that does not stain when exposed; yellow, nonstaining pores; and brownish stalk. Cap: 4–6 cm wide, pulvinate, becoming broadly convex to nearly plane; surface dry, unpolished, yellowish brown at first, becoming pale tan with a pinkish tint; flesh whitish, sometimes with a pinkish tint, not staining when exposed; odor and taste not distinctive. Pores and tubes: pale yellow to pale greenish yellow, not staining when bruised, depressed at the stalk; tubes 5–15 mm deep. Stalk: slightly enlarged downward or nearly equal, solid; surface glabrous, with or without longitudinal striations, sometimes weakly reticulate at the very apex, concolorous with the cap or paler, basal mycelium white; flesh white to pale tan, often with pinkish tints. Habitat and season: solitary, scattered, or in groups under oak and beech; summer and fall. Distribution and frequency: New England south to West Virginia and Kentucky, west to Ohio and Wisconsin; rare. Spore print: olive brown. Microscopic features: spores 9–12 × 4–5 µm, ellipsoid to subfusiform, smooth, very pale yellow to nearly hyaline.

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Edibility: reported to be edible but too rare to collect for the table. Lookalikes: This small and seldom-encountered bolete might be confused with the similar bolete Aureoboletus roxanae (p. 67), which has a yellow-brown to reddish-brown cap that becomes dull orange to dark orange yellow with age and pores that are whitish at first but then become pale yellow and finally pale golden yellow. The stalk of Aureoboletus roxanae is yellowish brown to reddish brown, often with a dull-orange zone at the apex, and is longitudinally striate, at least on the upper portion.

Boletus atkinsonii (A)

Boletus atkinsonii (B)

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Boletus atkinsonii Peck Common name(s): none. Overview: This bolete was named in honor of American mycologist George F. Atkinson (1854–1918). The roughened cap surface with tufts of hyphae is a noteworthy feature. NH4OH stains the cap purplish. Cap: 4–20 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, roughened with tufts of hyphae that are best observed with a hand lens, minutely cracked at maturity, grayish brown or yellowish brown; flesh white, unchanging when exposed; odor and taste somewhat nutty sweet or not distinctive. Pores and tubes: white at first, soon yellow to pale ochraceous, sometimes depressed at the stalk at maturity; tubes 6–12 mm deep. Stalk: slightly enlarged downward or nearly equal, solid; surface dry, dingy white, with fine brownish reticulation; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in mixed broadleaf woods, usually with beech or oak; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, western distribution limits yet to be determined; occasional. Spore print: ochraceous brown. Microscopic features: spores 10–13 × 4–5 µm, ellipsoid to subfusiform, smooth, pale yellow; hyphae of the cap cuticle roughened and in fascicles. Edibility: edible. Lookalikes: Boletus nobilis (p. 121) has a paler, more ochraceous cap that is felt-like to nearly smooth, lacks tufts of hyphae, and has larger spores that measure 12.5–15 × 5–6 µm. Boletus variipes (p. 167) has variable cap colors ranging from creamy tan to dark brown, coarser and more prominent reticulation, and larger spores that measure 12–18 × 4–6 µm.

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Boletus aurantiosplendens

Boletus aurantiosplendens Baroni Common name(s): Showy Orange Bolete Overview: The term aurantiosplendens means “showy orange.” This bolete’s key identification features include the orange to brownish-yellow cap, pale-yellow flesh, and obscurely to distinctly reticulate, yellow to orange stalk. Cap: 3–11 cm wide, convex, becoming broadly convex to nearly plane in age, margin inrolled at first, becoming decurved, even; surface dry, somewhat velvety, color variable, orange to brownish orange or brownish yellow, sometimes duller yellowish brown or pale brown to rusty brown with a brighter yellowish-orange margin; flesh pale yellow to yellow, slowly darkening when exposed; odor pungent when fresh; taste not distinctive. Pores and tubes: yellow, becoming dull brownish yellow in age, unchanging or staining brownish orange when bruised, sometimes depressed at the stalk at maturity; tubes 5–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, glabrous, obscurely to distinctly reticulate, yellow to apricot or orange, with tawny to reddish-brown streaks over the midportion, with yellow basal mycelium; flesh concolorous with the cap flesh.

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Habitat and season: scattered or in groups in mixed broadleaf and pine woods; summer and fall. Distribution and frequency: Mississippi and North Carolina, distribution limits yet to be established; uncommon. Spore print: olive brown. Microscopic features: spores 9.5–12 × 4–5.5 µm, subfusiform, smooth, pale olivaceous yellow. Edibility: unknown. Lookalikes: This beautiful bolete is sometimes confused with Pulveroboletus auriflammeus (p. 291), which has a brilliant yellow-orange powdery cap, a stalk that stains fingers after being held, and yellow-orange pores that become olive yellow to greenish yellow and sometimes develop bright-crimson to crimson-orange tints in age. Boletus melleoluteus (Snell, E. A. Dick & Hesler) T. J. Baroni & Halling (not illustrated), known only from Tennessee and North Carolina south to Georgia, has a bright-yellow cap that becomes dull yellow ochraceous in age, a pale-yellow stalk that lacks reticulation, and pale-yellow flesh that blues slightly when exposed. Also compare with Boletus aureissimus var. aureissimus (p. 92), which has a bright-yellow to honey-yellow or yellow-ocher cap, a yellow to brightyellow stalk with or without delicate reticulation on the upper portion, and mild-tasting flesh.

Boletus aureissimus var. aureissimus

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Ceriomyces aureissimus var. castaneus

Boletus aureissimus var. aureissimus (Murrill) Murrill = Boletus auripes var. aureissimus (Murrill) Singer = Ceriomyces aureissimus Murrill Common name(s): none Overview: The term aureissimus means “golden yellow.” This bolete’s key identification features include the yellow cap, yellow and unchanging flesh, yellow pores, and yellow stalk that typically has delicate yellow reticulation on the upper portion. Cap: 4–14 cm wide, obtuse to convex, becoming broadly convex to nearly plane in age, margin even; surface dry, glabrous, honey yellow to bright yellow or yellow ocher, often duller in age; flesh yellow, not staining blue when exposed; odor and taste not distinctive.

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Pores and tubes: pale yellow to yellow, becoming olive yellow at maturity, not staining blue when bruised, often depressed at the stalk in age; tubes 1–2.5 cm deep. Stalk: typically bulbous to clavate when young, becoming nearly equal at maturity, solid; surface dry, yellow to bright yellow, with delicate yellow reticulation on the upper portion or sometimes lacking reticulation; flesh yellow, unchanging when exposed. Habitat and season: solitary, scattered, or in groups under oaks; summer and fall. Distribution and frequency: Piedmont region of North Carolina south to Florida, west to Texas; occasional. Spore print: yellow brown to olive brown. Microscopic features: spores 10–14 × 3–5 µm, ellipsoid-cylindric to subfusoid, smooth, yellowish. Edibility: edible. Lookalikes: Ceriomyces aureissimus var. castaneus Murrill (see photo, p. 92) is a nearly identical beautiful bolete with a velvety, purplish-brown cap surface. Boletus auripes (p. 94) has a yellow-brown to chestnut-brown or grayish-brown cap and a golden-yellow stalk with yellow reticulation. Retiboletus ornatipes (p. 300) has a darker-yellow or gray cap, coarser and more prominent reticulation, and bitter-tasting flesh. Boletus aurantiosplendens (p. 90) has a darker-orange to brownish-orange or brownish-yellow cap, a yellow to apricot or orange stalk with tawny to reddish-brown streaks over the midportion and obscure to distinct reticulation, and yellow flesh that darkens or becomes bright orange when exposed.

Boletus auripes (A)

94  ·  Boletes of Eastern North America

Boletus auripes (B)

Boletus auripes Peck Common name(s): none. Overview: The term auripes means “golden-yellow foot.” The rich brown and yellow colors of this handsome bolete and the lack of blue staining anywhere when bruised are its distinctive features. Cap: 4–13 cm wide, obtuse to convex, becoming broadly convex to nearly plane, with a narrow sterile margin; surface dry, finely velvety to nearly glabrous, yellowish brown to chestnut brown or grayish brown, becoming paler in age; flesh bright yellow, color intensifying when rubbed, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: pale yellow to yellow, becoming olive yellow at maturity, unchanging or staining yellow orange when bruised, often depressed at the stalk in age; tubes 1–2.5 cm deep. Stalk: often bulbous to clavate when young, clavate to nearly equal with age, solid; surface dry, golden yellow with yellow reticulation, at least on the upper portion; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak and beech, or sometimes in grassy places where oak is present; late spring, summer, and fall.

Boletus   ·  95

Distribution and frequency: eastern Canada south to Florida, west at least to New York and North Carolina, western distribution limits yet to be established; fairly common. Spore print: yellow brown to olive brown. Microscopic features: spores 10–14 × 3–5 µm, ellipsoid-cylindric to subfusoid, smooth, yellowish. Edibility: edible. Young, firm specimens are excellent. Lookalikes: Boletus aureissimus var. aureissimus (p. 92) is very similar. It has a honey-yellow to bright-yellow or yellow-ocher cap and more delicate reticulation on its stalk. Ceriomyces aureissimus var. castaneus Murrill (see photo, p. 92) has a velvety purplish-brown cap.

Boletus billieae

Boletus billieae Both, Bessette & W. J. Neill Common name(s): Billie’s Bolete Overview: This rare and beautiful bolete is currently unknown beyond Cape Cod, Massachusetts. It is named in honor of Billie Both (1935–2007), wife of the eminent American boletologist Ernst E. Both (1930–2012), who first discovered it. The dark chocolate-brown to very dark purplish-brown cap and red stalk with yellow reticulation are its distinctive features. The cap surface stains dark amber and then red brown with KOH, amber with NH4OH, and bluish to grayish green with FeSO4.

96  ·  Boletes of Eastern North America

Cap: up to 16 cm wide, broadly convex, margin wavy, strongly incurved, even; surface dry, covered with tiny matted fibers, opaque to subshiny, dark chocolate brown to dark purplish brown, staining dingy reddish when bruised; flesh pale yellow, slowly staining blue when exposed; odor not distinctive when fresh, described as reminiscent of chocolate-chip cookies when dried; taste sweetish or not distinctive. Pores and tubes: golden yellow at first, becoming greenish yellow and finally olivaceous in age, quickly staining blue when bruised; tubes 8–10 mm deep. Stalk: enlarged downward with a pinched base, solid; surface dry, blood red on the upper portion with a hint of purplish red, pinkish red on the lower portion, very dark red at the base, covered with a yellow reticulum extending from the apex to the base, with a white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, in sandy soil under bear oak and pitch pine; summer, seasonal range yet to be determined. Distribution and frequency: known only from Cape Cod, Massachusetts; rare. Spore print: olivaceous brown. Microscopic features: spores 12–16 × 4–6 µm, subfusoid to ellipsoid, smooth, hyaline. Edibility: unknown. Lookalikes: none.

Boletus carminiporus

Boletus   ·  97

Boletus carminiporus Bessette, Both & Dunaway Common name(s): none. Overview: The red cap, red stalk, red pores, and white to yellow flesh that does not stain blue are the key features for identifying this bolete. The term carminiporus means “carmine-red pores.” Cap: 3–14 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved to inrolled at first, becoming decurved at maturity, with a narrow band of sterile tissue; surface dry to slightly viscid, glabrous or nearly so, dull red at first, becoming pinkish red to orange red and fading to reddish orange to dull golden orange in age; flesh whitish to pale yellow, becoming darker yellow when exposed or in age, not staining blue at all; odor and taste not distinctive. Pores and tubes: yellow when very young, soon becoming dark red to brownish red, fading to dull red or orange red in age, staining bluish green and then dull olive when bruised, depressed at the stalk at maturity; tubes 3–12 mm deep, yellow, staining bluish green when cut. Stalk: enlarged downward to a pinched base, rarely tapered downward or nearly equal, solid; surface dry, distinctly reticulate overall or at least on the upper portion, rose pink at first, soon becoming dark red at the apex and paler red below, staining brownish red or slowly olive green to olive yellow when handled or bruised; flesh pale yellow to yellow, darker than the cap flesh, unchanging when exposed, becoming dull red around larval tunnels. Habitat and season: solitary, scattered, or in groups in mixed broadleaf forests, especially with beech, hickory, and oak, or in mixed woods with oak and pine; summer and fall. Distribution and frequency: southeastern United States from North Carolina south to Florida, west to Arkansas and Louisiana, northern distribution limits yet to be established; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 8–11 × 3–4 µm, subfusoid, smooth, ochraceous. Edibility: unknown. Lookalikes: Boletus flammans (p. 107) has yellowish flesh that quickly stains blue when exposed, and it grows under conifers, especially spruce, hemlock, and pine. It also has larger spores, measuring 10–16 × 4–5 µm. Suillellus subluridus (p. 314) is also very similar but lacks reticulation on the stalk. Lanmaoa borealis (p. 229), known from New England west to Michigan, has an evenly brick-red to dark rusty-rose cap. Its stalk is concolorous with the cap and lacks reticulation or is reticulate only at the very apex. The cap flesh is yellow to pale yellow. The stalk flesh is lemon yellow and slowly stains blue at the apex when exposed. Its pores are dull orange to orange red and stain blue when bruised. Its spores measure 11–15 × 4–5 µm.

98  ·  Boletes of Eastern North America

Boletus chippewaensis

Boletus chippewaensis A. H. Sm. & Thiers Common name(s): none. Overview: The term chippewaensis is a reference to Chippewa County, Michigan, where this bolete was first collected. Although it has a few distinctive features, including a pale lemon-yellow cap splashed with brick red to rusty red, pores that stain pinkish cinnamon, and wide spores, it is clearly a member of the Boletus edulis group and has been identified and illustrated in field guides as Boletus edulis (p. 101). We have decided to treat Boletus chippewaensis as a distinct species, realizing, however, that in the future molecular analysis may very well indicate that it should be placed in synonymy with Boletus edulis. Cap: 6–15 cm wide, convex to broadly convex, becoming nearly plane in age, margin even; surface dry, smooth, pale lemon yellow splashed irregularly with brick red to bright rusty red, finally brick red overall; flesh white, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: white when very young, soon turning creamy lemon yellow, finally olive brown in age; staining pinkish cinnamon or sometimes grayish olive when bruised; tubes 1–2 cm deep. Stalk: nearly equal or enlarged downward, sometimes bulbous, often with a pinched base, solid; surface dry, pinkish tan to cinnamon brown on the upper portion or nearly overall, whitish at the base, with a white reticulum on the upper one-third or more; flesh concolorous with the cap flesh.

Boletus   ·  99

Habitat and season: solitary, scattered, or in groups under conifers, especially eastern white pine, or in mixed broadleaf and conifer woods; summer and fall. Distribution and frequency: New England and New York, west to Michigan, distribution limits yet to be established; uncommon to rare. Spore print: yellow brown. Microscopic features: spores 11–16 × 5–7 µm, ellipsoidal to fusiform, smooth, pale brown. Edibility: edible and choice. Lookalikes: Boletus edulis (p. 101), Boletus huronensis (p. 110), and Boletus subcaerulescens (p. 157) have brown caps that lack the pale lemon-yellow coloration in all stages of development.

Boletus durhamensis

Boletus durhamensis B. Ortiz, Bessette & McConnell Common name(s): none. Overview: The term durhamensis is a reference to Durham, North Carolina, where this bolete was first collected by American mycologist Owen McConnell. The cap surface stains red orange with KOH and slightly dull vinaceous with NH4OH and is negative with FeSO4. Cap: 2–12(17) cm wide, rounded at first, becoming convex to broadly convex, margin with a narrow overlapping band of sterile tissue, incurved when young; surface dry, felt-like to somewhat velvety, medium brown to rusty brown, becoming paler brown to yellowish brown with age, typically

100  ·  Boletes of Eastern North America

yellowish along the margin; flesh white, slightly tinged with yellowish tan, unchanging when exposed; odor and taste not distinctive. Pores and tubes: uniformly dark cinnamon brown, unchanging or slightly darker when bruised, attached to narrowly depressed near the stalk; tubes 2–6(11) mm deep, straw-colored, not staining when bruised. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, smooth, yellow with cinnamon-brown pruinosity that is sparse on young specimens and heavier on mature ones, typically with fine cinnamon-brown reticulation on the upper portion or at least near the apex, with white basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in broadleaf woods or in sparse grassy areas near willow oak and white oak, sometimes with red maple, sweetgum, and dogwood also present; summer and early fall. Distribution and frequency: reported only from the eastern Piedmont of North Carolina, distribution limits yet to be established; occasional. Spore print: olivaceous gray brown. Microscopic features: spores 7–12 × 3.7–5 µm, ellipsoid, smooth, hyaline; pleurocystidia and cheilocystidia present. Edibility: unknown. Lookalikes: Boletus vermiculosoides (p. 169) and Boletus vermiculosus Peck (see photo, p. 169) are similar, but both bruise dark blue on the pores, and their cut flesh instantly stains blue. Tylopilus tabacinus (p. 398) has a pinkish-brown to reddish-brown spore print, more pronounced reticulation, and typically bitter-tasting flesh, and it lacks cheilocystidia.

Boletus edulis (A)

Boletus   ·  101

Boletus edulis (B)

Boletus edulis Bull. Common name(s): Cep, King Bolete, Porcini, and Steinpilz Overview: The term edulis means “edible,” which is an understatement when referring to this truly delicious bolete. In fact, the Cep is arguably the most popular and coveted of all edible wild mushrooms wherever it exists in the world. Several different color forms and varieties occur in eastern North America. The key identification features for members of this group include white pores that become yellow at maturity, white reticulation on the stalk, mild-tasting flesh, and association with conifers. Cap: 4–20 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry or tacky, smooth or somewhat uneven and slightly pitted, brown to cinnamon brown or rusty red to tan; flesh white, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: white when young, becoming yellow to olive yellow and then brownish yellow to brown in age, typically unchanging when bruised, but occasionally bruising dull orange cinnamon to pale yellowish brown; tubes 1–3 cm deep. Stalk: enlarged downward and sometimes bulbous, thick, firm and solid; surface dry, white to pale brown, with a fine white reticulum overall or at least on the upper one-third; flesh white, unchanging when exposed. Habitat and season: solitary, scattered, or in groups under conifers, especially Norway spruce and pine; late spring through summer and fall.

102  ·  Boletes of Eastern North America

Distribution and frequency: eastern Canada south to Georgia, west to Minnesota and Arkansas; occasional to locally quite common. Spore print: olive brown. Microscopic features: spores 13–19 × 4–6.5 µm, ellipsoidal, smooth, pale yellowish brown. Edibility: edible and choice. Dried Ceps, better known as King Boletes, have a rich, concentrated flavor and are frequently available in supermarkets. Lookalikes: Boletus chippewaensis (p. 98) is nearly identical but has a pale lemon-yellow cap splashed irregularly with brick red to bright rusty red. Boletus subcaerulescens (p. 157) has more vinaceous tones on the cap and stalk, flesh that often has an odor of lemon, and pores that bruise blue gray. Boletus pseudopinophilus (p. 137) has a conspicuous brownish reticulum that darkens when handled. Boletus huronensis (p. 110) is a similar but dangerously poisonous species that can cause severe digestive distress and may have been responsible for at least one fatality. It has pale-yellow flesh that typically stains blue when exposed, sometimes slowly or occasionally not at all. Its yellow to brownish pores slowly stain greenish blue, and it lacks reticulation on the stalk or is reticulate only at the very apex. Boletus huronensis is associated with hemlock. Tylopilus felleus (p. 375) is sometimes mistaken for Boletus edulis but differs in having very bitter-tasting flesh, brown reticulation on the stalk, and a pinkish-brown spore print.

Boletus fairchildianus (A)

Boletus fairchildianus (B)

Boletus fairchildianus (C)

104  ·  Boletes of Eastern North America

Boletus fairchildianus (Singer) Singer = Boletus rubricitrinus var. fairchildianus Singer Common name(s): none. Overview: The term fairchildianus is a reference to the Fairchild Tropical Botanical Garden in Miami, Florida, where it was originally collected in 1942. The key identification features include the variable red cap that often bruises blue, yellow flesh that stains blue, red to orange-red pore surface, red or yellow stalk, and southern distribution. Cap: 3–14 cm wide, pulvinate to convex, becoming broadly convex to nearly plane or slightly depressed in age, margin incurved at first, with a narrow band of sterile tissue; surface dry, glabrous to somewhat velvety, color variable, dull rose red to dull brick red, brownish orange to cinnabar red, or vinaceous red, fading to tawny olive or reddish brown to dull brown with age, sometimes with yellow tints, often staining blue when bruised; flesh yellow, strongly staining blue when exposed; odor and taste not distinctive. Pore and tubes: at first red to dull red or orange red on a yellow ground color, often duller in age, bruising blue; tubes 8–20 mm deep. Stalk: tapered in either direction or nearly equal, with a ventricose base, solid; surface dry, yellow with red tints and streaks or dots especially near the base, sometimes with red over much of its length, usually weakly reticulate at the extreme apex, often longitudinally striate; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups with oak or in oak and pine woods; summer and fall. Distribution and frequency: Florida west to Texas, northern limits yet to be established. Spore print: olive brown. Microscopic features: spores 13–19 × 5–8 µm, fusoid to subfusoid-ellipsoid, smooth, yellowish. Edibility: unknown. Lookalikes: Boletus rubricitrinus (p. 145) has yellow to dull-yellow or olive-yellow pores and less red on the stalk. .

Boletus   ·  105

Boletus ferrugineus

Boletus ferrugineus Schaeff. = Boletus citrinovirens Watling = Boletus spadiceus Krombh. = Boletus spadiceus var. furcatus T. J. Baroni, Largent & Thiers = Boletus spadiceus var. gracilis A. H. Sm. & Thiers = Boletus spadiceus var. rufobrunneus Thiers = Xerocomus ferrugineus (Schaeff.) Alessio = Xerocomus spadiceus (Fr.) Quél. Common name(s): Rusty Iron Bolete Overview: The term ferrugineus means “rusty iron,” a reference to the color of this bolete’s cap. The cap surface displays a green flash and then stains reddish brown with NH4OH. The flesh stains olive green with FeSO4. Cap: 5–11 cm wide, convex, becoming broadly convex or sometimes nearly plane in age, margin sometimes with a narrow band of sterile tissue or even; surface dry, somewhat velvety, occasionally cracked in age, dark olive to olive yellow with reddish tints, yellow brown, or reddish brown; flesh yellow with a reddish line beneath the cuticle, pinkish around larval tunnels, unchanging or staining slightly blue when exposed; odor mild to slightly pungent; taste not distinctive. Pores and tubes: yellow to olive yellow, usually but not always staining blue or blue green when bruised; tubes 8–10 mm deep.

106  ·  Boletes of Eastern North America

Stalk: nearly equal or narrowed downward, typically with a pinched base, solid; surface dry, typically with raised longitudinal lines that may form a net-like pattern that mimics reticulation at the apex or nearly overall, pale yellow or whitish, often streaked with brown, whitish near the base, with a whitish to yellowish basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in small groups in mixed woods and along road banks and trails, usually under conifers; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; fairly common. Spore print: olive to pale olive brown. Microscopic features: spores 10–14 × 4.5–5 µm, oblong to ventricose, smooth, pale brown. Edibility: edible. Lookalikes: Boletus subtomentosus var. subtomentosus (p. 165) is very similar, but its pores less readily stain blue when bruised, and its cap surface instantly stains reddish brown with NH4OH. Xerocomellus chrysenteron (p. 416) has a dark-olive to olive-brown or grayish-brown cap. It often has a reddish margin and is conspicuously cracked, with reddish color visible in the cracks.

Boletus flammans

Boletus   ·  107

Boletus flammans E. A. Dick & Snell = Boletus rubroflammeus A. H. Sm. & Thiers Common name(s): none. Overview: The term flammans means “flaming or fiery,” a reference to this bolete’s overall red colors. The key identification features include the red cap that stains blue, yellow flesh that quickly stains blue when exposed, bright-red to orange-red pores that bruise blue, and a red, reticulated stalk that bruises blue. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin even or nearly so; surface dry to slightly viscid when moist, somewhat velvety to glabrous, dark red to brick red, rosy red, or brownish red, sometimes with tan areas near the margin in age, staining blue when bruised; flesh yellowish, quickly staining blue when exposed; odor and taste not distinctive. Pores and tubes: bright red to orange red, depressed at the stalk at maturity, staining blue when bruised; tubes 8–20 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, concolorous with the cap, sometimes yellow near the base, finely reticulate on the upper half or at least near the apex, staining blue when handled or bruised; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under conifers, especially spruce, hemlock, and pine, or sometimes under broadleaf trees, especially beech; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Texas; occasional. Spore print: olive brown. Microscopic features: spores 10–16 × 4–5 µm, cylindric to ellipsoid, smooth, yellowish. Edibility: unknown. Lookalikes: Rubroboletus rhodosanguineus (p. 304) has a similarly colored cap that first stains dark red to purplish then gray when bruised. It has flesh with an odor of overripe fruit and a sweet taste as well as a more prominently reticulate stalk, and it grows with red oak. Boletus carminiporus (p. 97) has whitish to pale-yellow flesh that does not stain blue when exposed; grows in mixed broadleaf forests, especially with beech, hickory, and oak, or in mixed woods with oak and pine; and has smaller spores that measure 8–11 × 3–4 µm. Suillellus subluridus (p. 314) is similar, but its stalk lacks reticulation. Lanmaoa borealis (p. 229), known from New England west to Michigan, has a dark-red to bright apple-red cap that becomes brick red to dull rusty rose in age. It has yellow to pale-yellow flesh that slowly stains blue at the stalk apex when exposed, orange-red to red pores that stain greenish blue when bruised, a stalk that is concolorous with the cap and lacks reticulation or is reticulate only at the very apex, and spores that measure 11–15 × 4–5 µm.

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Boletus gertrudiae

Boletus gertrudiae Peck Common name(s): Gertrude’s Bolete Overview: This bolete was named in honor of the late Gertrude Wells of Connecticut. The yellow coloration on the upper portion of the stalk is most distinctive. Cap: 5–17 cm wide, convex, becoming broadly convex in age, margin even or nearly so; surface dry or moist, soft, glabrous, pale brownish yellow to orange yellow and usually brighter yellow toward the margin when young,

Boletus   ·  109

becoming yellow to bright yellow with a paler disc at maturity; flesh white, unchanging when exposed; odor and taste not distinctive. Pores and tubes: white at first, becoming bright yellow and then brownish yellow to rusty yellow at maturity, attached or slightly depressed near the stalk; tubes up to 2 cm deep. Stalk: nearly equal down to an enlarged base, solid; surface dry, reticulate over the upper one-third or at least near the apex and glabrous below, white overall when young, becoming bright yellow over the upper portion and gradually extending downward until the stalk is yellow nearly overall; flesh white or sometimes yellowish near the apex, not staining when exposed. Habitat and season: solitary, scattered, or in groups in broadleaf forests or mixed woods, especially with beech or oak; summer and fall. Distribution and frequency: New England and western New York south to North Carolina, western limits yet to be established; uncommon. Spore print: color unknown, not provided in the original description. Microscopic features: spores 10–17 × 4–6 µm, subfusiform, smooth, hyaline to pale ochraceous. Edibility: unknown. Lookalikes: Boletus separans (p. 153) sometimes has a yellow zone near the stalk apex but usually has lilac tones on the cap and stalk. Boletus nobilis (p. 121) is similar but has an ochraceous to olive-ochraceous or reddish-ochraceous cap at maturity and a white to pale-ochraceous stalk that lacks bright-yellow coloration.

Boletus huronensis (A)

110  ·  Boletes of Eastern North America

Boletus huronensis (B)

Boletus huronensis A. H. Sm. & Thiers Common name(s): none. Overview: The term huronensis means “Huron,” a reference to the Canyon Lake area of the Huron Mountains of Michigan, where this species was first collected. The diagnostic features of this robust bolete include a dull yellow-brown to pale cinnamon-brown cap; pale-yellow flesh that typically stains blue when exposed; yellow to brownish-yellow pores that stain greenish blue, sometimes slowly; a thick stalk that is enlarged downward and lacks reticulation or is reticulate only at the apex; and association with hemlock. Cap: 5–25.5 cm wide, convex, becoming broadly convex or irregularly expanded in age, margin with a narrow band of sterile tissue, incurved to inrolled at first, often wavy to irregular in age; surface dry, somewhat velvety or glabrous, dull yellow brown, ocher brown, or pale cinnamon brown; flesh pale yellow, typically staining blue when exposed but sometimes slowly or not at all, with a reddish-brown line under the cuticle; odor of mercaptan or not distinctive; taste not distinctive. Pores and tubes: yellow at first, becoming olive yellow to brownish yellow at maturity, often depressed at the stalk in age, sometimes slowly or erratically staining greenish blue and then reddish brown when bruised; tubes 1–3.5 cm deep.

Boletus   ·  111

Stalk: enlarged downward, sometimes enlarged in the middle portion and tapered above and below, 2–7.5 cm thick, dense and heavy, solid; surface dry, yellow to yellowish at the apex, paler yellow to whitish below, sometimes streaked ocher to brownish or reddish brown, staining brownish where handled, sometimes reticulate only at the very apex; flesh mostly whitish, watery-streaked and grayish yellow in the base, brownish orange around larval tunnels. Habitat and season: solitary, scattered, in groups, or sometimes in fused clusters, under hemlock; summer and fall. Distribution and frequency: eastern Canada south to New York and Connecticut, west to Michigan; infrequent. Spore print: olive brown. Microscopic features: spores 12–15 × 4–5 µm, ellipsoidal to oblong, smooth, pale yellowish brown. Edibility: not recommended. Although eaten by some individuals without adverse effects, this bolete has caused numerous cases of severe gastrointestinal distress. Lookalikes: Boletus edulis (p. 101) has white flesh that does not stain blue when exposed, fine white reticulation that covers most of the stalk or at least the upper one-third, and white to brownish-yellow pores that do not stain blue, and it is associated with Norway spruce and pine.

Boletus longicurvipes

112  ·  Boletes of Eastern North America

Boletus longicurvipes Snell & A. H. Sm. = Boletus viscidocorrugis Both Common name(s): none. Overview: This typically small- to medium-size bolete is easily recognized. It lives up to the name longicurvipes, which means “long curved foot” in reference to a relatively thin stalk that is frequently curved near the base. The viscid yellow-orange to brownish-orange or ochraceous cap, yellow pores that do not stain blue when bruised, and the long, curved, punctate stalk are distinctive features. The cap surface stains cherry red to reddish orange with NH4OH and bright cherry red with KOH. Boletus viscidocorrugis was originally described as a distinct species that differed from Boletus longicurvipes by having white flesh that slowly to rapidly stains pink to red and then dingy slate, whitish punctae that do not darken in age, and white stalk flesh that stains bright blue in the base or erratically throughout. Molecular analysis has shown that the ITS sequence of Boletus viscidocorrugis is identical with that of Boletus longicurvipes (Manfred Binder, personal communication, November 5, 2014), and therefore we have placed it in synonomy. Cap: 2.5–11 cm wide, convex, becoming broadly convex at maturity, margin even; surface smooth to slightly wrinkled or pitted, viscid to tacky or glutinous, shiny, yellow orange to brownish orange or ochraceous, often developing green tones in wet weather or in age; flesh white to pale yellow, usually unchanging when exposed or sometimes slowly to rapidly staining pink and then brick red and finally dingy slate; odor not distinctive; taste slightly acidic or not distinctive. Pores and tubes: pale yellow at first, becoming dull greenish yellow to greenish gray, sometimes staining yellow or brownish when bruised; tubes 8–12 mm deep. Stalk: enlarged downward or nearly equal, typically curved on the lower portion, solid; surface dry, whitish to yellowish, sometimes with reddish or pinkish tan tones, with white to yellowish punctae that usually darken to reddish brown in age, with white basal mycelium; flesh white to pale yellow, usually unchanging when exposed or sometimes slowly to rapidly staining pink and then brick red and finally dingy slate, sometimes staining bright blue in the base or erratically overall. Habitat and season: solitary or scattered in broadleaf and mixed woods with oak, pine, and hemlock; summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Minnesota and Arkansas; fairly common. Spore print: olive brown. Microscopic features: spores 13–17 × 4–5 µm, oblong, smooth, pale brown. Edibility: edible and good when young, taste sweet and nutty.

Boletus   ·  113

Lookalikes: Because its stalk has whitish to yellow punctae that darken to reddish brown in age, Boletus longicurvipes could be mistaken for some Leccinum or Hemileccinum species. Hemileccinum subglabripes (p. 220), although similar, does not stain bright cherry red on the cap surface with the application of KOH.

Boletus luridellus (A)

Boletus luridellus (B)

114  ·  Boletes of Eastern North America

Boletus luridellus (Murrill) Murrill = Boletus subsensibilis (Murrill) Murrill = Ceriomyces subsensibilis Murrill Common name(s): none. Overview: The term luridellus means “dirty brownish,” a reference to the color of this bolete’s cap. Because of the variable cap and stalk color combinations, this species may be difficult to identify. The brown cap, yellow flesh that quickly stains blue, yellow pores that stain blue, brown stalk reticulation, and association with oak and pine are its key identification features. Cap: 4–12 cm wide, pulvinate, becoming broadly convex to nearly plane in age, margin with a narrow band of sterile tissue; surface dry, somewhat velvety to nearly glabrous, yellow brown to amber brown, reddish brown, or streaked dull brown over a yellow ground color, becoming ochraceous tawny to hazel in age; flesh yellow, quickly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow at first, becoming greenish yellow to olive yellow in age, depressed at the stalk at maturity, quickly staining blue when bruised; tubes 8–16 mm deep. Stalk: nearly equal or tapered in either direction, sometimes swollen near the middle, solid; surface dry, color variable, yellow overall or only near the apex, sometimes brownish red on the lower portion or overall, usually brownish punctate and streaked dark red at the base, quickly staining blue when handled or bruised, with brown or yellow reticulation at least over the upper portion; flesh yellow down to a red base, quickly staining blue when exposed. Habitat and season: scattered or in groups in grassy areas or in woods under oak and pine; summer and fall. Distribution and frequency: North Carolina south to Florida, west to Texas, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 12–17 × 4–6 µm, fusoid, smooth, pale yellow brown. Edibility: edible. Lookalikes: Lanmaoa pseudosensibilis (p. 233) has a dull-brown cap, bright-yellow flesh that stains blue when exposed, yellow pores that bruise blue, and a yellow stalk that is sometimes tinged reddish to purplish and lacks reticulation.

Boletus   ·  115

Boletus mahogonicolor

Boletus mahogonicolor Bessette, Both & Dunaway Common name(s): Mahogany Bolete Overview: The term mahogonicolor refers to the color of the wood of the mahogany tree, Swietenia mahagoni, which is the color of this handsome bolete’s cap. The cap surface stains yellowish with NH4OH, and the flesh stains orange with NH4OH. Cap: 2.5–9 cm wide, convex, becoming broadly convex in age, margin incurved at first and remaining so well into maturity, with a narrow band of sterile tissue; surface dry, glabrous to silky and shiny, mahogany to reddish brown or rose brown, paler toward the margin, staining blue along the margin when bruised; flesh yellow, rapidly staining blue when exposed and then changing to pale reddish brown; odor and taste not distinctive. Pores and tubes: bright yellow at first, becoming greenish yellow and then brownish yellow at maturity, staining dark blue to bluish black when bruised; tubes very shallow, 2–8 mm deep. Stalk: enlarged downward, solid; surface dry, minutely scurfy to nearly glabrous, bright yellow on the upper portion, with orange-red to

116  ·  Boletes of Eastern North America

brownish-red tones on the lower half or more, staining blue when handled or bruised, basal mycelium whitish to very pale yellow; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered in mixed woods under oak and pine; late spring and summer. Distribution and frequency: currently known only from Mississippi, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 10–13 × 3.5–4 µm, ellipsoidal, smooth, ochraceous. Edibility: unknown. Lookalikes: Boletus rubricitrinus (p. 145) has a more reddish cap, different macrochemical test reactions, longer tubes, and much larger spores, 13–19 × 5–8 µm. Boletus oliveisporus (p. 127) has a dark-fulvous cap tinged with bay that instantly stains blue black when bruised as well as much longer tubes, up to 2 cm deep, and larger spores, 11–17 × 4–6 µm.

Boletus miniato-olivaceus (A)

Boletus   ·  117

Boletus miniato-olivaceus (B)

Boletus miniato-olivaceus Frost Common name(s): Red and Olive Velvet-cap Overview: The term miniato-olivaceus means “colored with red and olive,” a reference to the cap’s colors. The cap surface stains olive yellow and then rapidly olive green to olive bronze with KOH and grayish olive with FeSO4. The flesh stains pale orange with KOH and pale yellowish with FeSO4 and is negative with NH4OH. Cap: 5–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, even or with a narrow band of sterile tissue; surface dry, velvety, red to rosy red when young, becoming pale rose pink and developing olive tones, finally becoming rosy tan to olive yellow with rosy tints in age, often staining dark blue when bruised; flesh dull white to pale yellow, reddish under the cuticle, slowly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow at first, becoming dingy yellow to olive yellow and sometimes developing reddish tints in age, depressed at the stalk at maturity, quickly staining blue and then slowly brownish when bruised; tubes 6–16 mm deep. Stalk: nearly equal or tapered in either direction, solid, often with a pinched base; surface dry, somewhat roughened with delicate reddish punctae,

118  ·  Boletes of Eastern North America

sometimes longitudinally striate, with reddish tints over a yellow ground color, possibly reticulate on the upper portion, basal mycelium yellowish, often staining dark blue when bruised; flesh yellow, usually darker than the cap flesh, reddish to brownish around larval tunnels, sometimes slowly staining blue. Habitat and season: solitary, scattered, or in groups in conifer, broadleaf, or mixed woodlands, often with hemlock, birch, oak, or beech; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to the Great Lakes region; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 10–15(18) × 4–6(7) µm, fusoid to subellipsoidal, smooth, ochraceous. Edibility: reported to be poisonous, causing gastrointestinal distress. Lookalikes: Boletus sensibilis (p. 150) is very similar, but its cap does not develop olive tones with age. Its flesh has a distinctive odor that has been variously described as fruity, like maple syrup, fenugreek, curry, or licorice. In addition, it has smaller spores, 10–13 × 3.5–4.5 µm.

Boletus miniatopallescens (A)

Boletus   ·  119

Boletus miniatopallescens (B)

Boletus miniatopallescens A. H. Sm. & Thiers Common name(s): none. Overview: The term miniatopallescens means “red becoming pale,” a reference to this bolete’s cap color. The key identification features include the red to brick-red cap, pale-yellow flesh that stains blue, yellow pores that stain greenish blue, and yellow stalk with reddish-brown tones on the lower portion. Cap: 8–18 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, smooth or somewhat velvety, often cracked in age, red to brick red, soon fading to reddish orange or orange yellow; flesh pale yellow, staining blue when exposed; odor somewhat sweetish or not distinctive; taste not distinctive. Pores and tubes: yellow, sometimes with a pale-orange tinge when young, becoming dingy yellow or tinged reddish orange with age, quickly bruising greenish blue; tubes 3–15 mm deep. Stalk: nearly equal or tapered downward, solid; surface dry, yellow overall with a pinkish bloom when young, becoming yellow on the upper portion, orange to reddish below, and developing reddish-brown tones or streaks, especially over the lower portion, occasionally finely reticulate at the apex, often staining blue when bruised; flesh concolorous with and staining like the cap flesh.

120  ·  Boletes of Eastern North America

Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oaks; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: olive brown. Microscopic features: spores 11–17 × 3–5 µm, narrowly fusiform, smooth, pale brown. Edibility: unknown. Lookalikes: Boletus sensibilis (p. 150) has a dark to pale brick-red cap that fades to dull rose or cinnamon with age, not to reddish orange. Its stalk is brighter yellow with less red and quickly stains blue when handled or bruised. It has yellow flesh that instantly stains blue when exposed and has an odor variously described as fruity, like maple syrup, fenugreek, curry, or licorice.

Boletus nobilis (A)

Boletus   ·  121

Boletus nobilis (B)

Boletus nobilis Peck Common name(s): Noble Bolete Overview: The term nobilis means “noble or grand,” in reference to this bolete’s sizeable stature when mature. The key identification features include the yellowish-brown to olive-ochraceous cap, white unchanging flesh, white to yellow pore surface, and the white to pale-ochraceous and finely reticulated stalk. A new genus, Alloboletus, has been proposed for this species. Cap: 7–20 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, smooth or sometimes shallowly pitted, glabrous, yellowish brown or reddish brown at first, becoming ochraceous to olive ochraceous or reddish ochraceous with age; flesh white, becoming yellowish near the tubes, not staining when exposed; odor and taste not distinctive. Pores and tubes: white at first, becoming yellow to pale ochraceous or brownish yellow, often slightly depressed at the stalk in age; tubes 8–25 mm deep. Stalk: nearly equal or sometimes swollen near the middle to slightly enlarged near the base, solid; surface dry, white to pale ochraceous, finely reticulate

122  ·  Boletes of Eastern North America

on the upper half or at least near the apex and glabrous below, rarely glabrous overall; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak and beech; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally fairly common. Spore print: dull ocher brown to dull rusty brown. Microscopic features: spores 11–16 × 4–5 µm, subfusiform, smooth, pale olivaceous. Edibility: edible but not as desirable as its lookalikes. Lookalikes: Boletus nobilis may be confused with Boletus edulis (p. 101), which has more prominent reticulation on its stalk and grows with conifers, especially Norway spruce and pines. Boletus gertrudiae (p. 108) has a pale brownish-yellow to orange-yellow cap, and its stalk becomes bright yellow over the upper portion or nearly overall. Boletus variipes (p. 167) has a darker cap and more prominent reticulation that typically covers most of the stalk. Boletus nobilissimus (p. 123) has a darker, strongly pitted to corrugated cap and a pale-brown stalk with conspicuous reticulation that is raised in places.

Boletus nobilissimus

Boletus   ·  123

Boletus nobilissimus Both & R. Riedel Common name(s): none. Overview: The term nobilissimus means “most dignified,” a reference to this bolete’s stately appearance. The strongly pitted to corrugated and yellow-brown to vinaceous-brown cap, club-shaped to strongly bulbous pale-brown stalk with strongly raised reticulation, and the white to yellow pores that remain unchanged when bruised are its key identification features. Cap: 9.5–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, becoming decurved, projecting as a narrow band of sterile tissue; surface dry, opaque, with tiny hairs in small scalelike bundles, strongly pitted to corrugated, rarely only weakly pitted, yellow brown to deeper vinaceous brown, rarely beige, at times somewhat mottled in these colors; flesh white, unchanging when exposed; odor fragrant; taste mild. Pores and tubes: white and stuffed at first, soon yellow, becoming greenish yellow to greenish olivaceous in age, unchanging when bruised; tubes 8–15 mm deep. Stalk: clavate to strongly bulbous, reticulate nearly the entire length with large, compound, elongated meshes that are raised in places, apex and base white, midsection pale brown, not changing color when bruised; flesh white, with some brown stains in the basal area and around larval tunnels. Habitat and season: solitary, scattered, or in groups in mixed woods of red oak and pine; summer and fall. Distribution and frequency: New England and New York, distribution limits yet to be established; uncommon. Spore print: brownish olivaceous. Microscopic features: spores 11–15.5 × 4.4–5.5 µm, subfusiform, smooth, pale yellow in Melzer’s. Edibility: edible. Lookalikes: Boletus nobilis (p. 121) has a much smoother cap surface and a whitish stalk with delicate reticulation that is not raised. Boletus edulis (p. 101) grows with conifers, especially Norway spruce or pine, and its cap is not strongly pitted or corrugated.

Boletus ochraceoluteus (A)

Boletus ochraceoluteus (B)

Boletus   ·  125

Boletus ochraceoluteus Bessette, Both & A. R. Bessette Common name(s): none. Overview: The term ochraceoluteus means “ochraceous and yellow,” referring to the range of this bolete’s cap colors. The cap surface stains bluish green with NH4OH and exhibits an orange flash that soon becomes reddish brown with KOH. The flesh stains grayish with FeSO4. Cap: 2–7 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved, with a narrow band of sterile tissue when young, becoming decurved and even at maturity; surface dry to slightly viscid, somewhat velvety at first, becoming glabrous at maturity, finely cracked in age, ochraceous yellow to orange ocher or pale pinkish cinnamon with yellow on the margin at first, becoming dull brownish yellow in age; flesh pale to bright yellow, staining reddish and then bluish green within five minutes after being exposed; odor and taste not distinctive. Pores and tubes: bright yellow and attached to subdecurrent at first, becoming dull yellow in age, slowly staining blue when bruised, somewhat depressed at the stalk in age; tubes 8–13 mm deep. Stalk: nearly equal or tapered in either direction, solid; surface dry, pale yellow, staining reddish from the base upward when bruised or with age, sometimes slowly staining blue, apex bright yellow with brownish connecting lines that form a partial network, pruinose, with a scant yellow basal mycelium; flesh yellow, staining reddish and then slowly bluish green when exposed. Habitat and season: scattered or in groups in mixed oak and pine woods; summer and fall. Distribution and frequency: Georgia and Florida, west to Mississippi; uncommon. Spore print: olive brown to yellowish brown. Microscopic features: spores 10–14 × 4.5–6 µm, subfusiform to subellipsoidal, smooth, yellow, bluish gray slowly becoming dull yellow in Melzer’s, lacking an apical pore. Edibility: unknown. Lookalikes: Xerocomus hypoxanthus (p. 422) has flesh that does not stain reddish, and its cap surface stains blue to greenish blue with the application of NH4OH or KOH. Xerocomus hemixanthus (Singer) Murrill (not illustrated) has a dark-brown to reddish-brown or chestnut-brown cap that stains dark reddish brown with KOH and tan with NH4OH. It also has reddish-brown, raised ribs near the stalk apex and a white to yellowish-white basal mycelium.

Boletus oliveisporus (A)

Boletus oliveisporus (B)

Boletus   ·  127

Boletus oliveisporus (Murrill) Murrill = Ceriomyces oliveisporus Murrill Common name(s): none. Overview: All parts of this widely distributed but uncommon bolete stain dark blue when handled or bruised. The term oliveisporus means “olive-colored spores.” Cap: 5–18 cm wide, convex, becoming broadly convex to nearly plane in age, margin with a narrow band of sterile tissue; surface dry, slightly velvety, often finely cracked in age, reddish orange overall, sometimes paler yellow orange near the margin when young, becoming dark fulvous tinged with bay or cinnamon brown to dull brown and somewhat shiny in age, instantly staining blue black when bruised; flesh yellow, staining blue when exposed, sometimes slowly; odor and taste not distinctive. Pores and tubes: yellow to greenish yellow, becoming olive yellow at maturity, depressed at the stalk in age, quickly staining blue when bruised; tubes 5–20 mm deep. Stalk: nearly equal or tapered in either direction, solid; surface dry, covered nearly overall with distinct brown punctae or sometimes nearly glabrous, yellow with reddish tinges when young, becoming olive brown from the base upward as it matures, instantly staining blue black when bruised, sometimes reticulate on the upper portion, conspicuously longitudinally striate; flesh concolorous with and staining like the cap flesh, reddish brown around larval tunnels and at the very base. Habitat and season: solitary, scattered, or in groups on the ground or on stumps under pines or sometimes in mixed oak and pine woods; summer and fall. Distribution and frequency: New England south to Florida, west to Texas; occasional. Spore print: olive brown. Microscopic features: spores 11–17 × 4–6 µm, fusoid, smooth, ochraceous. Edibility: unknown. Lookalikes: Boletus mahogonicolor (p. 115) has a silky and shiny, mahogany to reddish-brown or rose-brown cap that stains blue along the margin when bruised; very shallow tubes, 2–8 mm deep; and smaller spores, 10–13 × 3.5–4 µm. Cyanoboletus pulverulentus (p. 202) has a smaller, darker-brown cap and raised longitudinal ridges on its stalk, and its cap surface displays a green flash when NH4OH is applied. Suillellus subvelutipes (p. 315) has red to orange pores.

Boletus pallidoroseus (A)

Boletus pallidoroseus (B)

Boletus   ·  129

Boletus pallidoroseus Both Common name(s): Pale Rose Bolete Overview: The term pallidoroseus means “pale rose,” a reference to this beautiful bolete’s cap color. The pale rose-pink to pale purplish-pink cap with a yellow margin, yellow stalk with pink to reddish pink near the base, reticulation at the stalk apex, and yellow to golden-yellow flesh that has an odor of beef or chicken bouillon are its distinctive features. The cap surface stains crimson and then dull dark olive with the application of NH4OH. Cap: 6–17 cm wide, rounded at first, becoming convex to nearly plane in age, margin incurved at first, even; surface dry, glabrous to somewhat velvety or with small scale-like bundles of fibers, initially pale rose pink, purplish pink, or mottled with darker-pink tones, becoming pale orange brownish in age, golden yellow or paler yellow along the margin; flesh yellowish to yellowish white, slowly staining blue when exposed; odor reminiscent of beef or chicken bouillon or of spoiled meat with age; taste not distinctive. Pores and tubes: yellow to bright yellow at first, becoming olivaceous brownish and narrowly depressed at the stalk in age, instantly staining blue when bruised; tubes 4–8 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, typically smooth but sometimes reticulate at the very apex, with fine matted fibers or pruinose when young, becoming nearly glabrous in age, yellow on the upper portion, becoming pink to reddish pink and often mottled downward, slowly staining pale blue when handled or bruised, with a white basal mycelium; flesh deep golden yellow and darker than the cap flesh, especially toward the base, irregularly staining blue or not staining blue at all when exposed. Habitat and season: solitary, scattered, or in groups under oak and beech, sometimes in mixed woods; summer and early fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: brownish olivaceous. Microscopic features: spores 9–12 × 3.5–5.5 µm, subfusiform to somewhat inequilateral, smooth, yellowish in Melzer’s. Edibility: edible, but not worth the effort. Lookalikes: The Baorangia bicolor complex (p. 74) is very similar, but its cap and stalk are deep red, and the odor of its flesh is not distinctive. Compare with Boletus sensibilis (p. 150), which has a dark to pale brick-red cap that fades to dull rose or sometimes dingy cinnamon in age and stains blue when bruised. It has pale-yellow cap flesh that quickly stains blue when exposed, a mostly yellow stalk, and an odor variously described as faintly fruity, like maple syrup, fenugreek, curry, or licorice.

Boletus pallidus (A)

Boletus pallidus (B)

Boletus   ·  131

Boletus pallidus Frost Common name(s): Pallid Bolete Overview: This small- to medium-size, pale-colored bolete is typically found in groups under oak trees. The dull creamy-white to pale-yellow pores readily bruise bluish green on mature specimens. The term pallidus means “pallid,” a reference to this bolete’s overall pale colors. The whitish to buff or pale gray-brown cap, whitish to yellow or greenish-yellow pores that stain greenish blue to grayish blue, mild- to slightly bitter-tasting flesh, and whitish stalk separate it from other similarly colored boletes. The flesh stains pale rusty orange with KOH, blue green with NH4OH, and bluish green to grayish with FeSO4. Cap: 4.5–15 cm wide, convex with an incurved margin when young, becoming broadly convex to nearly plane in age, sometimes slightly depressed, margin even; surface dry, smooth and glabrous or nearly so, often cracked at maturity, whitish to buff or pale brownish when young, becoming dingy brown with rose or salmon tints in age; flesh thick, whitish or pale yellow, unchanging or sometimes slowly staining bluish or pinkish when exposed; odor not distinctive; taste mild or slightly bitter. Pores and tubes: whitish to pale yellow when young, becoming yellow to greenish yellow in age, quickly bruising greenish blue and then grayish brown, immature specimens possibly not staining or staining grayish brown right away; pores minute; tubes 1–2 cm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, smooth overall or slightly reticulate at the apex, whitish when young, often developing brownish streaks, especially toward the base, sometimes yellow at the apex, with occasional reddish flushes near the base in age, often with a white basal mycelium, sometimes slightly staining blue when bruised; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups on the ground or on decaying wood with broadleaf trees or in mixed woods; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to fairly common. Spore print: olive or olive brown. Microscopic features: spores 9–15 × 3–5 µm, narrowly oval to subfusoid, smooth, pale brown. Edibility: edible and good, though sometimes bitter. Lookalikes: Tylopilus intermedius (p. 382) is somewhat similar but has grayish-buff to dull-pinkish pores and intensely bitter-tasting flesh.

132  ·  Boletes of Eastern North America

Boletus paluster

Boletus paluster Peck = Boletinellus paluster (Peck) Murrill = Boletinus paluster (Peck) Peck = Fuscoboletinus paluster (Peck) Pomerleau & A. H. Sm. = Suillus paluster (Peck) Kretzer & T. D. Bruns Common name(s): Larch Bolete, Red Bog Bolete Overview: The term paluster means “of the bog or marsh,” a reference to this colorful little bolete’s habitat. Although normally found growing among sphagnum mosses, it may also fruit on moss-covered, prostrate tree trunks. Cap: 2–7 cm wide, broadly convex to nearly plane or slightly depressed, with or without an umbo, margin incurved when young, sometimes with tiny universal veil remnants on immature specimens; surface dry, covered with tiny hairs or scales, pale pinkish purple to reddish purple; flesh yellowish white to yellow, reddish under the cuticle, thin, soft, unchanging when exposed; odor somewhat mealy or not distinctive; taste slightly acidic or not distinctive. Pores and tubes: pale yellow when young, becoming golden yellow and finally brownish yellow in age, unchanging when bruised; pores large, angular and radially arranged when young, becoming gill-like with crosswalls at maturity, strongly decurrent, covered by a fibrillose veil when young; tubes very shallow, 1.5–3 mm deep.

Boletus   ·  133

Stalk: nearly equal, often cracked, solid; surface dry, sheathed from the base upward with a torn, fibrillose veil, reticulate to poroid and yellow at the apex, pale purple to reddish purple below, very rarely leaving a ring; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups, usually among sphagnum mosses in bogs, fens, or other wet areas under larch trees; summer and fall. Distribution and frequency: eastern Canada south to Pennsylvania, west to Minnesota; occasional to fairly common. Spore print: purple brown to pinkish brown. Microscopic features: spores 7–10 × 3–4 µm, ellipsoidal to subellipsoidal, smooth, pale brown. Edibility: apparently edible although seldom collected for the table. Lookalikes: Suillus cavipes (p. 326) is somewhat similar and can be found in the same habitat, but it has a yellow-brown to reddish-brown or darkbrown cap and a concolorous stalk that is hollow in the lower half at maturity.

Boletus patrioticus

Boletus patrioticus T. J. Baroni, Bessette & Roody Common name(s): Patriotic Bolete Overview: The term patrioticus means “patriotic,” a reference to the red, white, and blue colors displayed by this bolete’s exposed flesh. The cap

134  ·  Boletes of Eastern North America

surface stains olive brown with KOH, olive amber with NH4OH, and olive gray with FeSO4. Cap: 3–13 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, becoming decurved, with a narrow band of sterile tissue; surface dry, somewhat velvety, olive when very young, soon pinkish to brick red or dark red, often brownish red toward the margin, usually with olive to tarnished-brass tints; flesh white to pale yellowish, pinkish red to purplish red under the cuticle or extending throughout the cap, slowly staining blue beneath the reddish area when exposed; odor not distinctive, taste tart to acidic. Pores and tubes: pale yellow at first, becoming olive yellow at maturity, depressed at the stalk in age, staining blue when bruised; tubes 3–15 mm deep. Stalk: usually enlarged downward, sometimes tapered downward or nearly equal, base often pinched, solid; surface dry, pruinose to scurfy, color variable, usually rosy red on the upper portion or nearly overall and olive toward the base, often a mixture of these colors over a yellow ground color, with a whitish to pale-yellow basal mycelium; flesh whitish to pale yellow, becoming dingy yellow to brownish toward the base, staining bluish to greenish and sometimes rosy red when exposed. Habitat and season: solitary, scattered, or in groups, often in grassy areas under oak or in mixed woods with oak present; spring, summer, and fall. Distribution and frequency: North Carolina south to Florida, west to Ohio and Texas; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 10–13 × 4–5.5 µm, subfusiform, smooth, deep golden brown. Edibility: edible. Lookalikes: Boletus miniato-olivaceus (p. 117) has a similarly colored cap that often stains dark blue when bruised, cap flesh that does not taste tart or acidic, a stalk that often stains dark blue when bruised, and a wider distribution that extends into Canada. Its spores measure 10–15(18) × 4–6(7) µm.

Boletus   ·  135

Boletus pseudo-olivaceus

Boletus pseudo-olivaceus A. H. Sm. & Thiers Common name(s): none Overview: The term pseudo-olivaceus means “falsely olive-colored,” a reference to the cap’s pale brownish-olive to yellowish-olive colors. The key identification features include the pale brownish-olive to yellowish-olive cap that bruises blue and then fuscous, rapidly bluing flesh, dark maroon-red to reddish-brown pores, and yellow pruinose stalk. Cap: 3–10 cm wide, convex, becoming broadly convex in age, margin incurved at first, even; surface dry, covered with tiny, matted fibers when young, becoming scurfy at maturity, pale brownish olive to yellowish olive, staining blue and then fuscous when handled or bruised; flesh ochraceous, rapidly staining blue and then slowly olive fuscous when exposed; odor of young specimens not distinctive, usually becoming faintly chlorine-like with maturity; taste not distinctive. Pores and tubes: dark maroon red at first, becoming reddish brown, staining blue when bruised, depressed at the stalk in age; tubes 6–10 mm deep. Stalk: enlarged downward to nearly equal, solid; surface dry, yellow with olivaceous tones at the base, pruinose with reddish pruina that often form a pattern that simulates reticulation on the upper portion, lacking dark-reddish hairs at the base; flesh concolorous with and staining like the cap flesh.

136  ·  Boletes of Eastern North America

Habitat and season: solitary or scattered under broadleaf trees, especially beech or oak; summer and fall. Distribution and frequency: New England south to North Carolina, west into the Great Lakes region, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 12–16 × 5–7 µm, subfusoid, with a minute apical pore, smooth, yellow. Edibility: unknown. Lookalikes: Boletus vermiculosoides (p. 169) has dark-brown pores when young that become paler in age and stain blackish blue when bruised, yellow flesh that tastes astringent, brownish pruina that do not form a reticulate pattern on the stalk, and smaller spores, 9–12 × 3–4 µm. Boletus vermiculosus Peck (see photo, p. 169) has a darker-brown cap when young that becomes dull cinnamon brown in age and dark-brown pores that become reddish brown or brownish orange in age. Its whitish to pale-yellow flesh lacks a distinctive odor or taste, and the brownish pruina on its stalk do not form a reticulate pattern.

Boletus pseudopinophilus (A)

Boletus   ·  137

Boletus pseudopinophilus (B)

Boletus pseudopinophilus Siegel, Frank, Bessette, A. R. Bessette & Craine, nom. prov. Common name(s): Pine-loving King Bolete Overview: The term pseudopinophilus means “false pinophilus,” a reference to Boletus pinophilus Pilát & Dermek, which is a European species. Boletus edulis f. pinicola (Vittad.) Vassilkov and Boletus pinicola (Vittad.) A. Venturi are misapplied names. The conspicuous brownish reticulum that darkens when handled is a distinctive feature of Boletus pseudopinophilus. Molecular analysis clearly demonstrates that Boletus pinophilus and Boletus pseudopinophilus are distinct species that nest in separate clades. Cap: 5–15 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, margin incurved at first, with a narrow band of sterile tissue or nearly even; surface dry, typically wrinkled or shallowly pitted, reddish brown to dark rusty brown; flesh whitish, pinkish brown under the cuticle, not staining when exposed; odor spicy or not distinctive; taste not distinctive. Pores and tubes: whitish at first, becoming yellow to olive yellow or brownish yellow, bruising rusty brown, depressed at the stalk in age; tubes 8–25 mm deep. Stalk: enlarged downward, sometimes bulbous, solid; surface dry, white to pale yellow near the apex, pale brown to brown below, darkening when

138  ·  Boletes of Eastern North America

handled or bruised, reticulate overall or at least on the upper two-thirds, reticulum whitish near the apex, becoming brownish below, darkening when handled; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under conifers, especially pine, sometimes buried in duff; spring, summer, and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: dark olive to olive brown. Microscopic features: spores 15–20 × 4–6 µm, fusoid to cylindrical, smooth, yellowish. Edibility: edible and choice. Lookalikes: Boletus edulis (p. 101) has a whitish to pale-brown stalk and a whitish reticulum. Boletus subcaerulescens (p. 157) has more vinaceous tones on the cap and stalk, flesh that typically has a lemony odor, pores that stain bluish gray when bruised, and a cap surface that stains bluish green, then bright orange surrounded by dull grayish green, and then entirely yellow orange with NH4OH.

Boletus purpureorubellus (A)

Boletus   ·  139

Boletus purpureorubellus (B)

Boletus purpureorubellus T. J. Baroni, Yetter & Norar. Common name(s): none. Overview: The viscid and shiny purplish-red cap, yellow flesh, pores that stain bluish when bruised, reddish stalk, very small spores (atypical for boletes), and peculiar habitat are the distinctive features of this most unusual bolete. The cap surface stains yellow with KOH. The term purpureorubellus means “purplish red.” Cap: 7.5–12.5 cm wide, convex, becoming broadly convex to nearly plane in age, margin undulating, with a narrow band of sterile tissue; surface viscid when wet, shiny when dry, glabrous or nearly so, color variable, dark red, purplish to blood red, or reddish orange, yellow on the margin; flesh yellow with a reddish line under the cuticle, becoming pale reddish vinaceous above the tube layer; yellow areas stain blue and then slowly whitish when exposed; odor not distinctive; taste slightly acidic or not distinctive. Pores and tubes: bright yellow to golden yellow, staining dark bluish gray to blackish blue and eventually brown when bruised, slightly depressed with a decurrent tooth; pores conspicuously gill-like near the stalk; tubes 8–10 mm deep, staining fuscous blue when cut. Stalk: nearly equal or tapered downward, solid; surface dry, with decurrent lines that form a pseudoreticulum at the apex, delicately pruinose in the red

140  ·  Boletes of Eastern North America

areas, glabrous elsewhere, apex yellow to pale cream, streaked downward with pale red to red, becoming dull red to brown or golden yellow at the base, with a bright-yellow basal mycelium; flesh yellow at the apex, reddish vinaceous and yellow below, yellow areas staining dark blue when exposed. Habitat and season: scattered or in groups in swamps, growing in mats of moss with tree roots under loblolly bay, leatherwood, red maple, and cabbage palm, also found on trunks of Atlantic white cedar; summer and fall. Distribution and frequency: recorded from Florida and Georgia, also Cape Cod, Massachusetts, distribution limits yet to be established; rare. Spore print: olive to brownish olivaceous. Microscopic features: spores 5–7.3 × 3–4 µm, short-ellipsoid, smooth, dull yellow. Edibility: unknown. Lookalikes: Hortiboletus rubellus (p. 225) has a dry, somewhat velvety cap and much larger spores.

Boletus roodyi (A)

Boletus   ·  141

Boletus roodyi (B)

Boletus roodyi B. Ortiz, D. P. Lewis & Both Common name(s): Roody’s Bolete Overview: This red-capped bolete with a bright-yellow stalk and pores is distinct from similarly colored boletes by features that it lacks. There is no staining on any part when cut or bruised and no reticulation on the stalk. Boletus roodyi was named in honor of American mycologist William C. Roody. It is characterized by its blood-red to pinkish- or purplish-red cap, yellow pores, smooth stalk stained with red, and lack of any blue staining reaction. The addition of KOH or NH4OH to the cap surface develops a slate-blue flash that quickly changes to yellow ocher. KOH stains the flesh pale bluish. Cap: 5–16 cm wide, convex, becoming nearly plane or slightly depressed in the center when mature, margin incurved at first, becoming decurved, with a narrow projecting band of sterile tissue; surface dry, smooth and glabrous to faintly velvety, sometimes cracked in age, with a faint whitish pruina when immature, uniformly pinkish purplish red to blood red or dark red; flesh very pale yellow to nearly white, with a very narrow red line under the cap surface, unchanging when exposed or sometimes developing reddish stains; odor not distinctive when fresh, strongly vanilla scented when dried; taste slightly astringent or not distinctive.

142  ·  Boletes of Eastern North America

Pores and tubes: attached to narrowly depressed, bright yellow to pale golden yellow, becoming greenish yellow to greenish olivaceous in age, not staining when bruised; tubes 5–10 mm deep. Stalk: equal down to a narrowed base; surface dry, glabrous to finely pruinose, pale golden yellow near the apex, paler yellow downward, irregularly streaked, mottled, or flecked with red that is concolorous with the cap mainly on the lower half or sometimes only at the base, lacking reticulation or only slightly reticulate at the apex, basal mycelium white; flesh whitish to very pale yellow, golden yellow in larval tunnels, sometimes red in the base, unchanging when exposed. Habitat and season: solitary to scattered, in groups, or rarely caespitose, on the ground with various species of oak in mixed oak and pine woods or in broadleaf forests of oak, hickory, and beech; summer and early fall. Distribution and frequency: New England south to Arkansas and eastern Texas; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 9.5–16.2 × 3.6–4.5 µm, fusoid, smooth, with grayish-yellow or greenish-yellow contents in KOH; dextrinoid or with pale grayish-blue contents in Melzer’s. Edibility: edible. Lookalikes: Boletus rubissimus A. H. Sm. (not illustrated), reported from Michigan and New York, shares the same overall colors. However, its cap has a sulfur-yellow to ochraceous-yellow margin, and its pores stain grayish blue when bruised. It has white flesh that is unchanging or sometimes weakly and erratically staining blue when exposed. Its stalk has fine reticulation on the upper portion and yellow basal mycelium. Microscopically, it has smaller spores that measure 9–11 × 3–4.5 µm. The Baorangia bicolor complex (p. 74) is also similar, but its flesh slowly stains blue when exposed.

Boletus   ·  143

Boletus roseolateritius

Boletus roseolateritius Bessette, Both & Dunaway Common name(s): Rosy Brick-red Bolete Overview: The term roseolateritius means “rosy brick red,” a reference to the color of this bolete’s mature cap. The cap surface stains black with FeSO4 and is negative with KOH or NH4OH. Cap: 4–15.5 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first and remaining so well into maturity, with a narrow band of sterile tissue; surface dry, somewhat velvety with a grayish bloom when young, nearly glabrous at maturity, dark reddish salmon or burnt orange at first, becoming rosy brick red to reddish brown at maturity, fading to brownish orange with yellow tints and finally to dull dingy yellow in age, whitish along the margin, instantly staining dark blue when bruised; flesh pale lemon yellow, instantly staining blue when exposed and then fading to pale lemon yellow; odor and taste not distinctive. Pores and tubes: orange red at first, becoming dull orange and finally orange yellow in age, instantly staining blue when bruised; tubes 5–12 mm deep. Stalk: nearly equal or tapered in either direction, often with a pinched base, solid; surface dry, faintly longitudinally striate, lacking reticulation, pale lemon yellow, instantly staining blue when bruised, slowly developing

144  ·  Boletes of Eastern North America

rusty-brown stains in age or when bruised; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in river bottomland under beech, with oak and hickory nearby; summer. Distribution and frequency: currently reported only from McComb, Mississippi, distribution range yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 8.5–11 × 3.5–4.5(5.5) µm, narrowly ellipsoid to subfusoid, smooth, pale ochraceous. Edibility: unknown. Lookalikes: Boletus fairchildianus (p. 104) is similar, but its pore surface is darker red; its cap stains olive with NH4OH and deep maroon or orange yellow to brown with KOH; and it has much larger spores, 13–19 × 5–8 µm.

Boletus rubricitrinus (A)

Boletus   ·  145

Boletus rubricitrinus (B)

Boletus rubricitrinus (Murrill) Murrill = Ceriomyces rubricitrinus Murrill Common name(s): none. Overview: The term rubricitrinus means “red and lemon yellow,” a reference to the colors exhibited by this attractive bolete. The key identification features include the variable, dull rose-red to tawny-cinnamon cap that usually stains bluish black, yellow flesh that quickly stains blue, yellow pores that stain blue, and yellow stalk that quickly stains blue green to bluish black. Cap: 3–15 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, sometimes slightly depressed, margin incurved at first with a band of sterile tissue; surface dry, acidic or tart tasting, flesh glabrous to slightly velvety, color variable, dull rose red to dull brick red or reddish brown to tawny cinnamon when fresh, fading to tawny olive or dull brown in age, sometimes with yellow tints, usually bruising bluish black; flesh pale yellow to yellow, quickly staining blue when exposed; odor not distinctive; taste of flesh slightly tart to acidic. Pores and tubes: yellow at first, becoming dull yellow to olive yellow, depressed at the stalk in age, staining blue when bruised; tubes 8–20 mm deep. Stalk: nearly equal or tapered in either direction, sometimes swollen near the base, solid; surface dry, yellow, typically with dull reddish to reddish-brown streaks and dots, especially toward the base, quickly staining blue green to bluish black when bruised, sometimes weakly reticulate only at the very

146  ·  Boletes of Eastern North America

apex, typically longitudinally striate nearly overall; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in oak or mixed oak and pine woods; spring, summer, and fall, also early winter on the Gulf Coast. Distribution and frequency: New Jersey south to Florida, west to Texas; fairly common. Spore print: olive brown. Microscopic features: spores 13–19 × 5–8 µm, fusoid to subfusoid-ellipsoid, smooth, yellowish. Edibility: edible, tart to acidic but good. Lookalikes: Boletus fairchildianus (p. 104), an uncommon species reported from Florida west to Texas, is similar but has dark-red pores and more red on the stalk.

Boletus rufocinnamomeus

Boletus rufocinnamomeus A. H. Sm. & Thiers Common name(s): none. Overview: The term rufocinnamomeus means “reddish cinnamon,” a reference to mature specimens’ cap color. The key identification features include the velvety, reddish-brown cap, yellow flesh that instantly stains blue, maroonred to orange-red pores, and orange stalk coated with orange-cinnamon pruina.

Boletus   ·  147

Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, even or nearly so; surface dry, somewhat velvety, dull brick red with an ochraceous overtone from an evanescent bloom when young, becoming reddish brown to dull cinnamon brown in age; flesh yellow, instantly staining blue when exposed; odor not distinctive; taste acidic or not distinctive. Pores and tubes: maroon red when young, becoming orange red at maturity, quickly staining blue when bruised, depressed at the stalk in age; tubes 8–15 mm deep. Stalk: nearly equal, solid; surface dry, yellow and glabrous on the upper portion, coated with orange-cinnamon pruina toward the base, with yellow basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered in broadleaf forests, especially with hazelnut, oak, and beech; summer and fall. Distribution and frequency: New England south to North Carolina, west to the Great Lakes region, distribution limits yet to be established; uncommon. Spore print: olive brown. Microscopic features: spores 14–17 × 4.5–6.5 µm, subfusoid with a minute apical pore, smooth, dull yellowish. Edibility: unknown. Lookalikes: Boletus subluridellus (p. 163) is similar, but all parts instantly stain blackish blue when bruised. It has a pale-yellow stalk that darkens from the base upward in age or where handled and is nearly smooth to slightly pruinose-scurfy. It also has slightly smaller spores, 11–15 × 4–6 µm.

Boletus rufomaculatus (A)

148  ·  Boletes of Eastern North America

Boletus rufomaculatus (B)

Boletus rufomaculatus Both Common name(s): none. Overview: The term rufomaculatus means “spotted reddish,” a reference to the reddish spots and splashes on this bolete’s mottled cap. The cap and flesh stain yellow with NH4OH and dark amber with KOH. Boletus rufomaculatus is very closely related to the Baorangia bicolor (p. 74) complex and nests in the same clade (Wu et al. 2014). It is possible that it will be renamed Baorangia rufomaculatus at some future time. Cap: 6.5–14 cm wide, rounded at first, becoming broadly convex to nearly plane in age, margin incurved at first, with a narrow band of sterile tissue; surface dry to subviscid when wet, granulose to tomentose at first, becoming matted to fibrillose with age, color variable, red to dull red, dull rusty brown to ocher brown or honey yellow, becoming cracked, mottled with brick red or brownish red in age; flesh pale yellow, slowly staining blue when exposed, especially above the tubes, at times staining blue only weakly and erratically; odor and taste unpleasant or not distinctive. Pores and tubes: pale yellow at first, becoming brighter lemon yellow and then deeper yellow and finally yellowish olive, shallowly depressed at the stalk at maturity, sometimes with rusty spots or reddish tints in age, staining blue when bruised; tubes 6–14 mm deep. Stalk: nearly equal or somewhat swollen in the middle and tapered in both directions, solid; surface dry, nearly glabrous, somewhat ribbed to

Boletus   ·  149

pseudoreticulate, golden yellow near the apex, pale yellow below, strongly dotted or mottled with burgundy red on the lower half and sometimes nearly overall, staining blue when bruised or handled, with a white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under beech in mixed woods; early summer through fall. Distribution and frequency: New England and New York south to Georgia, west to Mississippi; occasional. Spore print: dark olive. Microscopic features: spores 10–13 × 3–4.5 µm, subfusiform, smooth, yellowish in Melzer’s. Edibility: unknown. Lookalikes: The Baorangia bicolor complex (p. 74) has a glabrous, red cap with an even margin at all stages of development, a glabrous stalk that typically is red on the lower two-thirds or more, and darker golden-yellow stalk flesh, and it grows under oaks. The caps of the Baorangia bicolor complex typically stain blackish with FeSO4 and are negative with NH4OH or KOH. Their flesh usually stains pale yellow to pale orange with KOH and is negative with NH4OH.

Boletus sensibilis (A)

150  ·  Boletes of Eastern North America

Boletus sensibilis (B)

Boletus sensibilis Peck = Boletus miniato-olivaceus var. sensibilis (Peck) Peck = Boletus sensibilis var. subviscidus A. H. Sm. & Thiers Common name(s): Sensitive Bolete Overview: The term sensibilis means “sensitive,” a reference to this bolete’s easily bruised tissue. All parts instantly stain blue when handled or bruised. The cap surface stains yellow with KOH or NH4OH and greenish gray with FeSO4. Cap: 5–16 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety when young, becoming nearly glabrous in age, orangish red or dark to pale brick red, fading to dull rose or sometimes dingy cinnamon in age, staining blue when bruised; flesh pale yellow, staining blue when exposed; odor variously described as faintly fruity or like maple syrup, fenugreek, curry, or licorice; taste not distinctive. Pores and tubes: yellow when young, becoming duller or browner in age, instantly staining blue when bruised; tubes 8–12 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, mostly yellow with pink or red on the lower portion, sometimes with white basal mycelium, occasionally finely reticulate at the apex, quickly staining blue when handled; flesh bright yellow, more intensely yellow than the cap flesh, usually staining blue quickly when exposed.

Boletus   ·  151

Habitat and season: scattered or in groups in woods, usually under broadleaf trees, especially oak; late spring, summer, and fall. Distribution and frequency: eastern Canada south to South Carolina, west to Minnesota; fairly common. Spore print: olive brown. Microscopic features: spores 10–13 × 3.5–4.5 µm, suboblong to slightly ventricose, smooth, pale brown; hymenial cystidia fusoid-ventricose with an elongated neck. Edibility: this bolete has long been considered to be a poisonous species. However, we have received several recent reports from individuals who have eaten it without ill effects. Lookalikes: Lanmaoa pseudosensibilis (p. 233) has a darker reddish-brown cap and bright yellow flesh that stains blue when exposed. A drop of NH4OH stains the cap surface greenish blue. The cap of Boletus miniato-olivaceus (p. 117) develops olive tones in age; its flesh lacks a distinctive odor; and it has larger spores, 10–15(18) × 4–6(7) µm. Boletus pallidoroseus (p. 129) has a pink to reddish-pink stalk with yellow near the apex, yellowish to yellowish-white flesh that slowly and erratically stains blue when exposed, and an odor reminiscent of beef or chicken bouillon. Boletus miniatopallescens (p. 119) has a brick-red cap that fades to reddish orange or orange yellow in age, a more reddish stalk, and pale-yellow flesh that lacks a distinctive odor. The Baorangia bicolor complex (p. 74) has red over most of the stalk, flesh that is unchanging or only weakly and erratically staining blue, and shorter tubes.

Boletus separans (A)

Boletus separans (B)

Boletus separans (C)

Boletus   ·  153

Boletus separans Peck = Boletus pseudoseparans Grand & A. H. Sm. = Xanthoconium separans (Peck) Halling & Both Common name(s): Lilac Bolete Overview: This medium to large bolete is a beautiful mushroom that is much sought after as an edible. Its superb flavor is often compared with its better-known cousin, the King Bolete or Cep, Boletus edulis (p. 101). The color of the cap and stalk is variable, but lilac tones are usually present in both young and mature specimens. Lilac areas of the cap or stalk surface stain aquamarine to deep blue with NH4OH. The term separans refers to how the tube layer sometimes pulls away from the stalk as the cap expands. A new genus, Alloboletus, has been proposed for this species. Cap: 5–15 cm wide, convex to broadly convex, margin even; surface dry, somewhat velvety or glabrous, often pitted, wrinkled, or lumpy and uneven; color varying from creamy white tinged with lilac to pinkish brown, lilac brown, reddish brown, or sometimes dark purple, often paler near the margin and becoming yellowish brown to almost bronze colored in age; flesh white, unchanging when exposed; odor not distinctive; taste sweet and nutty or not distinctive. Pores and tubes: white at first, becoming yellowish to ocher brown at maturity, unchanging when bruised; tubes 1–3 cm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, concolorous with the cap or paler, often with lilac tones in the midportion, sometimes purplish overall, occasionally with yellow coloration near the apex, with fine white reticulation over most of its length or at least over the upper half; reticulation occasionally very delicate and obscure to nonexistent; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in clusters in mixed woods, oak woods, or occasionally with Norway spruce or pine; late spring, summer, and fall, also winter in the southern part of its range. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: brownish ocher to pale reddish brown. Microscopic features: spores 12–16 × 3.5–5 µm, narrowly subfusiform, smooth, pale brown. Edibility: edible and choice. Lookalikes: Compare with Xanthoconium purpureum (p. 412), which has a darker cap and stalk without lilac tones. Its pores stain yellow brown to brown when bruised.

Boletus speciosus (A)

Boletus speciosus (B)

Boletus   ·  155

Boletus speciosus (C)

Boletus speciosus Frost = Ceriomyces speciosus Murrill Common name(s): Showy Bolete Overview: The term speciosus means “showy.” The flesh stains orange with KOH and grayish with FeSO4. According to Ernst Both (1993), Walter Snell and Esther Dick’s illustration at the bottom of plate 39 of The Boleti of Northeastern North America (1970) is most likely Boletus rubissimus A. H. Sm. (not illustrated), not Boletus speciosus as labeled. The wide range of cap colors, including raspberry red (rosolane purple), that Snell and Dick give in their description of Boletus speciosus may represent two or more species, including Butyriboletus roseopurpureus (p. 184). Cap: 7–16 cm wide, rounded to convex, becoming broadly convex in age, margin even, incurved at first and remaining so well into maturity; surface dry, coated with tiny hairs or nearly glabrous, bright rose red to rose pink, often retaining this color well into maturity, becoming orange red and then pinkish brown to olive brown in age, sometimes with fine cracks over the disc on mature specimens, bruising slowly blackish blue; flesh pale yellow, soon staining grayish blue when exposed; odor sometimes like iodine or not distinctive; taste not distinctive.

156  ·  Boletes of Eastern North America

Pores and tubes: yellow at first, becoming dull yellow to olive yellow in age, rapidly staining blue when bruised and then eventually reddish brown; tubes 8–15 mm deep. Stalk: nearly equal or enlarged downward, base often abruptly narrowed, solid; surface dry, finely reticulate overall or at least on the upper half, yellow on the upper portion, pinkish red to red or pale pinkish brown on the lower portion, staining blue when bruised or handled; flesh pale or medium yellow, staining like the cap flesh. Habitat and season: solitary or scattered in broadleaf forests, conifer woods, or mixed woods; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; uncommon. Spore print: olive brown. Microscopic features: spores 11–15 × 3–5.5 µm, narrowly oblong to subfusoid, smooth, ochraceous. Edibility: edible but too uncommon to collect for the table. Lookalikes: Butyriboletus brunneus (p. 181) has a reddish-brown to yellowbrown or olive-brown cap. The cap of Butyriboletus roseopurpureus (p. 184) is strikingly pinkish purple in fresh specimens and becomes darker purplish pink to dark purplish red as they age or at times is mottled in these colors. Boletus rubissimus A. H. Sm. (not illustrated), known from Michigan and New York, is also very similar, but it has a dark-pink cap, pale-yellow flesh that does not blue or only weakly blues when exposed, yellow pores that stain grayish blue, a yellow stalk flushed red over the lower portion and finely reticulate near the apex, and shorter spores, 9–11 µm long.

Boletus subcaerulescens (A)

Boletus   ·  157

Boletus subcaerulescens (B)

Boletus subcaerulescens (E. A. Dick & Snell) Both, Bessette & A. R. Bessette = Boletus edulis ssp. subcaerulescens E. A. Dick & Snell Common name(s): none. Overview: The term subcaerulescens means “somewhat bluing,” a reference to the pale bluish-gray color that develops on this bolete’s pores when they are bruised. The cap surface stains amber orange with KOH, stains bluish green and then bright orange surrounded by dull grayish green and finally entirely yellow orange with NH4OH, and is negative with FeSO4. Cap: 4–18 cm wide, rounded at first, becoming broadly convex to nearly plane in age, margin incurved at first, becoming decurved, with a narrow band of sterile tissue; surface dry, covered with tiny fibrils or nearly glabrous, often wrinkled to shallowly pitted, rusty cocoa brown to pinkish brown with a more cinnamon center when young, becoming reddish brown to chestnut brown or rusty cinnamon at maturity, mottled dingy yellowish at times, fading to more yellow tones and always flushed or mottled pale cinnamon in age; flesh white, dull vinaceous above the tubes, becoming vinaceous near the cuticle, unchanging or sometimes slowly and weakly staining blue when exposed; odor reminiscent of lemon or not distinctive; taste nutty, sweet. Pores and tubes: whitish cream at first, becoming pale yellow and then yellowish olive to ocher olive in age, narrowly depressed at the stalk at maturity, staining bluish gray and then ochraceous to brownish when bruised; tubes up to 1.5 cm deep.

158  ·  Boletes of Eastern North America

Stalk: nearly equal to enlarged downward, at times somewhat bulbous, base sometimes with a pointed root-like extension, solid; surface dry, conspicuously reticulate nearly overall or at least over the upper two-thirds, whitish near the apex and rusty cinnamon to brownish vinaceous over the lower one-third, typically white at the base, with a white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under pines or Norway spruce; spring, summer, and fall. Distribution and frequency: eastern Canada and New England south to West Virginia and west to Minnesota; occasional. Spore print: deep, smoky olive. Microscopic features: spores 13–19 × 4–5 µm, subfusoid to cylindrical, smooth, pale ochraceous. Edibility: edible and choice, one of the best edible boletes. Lookalikes: Boletus subcaerulescens is sometimes mistaken for Boletus edulis (p. 101) because they are quite similar in appearance and can sometimes be found in the same habitat. However, the pores of Boletus edulis do not bruise bluish, and its cap surface lacks the initial bluish-green staining phase with NH4OH and slowly stains pale grayish green with FeSO4. Boletus pseudopinophilus (p. 137) is also similar, but it has a reddish-brown to dark rusty-brown cap and pores that do not stain blue when bruised.

Boletus subfraternus (A)

Boletus   ·  159

Boletus subfraternus (B)

Boletus subfraternus Coker & Beers = Boletellus subfraternus (Coker & Beers) Singer Common name(s): none. Overview: The term subfraternus means “near Boletus fraternus,” which is currently a synonym of Hortiboletus rubellus (p. 225). The key identification features include the red cap that does not become cracked, pale-yellow flesh that stains blue, yellow pores that stain greenish blue, and a pale-red, pruinose stalk. Cap: 2–5 cm wide, broadly convex, becoming nearly plane in age, margin incurved at first, even; surface dry, somewhat velvety, not becoming cracked or only very slightly so in age, dull rose red to bright orange red at first, becoming brick red to brownish red and finally orange buff with reddish tinges in age; flesh pale yellow, staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow to olive yellow at first, becoming red brown in age, staining greenish blue when bruised, depressed at the stalk at maturity; pores irregular to angular, becoming elongated and radially arranged to somewhat gill-like near the stalk at maturity; tubes up to 1 cm deep.

160  ·  Boletes of Eastern North America

Stalk: nearly equal, usually with an abruptly narrowed base, solid; surface dry, concolorous with the cap but paler, ochraceous at the apex, longitudinally streaked and finely pruinose; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups in sandy soil along road banks, in mixed woods, or along the edges of mixed woods; summer and fall. Distribution and frequency: Maine south to North Carolina, west to Michigan, distribution limits yet to be established; infrequent. Spore print: olive brown. Microscopic features: spores 9–14 × 4–6 µm, subfusiform, smooth, ochraceous. Edibility: unknown. Lookalikes: Hortiboletus campestris (p. 223) is very similar, but its cap becomes conspicuously cracked at maturity, and it has circular to angular pores that are not elongated and not radially arranged to somewhat gill-like near the stalk. Hortiboletus rubellus (p. 225) has a cap that becomes finely cracked at maturity, stalk flesh that is yellow on the upper portion and reddish orange in the base, and angular pores that are not elongated and not radially arranged to somewhat gill-like near the stalk.

Boletus subgraveolens

Boletus   ·  161

Boletus subgraveolens A. H. Sm. & Thiers Common name(s): none. Overview: The term subgraveolens means “somewhat strong and disagreeable,” a reference to the odor emitted by dried specimens of this bolete. The cap surface displays a blue flash and then slowly stains vinaceous with NH4OH. Cap: 4–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin even, incurved at first; surface slightly viscid or dry, glabrous or nearly so, becoming conspicuously cracked in hot, dry weather, dull yellow brown overall or mottled with darker or paler shades of yellow or brown, usually staining dark blue and then darker brown when fresh; flesh yellowish white to pale yellow, staining blue when exposed; odor pungent; taste not distinctive. Pores and tubes: brownish yellow when very young, soon becoming bright yellow brown and then paler in age, depressed near the stalk, bruising dark blue and then slowly brown; tubes pale yellow, staining bluish green when cut, 6–13 mm deep. Stalk: tapered downward and typically pinched or conspicuously pointed at the base, solid; surface dry, bright yellow at the apex, dull pale yellow and pruinose below, with a dark vinaceous-red to maroon base, developing yellowish-brown tints from the base upward in age or when handled, sometimes bruising bluish; flesh pale yellow to orange yellow with pinkish tints on the upper portion, dark vinaceous red to maroon or reddish cinnamon near the base, staining blue when exposed. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially aspen and oak; summer and fall. Distribution and frequency: reported from Michigan and Mississippi, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 9–13 × 3.5–4.5 µm, subfusiform, smooth, yellowish brown. Edibility: unknown. Lookalikes: Both Boletus vermiculosoides (p. 169) and Boletus vermiculosus Peck (see photo, p. 169) are similar but lack the reddish coloration at the stalk base. Boletus fagicola A. H. Sm. & Thiers (not illustrated) is also similar but has reticulation on its stalk.

Boletus subluridellus (A)

Boletus subluridellus (B)

Boletus   ·  163

Boletus subluridellus A. H. Sm. & Thiers Common name(s): none. Overview: All parts of this bolete instantly stain blue when bruised. The key identification features include a bright-red to orange-red or brick-red cap, a yellow stalk that darkens when handled or in age, and pinkish-red to rose-red or dark-red pores that fade to orange red or brownish in age. This bolete’s flesh has a peculiar and somewhat pungent but distinctive odor. The term subluridellus means “nearly drab yellow to dirty brown.” Cap: 5–12 cm wide, convex, becoming broadly convex in age, margin with a narrow band of sterile tissue, incurved at first; surface dry, somewhat velvety, purplish red, brownish red, pinkish red, or orange red, instantly staining blackish blue when bruised; flesh bright yellow, quickly staining dark blue when exposed; odor pungent; taste not distinctive. Pores and tubes: rosy red to pinkish red or dark red, becoming orange red in age, quickly staining blue when bruised, depressed at the stalk in age; tubes 6–10 mm deep. Stalk: nearly equal, solid; surface dry, pale yellow, darkening from the base upward in age or where handled, faintly pruinose-scurfy overall; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under broadleaf trees, especially oak, or in mixed woods usually with oak present; summer and fall. Distribution and frequency: New England and New York south to Georgia, west to Minnesota; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 11–15 × 4–6 µm, subfusoid with a minute apical pore, smooth, dull yellowish. Edibility: unknown. Lookalikes: Boletus rufocinnamomeus (p. 146) has a dull brick-red cap with an ochraceous overtone from an evanescent bloom when young. The cap becomes dull cinnamon brown in age and does not stain blackish blue when bruised. The stalk is yellow and glabrous on the upper portion and is coated with orange-cinnamon to brown pruina toward the base, and it has slightly larger spores, 14–17 × 4.5–6.5 µm. Suillellus subvelutipes (p. 315) has a more orange to brownish cap and often bristle-like red hairs on its stalk base. Suillellus subluridus (p. 314) has a variably colored cap ranging from yellow to orange with purple stains or is entirely purplish red. Its yellow flesh stains blue when exposed, and the purple-red to dark-red pores also bruise blue.

Boletus subtomentosus var. subtomentosus

Boletus subtomentosus var. perplexus

Boletus   ·  165

Boletus subtomentosus var. subtomentosus L. = Ceriomyces subtomentosus (L.) Murrill = Xerocomus subtomentosus (L.) Quél. Common name(s): Brown and Yellow Bolete, Suede Bolete, Yellow-cracked Bolete Overview: The term subtomentosus means “somewhat coated with fine hairs,” a reference to the surface of this bolete’s cap. The cap surface instantly stains reddish brown with NH4OH. Cap: 5–18 cm wide, convex, becoming broadly convex in age, margin somewhat incurved at first, with a narrow band of sterile tissue that is obvious at least on young specimens; surface dry, covered with fine hairs, often cracked in age, olive brown to yellow brown; flesh whitish, unchanging or slightly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow at first, becoming dull yellow in age, staining slightly and slowly greenish and then brownish when bruised; tubes 1–2.5 cm deep. Stalk: nearly equal or tapered slightly downward, solid; surface dry, scurfy, sometimes with raised longitudinal lines that may form a net-like pattern near the apex, predominantly yellow with reddish-brown streaks and yellowish mycelium at the base, staining brownish to reddish brown when handled or in age; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered under broadleaf or conifer trees; summer and fall, also early winter in the southern part of its range. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; fairly common. Spore print: olive brown. Microscopic features: spores 10–15 × 3–5 µm, subfusoid to oblong, smooth, yellow. Edibility: edible. Lookalikes: Boletus subtomentosus var. perplexus A. H. Sm. & Thiers (see photo, p. 164) is very similar but has a darker olive-brown cap and a pruinose, reddish-brown to cinnamon stalk, and its pores more readily stain greenish blue when bruised. Boletus ferrugineus (p. 105) is similar, but its pores more readily stain blue when bruised, and its cap surface displays a blue or blue-green flash and then stains reddish brown with NH4OH. Boletus alutaceus (p. 87) has been reported from New England south to Kentucky, west to Ohio and Wisconsin, and it grows in broadleaf and mixed woods, especially oak and beech. It has a yellowish-brown cap that becomes pale tan with a pinkish tint, pale greenish-yellow pores that do not stain when bruised, and whitish to pale-tan flesh that does not stain when exposed. Its spores measure 9–12 × 4–5 µm. Xerocomellus chrysenteron (p. 416) has a dark-olive to olive-brown or grayish-brown cap that often has a reddish zone at the margin and soon becomes cracked, with red showing in the cracks.

Boletus variipes (A)

Boletus variipes (B)

Boletus   ·  167

Boletus variipes (C)

Boletus variipes Peck = Boletus variipes var. variipes Peck = Boletus variipes var. fagicola A. H. Sm. & Thiers Common name(s): Variable-stalk Bolete Overview: The term variipes means “variable foot,” referring to the color, pattern, and amount of reticulation on this bolete’s stalk. Cap: 6–20 cm wide, convex when young, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety to nearly glabrous, color variable, creamy tan to yellowish tan, grayish brown to yellow brown or dark brown, often cracked at maturity; flesh white, unchanging or slowly staining pale pinkish gray when exposed; odor and taste not distinctive. Pores and tubes: white at first, becoming yellowish to yellowish olive in age, unchanging when bruised; tubes 1–3 cm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, whitish to yellow brown or grayish brown, usually distinctly reticulate or sometimes less conspicuously so, reticulation white or brown; flesh concolorous with the cap flesh.

168  ·  Boletes of Eastern North America

Habitat and season: scattered or in groups under mixed broadleaf trees, especially oak, beech, or aspen, and sometimes with conifers, especially Norway spruce; spring, summer, and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 12–18 × 4–6 µm, subfusoid, smooth, yellow. Edibility: edible and very good. Lookalikes: Tylopilus felleus (p. 375) has bitter-tasting flesh, and its pores become dull pinkish as the spores mature. Boletus edulis (p. 101) has a reddish-brown cap and fine whitish reticulation. Boletus atkinsonii (p. 89) is also similar, but its cap surface is roughened with tufts of hyphae that are best observed with a hand lens, a dingy-white stalk with fine brownish reticulation, and smaller spores, 10–13 × 4–5 µm. Boletus nobilis (p. 121) has a taller stature, a paler-colored cap that is often somewhat pitted, a white to pale-ocher stalk that is sometimes streaked with pale brown, and delicate stalk reticulation that is normally limited to the upper portion. Compare with Boletus nobilissimus (p. 123).

Boletus vermiculosoides

Boletus   ·  169

Boletus vermiculosus

Boletus vermiculosoides A. H. Sm. & Thiers Common name(s): Wormy Bolete Overview: The Wormy Bolete and its lookalikes are distinctive because of their rich-brown pores that instantly bruise blackish blue. The term vermiculosoides means “resembling Boletus vermiculosus” (see photo, p. 169). The term vermiculosus means “infested with insect larvae,” a condition commonly observed in both of these species. Cap: 4–12 cm wide, convex, becoming broadly convex at maturity, margin even; surface dry, covered with tiny matted fibers, yellow when young or brownish with a yellow margin, becoming brown with maturity; flesh yellow, fading to pale yellow in age, quickly staining blue when exposed; odor pungent; taste astringent or not distinctive. Pores and tubes: dark brown when young, paler in age, staining blackish blue and then very slowly fading to dull brownish orange when bruised; tubes 6–20 mm deep. Stalk: nearly equal, solid; surface dry, brownish pruinose over a dull-whitish or pale-yellow ground color with olive tints, staining blue and then dark brown when bruised; flesh concolorous with and staining like the cap flesh.

170  ·  Boletes of Eastern North America

Habitat and season: solitary, scattered, or in groups under oak; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: olive brown. Microscopic features: spores 9–12 × 3–4 µm, ellipsoidal, smooth, pale brown. Edibility: unknown; however, one of the authors has eaten this bolete several times without ill effect. Lookalikes: Boletus fagicola A. H. Sm. & Thiers (not illustrated) is nearly identical but has reticulation on the stalk, and its cap cuticle is a trichodermium rather than a cutis. Boletus vermiculosus Peck (see photo, p. 169) is very similar but has a dark-brown to grayish-brown or reddish-brown cap that stains vinaceous with KOH and has larger spores, 11–15 × 4–6 µm. Compare also with Boletus subgraveolens (p. 161), which has a stalk that is bright yellow at the apex, dull pale yellow and pruinose below, and reddish at the base. The stalk develops yellowish-brown tints from the base upward when handled or bruised and sometimes stains blue, and the flesh in its stalk base is reddish cinnamon.

Boletus weberi

Bothia   ·  171

Boletus weberi Singer Common name(s): none. Overview: The dry, cracked, and scaly brownish cap, reddish pores, reddish and somewhat scaly stalk, and tart-tasting flesh that does not stain when exposed are key features of this unusual Gulf Coast bolete. The cap surface stains reddish brown with NH4OH and is negative with FeSO4. The flesh quickly stains greenish blue with NH4OH and is negative with FeSO4. Cap: 4–7 cm wide, convex, becoming broadly convex to nearly plane in age; surface dry, cracked and scaly with pale-yellow flesh showing in the cracks, brownish olive at first, becoming olive brown at maturity, sometimes with reddish tints, not staining when bruised, margin incurved at first, becoming decurved at maturity, even; flesh pale yellow, not staining blue when exposed; odor not distinctive; taste tart and somewhat lemony. Pores and tubes: dull red at first, becoming dull orange red and finally reddish orange, depressed at the stalk in age, not staining blue but sometimes slowly staining brown when bruised; tubes 3–6 mm deep. Stalk: enlarged downward to a conspicuously pinched base, solid; surface dry, fibrillose-punctate on the upper portion, becoming somewhat scaly-punctate on the lower portion, pinkish red to dull purplish red over a pale-yellow ground color on the upper portion, olive to brownish olive or olive brown over a yellow ground color on the lower portion, with tiny darkbrown punctae; flesh pale yellow, becoming olive yellow to dull mustard yellow at the base, with red or brown stains around larval tunnels. Habitat and season: solitary, scattered, or in groups under longleaf pine, often with bluejack oak (Quercus incana) nearby; late spring and summer. Distribution and frequency: Florida west along the Gulf Coast into Texas; occasional. Spore print: olive brown. Microscopic features: spores 10–13 × 4–6 µm, narrowly ellipsoid to subfusoid, smooth, pale ochraceous. Edibility: unknown. Lookalikes: Caloboletus inedulis (p. 191) has yellow pores that bruise blue and bitter-tasting flesh. The flesh or pores of other similar red-pored boletes stain blue when cut or bruised. Gen us B o t h i a H a l l i ng, Ba ron i & Bi n der The genus Bothia was erected in 2007 to accommodate a single species that had formerly been classified in several genera, including Boletinellus, Boletinus, and Suillus. The genus was named in honor of the late Ernst E. Both (1930–2012), a distinguished boletologist, researcher, and author. Bothia is based on the species Boletinus castanellus, which was originally described by

172  ·  Boletes of Eastern North America

Charles H. Peck (1833–1917). At this time, the genus has only one species, Bothia castanella. The key identifying characteristics include a dry, somewhat velvety cap, whitish flesh that does not stain blue when exposed or bruised, shallow and decurrent boletinoid pores, and a dry stalk that is often reticulate, at least at the apex.

Bothia castanella

Bothia castanella (Peck) Halling, Baroni & Manfr. Binder = Suillus castanellus (Peck) A. H. Sm. & Thiers = Boletinus castanellus Peck = Xerocomus castanellus (Peck) Snell & E. A. Dick = Boletinus squarrosoides Snell & E. A. Dick Common name(s): Both’s Bolete Overview: Bothia castanella has a combination of macroscopic and microscopic features that prompted past classification in seven different genera. The term castanella means “small chestnut,” a reference to the cap surface color. Cap: 2.5–10 cm wide, convex, becoming nearly plane or shallowly depressed in age; surface dry, covered with densely matted hairs when young, becoming

Buchwaldoboletus   ·  173

nearly glabrous at maturity, brown to yellow brown, reddish brown, or burgundy; flesh soft-textured, white to dingy whitish, not staining or sometimes slowly becoming pale rusty; odor not distinctive; taste not distinctive or weakly acidic. Pores and tubes: subdecurrent to decurrent, with coarse and angular to elongated, often compound, radially arranged pores, pinkish brown to golden brown or pale cinnamon brown with pinkish tones, staining ochraceous tawny to dark rusty brown, not staining blue when bruised; tubes 4–6(10) mm deep. Stalk: variable from nearly equal to tapered downward or less often enlarged toward the base, solid; surface dry, concolorous with the cap but paler, reticulate at least on the upper portion, with a white basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary or in groups, sometimes in caespitose clusters, under oak in woods and parklands; summer and fall. Distribution and frequency: Maine south to South Carolina, west to Ohio; infrequent but possibly locally common. Spore print: yellow brown. Microscopic features: spores 8–11 × 4.5–5.5 µm, ovate to narrowly ellipsoidal, smooth, yellowish. Edibility: unknown. Lookalikes: Suillus decipiens (p. 330) has a partial veil and ring zone. Gen us B uc h wa l d ob ol e t u s P i l át The genus Buchwaldoboletus was erected in 1969 to accommodate species formerly classified in the genus Boletus. Buchwaldoboletus is a small genus of about a dozen species distributed worldwide. Three species have been reported to date from eastern North America. Boletes in this genus are not mycorrhizal and are saprotrophic and lignicolous or parasitic. They have dry caps that often become cracked in age and yellow flesh that stains blue when exposed. Their stalks are solid and nearly equal or tapered in either direction, and they sometimes have reticulation at least on the upper portion.

174  ·  Boletes of Eastern North America

Buchwaldoboletus hemichrysus

Buchwaldoboletus hemichrysus (Berk. & M. A. Curtis) Pilát = Boletus hemichrysus Berk. & M. A. Curtis = Pulveroboletus hemichrysus (Berk. & M. A. Curtis) Singer Common name(s): Half Yellow Powdery Bolete Overview: The term hemichrysus means “half yellow,” a reference to the cap color. The key identification features include the bright-yellow to golden-yellow, powdery cap, yellow flesh that sometimes stains blue, yellow pores that become reddish brown, and mostly reddish-brown stalk. Cap: 3–8 cm wide, obtuse to convex, becoming broadly convex at maturity, margin even, incurved at first and remaining so well into maturity; surface dry, floccose-scaly to somewhat velvety, often cracked in age, usually coated with powder, bright yellow to golden yellow; flesh yellow, staining blue, sometimes slowly, or unchanging; odor not distinctive; taste not distinctive or slightly acidic. Pores and tubes: yellow at first, soon becoming reddish brown; attached to subdecurrent; tubes up to 1 cm deep. Stalk: 3–7 cm long, nearly equal or tapered in either direction, often with a narrowed base, dry, solid; color variable, like the cap with reddish or reddish-brown tints or reddish brown overall, often with yellowish basal mycelium; flesh concolorous with the cap flesh.

Buchwaldoboletus   ·  175

Habitat and season: solitary or scattered on the ground attached to buried roots or on pine wood; summer through early winter. Distribution and frequency: North Carolina south to Florida, west to Texas, distribution limits yet to be determined; rare to uncommon. Spore print: olive brown. Microscopic features: spores 6–9 × 2.5–4 µm, ellipsoid to subfusoid, smooth, yellowish. Edibility: unknown. Lookalikes: Buchwaldoboletus sphaerocephalus (p. 179) has a sulfur-yellow cap when young that becomes pale yellow to whitish in age, yellow pores that become dull yellow to brownish yellow at maturity, and a yellow stalk without any reddish or brown tones. Buchwaldoboletus lignicola (p. 177) has a scaly or velvety and suede-like, reddish-brown to yellow-brown cap and grows on trunks, stumps, or roots of pine trees.

Buchwaldoboletus lignicola (A)

Buchwaldoboletus lignicola (B)

Buchwaldoboletus   ·  177

Buchwaldoboletus lignicola (C)

Buchwaldoboletus lignicola (Kallenb.) Pilát = Boletus lignicola Kallenb. = Pulveroboletus lignicola (Kallenb.) E. A. Dick & Snell Common name(s): none. Overview: This unusual and seldom-encountered bolete is always a treat to find. Its growth on pine wood, apparent association with the polypore Phaeolus schweinitzii (Fr.) Pat., and general morphology make it rather easy to identify. The term lignicola means “growing on wood.” The cap surface immediately stains very intense black with NH4OH and is negative with FeSO4. Because this bolete grows on wood, it could be mistaken for a stalked polypore. Phaeolus schweinitzii is a polypore that often fruits with this bolete. Evidence strongly suggests that Buchwaldoboletus lignicola is not saprotrophic but instead is parasitic on Phaeolus schweinitzii (Nuhn et al. 2013). Cap: 2.5–12 cm wide, convex, becoming broadly convex in age, margin wavy, with a narrow band of sterile tissue, inrolled when young and remaining so well into maturity; surface dry, scaly or velvety and suede-like, sometimes finely cracked in age, reddish brown to yellow brown, occasionally darker brown at the margin, sometimes staining darker brown when bruised; flesh firm, pale yellow, quickly or sometimes slowly staining blue when cut; odor sweet, somewhat minty or citrus-like; taste weakly acidic or not distinctive.

178  ·  Boletes of Eastern North America

Pores and tubes: yellow to golden yellow at first, becoming greenish yellow in age, staining dark greenish blue to blackish blue and then fading to reddish brown when bruised, attached to subdecurrent; tubes 3–12 mm deep. Stalk: nearly equal or tapered in either direction, usually narrowed at the base, frequently eccentric, solid; surface dry, somewhat velvety, golden mustard yellow at the apex and reddish brown to yellow brown below, staining darker brown when bruised, often finely reddish reticulate at the apex, basal mycelium yellow; flesh rusty toward the base, otherwise concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered on pine trees, stumps, or roots, sometimes on the ground near stumps, in pine or mixed oak and pine woods; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Michigan, distribution limits yet to be established; rare. Spore print: olive brown. Microscopic features: spores 6–10 × 3–4 µm, ellipsoid, smooth, yellowish. Edibility: unknown. Lookalikes: Buchwaldoboletus hemichrysus (p. 174) is similar but has a brighter-yellow to golden-yellow and very powdery cap, flesh that lacks a distinctive odor, taste that is not distinctive or slightly acidic, and yellow pores that soon become reddish brown. Buchwaldoboletus sphaerocephalus (p. 179) is similar but has a sulfur-yellow cap when young that becomes pale yellow to whitish in age, yellow pores that become dull yellow to brownish yellow at maturity, and a yellow stalk without any reddish or brown tones.

Buchwaldoboletus sphaerocephalus

Buchwaldoboletus   ·  179

Buchwaldoboletus sphaerocephalus (Barla) Watling & T. H. Li = Boletus sphaerocephalus Barla = Boletus hemichrysus f. sphaerocephalus (Barla) Estadès & Lannoy = Buchwaldoboletus sulphureus (Quél.) Watling & N. M. Greg. = Phlebopus sulphureus Singer Common name(s): none. Overview: The term sphaerocephalus means “round cap.” This bolete’s key identification features include the yellow cap that becomes cracked in age, yellow flesh that typically stains blue, yellow pores that stain blue and then slowly brownish, and yellow stalk. Cap: 4–12 cm wide, nearly spherical at first, becoming convex to broadly convex at maturity, margin incurved and remaining so well into maturity, even or nearly so, often wavy or lobed in age; surface dry to slightly viscid, glabrous, often cracked in age, sulfur yellow when young, becoming pale yellow to whitish at maturity; flesh sulfur yellow to pale yellow, typically staining blue when exposed but sometimes slowly or not at all; odor not distinctive; taste somewhat bitter or not distinctive. Pores and tubes: yellow at first, becoming dull yellow to brownish yellow, depressed at the stalk in age, staining blue and then slowly brownish when bruised; tubes 8–15 mm deep. Stalk: nearly equal down to a tapered base, solid; surface dry, nearly glabrous, yellow, with yellow basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups or clusters in pine sawdust, on pine stumps, or in the surrounding soil or sometimes fruiting in woodsheds or woodstalls that contain pine wood; summer and fall or yearround in woodsheds or woodstalls. Distribution and frequency: eastern Canada south to Florida, west along the Gulf Coast to Texas; rare. Spore print: olive brown. Microscopic features: spores 7–9 × 3–4 µm, oblong to ellipsoid, smooth, yellow. Edibility: unknown. Lookalikes: Buchwaldoboletus lignicola (p. 177) is similar but has a yellow-brown to reddish-brown, scaly to velvety, and suede-like cap; a yellow-brown to reddish-brown stalk; pale-yellow flesh that has a sweet, citrus-like odor; and yellow to golden-yellow pores that quickly stain dark greenish blue to blackish blue.

180  ·  Boletes of Eastern North America

Gen us B u t y r i b ol e t u s A ror a & J. L . Fr a n k The genus Butyriboletus was erected in 2014 to accommodate species formerly classified in the genus Boletus, section Appendiculati, commonly known as the “butter boletes.” The name Butyriboletus means “butter bolete.” The genus presently contains fourteen species from Asia, Europe, North Africa, and North America. Three species are known to occur in eastern North America. Molecular data and macroscopic features place this group in a clade separate from Boletus sensu stricto. Species in this genus have a yellow tube layer that often bruises blue, a yellow reticulate stalk, and firm, yellow-tinged flesh that may or may not turn blue when exposed.

Butyriboletus brunneus (A)

Butyriboletus   ·  181

Butyriboletus brunneus (B)

Butyriboletus brunneus (Peck) Arora & Frank = Boletus speciosus var. brunneus Peck = Boletus pseudopeckii A. H. Sm. & Thiers Common name(s): Showy Reddish Brown Bolete Overview: Butyriboletus brunneus is one of the earliest boletes to appear in spring, often in early June. It is an excellent edible species, although the flesh is often riddled with insect larvae or eaten by slugs, so unblemished specimens are not often found. The term brunneus means “reddish brown,” a reference to this handsome bolete’s cap color. Cap: 4–14 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, even; surface dry, glabrous or nearly so, reddish brown or yellow brown to olive brown; flesh pale yellow, quickly staining blue when exposed; odor and taste not distinctive. Pores and tubes: bright yellow to yellow when fresh, becoming dull yellow to olive yellow in age, instantly staining blue when bruised, depressed at the stalk; tubes 8–22 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, finely reticulate overall or at least over the upper half, yellow on the upper portion, pinkish red to purplish red or at least tinged reddish on the lower portion, especially near the base, staining blue when bruised; flesh concolorous with the cap flesh or deeper yellow, staining like the cap flesh when exposed.

182  ·  Boletes of Eastern North America

Habitat and season: solitary, scattered, or in small groups under broadleaf trees or conifers, especially hemlock, or in mixed woods; late spring, summer, and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 10–15 × 3–5.5 µm, narrowly oblong to subfusoid, smooth, ochraceous. Edibility: edible and excellent. Lookalikes: Boletus speciosus (p. 155) is very similar but has a bright rose-red to rose-pink cap that becomes pinkish with olive tones or pinkish brown to olive brown at maturity. Alessioporus rubriflavus (p. 56) has a dark wine-red cap when young that becomes olive to brownish olive with red streaks and splashes at maturity, conspicuous reticulation over at least the upper half of the stalk, dark wine-red flesh near the stalk base, and larger spores.

Butyriboletus peckii

Butyriboletus peckii (Frost) K. Zhao, Z. L. Yang & Halling = Boletus peckii Frost = Caloboletus peckii (Frost) Vizzini Common name(s): Peck’s Bolete Overview: The species name honors American mycologist Charles H. Peck (1833–1917). For an interesting account of how this bolete may have been described, see Both 1993. The key identification features include the somewhat velvety reddish cap, whitish to pale-yellow flesh that stains slightly blue, yellow pores that stain blue and then brownish, and a red stalk that is typically reticulate overall. Cap: 2.5–9.5 cm wide, convex, becoming broadly convex to nearly plane at maturity, margin with a narrow band of sterile tissue when young, becoming even with maturity; surface dry, somewhat velvety, deep red to dull red

Butyriboletus   ·  183

or rose red, fading to brownish rose or pale tan to buffy brown from the disc outward; flesh whitish to pale yellow, staining slightly and sometimes slowly blue when exposed, burgundy red around larval tunnels; odor not distinctive; taste variously described as unpleasant, somewhat bitter, or tart. Pores and tubes: bright yellow to golden yellow at first, becoming dingy olive yellow to brownish yellow in age, staining blue and then slowly brownish when bruised; tubes 4–15 mm deep. Stalk: enlarged downward to nearly equal, solid; surface dry, minutely and densely pruinose, rosy red to darker red or purplish red overall or with yellow at the apex, sometimes with olive or brownish tinges near the base, typically reticulate nearly overall or at least on the upper portion, basal mycelium whitish to pale yellow; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups in broadleaf forests and mixed woods, especially with beech and oak, summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Tennessee and Indiana, distribution limits yet to be established; occasional. Spore print: ochraceous brown. Microscopic features: spores 10–14 × 3.5–4 µm, ellipsoidal, smooth, yellowish. Edibility: unknown. Lookalikes: The Baorangia bicolor complex (p. 74) is very similar but lacks reticulation on the stalk or is rarely reticulate only at the apex.

Butyriboletus roseopurpureus (A)

184  ·  Boletes of Eastern North America

Butyriboletus roseopurpureus (B)

Butyriboletus roseopurpureus (Both, Bessette & Roody) K. Zhao, Z. L. Yang & Halling = Boletus roseopurpureus Both, Bessette & Roody Common name(s): none. Overview: The term roseopurpureus means “pinkish purple,” a reference to this bolete’s cap color. Except for the cap color, this mushroom is all yellow. The flesh and pores stain rapidly blue when cut or bruised. The stalk, which also bruises blue, is conspicuously reticulate. When NH 4OH is applied to the cap, it produces a dingy pale-yellowish to dull-orange stain. Cap: 7–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, later decurved, projecting as a sterile band of tissue; surface viscid-tacky in wet weather but drying quickly, somewhat velvety or covered with matted fibers, strikingly pinkish purple when fresh, becoming darker purplish pink to dark purplish red, at times mottled in these colors, marginal areas becoming grayish in age; flesh pale yellow, instantly staining dark blue when exposed but soon fading to pale-slate color; odor fragrant; taste very sour, like lemon. Pores and tubes: lemon yellow or more golden at first, becoming yellowish olive to greenish olive at maturity, instantly staining dark blue to greenish

Caloboletus   ·  185

blue when bruised, soon fading to pale greenish blue or greenish gray; tubes 5–10 mm deep. Stalk: equal or slightly tapered downward; reticulate nearly the entire length or at least over the upper two-thirds, sometimes strongly raised, covered with tiny matted fibers on the lower portion, bright yellow nearly overall, occasionally with burgundy-red areas near the base, with a white base and a white basal mycelium, instantly staining dark blue when bruised; flesh deep yellow, sometimes burgundy red in the basal area, instantly staining blue when exposed but soon fading to a pale-slate color. Habitat and season: solitary or in small groups in mixed woods of oak, beech, hemlock, and maple, apparently associated with red oak; summer and early fall. Distribution and frequency: New England south to Florida, distribution limits yet to be established; occasional. Spore print: light brownish olive. Microscopic features: spores 9.4–13 × 2.7–3.5 µm, narrowly ellipsoid to subfusoid, smooth, pale ochraceous in Melzer’s. Edibility: unknown. Lookalikes: This Bolete may have been misidentified in the past as Boletus speciosus Frost (p. 155). However, Charles Frost described the cap color of his species as “scarlet lake red,” a bright red without purplish tones. In addition, the spores of Boletus speciosus are longer and wider, 11–15 × 3–5.5 µm. Boletus rubissimus A. H. Sm. (not illustrated) is similar to Butyriboletus roseopurpureus but differs sharply in the weak or absent blue staining of its exposed flesh, a poorly developed reticulum on the stalk, yellow mycelium at the stalk base, and shorter, somewhat broader spores, 9–11 × 3–4 µm. Gen us C a l ob ol e t u s Vi zzi n i The genus Caloboletus was erected in 2014 to accommodate species formerly classified in the genus Boletus. The name Caloboletus is derived from the Greek word calo, which means “beautiful or nice,” a reference to the pinkish or reddish tinges on the stalks of most of the species in the genus. The genus includes ten species worldwide, four of which occur in eastern North America. Species in this genus often have whitish to grayish caps and yellow pores and tubes. The exception is Caloboletus firmus, which has orange to red pores when young and whitish to yellow flesh that usually stains blue and tastes bitter. This genus has been supported by molecular analysis.

186  ·  Boletes of Eastern North America

Caloboletus calopus

Caloboletus calopus (Pers.) Vizzini = Boletus calopus Pers. = Boletus calopus var. frustosus (Snell & E. A. Dick) O. K. Mill. & Watling = Boletus frustosus Snell & E. A. Dick Common name(s): Bitter Beech Bolete, Scarlet-stemmed Bolete Overview: The common name “Bitter Beech Bolete” is a bit misleading because this species occurs primarily with hemlock in northeastern North America. The term calopus means “beautiful foot,” a reference to this handsome bolete’s stalk. Cap: 6–20 cm wide, rounded when young, becoming convex to broadly convex in age, margin incurved at first, with a narrow band of sterile tissue; surface dry, somewhat velvety, becoming nearly smooth at maturity, slightly to conspicuously cracked in age, light brown to grayish brown or olive brown at first, becoming paler olive brown to dark yellow brown as it ages; flesh whitish to pale yellow, rapidly staining blue when exposed; odor somewhat pungent or not distinctive; taste bitter. Pores and tubes: pale yellow at first, becoming olive yellow to brownish yellow at maturity, rapidly staining blue when bruised, typically depressed at the stalk in age; tubes up to 2 cm deep. Stalk: nearly equal or enlarged downward, sometimes with a pinched base, solid; surface dry, scurfy-punctate or nearly smooth, yellowish toward the apex, carmine red to nearly scarlet below, often reddish brown toward the base, typically staining blue when handled or bruised, conspicuously reticulate at least over the upper portion or sometimes nearly overall, reticulation

Caloboletus   ·  187

rarely inconspicuous or lacking; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in conifer woods, especially with hemlock or spruce, or less often in broadleaf woods with beech or oak, usually found at higher elevations; summer and fall. Distribution and frequency: eastern Canada south to New England and New York, west to Michigan, distribution limits yet to be established; uncommon. Spore print: olive brown. Microscopic features: spores 13–17(19) × (3)4.5–6 µm, ellipsoidal, smooth, yellowish; tube trama septa amyloid in Melzer’s reagent. Edibility: unknown but probably not edible because of the bitter-tasting flesh. Lookalikes: Caloboletus inedulis (p. 191) is typically smaller. It has a whitish to grayish-white or sometimes pale pinkish-gray cap that darkens to olivaceous brown and pale grayish brown in age, paler-pinkish to pinkish-red coloration on its stalk, and smaller spores that measure 9–13 × 3.3–4.5 µm; the septa of its tube trama are not amyloid in Melzer’s reagent. The photograph of Boletus roseipes (A) = Caloboletus roseipes (Bessette, Both & A. R. Bessette) Vizzini is mislabeled in North American Boletes (Bessette, Roody, and Bessette 2000, 308). It should be Caloboletus calopus. Caloboletus roseipes (p. 192) has an olive-brown to grayish-brown cap when young that becomes pale grayish brown to grayish tan on older specimens. Its stalk is bright yellow near the apex, with rosy-red or yellow with rosy-red areas below, and it has smaller spores.

Caloboletus firmus (A)

Caloboletus firmus (B)

Caloboletus firmus (C)

Caloboletus   ·  189

Caloboletus firmus (Frost) Vizzini = Boletus firmus Frost = Boletus piedmontensis Grand & A. H. Sm. = Boletus satanas var. americanus Coker & Beers Common name(s): Piedmont Bolete Overview: The term firmus means “firm,” a reference to this bolete’s sturdy, dense flesh, especially when it is young and fresh. The whitish to grayish, tan, or pale grayish-olive cap, whitish flesh that stains blue when exposed, red to red-orange pores that bruise blue, and the red reticulation on the stalk are its distinctive features. Cap: 6–15 cm wide, convex, becoming broadly convex in age, margin even, inrolled when young; surface dry, glabrous but appearing finely velvety, whitish to grayish, grayish brown, tan, pinkish tan, or pale grayish olive; flesh firm, whitish to pale yellow, staining blue when exposed, sometimes slowly or slightly; odor wax-like or not distinctive; taste mild to slightly bitter. Pores and tubes: pinkish at first, soon becoming red or red orange, staining blue when bruised or bruising yellow on very young specimens; tubes 8–20 mm deep. Stalk: nearly equal or tapered in either direction, solid; surface dry, nearly smooth, concolorous with the cap, sometimes with reddish tints, staining olive to brownish when handled or bruised, with fine red reticulation near the apex or nearly overall or sometimes completely lacking; flesh concolorous with and staining like the cap flesh or tinged pinkish near the base. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak and beech; late spring, summer, and fall. Distribution and frequency: eastern Canada south to Georgia, west to Mississippi; occasional. Spore print: olive brown. Microscopic features: spores 9–15 × 3.5–5 µm, subellipsoid, smooth, yellowish. Edibility: inedible because of the bitter taste. Lookalikes: Both Caloboletus inedulis (p. 191) and Caloboletus roseipes (p. 192) are similar, but they have yellow pores and red areas on their stalks.

Caloboletus inedulis (A)

Caloboletus inedulis (B)

Caloboletus   ·  191

Caloboletus inedulis (Murrill) Vizzini = Boletus inedulis (Murrill) Murrill = Boletus subclavatipes subsp. inedulis (Murrill) Singer = Ceriomyces inedulis Murrill Common name(s): Rosy-footed Oak Bolete Overview: The term inedulis means “not edible,” a reference to this bolete’s bitter-tasting flesh. The key identification features include a whitish to grayish cap, whitish to yellowish flesh that stains blue, yellow pores that stain blue, and a reticulated yellow stalk with red tinges. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved to inrolled at first, remaining so well into maturity, with a narrow band of sterile tissue; surface dry, covered with tiny, cottony, matted fibers at first, becoming conspicuously cracked, with whitish flesh showing in the fissures, color whitish to grayish white or pale pinkish gray, sometimes with pinkish-rose areas especially toward the margin, darkening to olivaceous brown to pale grayish brown in age; flesh whitish to yellowish, staining blue when exposed; odor not distinctive; taste bitter. Pores and tubes: pale yellow, becoming olive yellow at maturity, staining dark blue and then brownish when bruised, depressed at the stalk in age; tubes 8–16 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, pale yellow to yellow, at times with a pinkish zone at the apex or pinkish to pinkish red below, staining blackish brown near the base when handled, with reddish or brownish reticulation at least on the upper portion; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups in broadleaf or mixed woods, associated with oak, hickory, hemlock, or spruce; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; infrequent. Spore print: olive brown. Microscopic features: spores 9–13 × 3.3–4.5 µm, subfusoid, smooth, ochraceous; tube trama lacking amyloid septa. Edibility: inedible. Lookalikes: Caloboletus roseipes (p. 192) is similar but has an olive-brown to grayish-brown cap when young that becomes pale grayish brown to grayish tan on older specimens, a thicker stalk, and different macrochemical test reactions, and it is associated with hemlock. Caloboletus calopus (p. 186) is typically larger, with an olive-brown to gray-brown cap that darkens to dark yellow brown in age, darker-red coloration on its stalk, and larger spores that measure 13–19 × 5–6 µm.

192  ·  Boletes of Eastern North America

Caloboletus roseipes

Caloboletus roseipes (Bessette, Both & A. R. Bessette) Vizzini = Boletus roseipes Bessette, Both & A. R. Bessette Common name(s): Rosy-footed Hemlock Bolete Overview: The term roseipes means “rosy foot,” a reference to the rosy-red coloration on this bolete’s stalk. The key identification features include an olive-brown to grayish-tan cap, white to pale-yellow flesh that stains blue, yellow pores that stain blue, and a yellow stalk with red areas. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first and remaining so well into maturity, with a narrow band of sterile tissue; surface dry, somewhat velvety at first, becoming appressed-tomentose and somewhat cracked in age, olive brown to grayish brown when young, becoming pale grayish brown to grayish tan on older specimens; flesh white to pale yellow, staining blue when exposed; odor not distinctive; taste bitter. Pores and tubes: pale yellow, becoming olive yellow to brownish yellow at maturity, staining blue when bruised; attached or slightly depressed at the stalk; tubes 5–16 mm deep, concolorous with and staining like the pores. Stalk: enlarged downward to a pinched base or nearly equal, solid; surface dry, scurfy-punctate nearly overall, bright yellow near the apex, rosy red or yellow with rosy-red areas below, staining blue when bruised,

Chalciporus   ·  193

reticulation reddish and variable, distinct on the upper portion of some specimens and absent on others; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under hemlock or in mixed woods with hemlock and spruce; summer and fall. Distribution and frequency: eastern Canada south to New England and New York, distribution limits yet to be established; infrequent. Spore print: olive brown. Microscopic features: spores 10–14 × 3–4.5 µm, narrowly oblong to subfusoid, smooth, ochraceous in KOH or Melzer’s, lacking an apical pore; tube trama lacking amyloid septa. Edibility: inedible because of the bitter taste. Lookalikes: Caloboletus calopus (p. 186) is similar but has a darker-brown cap, a darker-red stalk with prominent reticulation, and larger spores. The photograph of Boletus roseipes (A) = Caloboletus roseipes (Bessette, Both & A. R. Bessette) Vizzini in North American Boletes (Bessette, Roody, and Bessette 2000, 308) is mislabeled and should be Caloboletus calopus. Caloboletus inedulis (p. 191) is also similar but has a whitish to grayish-white or pale pinkish-gray cap that becomes darker brown and conspicuously cracked in age and is associated with oaks. Gen us C h a l ci p oru s Bata i l l e The genus Chalciporus was erected in 1908 to accommodate species formerly classified in several genera, including Boletus and Suillus. The name Chalciporus means “copper-colored pores,” a reference to the color of the mature pores. The genus has less than ten species described from North America. They are small- to medium-size terrestrial boletes with mature pores that are pinkish, reddish, dull ocher to yellow brown, cinnamon, or orange to reddish brown. Their stalks are solid and smooth, often with bright-yellow mycelium at the base. Spore-print colors include yellow brown, cinnamon brown, brown, rusty cinnamon, and dark smoky olive.

Chalciporus piperatus

Chalciporus piperatoides (A)

Chalciporus   ·  195

Chalciporus piperatoides (B)

Chalciporus piperatus (Bull.) Bataille = Boletus piperatus Bull. Common name(s): Peppery Bolete Overview: The term piperatus means “peppery.” The combination of a brownish cap, a stalk with bright-yellow mycelium at the base, and peppery-tasting flesh makes this small bolete easy to recognize. Cap: 1.6–9 cm wide, convex, becoming nearly plane in age, margin even; surface smooth, dry or somewhat viscid, sometimes cracked in age, buff to yellow brown, orange brown, or reddish brown; flesh pale yellow or tinged reddish, becoming dingy purplish brown in age, not staining blue when cut or bruised; odor not distinctive; taste bitter, hot and peppery. Pores and tubes: attached to slightly depressed, with small angular pores that sometimes appear to radiate from the stalk, dull cinnamon, reddish cinnamon, or cinnamon brown, becoming darker reddish brown in age, sometimes staining brown when cut or bruised; tubes 3–10 mm deep. Stalk: equal or tapered downward, solid; surface dry, concolorous with the cap or paler, with a bright-yellow basal mycelium; flesh lemon yellow, unchanging when cut or exposed. Habitat and season: solitary, scattered, or in groups under conifers or broadleaf trees; spring, summer, and fall. Distribution and frequency: widely distributed; fairly common.

196  ·  Boletes of Eastern North America

Spore print: brown to cinnamon brown. Microscopic features: spores 9–12 × 4–5 µm, narrowly fusiform, smooth, pale brown. Edibility: unknown. Lookalikes: Chalciporus piperatoides (A. H. Sm. & Thiers) T. J. Baroni & Both (see photos, p. 194) is nearly identical, but its pores quickly bruise blackish blue. Chalciporus rubinellus (p. 198) has a broadly conical cap, pinkish-red pores that do not bruise blue, and mild-tasting flesh.

Chalciporus pseudorubinellus

Chalciporus pseudorubinellus (A. H. Sm. & Thiers) L. D. Gomez = Boletus pseudorubinellus A. H. Sm. & Thiers Common name(s): none. Overview: The term pseudorubinellus means “false rubinellus” in reference to this bolete’s similarity with Chalciporus rubinellus (p. 198). The key identification features include the variable yellowish or pinkish-red cap, pale-yellow flesh that does not stain when exposed, bright rose-red pores, and a rose-pink stalk with a yellow base. Cap: 1.5–6 cm wide, convex to broadly convex, sometimes broadly umbonate, margin even; surface dry, glabrous or nearly so, color variable, yellowish or pinkish red, becoming pinkish cinnamon in age; flesh pale yellow, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: bright rose red when fresh, fading to orangish or yellowish brown in age, not staining blue when bruised; tubes 6–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, finely pruinose, mostly rose pink, typically yellow near the base, with yellow basal mycelium; flesh concolorous with the cap flesh.

Chalciporus   ·  197

Habitat and season: scattered or in groups in conifer woods, often among mosses; summer and fall. Distribution and frequency: New England and New York west to Minnesota, distribution limits yet to be established; occasional. Spore print: brown. Microscopic features: spores 9–13 × 3–4 µm, fusoid, smooth, distinctly olivaceous. Edibility: unknown. Lookalikes: Chalciporus rubinellus (p. 198) has a redder cap, especially when immature, lacks yellow mycelium at the stalk base, and has larger spores, 12–15 × 3–5 µm. Chalciporus rubritubifer (p. 199), known only from the Myakka River State Park in Florida, has a dark orange-yellow cap when young that becomes orange brown to reddish brown at maturity, grayish-red to dull-red pores that become brownish orange to copper in age, and a stalk that is concolorous with the cap or paler, with a sparse pale-yellow basal mycelium.

Chalciporus rubinellus (A)

198  ·  Boletes of Eastern North America

Chalciporus rubinellus (B)

Chalciporus rubinellus (Peck) Singer = Boletus rubinellus Peck = Suillus rubinellus (Peck) Singer Common name(s): Purple-red Bolete Overview: The Purple-red Bolete is one of the smallest of the eastern boletes and the only one in the genus that lacks even a hint of yellow at the stalk base. The term rubinellus means “somewhat red” or “reddish,” alluding to this bolete’s overall colors. The cap surface stains blood red and then dingy orange with NH 4OH. Cap: 2–4.5 cm wide, broadly conical when young, becoming convex in age, margin even; surface dry, somewhat velvety when young, becoming cracked in age, red or reddish when young, more yellow in age; flesh bright yellow, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: bright rose red when young, becoming dull rose red to rose pink in age, not staining blue when bruised; tubes 6–10 mm deep. Stalk: nearly equal, solid; surface dry, initially red or reddish, sometimes mixed with yellow, lacking yellow mycelium at the base; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in mixed woods or conifer forests; summer and fall.

Chalciporus   ·  199

Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: brown. Microscopic features: spores 12–15 × 3–5 µm, subfusoid, smooth, pale dull ochraceous. Edibility: unknown. Lookalikes: Chalciporus pseudorubinellus (p. 196) has a less-red cap, especially when immature; yellow mycelium at the stalk base and smaller spores, 9–13 × 3–4 µm. Chalciporus rubritubifer (p. 199), known only from the Myakka River State Park in Florida, has a dark orange-yellow cap when young that becomes orange brown to reddish brown at maturity, grayish-red to dull-red pores when young that become brownish orange to copper in age, and a stalk that is concolorous with the cap or paler, with a sparse pale-yellow basal mycelium.

Chalciporus rubritubifer

Chalciporus rubritubifer (Kauffman) Singer = Boletus rubritubifer Kauffman Common name(s): none. Overview: The term rubritubifer means “red or reddish tubes,” a reference to the color of this bolete’s tubes and pores. The cap surface stains reddish brown with KOH or NH4OH. Cap: 1.3–5 cm wide, hemispherical to convex, becoming broadly convex in age, margin incurved at first, sometimes with a narrow band of sterile tissue; surface dry, somewhat velvety, becoming finely cracked in age with yellow flesh showing in the fissures, dark orange yellow and staining brown when

200  ·  Boletes of Eastern North America

young, becoming reddish brown to orange brown at maturity; flesh yellow, sometimes with grayish or orange hues, showing a reddish line or zone above the tube layer, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: grayish red to dull red when young, becoming brownish orange to copper at maturity, attached to slightly depressed at the stalk, not staining blue when bruised; tubes 2–5 mm deep. Stalk: tapered downward or nearly equal, often curved near the base, solid; surface dry, faintly longitudinally striate to finely punctate, concolorous with the cap or paler yellow, usually reddish at the apex, becoming dark reddish brown at the base, with a sparse pale ochraceous-yellow basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under oaks, pines, and palms; summer and fall. Distribution and frequency: known only from the Myakka River State Park in Florida; occasional. Spore print: yellow brown. Microscopic features: spores 9–18 × 4–6 µm, ellipsoidal to subfusoid, smooth, yellowish. Edibility: edible, with good flavor, according to the late Florida mycologist Robert S. Williams (1939–2014). Lookalikes: Chalciporus pseudorubinellus (p. 196) has a yellowish or pinkish-red cap when young that becomes pinkish cinnamon in age, lacks yellow basal mycelium, and is northern in distribution. Chalciporus rubinellus (p. 198) has a red to reddish cap when young that becomes more yellowish at maturity, a stalk with yellow mycelium at the base, and a northern distribution. Gen us C ya nob ol e t u s Gel a r di, Vi zzi n i & Si mon i n i The genus Cyanoboletus was erected in 2014 to accommodate three species formerly classified in the genus Boletus, one of which occurs in eastern North America. Species in this genus have yellow flesh that instantly discolors dark indigo blue to blackish blue when exposed. The name Cyanoboletus is derived from the Greek word cyano, which means “blue,” a reference to the color reaction of the exposed flesh. This genus is supported by molecular analysis.

Cyanoboletus pulverulentus (A)

Cyanoboletus pulverulentus (B)

202  ·  Boletes of Eastern North America

Cyanoboletus pulverulentus (Opat.) Gelardi, Vizzini & Simonini = Boletus pulverulentus Opat. Common name(s): none. Overview: The most apparent identifying feature of this small, dark bolete is that all parts quickly stain dark blue when handled or bruised. Also, the cap surface displays a green flash with the application of NH4OH. The term pulverulentus means “dusty or powdery,” a reference to this bolete’s cap, which often appears to be lightly coated with a layer of dust or powder. Cap: 4–12.5 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat dusty or powdery and slightly velvety at first, becoming glabrous and often somewhat shiny in age, dark yellow brown to blackish brown or dark cinnamon brown, often developing reddish tints in age, instantly staining blackish blue when bruised or handled; flesh yellow, instantly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow when young, darkening to golden yellow and then to brownish yellow at maturity, instantly staining dark blue and then dull brown when bruised; tubes 6–12 mm deep. Stalk: nearly equal or sometimes enlarged downward, solid; surface dry, pruinose at the apex, yellow on the upper portion and darker orange yellow on the lower portion, typically reddish brown and pruinose toward the base, at first quickly staining dark blue and then slowly to dull brown when handled, often with raised longitudinal ridges; flesh reddish brown in the base, yellow above the base, instantly staining blue when exposed. Habitat and season: scattered or in groups under conifers or broadleaf trees; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: dark olive to olive brown. Microscopic features: spores 11–15 × 4–6 µm, fusoid to ellipsoidal, smooth, yellowish. Edibility: edible. Lookalikes: Boletus oliveisporus (p. 127) is similar but typically is larger in size, and its fulvous to cinnamon-brown cap does not show a green flash reaction with NH4OH. Gen us E x su d op oru s Vi zzi n i, Si mon i n i & Gel a r di The genus Exsudoporus was erected in 2014 to accommodate three species that have pores often beaded with yellow drops when specimens are young and fresh and that were formerly classified in the genus Boletus. The term exsudoporus means “exuding droplets from the pores.” Species in this genus have red to

Exsudoporus   ·  203

reddish-brown, dry to slightly viscid caps, reddish-orange to dark-red or rarely yellow-orange pores, conspicuously reticulate stalks, yellow flesh that quickly stains dark blue when exposed, and an olive-brown spore print. Molecular analysis shows that they are closely related.

Exsudoporus floridanus (A)

Exsudoporus floridanus (B)

204  ·  Boletes of Eastern North America

Exsudoporus floridanus (Singer) Vizzini, Simonini & Gelardi = Boletus floridanus (Singer) Murrill = Boletus frostii ssp. floridanus Singer = Suillellus floridanus (Singer) Murrill Common name(s): none. Overview: The term floridanus means “Florida,” a reference to the location where this bolete was originally collected. The red cap, yellowish flesh that rapidly stains blue, red pores that stain blue when bruised, and red stalk with red reticulations are the diagnostic features of this strikingly beautiful bolete. The cap surface stains olive black with NH4OH. Cap: 4–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, even or nearly so; surface dry, somewhat velvety, pinkish red, rose red, purplish red, or brownish red, typically yellowish or whitish along the margin; flesh pale to bright yellow, quickly staining dark blue when exposed; odor and taste not distinctive. Pores and tubes: color variable, reddish orange to pinkish red, sometimes with yellowish tints, or yellow overall with reddish tints, often beaded with yellow drops when young and fresh, usually depressed at the stalk at maturity, rapidly staining blue; tubes 6–12 mm deep. Stalk: often bulbous when young, usually club-shaped or sometimes nearly equal at maturity, solid; surface dry, typically yellow at the apex and red below or sometimes red overall, with conspicuous longitudinally elongated red reticulation at least over the upper half; flesh pale to bright yellow, quickly staining dark blue when exposed. Habitat and season: scattered or in groups in sandy soil under oaks; summer and fall, also winter in the southern part of its range. Distribution and frequency: Tennessee and the Coastal Plain of North Carolina south to Florida, west to Texas. Spore print: olive brown. Microscopic features: spores 13–18 × 4–5 µm, ellipsoidal to subfusiform, smooth, pale yellow brown. Edibility: edible but of very poor quality. Lookalikes: Exsudoporus frostii (p. 205) has darker-red coloration overall and much coarser, raised reticulation.

Exsudoporus   ·  205

Exsudoporus frostii

Exsudoporus frostii (J. L. Russell) Vizzini, Simonini & Gelardi = Boletus frostii J. L. Russell Common name(s): Frost’s Bolete Overview: This is one of the most beautiful and distinctive boletes in eastern North America. The combination of a shiny, candy-apple-red cap, dark-red pores that exude golden moisture droplets when fresh, and a deeply lacerate-reticulate stalk is unmistakable. The species name honors Charles C. Frost (1805–1880), a Vermont shoemaker turned mycologist who was the first to describe several North American species of boletes. Cap: 5–15 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, margin incurved and even when young; surface smooth, shiny, dry or tacky when moist, deep blood red to candy-apple red, often with a narrow yellow band at the margin, sometimes developing yellowish areas in age; flesh pale yellow to dull white, quickly staining blackish blue when exposed; odor not distinctive; taste sour, acidic. Pores and tubes: dark red at first, becoming paler red in age, often exuding golden moisture droplets when young, instantly staining blackish blue when bruised, depressed at the stalk at maturity; pores minute; tubes 6–15 mm deep. Stalk: nearly equal or enlarged downward, often curved at the base, solid; surface dry, dark red to pinkish red with a white to yellowish base, staining

206  ·  Boletes of Eastern North America

blackish blue when handled or bruised, coarsely and deeply reticulate over the entire length. Habitat and season: solitary to scattered or in groups in broadleaf or mixed woods, especially with oak; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 11–17 × 4–5 µm, ellipsoidal to subfusiform, smooth, pale brown. Edibility: edible, with a pronounced lemony flavor. Although most people can consume this bolete without negative effects, there are reports of gastrointestinal upset from eating it. Lookalikes: Exsudoporus floridanus (p. 204) is similar but has a lighter-red cap and less-prominent reticulation on its stalk. Gen us F ro s t i e l l a Mu r r i ll The genus Frostiella was erected in 1942 to accommodate two species formerly classified in various genera, including Boletellus, Boletus, Ceriomyces, and Suillus. The name Frostiella honors Vermont mycologist Charles C. Frost (1805–1880). Two species—Frostiella betula and Frostiella russellii, which have fairly small caps on long, deeply grooved, and shaggy stalks that lack a partial veil or ring— were formerly assigned to this genus, but Frostiella betula has now been transferred to the genus Heimioporus, leaving only a single species. Frostiella russellii was formerly classified in the genus Boletellus, but molecular studies have shown that it does not belong there.

Frostiella russellii (A)

Frostiella   ·  207

Frostiella russellii (B)

Frostiella russellii (Frost) Murrill = Boletellus russellii (Frost) E. J. Gilbert = Boletus russellii Frost Common name(s): Russell’s Bolete Overview: Although not often encountered, this tall and stately bolete is easy to recognize by its dry, velvety to scaly brown cap and deeply grooved and shaggy stalk. The cap surface stains reddish with NH4OH and olive gray with FeSO4. The flesh stains reddish with KOH and blackish blue with FeSO4. The name russellii honors John Lewis Russell (1808–1873), American botanist and friend of Charles C. Frost, who named this distinctive bolete after him. Cap: 3–13 cm wide, rounded to convex, becoming broadly convex, margin strongly incurved and even; surface dry, velvety when young, becoming cracked or forming scale-like patches as it ages, yellow brown to reddish brown, cinnamon brown, or olive gray; flesh pale yellow to yellow, often brownish around larval tunnels, not staining blue when exposed but sometimes slowly staining brown; odor and taste not distinctive. Pores and tubes: yellow to greenish yellow, unchanging or becoming brighter yellow when bruised, typically somewhat depressed at the stalk at maturity; tubes up to 2 cm deep.

208  ·  Boletes of Eastern North America

Stalk: up to 20 cm long, equal or enlarging slightly downward, solid, often curved at the base; surface dry, often viscid at the base when moist, reddish brown to pinkish tan, deeply grooved and ridged for most or all of its length, branched or torn ridges creating a honeycomb or shaggy-bark effect; flesh concolorous with the cap flesh. Habitat and season: solitary or scattered on the ground and on humus under oak, hemlock, or pine; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Michigan; uncommon. Spore print: dark olive to olive brown. Microscopic features: spores 15–20 × 7–11 µm, ellipsoidal, longitudinally striate with deep grooves or wrinkled with a cleft in the wall at the apex, pale brown. Edibility: edible. Lookalikes: Heimioporus betula (p. 219) is somewhat similar but has a glabrous, shiny, red to orange cap and pitted spores. Gen us G y rop oru s Qu él. The genus Gyroporus was erected in 1886 to accommodate species formerly classified in the genus Boletus. Gyroporus is a fairly small genus with less than a dozen species reported from North America. The name Gyroporus means “round pores.” These species are small- to medium-size terrestrial boletes with dry caps that are typically somewhat velvety to floccose-scaly. Their stalks are hollow or have several cavities at maturity. They have white to pale-yellow flesh that is brittle and stains blue in some species and white to yellow, tiny, round pores. Their spore-print colors include pale to bright yellow or buff.

Gyroporus castaneus (A)

Gyroporus   ·  209

Gyroporus castaneus (B)

Gyroporus castaneus (Bull.) Quél. = Boletus castaneus Bull. Common name(s): Chestnut Bolete Overview: The Chestnut Bolete is fairly common, especially in central and northern woodlands, but it is easily overlooked owing to its small size and somewhat muted colors. The chestnut-brown to orange-brown cap, straw-colored pores, hollow or chambered stalk, and firm but brittle flesh are useful field characters. The cap surface stains amber orange with NH4OH and is negative with FeSO4. The epithet castaneus is derived from the genus name for chestnut trees, Castanea, and refers to the chestnut-brown color of the cap and stalk. Cap: 3–10 cm wide, convex to broadly convex, becoming nearly plane, sometimes slightly depressed, margin often split and flaring in age; surface velvety to nearly glabrous, dry, yellow brown to orange brown or reddish brown; flesh brittle, white, not staining when cut or bruised; odor not distinctive or faintly pungent; taste not distinctive. Pores and tubes: whitish to buff or yellowish, unchanging when bruised, depressed at the stalk to nearly free at maturity; tubes 5–8 mm deep.

210  ·  Boletes of Eastern North America

Stalk: nearly equal or often swollen in the midportion or below, sometimes constricted at the apex and base, brittle, stuffed with a soft pith, developing several cavities or becoming hollow in age; surface dry, uneven, concolorous with the cap or slightly paler toward the apex; flesh firm, brittle, white. Habitat and season: solitary, scattered, or in groups usually in broadleaf woods, especially with oak, but also in conifer woods; late spring, summer, and fall, also winter in the southern part of its range. Distribution and frequency: throughout eastern North America; fairly common. Spore print: pale yellow. Microscopic features: spores 8–13 × 5–6 µm, ellipsoidal to ovoid, smooth, hyaline. Edibility: edible by most accounts but not particularly good. Lookalikes: Gyroporus purpurinus (p. 213) is nearly identical except for the vinaceous to burgundy colors of the cap and stalk. Austroboletus gracilis var. gracilis (p. 70) differs by having pores that become pinkish brown at maturity, a more slender and solid stalk, softer flesh, and spores that are pinkish brown and pitted.

Gyroporus cyanescens

Gyroporus   ·  211

Gyroporus cyanescens (Bull.) Quél. = Boletus cyanescens Bull. = Gyroporus cyanescens var. cyanescens (Bull.) Quél. = Gyroporus cyanescens var. violaceotinctus Watling Common name(s): Staining Blue Bolete Overview: The term cyanescens means “becoming deep blue,” a reference to this bolete’s dramatic staining reaction. Although this species is commonly called the Staining Blue Bolete, a variety with flesh that does not stain blue when exposed has been repeatedly collected in North Carolina. Cap: 4–12 cm wide, convex to broadly convex, sometimes nearly plane, margin even, incurved when young; surface dry, covered with matted hairs or floccose scales, buff or straw yellow to pale olive, tan or yellowish, sometimes with darker streaks or an olive tinge, staining greenish blue to dark blue when bruised; flesh brittle, whitish to pale yellow, staining greenish blue to dark blue when exposed; odor and taste not distinctive. Pores and tubes: white to yellowish, yellowish green or tan, staining greenish blue to dark blue when bruised; tubes 5–10 mm deep. Stalk: nearly equal or swollen in the middle or below, brittle, stuffed with a soft pith, becoming hollow or chambered at maturity; surface dry, covered with matted hairs or scales when young, becoming smoother in age, concolorous with the cap or paler, staining greenish blue to dark blue when bruised or handled; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in broadleaf forests and mixed woods, especially under birch and poplar, also along roadcuts; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: pale yellow. Microscopic features: spores 8–10 × 5–6 µm, ellipsoidal, smooth, hyaline. Edibility: edible. Lookalikes: Gyroporus umbrinisquamosus Murrill (not illustrated), known only from Georgia and Florida west to Texas, is similar but has white flesh that does not change color when exposed. Gyroporus phaeocyanescens (p. 212) has a fulvous to yellow-brown cap, flesh that stains indigo blue and eventually fades to creamy buff when exposed, yellow pores that do not stain blue, and larger spores, 9–15 × 5–7 µm.

212  ·  Boletes of Eastern North America

Gyroporus phaeocyanescens

Gyroporus phaeocyanescens Singer & M. H. Ivory Common name(s): none. Overview: The term phaeocyanescens means “dark blue,” a reference to the indigo-blue staining reaction of this bolete’s exposed flesh. This species is typically coated with sand. The exposed flesh of water-logged specimens may not exhibit the blue staining reaction. The cap surface stains amber to orange brown with KOH and reddish brown with NH4OH and is negative with FeSO4. The flesh stains bright yellow with KOH, yellow with NH4OH, and pale yellow with FeSO4. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin even, incurved when young; surface dry, coated with a thick, matted covering of hairs at first, becoming appressed- fibrillose to fibrillose-scaly as the bolete matures, fulvous to yellow brown or grayish brown; flesh whitish, with a dark-brown zone beneath the cuticle, staining indigo blue when exposed, sometimes erratically, eventually fading to creamy buff; odor and taste not distinctive. Pores and tubes: whitish when young, becoming pale yellow and darkening somewhat at maturity, depressed or free near the stalk in age, not staining when bruised; tubes 5–15 mm deep. Stalk: enlarged downward or sometimes nearly equal, solid at first, becoming hollow and brittle at maturity; surface dry, coated with tiny fibrils, buff to

Gyroporus   ·  213

pale straw yellow, darkening with age or handling, especially near the base; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under oaks; summer and fall. Distribution and frequency: Georgia south to Florida, west to Texas, distribution limits yet to be established; uncommon. Spore print: pale yellow. Microscopic features: spores 9–15 × 5–7 µm, ellipsoid, smooth, pale yellow. Edibility: unknown. Lookalikes: Gyroporus cyanescens (p. 211) has a paler-colored cap and stains greenish blue to dark blue on all parts. Gyroporus umbrinisquamosus Murrill (not illustrated), known only from Georgia and Florida west to Texas, has a yellow-ocher cap with conspicuous yellow-brown fibrillose scales; white, unchanging, mild-tasting flesh; white to pale yellowish pores; and a stalk that is strongly enlarged downward with pinkish tints on the upper part and pale yellow brown below. It is associated with oaks.

Gyroporus purpurinus

Gyroporus purpurinus Singer ex Davoodian & Halling = Boletus castaneus f. purpurinus Snell Common name(s): Red Gyroporus Overview: The term purpurinus means “purplish red,” a reference to this beautiful bolete’s cap and stalk color.

214  ·  Boletes of Eastern North America

Cap: 2–8 cm wide, convex, becoming broadly convex to plane, margin even, often split in age; surface dry, somewhat velvety, sometimes finely cracked and slightly scaly in age, purplish red to burgundy, paler and sometimes yellowish at the margin, fading from the margin toward the disc in age; flesh white, unchanging when cut or bruised; odor and taste not distinctive. Pores and tubes: whitish to buff at first, becoming yellowish in age, typically depressed at the stalk at maturity; tubes 5–8 mm deep. Stalk: equal or enlarged downward, brittle, hollow in age; surface dry, scurfy to somewhat velvety, purplish red to burgundy overall or sometimes whitish on the upper portion; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups in mixed woods including hickory, oak, magnolia, and pine; summer and fall. Distribution and frequency: New England south to Florida, west to Minnesota; infrequent to rare. Spore print: yellow. Microscopic features: spores 8–11 × 5–7 µm, ovoid to ellipsoid, smooth, yellowish. Edibility: edible but too rare to be gathered as food. Lookalikes: Gyroporus castaneus (p. 209) is nearly identical except for the yellow-brown to orange-brown or reddish-brown colors of the cap and stalk. Austroboletus gracilis var. gracilis (p. 70) is similar, but its pores become pinkish brown at maturity; it has a pinkish-brown spore print, a solid stalk, and pitted spores. Some species in the genus Tylopilus are also similar, but they have solid stalks and pinkish-brown to reddish-brown spore prints.

Gyroporus subalbellus

Harrya   ·  215

Gyroporus subalbellus Murrill = Suillus subalbellus (Murrill) Sacc. & Trotter Common name(s): none. Overview: The term subalbellus means “somewhat whitish,” referring to this rather pale bolete’s overall colors. The variably colored cap, chambered to hollow stalk, whitish to pale-yellow pores, lack of blue staining reaction, and distribution along the Coastal Plain are some of its distinctive identification features. Cap: 2.5–12 cm wide, convex, becoming nearly plane in age, often shallowly depressed at maturity, margin even; surface dry, nearly glabrous, color varying from apricot buff to pinkish buff or pinkish cinnamon to orange cinnamon, sometimes pale yellow to whitish, darkening to brownish in age or when handled; flesh white, unchanging when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, becoming pale yellow and then dull yellow in age, unchanging or slowly staining pinkish cinnamon when bruised or in age, sometimes deeply depressed at the stalk; tubes 3–8 mm deep. Stalk: typically enlarged downward to a swollen or sometimes tapered base, hollow or chambered and brittle at maturity; surface dry, smooth, whitish at first, soon flushed pinkish to salmon orange, especially toward the base, frequently stained cinnamon to brownish or olivaceous; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups in oak and pine woods; summer and fall. Distribution and frequency: along the Atlantic Seaboard from Cape Cod, Massachusetts, south to Florida, west to Texas; occasional to fairly common. Spore print: yellowish buff. Microscopic features: spores 8–14 × 4–6 µm, ellipsoid to ovoid, smooth, hyaline. Edibility: edible, mild tasting and with a firm texture but often covered with sand. Lookalikes: Tylopilus peralbidus (p. 386) is similarly colored but has bitter-tasting flesh and a pinkish-brown spore print. Gen us H a r rya H a l l i ng, Nu h n & Osm u n dson The genus Harrya was erected in 2012 to accommodate species formerly classified in several genera, including Boletus, Leccinum, Suillus, and Tylopilus. The name Harrya honors American mycologist Harry D. Thiers (1919–2000). It is a small genus with only two species known worldwide to date. Harrya chromapes is the only species reported to occur in eastern North America.

Harrya chromapes (A)

Harrya chromapes (B)

Heimioporus   ·  217

Harrya chromapes (Frost) Halling, Nuhn, Osmundson & Manfr. Binder = Boletus chromapes Frost = Leccinum chromapes (Frost) Singer = Tylopilus chromapes (Frost) A. H. Sm. & Thiers Common name(s): Chrome-footed Bolete, Yellowfoot Bolete Overview: The term chromapes means “chrome-yellow foot.” This pretty bolete is easily recognized by its pinkish cap, pink to reddish dotted stalk, and the bright-yellow stalk base. Cap: 3–15 cm wide, convex, becoming broadly convex to nearly plane in age, sometimes slightly depressed at maturity; surface dry or slightly viscid when moist, pink to rose colored or pale grayish when young, fading to pinkish tan to dingy brown as it ages; flesh white, not staining when cut or bruised; odor not distinctive; taste not distinctive or slightly lemony. Pores and tubes: white when young, becoming pinkish to dingy pinkish tan in age, unchanging when bruised; tubes 8–14 mm deep, concolorous with the pores or paler. Stalk: nearly equal or tapered in either direction, often crooked at the base, solid; surface dry, white to pinkish, covered with pink to reddish or sometimes whitish scabers when young, with a bright chrome-yellow base; flesh white, becoming chrome yellow at the base. Habitat and season: solitary or in small groups in broadleaf and conifer woods; summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Minnesota and Texas; fairly common. Spore print: pinkish brown to reddish brown. Microscopic features: spores 11–17 × 4–5.5 µm, nearly oblong to narrowly oval, smooth, hyaline to pale brown. Edibility: edible. Lookalikes: none. Gen us H e i m iop oru s E. Hor a k The genus Heimioporus was erected in 2004 to accommodate species formerly classified in several genera, including Boletellus, Frostiella, and Heimiella. It was named in honor of the late French botanist and mycologist Roger Heim (1900– 1979). It is a small genus of mostly tropical and subtropical species reported from Asia, Australia, Belize, Costa Rica, Mexico, and the United States (one species). Their caps may be viscid or dry. They have dry stalks with white basal mycelium. Their spore-print color is olive brown.

Heimioporus betula (A)

Heimioporus betula (B)

Heimioporus   ·  219

Heimioporus betula (Schwein.) E. Horak = Austroboletus betula (Schweinitz) E. Horak = Frostiella betula (Schwein.) Murrill = Heimiella betula (Schwein.) Watling Common name(s): Shaggy-stalked Bolete Overview: This long-legged bolete is one of the most distinctive and beautiful of our eastern boletes. With its disproportionately small, brightly colored, smooth cap atop a long, slender, shaggy stalk, it is unlikely to be confused with any other bolete. The name betula refers to the genus name for birch trees, Betula, many of which have shaggy bark. Cap: 3–9 cm wide, somewhat rounded to convex, margin narrow and sterile; surface smooth, glabrous, viscid when moist to somewhat tacky and shiny; bright to dark red, orange, or a blend of red and orange or orange and yellow, margin often yellow; flesh soft, pale yellow, at times tinged orange beneath the cap surface, unchanging when exposed; odor not distinctive; taste lemony or sour/acidic. Pores and tubes: bright to pale yellow or greenish yellow, usually not staining when bruised but sometimes bluing, depressed at the stalk; tubes 1–1.5 cm deep. Stalk: thin, slender, often curved near the base, solid and very firm; surface dry, coarsely reticulate-shaggy, yellow at first, becoming reddish to yellow, with a massive pad of cottony white mycelium at the base; reticulation yellow or sometimes reddish, raised; flesh concolorous with the cap flesh or at times tinged pinkish. Habitat and season: solitary or scattered in mixed broadleaf and pine woods, also in moist riparian habitats with hemlock, rhododendron, and birch; summer and fall. Distribution and frequency: Pennsylvania south to Georgia, west to Michigan; occasional to fairly common, especially in the central Appalachians. Spore print: olive brown. Microscopic features: spores 15–19 × 6–10 µm, narrowly ellipsoidal, ornamented with a loose reticulum and scattered minute pits, typically with a distinct apical pore, pale brown. Edibility: edible and considered by some to be very good but only mediocre according to others. Although the cap flesh is soft and acidic and not enjoyed by all, the stalks cook up crisp and tart and pair well with fish dishes and vegetables such as green beans and asparagus. Use young caps and all stalks—very lemony and very good. Lookalikes: Frostiella russellii (p. 207) is similar in stature but not nearly as colorful; plus, it has a dry, scaly cap. Austroboletus subflavidus (p. 71) also bears a very slight resemblance to the Shaggy-stalked Bolete but is whitish

220  ·  Boletes of Eastern North America

in color and is known only from the Gulf Coast, the southeast Coastal Plain, and coastal areas north into New Jersey. Gen us H e m i l ecci n u m Šu ta r a The genus Hemileccinum was erected in 2008 to accommodate two European and one North American species that molecular analysis indicate do not belong in the genera Boletus, Leccinum, and Xerocomus. The name Hemileccinum means “half Leccinum.” Species in this genus have a dry, subtomentose to nearly smooth cap, yellow or white flesh that does not stain when exposed, yellow pores that do not stain when bruised, a dry stalk with pale scabers that darken only slightly in age, and an olive-brown spore print.

Hemileccinum subglabripes

Hemileccinum subglabripes (Peck) Halling = Boletus subglabripes Peck = Leccinum subglabripes (Peck) Singer Common name(s): none. Overview: The term subglabripes means “nearly smooth foot,” a reference to this bolete’s stalk. Macroscopically, this bolete may appear to be more closely related to the genus Boletus, but molecular studies indicate that it is not.

Hortiboletus   ·  221

Cap: 4.5–10 cm wide, convex to broadly convex or nearly plane in age, sometimes broadly umbonate, margin even; surface smooth to slightly wrinkled, chestnut brown, ocher, cinnamon, or reddish brown to bay brown; flesh pale yellow to whitish, rarely staining slightly blue when cut or bruised; odor not distinctive; taste mild to slightly acidic. Pores and tubes: bright yellow to yellow when fresh, duller or slightly greenish yellow in age, not staining when bruised; tubes 5–16 mm deep. Stalk: nearly equal, solid, sometimes rather stout; surface dry, minutely scurfy with a thin coating of tiny yellow scabers, yellow with occasional reddish or reddish-brown tinges, especially on the lower portion; flesh concolorous with the cap flesh. Habitat and season: scattered under broadleaf trees, especially birch, and occasionally under conifers; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Minnesota and Texas; fairly common. Spore print: olive brown. Microscopic features: spores 11–17(21) × 3–5(7) µm, narrowly fusoid, smooth, pale brown. Edibility: edible, flavor with a hint of lemon and very good. Lookalikes: Xerocomus hortonii (p. 421) is nearly identical, but it has a conspicuously pitted cap, and its stalk is typically glabrous or sometimes pruinose. It occasionally has delicate reticulation at the apex. Gen us H ort i b ol e t u s Si mon i n i, Vi zzi n i & Gel a r di The genus Hortiboletus was erected in 2015 to accommodate a single species formerly classified in the genus Xerocomellus. Since then three other species have been added worldwide to this genus. The name Hortiboletus is derived from the Latin word hortus, “garden,” and means “bolete of the gardens.” This genus differs from Xerocomellus by having reddish flesh in the stalk base, smooth spores that are never striate or truncate, and an average spore-quotient value (spore length divided by spore width) of less than 2.5. This new genus is supported by molecular analysis.

Hortiboletus campestris

Boletus harrisonii

Hortiboletus   ·  223

Hortiboletus campestris (A. H. Sm. & Thiers) Biketova & Wasser = Boletus campestris A. H. Sm. & Thiers Common name(s): none. Overview: This is one of a confusing group of smallish, red-capped boletes. The key identification features include a red cap that becomes conspicuously cracked in age, yellow flesh that stains greenish blue when exposed, yellow pores that quickly bruise blue, and a yellow stalk that is pruinose to punctate with dull reddish-orange dots and points. The term campestris means “growing in fields,” where this bolete frequently occurs. Cap: 2–5 cm wide, convex, becoming broadly convex, margin even; surface dry, somewhat velvety, usually becoming conspicuously cracked, especially toward the margin in age, rose red when young, becoming pinkish red with a hint of yellow showing in the fissures when mature; flesh pale yellow to yellow, staining greenish blue when exposed; odor and taste not distinctive. Pores and tubes: yellow at first, becoming greenish yellow to olive yellow at maturity, staining greenish blue when bruised, typically depressed at the stalk in age; pores circular to angular, one to two per millimeter; tubes 4–8 mm deep. Stalk: nearly equal, solid; surface dry, pale yellow near the apex, deeper yellow downward, pruinose to punctate with dull reddish-orange dots and points, especially toward the base, basal mycelium yellowish; flesh bright yellow throughout, staining greenish blue. Habitat and season: scattered or in groups on lawns and grassy places in parks, open woodlands, and roadsides; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: olive brown. Microscopic features: spores 11–15 × 4.5–7 µm, ellipsoidal to subfusoid, smooth, yellow ocher. Edibility: reported to be edible. Lookalikes: Hortiboletus rubellus (p. 225) has a stalk with dull reddish-orange punctae from the base up to a yellow apex, stalk flesh that is yellow on the upper portion and reddish orange in the base, and smaller spores that measure 10–13 × 4–5 µm. Boletus harrisonii A. H. Sm. & Thiers (see photo, p. 222), reported from Illinois, Michigan, Minnesota, and North Carolina, is very similar, but its stalk lacks the reddish-orange dots and points. It grows under broadleaf trees, especially oak, but also occurs with spruce. Microscopically, it differs from all similar species by having a cap cuticle whose last three to four cells are short and inflated and whose end cell is cystidioid to subglobose. Boletus subfraternus (p. 159) lacks conspicuous cracks on the surface of mature caps and has large, irregular to angular pores that become elongated and radially arranged to somewhat gill-like near the stalk.

Hortiboletus rubellus (A)

Hortiboletus rubellus (B)

Hortiboletus   ·  225

Hortiboletus rubellus (Krombh.) Simonini, Vizzini & Gelardi = Boletus fraternus Peck = Boletus rubellus Krombholz = Xerocomellus rubellus (Krombh.) Šutara = Xerocomus rubellus (Krombholz) Quél. Common name(s): none. Overview: The term rubellus means “reddish,” a reference to the colors of this bolete’s cap and stalk. There was formerly much confusion about the identification of this beautiful bolete and in separating it from very similar species. Thanks to molecular analysis, much of the confusion has been eliminated. Cap: 2–8 cm wide, convex, becoming nearly plane in age, margin even; surface dry, somewhat velvety, often becoming finely cracked in age, dark red, fading in age to brick red or olivaceous brown; flesh pale yellow, slowly staining bluish green on exposure; odor and taste not distinctive. Pores and tubes: yellow, staining bluish green when bruised, typically depressed at the stalk when mature; tubes 6–10 mm deep. Stalk: nearly equal, enlarged or tapered downward, often with a narrowed base, solid; surface dry, yellow at the apex, pruinose to punctate with dull reddish-orange dots and points, darkening to brown where handled; flesh yellow on the upper portion, reddish orange below or at least near the base. Habitat and season: solitary, scattered, or in groups in grassy areas, parks, gardens, flowerbeds, and disturbed roadsides as well as along paths or in broadleaf and mixed woods, especially under oak and beech; summer and fall. Distribution and frequency: widespread throughout eastern North America; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 10–13 × 4–5 µm, ellipsoidal, smooth, pale brown. Edibility: unknown. Lookalikes: Hortiboletus campestris (p. 223) is very similar, but its stalk flesh is bright yellow throughout, not reddish orange in the base, and it has somewhat larger spores that measure 11–15 × 4.5–7 µm. Its cap surface stains amber with KOH and dark green to blackish green with FeSO4. Its flesh stains orange with KOH and dull orange with FeSO4. Boletus harrisonii A. H. Sm. & Thiers (see photo, p. 222), reported from Illinois, Michigan, and Minnesota, is also very similar, but its stalk lacks reddish-orange dots and points. It grows under broadleaf trees, especially oak, but also occurs with spruce. Microscopically, it differs from all similar species by having a cap cuticle whose last three to four cells are short and inflated and whose end cell is cystidioid to subglobose. Boletus subfraternus (p. 159) has large, irregular to angular pores, and its cap does not become cracked with age.

226  ·  Boletes of Eastern North America

Gen us I m l e r i a Vi zzi n i The genus Imleria was erected in 2014 to accommodate a single species, Imleria badia, which was formerly classified in several genera, including Boletus, Suillus, and Xerocomus. This genus was named in honor of Belgian mycologist Louis Imler (1900–1993). It presently consists of five species worldwide.

Imleria badia (A)

Imleria badia (B)

Imleria   ·  227

Imleria badia (Fr.) Vizzini = Boletus badius (Fr.) Fr. = Xerocomus badius (Fr.) E.-J. Gilbert Common name(s): Bay Bolete Overview: This medium-size bolete is a conifer associate and not difficult to identify. It is an excellent edible that is well worth getting to know. The term badius means “bay brown,” which is chestnut brown or reddish brown. The cap surface stains green to blue with NH4OH, blackish with KOH, and bluish green with FeSO4. Cap: 4–9 cm wide, convex to broadly convex, becoming nearly plane at maturity, sometimes depressed in the center, margin upturned in age, even; surface dry to slightly viscid, smooth or somewhat velvety, chestnut brown, reddish brown, or yellow brown, sometimes with olive tones; flesh soft, white, often staining bluish, at least near the tubes, when cut; odor and taste not distinctive. Pores and tubes: pale yellow to greenish yellow, quickly staining greenish blue to grayish blue when bruised; tubes 8–15 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, smooth to somewhat pruinose, concolorous with the cap, with a white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or scattered under pines or other conifers, sometimes on decaying conifer stumps or in hardwoods; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; fairly common. Spore print: olive brown. Microscopic features: spores 10–14 × 4–5 µm, ellipsoidal to fusiform, smooth, yellow. Edibility: Edible and very good. Tubes from older specimens can be removed, dried, and powdered for seasoning. The flesh is not often damaged by insects. Lookalikes: Xanthoconium affine (p. 409) has a dark-brown to chestnut-brown or ocher-brown cap and white pores that become yellowish to dingy yellow brown at maturity and stain dull yellow to brownish, not blue, when bruised. Aureoboletus projectellus (p. 64) has similar colors but is typically more robust and has a coarsely reticulate stalk. Boletus glabellus Peck (not illustrated), reported from New England west to Ohio and Michigan, has a pale-brown to yellow-brown dry cap, sometimes with reddish tinges, that often becomes finely cracked in age. It has whitish flesh that stains blue when exposed, pale greenish-yellow pores that become ochraceous to brownish at maturity and stain blue when bruised, and a dullwhite to yellowish stalk with reddish to purple-red tinges, especially toward the base. Its stalk flesh is deep yellow toward the base and pale yellow

228  ·  Boletes of Eastern North America

above the base and stains blue when exposed. Its spores measure 9–13 × 3–5 µm, and it grows in grassy areas or in woods with oaks. Similarly colored species in the genus Tylopilus have white or pinkish-brown spores that do not stain blue when bruised and produce a pinkish-brown spore print. Gen us L a n m aoa G. Wu & Zh u L . Ya ng The genus Lanmaoa was erected in 2015 to accommodate species formerly classified in several genera, including Boletus, Leccinum, and Suillus. Lanmaoa was named in honor of Mr. Lan Mao (1397–1476), a Chinese botanist from the Ming dynasty who first used the Chinese term niuganjun for boletes in the Chinese literature. The species included in this genus do not share very many macroscopic characteristics. However, DNA analysis clearly demonstrates that they are closely related. The genus Lanmaoa includes two species from China, one from Costa Rica, and four from eastern North America, including Lanmaoa roseocrispans, a species only recently described from Florida. They are mediumto large-size terrestrial boletes with dry caps and solid stalks. Their flesh is pale yellow to bright yellow and stains blue in most species when exposed. Pores are typically yellow, bruising blue and sometimes brown. One non–North American species has dull-orange to orange-red pores.

Lanmaoa borealis

Lanmaoa   ·  229

Lanmaoa borealis (A. H. Sm. & Thiers) A. E. Bessette, M. E. Nuhn & R. E. Halling, comb. et stat. nov. MycoBank: MB811208 = Boletus bicolor var. borealis A. H. Sm. & Thiers, Boletes of Michigan (Ann Arbor): 277. 1971. Common name (s): none. Overview: We have elevated this variety to species rank based on the results of molecular analysis obtained by Gang Wu and his colleagues (2014). The key identification features include a red cap, yellow flesh that slowly stains blue at the stalk apex, orange-red to red pores that stain greenish blue, and a red stalk that may be reticulate at the apex. Cap: 2–12 cm wide, obtuse to convex at first; surface dry, glabrous or nearly so, dark red to bright apple red, becoming brick red to dull rusty rose in age; flesh yellow to pale yellow, slowly bluing at the stalk apex when exposed; odor and taste not distinctive. Pores and tubes: orange red to red, staining greenish blue when bruised, becoming dull brownish red and sometimes depressed near the stalk in age; tubes 6–10 mm deep. Stalk: tapered downward or nearly equal, solid; surface dry, nearly glabrous, colored like the cap, lacking reticulation or reticulate only at the apex; flesh yellow to pale yellow, slowly bluing at the apex, vinaceous brown at the base. Habitat and season: solitary, scattered, or in groups under broadleaf trees or in mixed woods; summer and fall. Distribution and frequency: recorded from Michigan east to New England, distribution limits yet to be determined; uncommon. Spore print: olive brown. Microscopic features: spores 11–15 × 4–5 µm, subfusoid, smooth, pale yellow. Edibility: poisonous, causing gastrointestinal distress. Lookalikes: Species in the Baorangia bicolor complex (p. 74) are similar, but they have yellow pores.

Lanmaoa carminipes (A)

Lanmaoa carminipes (B)

Lanmaoa   ·  231

Lanmaoa carminipes (A. H. Sm. & Thiers) G. Wu, Halling & Zhu L. Yang = Boletus carminipes A. H. Sm. & Thiers Common name(s): none Overview: The term carminipes means “carmine-red foot,” a reference to this beautiful bolete’s red stalk. The cap surface stains dark yellow brown with KOH. The flesh slowly stains bluish with FeSO4 and pale ochraceous tan when KOH is applied to blue areas. Cap: 5–12 cm wide, convex, becoming broadly convex, margin incurved at first, even; surface dry, somewhat velvety to nearly smooth, pinkish red to rose red initially, becoming dingy brownish and sometimes finely cracked in age, often yellowish or paler pinkish red along the margin, quickly staining dark blue when bruised or handled; flesh pale yellow, quickly staining dark blue when exposed; odor and taste not distinctive. Pores and tubes: bright yellow, quickly staining dark blue when bruised and then slowly reddish brown to dull brown, depressed at the stalk at maturity; tubes 4–6 mm deep. Stalk: enlarged downward, often with a pinched base, solid; surface dry, finely pruinose, yellow on the upper portion, carmine red to pinkish red over a yellow ground color on the lower portion and retaining this color even when dry, sometimes yellow overall, staining dark blue and then slowly brownish when handled or bruised, sometimes with white basal mycelium; flesh yellow, often reddish at the base, quickly staining dark blue. Habitat and season: solitary, scattered, or in groups under broadleaf trees, usually with oak; summer and fall. Distribution and frequency: eastern Canada west to Minnesota, south to North Carolina; infrequent. Spore print: olive brown. Microscopic features: spores 9–12 × 3–3.5 µm, narrowly oblong, smooth, ochraceous. Edibility: unknown. Lookalikes: The Baorangia bicolor complex (p. 74) is similar, but it has a dark-red to rose-red cap and stalk and pale-yellow cap flesh that is unchanging or slowly and weakly staining blue when exposed.

Lanmaoa pseudosensibilis (A)

Lanmaoa pseudosensibilis (B)

Lanmaoa   ·  233

Lanmaoa pseudosensibilis (C)

Lanmaoa pseudosensibilis (A. H. Sm. & Thiers) G. Wu, Halling & Zhu L. Yang = Boletus pseudosensibilis A. H. Sm. & Thiers Common name(s): none. Overview: The term pseudosensibilis means “resembling Boletus sensibilis.” Adding a drop of NH4OH to the cap surface produces a greenish-blue, then dull purplish, stain, differentiating this bolete from Boletus sensibilis (p. 150). The key identification features include a dull-brown cap, bright-yellow flesh that stains blue when exposed, yellow pores that bruise blue, and a yellow stalk that is sometimes tinged reddish to purplish. Cap: 6–14 cm wide, convex, becoming broadly convex to nearly plane in age, margin even, often bright yellow; surface dry, smooth, often becoming cracked in age, with yellow flesh showing in the fissures, dull rusty brown to dull reddish brown, fading to dull cinnamon or dingy yellow brown; flesh bright yellow, quickly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow to ocher yellow when young, becoming olive ocher in age, rapidly staining blue and then brown when bruised; tubes 5–8 mm deep.

234  ·  Boletes of Eastern North America

Stalk: nearly equal or sometimes slightly flared at the apex, solid; surface dry, typically yellow overall, sometimes with tinges of rusty red, pink, or purple, especially near the base; flesh bright yellow on the upper portion, more intensely yellow on the lower portion, staining blue, sometimes slowly, when exposed. Habitat and season: gregarious in broadleaf woods, especially under oaks and hickory, or sometimes under pines; summer and fall. Distribution and frequency: New England south to North Carolina, west to Minnesota, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 9–12 × 3–4 µm, subfusiform to oblong, smooth, pale brown. Edibility: Although this bolete is edible and has a good flavor, it may be confused with similar species, some of which are suspected to be poisonous; therefore, it is not recommended as an edible bolete. Lookalikes: The Baorangia bicolor complex (p. 74) has a redder cap, and its flesh slowly and weakly stains blue, if at all.

Lanmaoa roseocrispans

Lanmaoa   ·  235

Lanmaoa roseocrispans Bessette, A. R. Bessette, Nuhn & Halling Common name(s): none. Overview: The term roseocrispans means “rosy-pink folds,” a reference to the coloration between the folds of this bolete’s cap. This mushroom is extremely robust, hefty, and dense. The strongly tapered stalk with a ventricose or pinched base and the remarkably folded and lobed cap margin with dull pinkish or rosy tones within the folds are the most distinguishing features. The cap surface quickly stains pale gray with NH4OH. Cap: 6–16 cm wide, pulvinate and distinctly folded and lobed at first, becoming broadly convex in age, margin with a narrow band of sterile tissue, strongly incurved, undulating, pinched and lobed when young and remaining so well into maturity; surface dry, somewhat velvety, with irregular pits and depressions, sometimes cracked in age, color variable, brick red or rosy red over a yellow ground color when young, becoming yellow orange or mustard yellow with dull pinkish or rosy tones within the folds, with burgundy-red or brownish-red splashes on portions of mature specimens, slowly bruising greenish blue to grayish blue, especially along the margin; flesh thick, firm, pale yellow to yellow, becoming darker yellow when exposed, sometimes slowly staining grayish blue; odor and taste not distinctive. Pores and tubes: yellow when young, becoming olive yellow to olive brown at maturity, rapidly staining greenish blue to grayish blue when bruised, dull brownish red around marginal folds, conspicuously depressed at the stalk in age; tubes 4–13 mm deep. Stalk: strongly tapered downward, ventricose or with a pinched base, solid; surface scurfy-punctate nearly overall, sometimes smooth at the apex, often longitudinally striate, dull burgundy red to brownish red over a yellow ground color, darker at the base, typically yellow at the apex, staining greenish blue to grayish blue when bruised or handled, typically with pale-yellow basal mycelium; flesh concolorous with and staining like the cap flesh, usually orange yellow to orange near the base. Habitat and season: solitary, scattered, or in groups or clusters in grassy areas under oaks; late summer and fall. Distribution and frequency: currently known only from Dunnellon, Florida, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 11–17 × 3–5 µm, subellipsoid to subfusoid, smooth, hyaline to pale brownish yellow. Edibility: unknown. Lookalikes: The Baorangia bicolor complex (p. 74) is similar but is much less robust, hefty, and dense. It has a darker-red cap with an even margin that

236  ·  Boletes of Eastern North America

is not distinctly folded and lobed, a red stalk that is not tapered downward, and smaller spores, 8–12 × 3.5–5 µm. Gen us L ecci n e l lu m Br e si nsk y & M a n f r . Bi n der The genus Leccinellum was erected in 2003 to accommodate those species formerly classified in the genus Leccinum that have a cap cuticle consisting of a trichodermium of septate hyphae with short, ellipsoid to clavate or subglobose terminal elements or markedly inflated hyphae. Andreas Bresinsky and Manfred Binder recognized the cap cuticle of Leccinum species as having long and narrow hyphae or rarely short hyphae, with terminal elements that are not inflated. Leccinellum is a small genus of medium to large terrestrial boletes that have solid stalks that are ornamented with scabers, which may or may not darken as specimens mature. Four species of Leccinellum are currently known from eastern North America.

Leccinellum albellum (A)

Leccinellum   ·  237

Leccinellum albellum (B)

Leccinellum albellum (C)

238  ·  Boletes of Eastern North America

Leccinellum albellum (Peck) Bresinsky & Manfr. Binder = Leccinum albellum (Peck) Singer Common name(s): none. Overview: Although the term albellum means “whitish,” specimens with various colors, especially white, brown, and gray, commonly grow beside each other in the same group. Collections of white specimens with white scabers that remain white to maturity were made from several states and originally thought to be a new species. These specimens, along with collections of typical brown and gray specimens with dark scabers, were sent to Dr. Matthew Smith at the University of Florida for molecular analysis. He sequenced ITS and 28S rDNA from all of these specimens and verified their identity as Leccinellum albellum. Cap: 2–8 cm wide, obtuse to convex, becoming broadly convex in age, margin even or with a narrow band of sterile tissue; surface dry or moist, glabrous or somewhat velvety, often developing large, shallow depressions at maturity, often with a conspicuous gelatinous layer beneath the cuticle, sometimes cracked in age, color highly variable, white, whitish, buff to pinkish buff or pale tan, pale gray to pinkish gray, or pinkish brown to medium brown, sometimes tinged yellow or with a bluish cast; flesh white, not staining when cut; odor and taste not distinctive. Pores and tubes: initially white, becoming buff to pale brown or pale gray in age, not staining when bruised, depressed near the stalk at maturity; tubes 2–10 mm deep. Stalk: nearly equal to enlarged downward, sometimes with a bent, slightly bulbous base, solid overall or stuffed in the lower portion; surface dry, white to whitish or pale olive buff, with tiny white scabers that remain white or darken to brownish or grayish in age, sometimes staining orange yellow when handled; flesh white, unchanging when exposed. Habitat and season: scattered or in groups on the ground or among mosses under broadleaf trees, especially oak; summer and fall. Distribution and frequency: eastern Canada south to Florida, west to Missouri; fairly common. Spore print: brown to dark olive brown. Microscopic features: spores 14–22 × 4–6 µm, ellipsoidal to subfusiform, smooth, pale brownish yellow. Edibility: edible. Lookalikes: Leccinum holopus (p. 248) is a somewhat similar whitish bolete, at least when young, but it is normally larger, is associated with birch, and has a more northern distribution.

Leccinellum crocipodium (A)

Leccinellum crocipodium (B)

240  ·  Boletes of Eastern North America

Leccinellum crocipodium (Letell.) Watling = Leccinum nigrescens Singer Common name(s): none. Overview: The term crocipodium means “saffron foot,” a reference to the stalk’s ground color. Reliable field features for identifying this bolete include a wrinkled, pitted, dark-brown to blackish-brown cap, pale-yellow to dingy-yellow pores, and a stalk that is typically swollen near the middle or toward the base. The cap surface stains reddish brown with NH4OH and is negative with FeSO4. Cap: 4–15 cm wide, convex, becoming broadly convex, margin even, incurved at first and remaining so well into maturity; surface dry to moist but not viscid, wrinkled, uneven, pitted, often coarsely cracked in age, dark brown to blackish brown when young, fading to pale yellow brown in age; flesh pale yellow, staining pinkish gray to dull reddish or pale fuscous when exposed, sometimes slowly; odor and taste not distinctive. Pores and tubes: pale yellow to dingy yellow, often staining brownish when bruised, depressed at the stalk in age; tubes 8–15 mm deep. Stalk: swollen near the middle or toward the base and tapered in either direction, solid; surface dry, pale yellow and often dull reddish at the base, with brown scabers that darken to blackish brown in age. Habitat and season: solitary, scattered, or in groups in sandy soil or sometimes in grassy areas under broadleaf trees, especially oak; summer, fall, and early winter. Distribution and frequency: New England south to Florida, west to the Great Lakes region and Texas; occasional to fairly common in the southern part of its range. Spore print: honey yellow. Microscopic features: spores 14–20 × 6–9 µm, fusiform, smooth, yellowish; caulocystidia narrowly fusoid-ventricose with flexuous necks or broadly fusoid-ventricose to clavate-mucronate. Edibility: edible. Lookalikes: Leccinum rugosiceps (p. 262) has a paler orange-yellow cap when young that becomes yellow brown in age, a nearly equal stalk with paler scabers, and flesh that slowly stains reddish or burgundy when exposed and orange-red with KOH. Also compare with Leccinellum quercophilum (p. 244) and Leccinellum griseum (p. 242), both of which have whitish to grayish or grayish-brown pores.

Leccinellum griseum (A)

Leccinellum griseum (B)

242  ·  Boletes of Eastern North America

Leccinellum griseum (Quél.) Bresinsky & Manfr. Binder = Leccinum griseum (Quél.) Singer Common name(s): none. Overview: The term griseum means “grayish,” a reference to this bolete’s somber colors. The cap surface stains dark red brown with KOH and brown with NH4OH and is negative with FeSO4. The flesh stains light brown with KOH, very pale yellow and soon fuscous with NH4OH, and greenish blue with FeSO4. Cap: 3.5–9 cm wide, pulvinate when young, becoming convex to broadly convex or nearly plane in age, margin even, incurved at first; surface dry, somewhat velvety, distinctly wrinkled and pitted or sometimes nearly smooth, becoming conspicuously cracked at maturity, with whitish to buff flesh showing in the cracks, dull brown to blackish brown when young, typically developing olive-green tones and becoming a mixture of these colors in age; flesh whitish to buff, sometimes staining pinkish brown and then darkening to reddish cinnamon, developing bluish-gray tints and slowly darkening; odor and taste not distinctive. Pores and tubes: whitish to grayish buff at first, darkening at maturity, staining brown when bruised; tubes 3–15 mm deep. Stalk: enlarged downward or nearly equal, often with a pinched base, typically curved especially on the lower portion, solid; surface dry, with conspicuous dark-brown to blackish scabers over a whitish ground color, darkening when handled or bruised, sometimes reddish cinnamon near the base, with whitish basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under oaks; summer and fall. Distribution and frequency: Georgia south to Florida, west to Texas; occasional. Spore print: dark brown. Microscopic features: spores 16–19(22) × 5.5–7(8) µm, fusiform to subfusiform, smooth, yellowish to pale brown. Edibility: unknown. Lookalikes: Leccinellum quercophilum (p. 244) is similar, but its cap is dull orangish brown to tawny brown when young; its stalk has whitish to palebrown scabers that become darker brown in age; and all parts stain green when bruised or in age. Leccinellum crocipodium (p. 240) has pale-yellow to dingy-yellow pores and a stalk that is swollen near the middle or toward the base. Leccinum rugosiceps (p. 262) has a paler orange-yellow cap when young that becomes yellow brown in age, a nearly equal stalk with paler scabers, and flesh that slowly stains reddish or burgundy when exposed and stains orange red with KOH.

Leccinellum quercophilum (A)

Leccinellum quercophilum (B)

244  ·  Boletes of Eastern North America

Leccinellum quercophilum Kuo Common name(s): none. Overview: The term quercophilum means “oak loving.” The cap surface is negative with NH4OH or FeSO4 and stains yellowish or is negative with KOH; the flesh stains gray to greenish gray with KOH, greenish or negative with FeSO4, and pinkish or negative with NH4OH. Cap: 3–9 cm wide, obtuse to convex, becoming broadly convex, sometimes with a low broad umbo, margin even; surface dry, glabrous, wrinkled and pitted when young, becoming conspicuously cracked, with whitish to grayish flesh showing in the spaces of mature specimens, dull orange brown to tawny brown when young, becoming dark brown to grayish brown in age, discoloring bluish green near the margin with age; flesh very thick and soft, whitish but gradually becoming grayish when cut, with or without an initial faintly pinkish phase; odor and taste not distinctive. Pores and tubes: whitish to grayish or pale grayish brown when young, darkening in age, staining greenish when bruised or in age and then often slowly becoming dingy yellowish brown to dark brown, depressed at the stalk at maturity; tubes 1–2 cm deep, creamy whitish, slowly staining dark brown to blackish when bruised. Stalk: somewhat enlarged downward to slightly ventricose or nearly equal, solid; surface dry, whitish to grayish or pale brown, with whitish to pale-brown scabers that become darker brown in age, frequently stained greenish on the lower portion, base with white appressed mycelium; flesh whitish, slowly staining grayish when exposed, sometimes staining greenish near the base. Habitat and season: scattered or in groups in lawns or grassy areas under broadleaf trees, especially oak, or in woodlands where oak is present; summer and fall. Distribution and frequency: reported from Illinois and Michigan; occasional. Spore print: cinnamon brown. Microscopic features: spores 15–18(28) × 5–8 µm, fusiform to subfusiform, smooth, yellowish to pale brown. Edibility: edible but too uncommon to gather for the table. Lookalikes: Leccinellum griseum (p. 242) is similar, but it has a dull-brown to blackish-brown cap when young, and its pores do not stain green when bruised. Leccinellum crocipodium (p. 240) has pale-yellow to dingy-yellow pores and a stalk that is swollen near the middle or toward the base. Leccinum rugosiceps (p. 262) has a paler, orange-yellow cap, white flesh that slowly stains reddish or burgundy when cut, dull-yellow to dingy-yellowish pores, and a pale-yellow to brownish stalk with brown scabers.

Leccinum   ·  245

Gen us L ecci n u m S.F. Gr ay The genus Leccinum was erected in 1821 to accommodate species that have scabers on their stalks and were formerly classified in the genus Boletus. Leccinum species have a cap cuticle that is typically composed of long and narrow or rarely short hyphae that are not inflated. The cuticle of at least one species has terminal elements that are markedly inflated, a feature found in species of the similar genus Leccinellum. Leccinum is a large genus of medium to large terrestrial boletes with solid stalks that are ornamented with scabers that usually darken as specimens mature. A few species in other genera have stalks ornamented with raised dots and points called punctae, but the punctae do not darken at maturity. Leccinum species have a sterile cap margin that often extends beyond the tube layer on young specimens. Spore-print colors include dark olive buff to olive brown, cinnamon brown, yellow brown, and brown.

Leccinum chalybaeum

Leccinum chalybaeum Singer Common name(s): none. Overview: This southern bolete was first described by Rolf Singer in 1945. William Murrill thought that it was a variety of Leccinum scabrum (p. 265) and in 1948 published it as Leccinum scabrum var. chalybaeum,

246  ·  Boletes of Eastern North America

which it clearly is not. The flesh of Leccinum chalybaeum stains yellow with KOH, slowly pale greenish blue with NH 4OH, and greenish blue with FeSO4. The term chalybaeum means “steel gray,” a color often present on the cap. Cap: 4–13 cm wide, obtuse at first, becoming convex to broadly convex, margin even; surface viscid when fresh, somewhat shiny when dry, smooth to slightly subtomentose, sometimes finely cracked in age, buff to pinkish buff when young, becoming dingy yellow brown, sometimes with grayish-green to bluish-gray tints, especially near the margin; flesh stains pinkish and then slowly pinkish brown to purplish gray to blackish when exposed. Pores and tubes: whitish to beige, usually staining dingy olive to brown when bruised, deeply depressed at the stalk in age; tubes up to 1.6 cm deep. Stalk: fairly short and thick, up to 8 cm long and 2 cm thick, enlarged downward or nearly equal, solid, extremely firm, almost woody; surface dry whitish beneath dingy-white scabers that darken to brown at maturity; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups in sandy soil under oak or in mixed woods with oak and pine; summer through winter. Distribution and frequency: Georgia south to Florida, west to Texas; occasional. Spore print: olive brown. Microscopic features: spores 16–18 × 5–6 µm, fusoid, smooth, brownish; caulocystidia fusoid to fusoid-ampullaceous. Edibility: edible. Lookalikes: Leccinum scabrum (p. 265) has a more northern range and is associated with birch. It has a grayish-brown to yellow-brown or darkbrown cap, white flesh that usually does not stain when exposed, a longer and thinner stalk with blackish scabers, and sometimes green stains near the base.

Leccinum   ·  247

Leccinum glutinopallens

Leccinum glutinopallens A. H. Sm., Thiers & Watling Common name(s): none. Overview: The term glutinopallens means “slimy and pale,” a reference to the cap of this rather nondescript bolete. The key identification features include a slimy-viscid, vinaceous-buff to pale grayish-brown cap, white flesh that stains pinkish tan when exposed, whitish to grayish flesh that stains brown, and a whitish stalk with whitish scabers that become tan to brown in age. Cap: 5–9 cm wide, convex, becoming broadly convex at maturity, margin incurved at first, even; surface slimy-viscid when fresh, glabrous, dingy vinaceous buff to pale pinkish gray or pale grayish brown, sometimes paler along the margin; flesh very soft, white, slowly staining pinkish tan when exposed, and with a bluish-green tinge developing above the tubes of mature specimens; odor and taste not distinctive. Pores and tubes: whitish to grayish, staining pinkish brown and then slowly dark brown when bruised, depressed at the stalk in age; tubes pale brown, 8–15 mm deep. Stalk: nearly equal or enlarged downward, often club-shaped at the base, solid; surface dry, whitish, stained pinkish tan to brown when handled or bruised, ornamented at first with very fine whitish scabers that become pinkish tan or darker brown in age; flesh concolorous with and staining like the cap flesh, often developing greenish-blue areas.

248  ·  Boletes of Eastern North America

Habitat and season: scattered or in groups on the ground or among mosses in mixed woods of aspen, birch, balsam fir, and spruce; summer and fall. Distribution and frequency: eastern Canada west to Michigan, distribution limits yet to be determined; occasional. Spore print: brown. Microscopic features: spores 15–19 × 5–6.5 µm, broadly fusiform, smooth, pale brown. Edibility: unknown. Lookalikes: Leccinum holopus (p. 248) is similar, but its cap is paler and not slimy, and its pores remain unchanged or stain yellowish when cut or bruised.

Leccinum holopus

Leccinum holopus (Rostk.) Watling = Boletus holopus Rostkovius = Leccinum holopus var. americanum A. H. Sm. & Thiers = Leccinum holopus var. holopus (Rostk.) Watling Common name(s): Ghost Bolete, White Birch Bolete, White Bog Bolete Overview: When young, this ghostly white bolete that flourishes in wet woods and boggy places is not difficult to recognize. However, with

Leccinum   ·  249

age the cap becomes darker, having brown to greenish or olivaceous tones. It is essentially a northern species but extends southward into the central Appalachians, especially at higher elevations. The term holopus means “whole or entire foot,” likely a reference to this bolete’s solid stalk. The cap surface stains pinkish with NH 4OH and slowly grayish olive with FeSO4. Cap: 3–10 cm wide, acutely convex, becoming broadly convex to nearly plane in age, margin with a narrow band of sterile tissue; surface glabrous or nearly so, somewhat viscid when moist or in age, predominantly white or whitish when young, occasionally with gray, buff, tan, or pinkish tints, often darkening with age and developing a greenish tinge; flesh thick, white, unchanging when cut or rubbed or slowly and faintly staining pinkish or red; odor and taste not distinctive. Pores and tubes: whitish to slightly grayish or pale dingy brown, unchanging or staining yellowish or slowly brown when cut or bruised, depressed at the stalk when mature; tubes 1–2.5 cm deep. Stalk: equal or enlarging slightly downward, solid; surface dry or moist, whitish beneath the scabers that are whitish when young and darken to tan or darker in age, occasionally with greenish stains on the lower portion; flesh concolorous with the cap flesh. Habitat and season: solitary to scattered on the ground in and around bogs, cedar swamps, or woodlands where birch and hemlock are present; summer and fall. Distribution and frequency: eastern Canada south to New England and New York and central Appalachians, west to Minnesota; fairly common in the north woods, less frequent southward. Spore print: brown. Microscopic features: spores 14–20 × 5–6.5 µm, subfusoid, smooth, pale brown. Edibility: edible. Lookalikes: Leccinum glutinopallens (p. 247) has a slimy-viscid, dingy vinaceous-buff to pale pinkish-gray or pale grayish-brown cap, and its pores stain pinkish brown and then slowly dark brown when bruised. Also compare with Leccinellum albellum (p. 238), which is smaller and grows under oaks.

Leccinum insigne (A)

Leccinum insigne (B)

Leccinum   ·  251

Leccinum insigne (C)

Leccinum insigne A. H. Sm., Thiers, & Watling Common name(s): Aspen Bolete, Aspen Scaber Stalk Overview: The term insigne means “decorated,” a reference to the conspicuous dark scabers on this bolete’s stalk. The cap surface stains grayish with KOH and is negative with NH4OH or FeSO4. The flesh stains bluish with FeSO4 and grayish with KOH and is negative with NH4OH. Cap: 7.5–15 cm wide, convex to broadly convex, with a band of sterile tissue that extends beyond the tube layer and remains as hanging flaps on the margin of mature specimens; surface dry, somewhat granular to fibrous, orange to rusty cinnamon or brick red, becoming brownish orange in age; flesh white, staining purplish gray to purplish black and lacking a red stage when cut or broken; odor and taste not distinctive. Pores and tubes: whitish at first, becoming yellowish to olive brown with age, often staining brownish when bruised; tubes 1–2 cm deep. Stalk: equal or enlarged downward, often with a pinched base, solid; surface dry, whitish beneath a covering of brownish to blackish scabers; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups in woods and parklands beneath big-toothed aspen, quaking aspen, and birch; summer and fall.

252  ·  Boletes of Eastern North America

Distribution and frequency: eastern Canada, south to North Carolina, west to Minnesota; occasional to locally common. Spore print: yellow brown to olive brown. Microscopic features: spores 11–16 × 4–5 µm, ellipsoidal, smooth, yellowish. Edibility: edible with caution. This is a good edible in our experience, but some species of Leccinum with orange-red caps have caused gastrointestinal upset in some individuals. Lookalikes: Leccinum vulpinum (p. 274) is nearly identical, but it grows in association with conifers. Leccinum potteri A. H. Sm., Thiers & Watling (not illustrated) has been reported from Michigan east to New York and grows under aspen or oak. It has a dull-orange to brick-orange or tawny-orange cap that is first covered by a thin whitish veil that soon breaks up to form small, scattered patches that eventually disappear. Also compare with Leccinum versipelle (p. 271).

Leccinum insolens (A)

Leccinum   ·  253

Leccinum insolens (B)

Leccinum insolens A. H. Sm., Thiers & Watling = Leccinum insolens var. brunneo-maculatum A. H. Sm., Thiers & Watling = Leccinum insolens var. insolens A. H. Sm., Thiers & Watling Common name(s): none. Overview: The term insolens means “unusual in relation to others.” The flesh stains blue with FeSO4. The key identification features include a dull-white to buff or brownish cap, whitish flesh that stains grayish and then fuscous when exposed, whitish to buff pores that bruise yellow brown, and a whitish stalk. Cap: 5–15 cm wide, convex, becoming broadly convex to nearly plane, margin incurved, with a narrow band of sterile tissue; surface dry, fibrillose to minutely scaly, sometimes finely cracked, dull white to pale pinkish buff or pale grayish cinnamon buff with dull pinkish buff along the margin, typically darkening to pale yellow brown in age, sometimes staining dull brown when bruised; flesh whitish, staining pinkish gray to purple gray and then fuscous when exposed or sometimes lacking the preliminary pinkish coloration before staining purple gray to fuscous; odor and taste not distinctive. Pores and tubes: white when young, becoming olive buff and then pale yellow brown in age, staining yellow brown when bruised, depressed at the stalk at maturity; tubes 8–20 mm deep.

254  ·  Boletes of Eastern North America

Stalk: nearly equal or enlarged downward, solid; surface dry, whitish, with whitish scabers that become coarse and grayish brown to blackish brown in age on a whitish ground color; flesh concolorous with the cap flesh, often staining reddish only in the base or not at all, sometimes staining blue in the base. Habitat and season: solitary, scattered, or in groups under birch and aspen; summer and fall. Distribution and frequency: eastern Canada south to New England and New York, west to Minnesota, distribution limits yet to be established; uncommon. Spore print: olive brown to dark brown. Microscopic features: spores 12–15 × 4–8 µm, subellipsoidal to subfusiform, smooth, yellowish. Edibility: reported to be edible. Lookalikes: Leccinum chalybaeum (p. 245) is similar, but its cap often has grayish-green to bluish-gray tints, and it is found in the Southeast from Georgia south to Florida, west to Texas.

Leccinum luteum (A)

256  ·  Boletes of Eastern North America

Leccinum luteum (B)

Leccinum luteum A. H. Sm. & Thiers Common name(s): none. Overview: The term luteum means “yellow,” a reference to the color of this bolete’s cap. The typically wrinkled and pitted, yellow to olive-brown cap, whitish to olive-buff pore surface, and association with American hornbeam and ironwood are this uncommon bolete’s distinctive features. Cap: 3–7 cm wide, obtuse to convex when young, obtuse to broadly convex in age, margin even; surface slightly viscid or dry, glabrous, typically wrinkled and shallowly pitted at maturity, becoming cracked in age, pale yellow when young, darkening to yellow and finally olive brown in age; flesh whitish to pale yellow, staining pinkish gray to pinkish brown and eventually dark brown when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, becoming olive buff at maturity, staining pinkish gray when bruised, deeply depressed at the stalk in age; tubes 8–16 mm deep. Stalk: enlarged downward, often curved, solid; surface dry, whitish on the upper portion, pale yellow below, with grayish to blackish scabers; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under American hornbeam (Carpinus caroliniana); summer and fall.

Leccinum   ·  257

Distribution and frequency: eastern Canada south to West Virginia, west to Minnesota; uncommon. Spore print: olive brown. Microscopic features: spores 17–20 × 5–6.5 µm, subfusoid, smooth, yellowish to brownish. Edibility: edible. Lookalikes: Leccinum rugosiceps (p. 262) has a wrinkled and shallowly pitted, orange-yellow cap that becomes yellow brown in age and sometimes is dark tobacco brown on the disc. Its flesh slowly, sometimes faintly, stains reddish or burgundy when exposed, and its cap surface stains orange red with KOH.

Leccinum pseudoinsigne

Leccinum pseudoinsigne A. H. Sm. & Thiers Common name(s): none. Overview: The term pseudoinsigne means “resembling insigne,” a reference to the similar species Leccinum insigne (p. 251). The flesh stains olive gray with FeSO4. The key identification features include the bright-yellow, orange-yellow, or bright-orange to bright orange-red cap color and the preliminary change to reddish cinnamon when the flesh of young specimens is cut. Cap: 4–15 cm wide, obtuse to convex, becoming broadly convex in age, margin with flaps of sterile tissue; surface dry to slightly viscid, glabrous, smooth

258  ·  Boletes of Eastern North America

or nearly so, bright yellow, orange yellow, or bright orange to bright orange red, fading to orange buff in age; flesh of young specimens white, staining reddish cinnamon and then bluish fuscous when exposed, often staining violet gray without a reddish phase when the flesh of mature specimens is exposed; odor and taste not distinctive. Pores and tubes: whitish to pale olive buff at first, becoming brownish in age, staining olive to olive fuscous when bruised, depressed at the stalk in age; tubes 8–15 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, whitish, with whitish scabers that become orange brown and finally blackish in age; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under aspen and birch; summer and fall. Distribution and frequency: New England west to the Great Lakes region and south to the central Appalachians, distribution limits yet to be established; occasional. Spore print: dark yellow brown. Microscopic features: spores 13–17 × 3.5–5 µm, narrowly fusoid, smooth, ochraceous. Edibility: edible. Lookalikes: Leccinum insigne (p. 251) is similar, but it has an orange to rusty-cinnamon or brick-red cap, and its flesh stains purplish gray to purplish black without a reddish phase. Other similar Leccinum species have darker rusty-brown caps.

Leccinum roseoscabrum (A)

Leccinum   ·  259

Leccinum roseoscabrum (B)

Leccinum roseoscabrum Singer & R. Williams Common name(s): none. Overview: This southern bolete has a limited distribution and is not common. The term roseoscabrum means “rosy-pink scabers,” which alludes to the distinctive rose-pink scabers on the stalks of fresh specimens. With age or when a specimen is handled, the scabers darken but usually retain enough color to facilitate identification of this delicately beautiful species. The cap surface stains brown with KOH and is negative with FeSO4 or NH4OH. The flesh stains blue green with FeSO4 and pale orange with NH4OH and is negative with KOH. Cap: 2–7 cm wide, obtuse to convex, becoming broadly convex in age, margin with a narrow band of sterile tissue; surface dry to somewhat viscid, smooth to slightly wrinkled or somewhat shallowly pitted, dark brown to chestnut brown, fading in age, often with paler spots; flesh firm, white at first, slowly staining pink and then grayish and finally dark purple brown when exposed; odor not distinctive; taste not distinctive. Pores and tubes: white to whitish when young, becoming yellowish or grayish with a dull-orange tint, staining brownish when bruised, often depressed at the stalk in age; tubes up to 13 mm deep. Stalk: enlarged downward, often curved, sometimes swollen near the midportion, typically with a pinched base, solid; surface dry, white on the upper

260  ·  Boletes of Eastern North America

portion, pinkish on the lower, sometimes with green or yellow at the base, covered with coarse rosy-pink scabers that darken to brown or dark brown in age or when handled, basal mycelium sparse and white; flesh white at first, quickly staining yellow near the base, then pinkish and purplish, and finally dark purple brown; flesh of the upper portion stains pink, then slowly grayish, and finally dark purple brown. Habitat and season: scattered or in groups in grassy areas or under broadleaf trees, especially oak; summer, fall, and early winter. Distribution and frequency: reported from Florida, Georgia, and Mississippi; occasional. Spore print: yellowish brown. Microscopic features: spores 12–18 × 3.5–6 µm, fusoid to cylindric, smooth, pale ochraceous brown. Edibility: edible and of good flavor. Lookalikes: Leccinum scabrum (p. 265) has a larger brown cap that is often distinctly cracked at maturity, grayish-white pores that stain brownish, and a whitish to pale-tan stalk with dark-brown to blackish scabers, and it occurs with birch.

Leccinum rubropunctum

Leccinum   ·  261

Leccinum rubropunctum (Peck) Singer = Boletus rubropunctus Peck Common name(s): Ashtray Bolete. Overview: The term rubropunctum means “having red punctae.” The cap surface stains amber with KOH or NH4OH and pale olive with FeSO4. The flesh stains pale orange with KOH, displays a brief blue-green flash with NH4OH, and stains grayish olive with FeSO4. Cap: 2–9 cm wide, convex to broadly convex, margin even; surface conspicuously furrowed or wrinkled or sometimes smooth, slightly viscid or dry, glabrous, bay red to chestnut red or reddish brown; flesh yellowish, not staining when exposed; odor unpleasant, reminiscent of stale cigarette butts in an ashtray; taste unpleasant. Pores and tubes: bright golden yellow at first, becoming dull yellow to brownish yellow in age, not staining blue when cut or bruised, depressed at the stalk at maturity; tubes 9–15 mm deep. Stalk: tapered downward, often curved toward the base, solid; surface dry, covered with reddish dots and points on a yellow ground color, with pale-yellow basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in conifer or broadleaf woods; summer and fall, also winter in the southern part of its range. Distribution and frequency: eastern Canada south to Florida west to Minnesota and Mississippi; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 16–22 × 5.5–7.5 µm, subfusiform to somewhat oblong, smooth, pale brown. Edibility: edible but unpleasant tasting. Lookalikes: Hemileccinum subglabripes (p. 220) is similar, but its flesh lacks a distinctive odor, and it has a yellow stalk with a thinner coating of yellow to red punctae and smaller spores, 11–17 × 3–5 µm.

262  ·  Boletes of Eastern North America

Leccinum rugosiceps

Leccinum rugosiceps (Peck) Singer = Boletus rugosiceps Peck = Krombholzia rugosiceps (Peck) Singer = Krombholziella rugosiceps (Peck) Šutara Common name(s): Wrinkled Leccinum Overview: There is nothing particularly remarkable about this handsome bolete. Its scabrous stalk quickly puts it into the Rough Stalk group. The typically wrinkled cap, minute pale-yellow pores that do not stain at first when bruised but eventually stain yellowish-brown, the cream-colored flesh that discolors dark coffee brown where there is insect damage, especially in the stalk, and frequent occurrence under pin oak provide the best clues for identification. The term rugosiceps means “coarsely wrinkled cap.” The cap surface stains orange red with KOH, stains reddish or is negative with NH4OH, and is negative with FeSO4. Cap: 5–15 cm wide, convex to broadly convex, margin with a narrow band of sterile tissue; surface dry, distinctly wrinkled and shallowly pitted at maturity, often finely cracked in age, with pale-yellow flesh showing through the fissures, color orange yellow, becoming yellow brown in age, at times dark

Leccinum   ·  263

tobacco brown on the disc; flesh white or pale yellow, slowly staining reddish or burgundy when exposed, sometimes only faintly, with stains most visible at the juncture of the cap and stalk; odor not distinctive; taste not distinctive or somewhat astringent. Pores and tubes: dull yellow to dingy yellowish when young, sometimes ochraceous to dingy olive brown in age, unchanging or slowly staining yellowish-brown, rarely very slowly staining blue green when cut or bruised, deeply depressed at the stalk in age; tubes 8–14 mm deep, often longer than the thickness of the cap flesh. Stalk: nearly equal or tapered at the apex or base, solid; surface dry, pale yellow to brownish beneath pale-brown scabers that darken in age and sometimes form lines that mimic reticulation, not staining blue when handled or bruised; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or in groups under oaks, especially pin oak, in forests or shaded lawns; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally common, especially in areas landscaped with pin oak. Spore print: olive brown. Microscopic features: spores 15–21 × 5–6 µm, fusiform, smooth, pale brown. Edibility: edible when young and firm but possibly with a slightly unpleasant aftertaste. Some collections are very good. Lookalikes: Leccinum subgranulosum A. H. Sm. & Thiers (not illustrated) has a dry, granulose-scaly to fibrillose, dull yellow-brown cap that becomes minutely cracked in age; whitish flesh that slowly stains brownish when exposed; whitish pores that darken at maturity; and blackish scabers that are finer on the upper portion and coarser on the lower portion over a whitish ground color. Its spores measure 15–21 × 5.5–7.5 µm, and it grows in broadleaf woods, especially under birch. Leccinellum crocipodium (p. 240) has a darker cap, darker scabers on the stalk, and flesh that stains pinkish gray to dull reddish or pale fuscous when exposed. Also compare with Leccinellum quercophilum (p. 244) and Leccinellum griseum (p. 242), both of which have darker caps with more pronounced cracks or fissures.

Leccinum scabrum (A)

Leccinum scabrum (B)

Leccinum   ·  265

Leccinum scabrum (C)

Leccinum scabrum (Bull.) Gray = Leccinum oxydabile (Singer) Singer = Leccinum roseofractum Watling = Leccinum rotundifoliae A. H. Sm., Thiers & Watling Common name(s): Common Scaber Stalk, Birch Bolete, Brown Birch Bolete Overview: The term scabrum means “rough or covered with scabers.” The current description of Leccinum scabrum may differ somewhat from those published in other books because it includes the features of several taxa previously thought to be separate species (see the list of synonyms). The key identification features include a brown cap that is often distinctly cracked at maturity, grayish-white pores that stain brownish, a whitish to pale-tan stalk with dark-brown to blackish scabers, and an association with birch. Cap: 4–12 cm wide, obtuse to broadly convex, margin even, incurved at first; surface moist or dry, glabrous or with matted fibrils, breaking up into tiny, flattened scales and often distinctly cracked at maturity, grayish brown to yellow brown or dark brown; flesh whitish, unchanging or slowly staining reddish when exposed and finally brownish or pale grayish after several hours; odor and taste not distinctive.

266  ·  Boletes of Eastern North America

Pores and tubes: grayish white, often with brownish spots, staining brownish when bruised, becoming ochraceous, grayish, or brownish in age, depressed at the stalk at maturity; tubes 7–18 mm deep. Stalk: enlarged downward or nearly equal, solid; surface dry, whitish to pale tan, covered with dark-brown to blackish scabers over a whitish to grayish ground color, sometimes with blue or greenish-blue stains near the base; scabers fine at the apex and gradually coarser downward; flesh concolorous with and unchanging or staining like the cap flesh when exposed, sometimes with blue or greenish-blue stains near the base. Habitat and season: solitary, scattered, or in groups under birch; summer and fall. Distribution and frequency: eastern Canada south to northern Georgia, west to Minnesota and Arkansas; common. Spore print: brown. Microscopic features: spores 14–19(22) × (4)5–6.5 µm, fusiform, smooth, ochraceous; caulocystidia lageniform to clavate, hyaline or with grayish-brown content in KOH. Edibility: edible. Lookalikes: Leccinum subleucophaeum (p. 269) has a darker, often mottled cap, and its stalk flesh stains grayish to brownish, with or without a reddish intermediate phase. Leccinum chalybaeum (p. 245) is a southern species associated with oak. It has a viscid or shiny buff to pinkish-buff cap that becomes dingy yellow brown with age, often with grayish-green to bluish-gray tints, especially near the margin.

Leccinum snellii

Leccinum   ·  267

Leccinum snellii A. H. Sm., Thiers & Watling = Krombholziella snellii (A. H. Sm., Thiers & Watling) Šutara Common name(s): Snell’s Bolete, Snell’s Leccinum Overview: This small- to medium-size bolete often occurs in dark, moist woods or bog margins where birch is present. It is one of the first boletes to appear in spring and can be found throughout the mushroom season. The combination of a dark-brown to nearly black or sometimes paler-brown cap that is frequently mottled with pallid or cream areas, a dense coating of black stalk scabers, and the blue-green and reddish stains of the stalk flesh separate this bolete from other similar species. It was named in honor of the American boletologist Walter H. Snell (1889–1980). Cap: 3–9 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, margin even; surface dry, smooth, dark brown to blackish, typically with paler mottling, covered with tiny dark-brown to black fibrils when young, often fading to dark yellowish brown in age as the fibrils erode; flesh white, staining reddish, sometimes slowly, when exposed, especially at the juncture of the cap and stalk, darkening to purple gray or black after one hour or more; odor and taste not distinctive. Pores and tubes: whitish when young, becoming grayish to dingy grayish brown in age, unchanging or staining yellowish or brownish when bruised, deeply depressed at the stalk at maturity; tubes 10–15 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, whitish beneath a dense coating of gray to black scabers, often with blue-green stains near the base or where damaged; flesh concolorous with the cap flesh, typically staining blue green, at least near the base, and reddish at the apex. Habitat and season: solitary to scattered or in groups in mixed broadleaf forests, especially under yellow birch; spring, summer, and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; fairly common. Spore print: brown. Microscopic features: spores 16–22 × 5–7.5 µm, fusoid, smooth, pale brown. Edibility: edible, but its flesh, like that of many boletes in the genus Leccinum, is rather soft except in the button stage. The quality is mediocre, so this bolete is best used in a mix with more-flavored boletes. Lookalikes: Leccinum subleucophaeum (p. 269) is similar, but it lacks the conspicuous orange-red staining of the cut stalk flesh, and it has smaller spores, 13–18 × 4.5–6.5 µm.

Leccinum subleucophaeum (A)

Leccinum subleucophaeum (B)

Leccinum   ·  269

Leccinum subleucophaeum (C)

Leccinum subleucophaeum E. A. Dick & Snell = Krombholziella subleucophaeum (E. A. Dick & Snell) Šutara Common name(s): none. Overview: The term subleucophaeum means “somewhat pale grayish and dark brown,” in reference to the colors of this bolete’s cap and pores. This rather darkish bolete is recognized by its dark-brown to blackish-brown and often mottled cap that is sometimes paler toward the margin on young specimens, dark-brown to dark-gray scabers, and white stalk flesh that stains grayish to brownish, with or without a reddish intermediate phase. Cap: 3–14 cm wide, obtuse to convex, becoming broadly convex to nearly plane in age, margin even; surface dry or slightly viscid when moist, nearly glabrous, dark brown with paler mottling, often blackish brown over the disc, sometimes fading to yellowish brown at maturity; flesh white, very slowly staining gray to grayish black or unchanging when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, becoming grayish to dull grayish brown in age, staining yellowish to brownish when bruised, deeply depressed at the stalk in age; tubes 1–2 cm deep.

270  ·  Boletes of Eastern North America

Stalk: tapered in either direction or nearly equal, solid; surface dry, white, with dark-brown to blackish scabers, often staining bluish near the base; flesh white, staining grayish to brownish, with or without a reddish intermediate phase, sometimes staining bluish. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially birch or aspen; summer and fall. Distribution and frequency: eastern Canada south to New England and New York, west to Minnesota; occasional to fairly common. Spore print: brown. Microscopic features: spores 13–18 × 4.5–6.5 µm, fusiform, smooth, brownish. Edibility: edible. Lookalikes: Leccinum snellii (p. 267) is similar, but the cut flesh at the apex of its stalk soon stains conspicuously orange red, and it has larger spores, 16–22 × 5–7.5 µm. Leccinum griseonigrum A. H. Sm., Thiers & Watling (not illustrated), reported from eastern Canada and New England west to Michigan, has a bluish-brown to bluish-black cap when young that becomes yellowish brown to tan and then cinnamon buff and is often cracked in age. It has white flesh that does not stain reddish but rather slowly and erratically stains blue and then violaceous gray when exposed and whitish pores that become grayish to dingy brownish with age and bruise yellowish or brownish. Its stalk is often curved at the base and has brown to dark-gray scabers over a whitish ground color; its spores measure 13–16 × 4–6 µm; and it grows under quaking aspen. Leccinum flavostipitatum E. A. Dick & Snell (not illustrated) grows under spruce from eastern Canada and New England west to Michigan. It has a gray to blackish-gray or brownish cap, white flesh that stains either slowly pale salmon pink or quickly bright blue green when exposed, and brownish to blackish stalk scabers over a pale-yellow ground color. Its spores measure 9–16 × 3.5–5 µm.

Leccinum   ·  271

Leccinum versipelle

Leccinum versipelle (Fr. and Hök) Snell = Leccinum atrostipitatum A. H. Sm., Thiers & Watling = Leccinum testaceoscabrum Secr. ex Singer Common name(s): none. Overview: The term versipelle means “having skin,” a reference to the thin flaps of tissue on this bolete’s cap margin. The combination of a dull-orange cap with marginal flaps of sterile tissue, exceptionally dense black scabers on both young and mature specimens, white flesh that stains reddish and then purple gray to black, and association with birch is distinctive. Cap: 6–18 cm wide, convex, becoming broadly convex, margin with thin flaps of sterile tissue, at least when young; surface dry to slightly viscid, fibrillose, breaking up to form downy patches or small scales in age, dull orange at first, becoming tan or brownish in age; flesh white, staining pinkish and then purple gray to blackish when exposed, especially at the juncture of the cap and stalk; odor and taste not distinctive. Pores and tubes: whitish to buff or pale gray when young, becoming dingy white to pale brown, staining olive to dark brownish when bruised, sometimes slowly; tubes paler than the pores, 1–1.8 cm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, with a dense layer of scabers that are dark gray to coal black, rarely whitish, at both youth and maturity, whitish to dingy tan beneath the scabers, sometimes with bluish to blue-green stains on the lower portion; flesh white, staining

272  ·  Boletes of Eastern North America

reddish and then purple gray to black when exposed, sometimes with bluish to blue-green stains at the base. Habitat and season: solitary or scattered in broadleaf forests, under birch; summer and fall. Distribution and frequency: eastern Canada south into New England and New York, west to Minnesota; occasional to fairly common. Spore print: brown. Microscopic features: spores 11.5–17 × 4–5 µm, subfusiform, smooth, pale brown. Edibility: edible with caution. Some large orange-red Leccinum species have caused gastrointestinal upset in some individuals. The flesh turns black on cooking. Lookalikes: Leccinum insigne (p. 251) is similar, but its white flesh stains purplish gray to purplish black without a pinkish phase; its stalk has brownish scabers that darken to blackish at maturity; and it grows with aspen or birch. Leccinum subtestaceum A. H. Sm., Thiers & Watling (not illustrated) has a dull rusty-red or liver-colored cap, white flesh that lacks a reddish stage but stains purple gray to blackish, and cinnamon-brown pores that become paler in age. Its tubes slowly stain vinaceous gray when cut; it has dark-brown to black scabers, even on young specimens, and white flesh that sometimes stains bluish near the base. It occurs under aspen and birch from Massachusetts to the Great Lakes region.

Leccinum vulpinum (A)

Leccinum vulpinum (B)

Leccinum vulpinum (C)

274  ·  Boletes of Eastern North America

Leccinum vulpinum Watling = Boletus vulpinus (Watling) Hlaváček = Krombholziella vulpina (Watling) Šutara = Leccinum aurantiacum var. vulpinum (Watling) Pilát = Leccinum piceinum Pilát & Dermek Common name(s): Brick-red Conifer Scaber Stalk, Foxy Bolete Overview: This common and robust bolete has been misinterpreted and confused in the literature for years. Most American authors have assigned the European name Leccinum aurantiacum (Bull.) Gray when describing Leccinum vulpinum. According to Henk den Bakker and Machiel Noordeloos (2005), Leccinum vulpinum is the correct name for this coniferassociated bolete. The term vulpinum means “fox,” a reference to the cap color. Cap: 5–20 cm wide, convex, becoming broadly convex to nearly plane in age, with a conspicuous sterile margin that splits into hanging flaps on immature caps and is sometimes present on mature specimens; surface dry to slightly viscid, fibrillose, at times becoming finely cracked in age, dull orange to brick red, rusty red, or reddish brown to dark reddish brown; flesh white on exposure, slowly staining grayish to fuscous, with or without a preceding pinkish phase; odor and taste not distinctive. Pores and tubes: whitish when young, becoming dingier and olive buff to pale grayish brown or pale olive brown in age, slowly staining darker brown to olivaceous when bruised, often depressed at the stalk; tubes concolorous with the pores, 1–2 cm deep. Stalk: nearly equal or enlarged downward, solid; surface whitish, with brown scabers that become blackish in age; flesh concolorous with and staining like the cap flesh in the upper half or more, sometimes with bluish stains near the base. Habitat and season: solitary, but more often scattered or in groups under conifers, especially pines and Norway spruce; summer and fall. Distribution and frequency: widely distributed across the northeast from eastern Canada to Minnesota, south to Tennessee and North Carolina; occasional to locally common. Spore print: brown. Microscopic features: spores 10–18 × 3.5–5 µm, fusiform to subfusiform, smooth, pale brown in water, yellowish in KOH. Edibility: edible with caution. This bolete is a good edible in our experience, but some species of Leccinum with orange-red caps have caused gastrointestinal upset in some individuals. Lookalikes: The Sand-loving Leccinum, Leccinum arenicola Redhead & Watling (not illustrated), reported from New Brunswick south to Cape

Neoboletus   ·  275

Cod, Massachusetts, is normally partially buried and coated with sand. It grows with beach grass, beach heather, and sedges in coastal sand dunes and their margins. Its cap is 5–15 cm wide and orange to yellowish orange when young, fading to dull cinnamon or ochraceous in age, and its white flesh stains vinaceous and then slowly vinaceous gray to purplish gray. It has whitish to tan or pale yellow-brown pores, and its stalk has hazel to vinaceous-buff scabers on a buff ground color. Also compare with Leccinum insigne (p. 251), which is mycorrhizal with aspens and birch. Gen us N eob ol e t u s Gel a r di, Si mon i n i & Vi zzi n i The genus Neoboletus was erected in 2014 to accommodate species formerly classified in two genera, Boletus and Gastroboletus. The name Neoboletus means “new Boletus.” The genus has nine species worldwide, including two species currently known to occur in eastern North America. Although the species in this genus vary widely in macroscopic and microscopic features, molecular analysis clearly shows that they are closely related.

Neoboletus luridiformis (A)

Neoboletus luridiformis (B)

Neoboletus luridiformis (C)

Neoboletus   ·  277

Neoboletus luridiformis (D)

Neoboletus luridiformis (Rostk.) Gelardi, Simonini & Vizzini = Boletus discolor (Quél.) Boud. = Boletus erythropus f. michiganensis A. H. Sm. & Thiers = Boletus erythropus ssp. discolor (Quél.) Dermek, Kuthan & Singer = Boletus luridiformis Rostk. = Boletus luridiformis ssp. discolor (Quél.) Rauschert Common name(s): Slender Red-pored Bolete Overview: The deep, rich, velvety dark-brown cap and orange-red pores that instantly bruise blackish blue are distinctive features of this beautiful bolete. The term luridiformis means “appearing wan or dull,” a reference to its cap colors. Cap: 5–12 cm wide, convex, becoming broadly convex in age, margin even, incurved at first; surface dry, somewhat velvety, dark brown to nearly blackish brown when young, becoming reddish brown to olive brown at maturity, staining dark blue to blackish brown when bruised; flesh greenish yellow to yellow, instantly staining blue when exposed; odor and taste not distinctive. Pores and tubes: orange red to dull orange, sometimes yellow when very young, especially at the margin, staining blackish blue when bruised, depressed at the stalk; pores minute and stuffed at first; tubes pale yellow, 8–14 mm deep.

278  ·  Boletes of Eastern North America

Stalk: nearly equal or enlarged in either direction, solid; surface dry, with reddish to orange-cinnamon punctae over a yellow ground color, often reddish at the base, staining blue when bruised, basal mycelium absent, not reticulate or minutely so at the very apex, lacking reddish hairs at the base; flesh yellow, reddish in the base, rapidly staining blue when exposed. Habitat and season: solitary, scattered, or in groups under conifers or broadleaf trees; summer and fall, also early winter in the southern part of its range. Distribution and frequency: eastern Canada south to South Carolina, west to Minnesota, distribution limits yet to be determined; occasional. Spore print: olive brown. Microscopic features: spores 12–16 × 4.5–6 µm, subfusoid, smooth, ochraceous. Edibility: unknown but reportedly causing gastrointestinal distress. Lookalikes: Suillellus hypocarycinus (p. 310) is very similar, but its stalk has white basal mycelium, and it has different macrochemical test reactions and smaller spores, 8–12 × 3–4 µm. Suillellus luridus (p. 312) has a reticulate stalk. Suillellus subvelutipes (p. 315) often has reddish hairs on the stalk base and a brighter, more orange-colored cap.

Neoboletus pseudosulphureus (A)

Neoboletus   ·  279

Neoboletus pseudosulphureus (B)

Neoboletus pseudosulphureus (Kallenb.) W. Klofac = Boletus pseudosulphureus Kallenb. = Boletus pseudosulphureus var. pallidus Grund & K. A. Harrison Common name(s): none. Overview: This striking bright-yellow bolete that dramatically stains deep blue on all parts is easily identified and a pleasure to encounter. The term pseudosulphureus means “resembling Boletus sulphureus Fr.,” an older name for Buchwaldoboletus sphaerocephalus (p. 179). The bright-yellow cap that becomes reddish brown to brown in age, bright-yellow flesh, bright-yellow pores and stalk, and association with oak and pine are this bolete’s key identifying features. The cap surface stains orange to reddish with KOH, stains grayish to grayish olive with FeSO4, and is negative with NH4OH. Cap: 4–20 cm wide, rounded to convex at first, becoming broadly convex in age, margin even or nearly so; surface dry, covered with tiny, matted fibrils when young, becoming nearly glabrous in age, dull or shiny at maturity, bright yellow, becoming duller yellow to tawny at maturity, typically developing brown to brownish-red tints, especially over the disc in age, quickly staining bluish to bluish black when bruised; flesh bright yellow to greenish yellow, quickly staining blue when exposed; odor not distinctive; taste acidic, astringent, or not distinctive.

280  ·  Boletes of Eastern North America

Pores and tubes: bright yellow at first, becoming greenish yellow and then brownish yellow in age, quickly staining blue and then slowly becoming brownish when bruised, depressed at the stalk at maturity; tubes 8–12 mm deep. Stalk: up to 5 cm thick, enlarged downward, solid; surface dry, bright yellow to yellow, quickly staining blue and then slowly brownish when bruised, sometimes with reddish tints, especially near the base, typically lacking reticulation but sometimes reticulate on the upper portion; flesh yellow on the upper portion, dark red at the base, quickly staining blue at the base or overall when exposed. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak, or in mixed oak and pine woods, sometimes under pines; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional to locally fairly common. Spore print: olive brown. Microscopic features: spores 10–16 × 4–6 µm, fusoid to ellipsoid-fusoid, smooth, brownish yellow. Edibility: edible. Lookalikes: The mature cap of Alessioporus rubriflavus (p. 56) has streaks and splashes of various shades of wine red, red brown, and ocher over a yellow ground color and becomes olive to brownish over the disc in age, but the cap of young specimens is wine red. Its stalk is distinctly reticulated, at least over the upper half. Compare with Lanmaoa carminipes (p. 231). Gen us Pa r ag y rod on Si nger The genus Paragyrodon was erected in 1942 to accommodate a single species formerly classified in several genera, including Boletus, Gyrodon, and Suillus. This bolete has an unusual combination of characters: a thick double veil, decurrent pores, subglobose spores, and an association with oak. The spore-print color is dark yellow brown to olive brown. The name Paragyrodon means “close to Gyrodon,” a genus in which the Ash Tree Bolete, Boletinellus merulioides (p. 82), was formerly classified.

Paragyrodon sphaerosporus (A)

Paragyrodon sphaerosporus (B)

282  ·  Boletes of Eastern North America

Paragyrodon sphaerosporus (Peck) Singer = Boletus sphaerosporus Peck = Gyrodon sphaerosporus (Peck) Singer = Suillus sphaerosporus (Peck) A. H. Sm. & Thiers Common name(s): Double-veiled Bolete Overview: The term sphaerosporus means “rounded spores.” According to Michael Kuo and Andrew Methven (2014), dried specimens yielded the following reactions: the cap surface stains red to pinkish with KOH and grayish with FeSO4 and is negative with NH4OH; the flesh stains red to pinkish with KOH and grayish with FeSO4 and is negative with NH4OH. The presence of a veil excludes this mushroom from the genus Boletus. The combination of subglobose spores and hymenial cystidia separates it from Suillus species, which have ellipsoidal to fusiform spores and typically have resinous dots and smears containing cystidia in bundles on the stalk. Cap: 4–18 cm wide, convex, becoming broadly convex to nearly plane, sometimes shallowly depressed in age, margin incurved at first, becoming decurved to uplifted and irregular, and fringed with remnants of the partial veil; surface viscid to glutinous, glabrous, buff yellow to golden yellow or ocher, becoming dingy yellow brown to reddish brown in age, staining brown when bruised; flesh whitish to yellowish, staining vinaceous brown when exposed; odor and taste not distinctive. Pores and tubes: yellow, becoming brownish in age, staining tawny to cinnamon when bruised, attached to subdecurrent, covered by a partial veil when young; tubes 4–10 mm deep. Stalk: nearly equal, solid; surface dry, yellow above the ring, white to yellowish below, staining brownish when bruised; partial veil whitish, sheathing the entire stalk and attached to the cap margin, tough, fibrous-membranous with an inner gelatinous layer, tearing to form a median ring on the stalk and leaving remnants on the cap margin; ring double, fibrous-membranous on the outside, gelatinous on the inside; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or in groups under broadleaf trees, especially oak, and sometimes found in suburban areas beneath shade trees; summer and fall. Distribution and frequency: eastern Canada, Michigan, Minnesota, and Wisconsin; infrequent. Spore print: dark yellow brown to olive brown. Microscopic features: spores 6–9 × 6–8 µm, globose to subglobose, smooth, ochraceous to brownish. Edibility: unknown. Lookalikes: none.

Phylloporus   ·  283

Gen us P h y l l op oru s Qu él. The genus Phylloporus was erected in 1888 to accommodate species formerly classified in several genera, including Agaricus, Boletus, and Gomphidius. The name Phylloporus means “leaf pores,” a reference to the gill-like fertile surface. Phylloporus is a small, mostly tropical genus, with only three species known from eastern North America. Species in this genus are small to medium terrestrial boletes with a typically decurrent gill-like to poroid spore-bearing layer. They have dry caps with white to yellowish flesh, which in most species does not stain blue when exposed. Spore-print colors are yellowish ochraceous to olive brown.

Phylloporus boletinoides (A)

Phylloporus boletinoides (B)

Phylloporus leucomycelinus (A)

Phylloporus   ·  285

Phylloporus leucomycelinus (B)

Phylloporus boletinoides A. H. Sm. & Thiers Common name(s): none. Overview: The term boletinoides means “having radially arranged and elongated pores.” The cap surface stains reddish to red brown with NH4OH and is negative with FeSO4. Cap: 2–10 cm wide, broadly convex when young, becoming nearly plane and sometimes shallowly depressed in age, margin strongly incurved when young and remaining so at maturity, with a narrow band of sterile tissue; surface dry, somewhat velvety to minutely scaly, becoming nearly smooth at maturity, cinnamon to dark pinkish brown, fading to dull yellow brown in age; flesh white to whitish, slowly staining bluish gray to gray when exposed; odor not distinctive; taste not distinctive or slightly acidic. Pores and tubes: gill-like with numerous crossveins or poroid, radially arranged and strongly decurrent; pale olive buff when young, becoming dark olive buff at maturity, at times with a bluish-green tinge near the margin in age, usually unchanging but sometimes slowly staining bluish green to dark blue when bruised; tubes 3–5 mm deep. Stalk: tapered downward to nearly equal, solid or hollow in the base; surface dry, smooth, pale yellow at the apex, pale cinnamon below, with a sparse layer of white basal mycelium; flesh colored like the cap flesh or darker cinnamon toward the base. Habitat and season: solitary or scattered in mixed pine and oak woods; summer, fall, and early winter. Distribution and frequency: Maine south to Florida, west to Texas; infrequent to fairly common, especially in the southern part of its range. Spore print: olive brown.

286  ·  Boletes of Eastern North America

Microscopic features: spores 11–16 × 5–6.5 µm, subcylindrical to narrowly oval, smooth, pale brown. Edibility: unknown. Lookalikes: Phylloporus leucomycelinus (see photos, p. 284) and Phylloporus rhodoxanthus (p. 286) are similar but have redder caps and yellow to golden-yellow gill-like pores.

Phylloporus rhodoxanthus

Phylloporus rhodoxanthus (Schwein.) Bres. = Phylloporus foliiporus (Murrill) Singer = Phylloporus rhodoxanthus ssp. foliiporus (Murrill) Singer = Xerocomus rhodoxanthus (Schwein.) Bres. & Manfr. Binder Common name(s): Gilled Bolete Overview: This mushroom has the general aspect of a bolete from above and of a gilled mushroom from below. The term rhodoxanthus means “rose yellow.” The cap surface stains blue with NH4OH. Phylloporus foliiporus was once thought to be a distinct species because its pores stain greenish blue when cut or bruised. Cap: 4–10 cm wide, convex, becoming broadly convex to nearly plane or shallowly depressed in age, margin incurved at first; surface dry, somewhat velvety, often cracked, with yellowish flesh showing in the cracks, dull red to reddish brown, reddish yellow, or olive brown; flesh thick near the center, pale yellow; odor and taste not distinctive. Pores and tubes: strongly decurrent, deep yellow to olivaceous yellow, infrequently staining bluish when bruised; fertile portion 8–16 mm deep, occasionally somewhat poroid but typically gill-like with crossveins, radially

Pseudoboletus   ·  287

arranged, often wrinkled and sometimes forked, not easily separable from the cap. Stalk: usually tapered downward, with a swollen base or sometimes nearly equal, firm, solid; surface dry, scurfy or punctate with small reddish-brown dots and points, usually with lines extending down from the gill-like pores, yellow with reddish tinges, with yellow basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in broadleaf woods, especially with oak or beech, sometimes in conifer woods; summer and fall. Distribution and frequency: throughout eastern North America; occasional to fairly common. Spore print: olivaceous yellow brown. Microscopic features: spores 8–14 × 3–5 µm, ellipsoid to fusoid, smooth, pale yellowish. Edibility: edible and usually good, although some collections have a bitter component that makes them unpalatable. Lookalikes: Phylloporus leucomycelinus (see photos, p. 284) is nearly identical but has white basal mycelium. Its pores sometimes stain bluish when cut or bruised. Gen us P s eu d ob ol e t u s Šu ta r a The genus Pseudoboletus was erected in 1991 to accommodate a single species formerly classified in several genera, including Boletus, Suillus, and Xerocomus. This unique bolete, Pseudoboletus parasiticus, only grows attached to the earthball, Scleroderma citrinum. It has a dry and shiny cap and a dry, smooth stalk. It is the only species in this genus that occurs in eastern North America.

Pseudoboletus parasiticus

288  ·  Boletes of Eastern North America

Pseudoboletus parasiticus (Bull.) Šutara = Boletus parasiticus Bull. = Xerocomus parasiticus (Bull.) Quél. Common name(s): Parasitic Bolete Overview: The term Parasiticus means “parasitic,” a reference to the relationship this bolete has with its mycorrhizal associate. This unusual bolete is easy to identify because of its specific association with the earthball Scleroderma citrinum. It is interesting that the Parasitic Bolete is apparently edible, but its earthball host is poisonous. Cap: 2.5–6.5 cm wide, rounded to convex, margin incurved when young; surface dry, somewhat velvety, becoming glabrous and shiny in age, ocher brown to tawny olive or like tarnished brass; flesh pale yellow, unchanging when exposed; odor and taste not distinctive. Pores and tubes: yellow, becoming olivaceous with age, sometimes with reddish or rusty stains, unchanging or very rarely slightly bluish when bruised, often somewhat decurrent; tubes 3–6 mm deep. Stalk: nearly equal, usually curved, solid; surface dry, matted- fibrillose, concolorous with the cap, with white to very pale-yellow mycelium at the base; flesh concolorous with the cap flesh. Habitat and season: solitary, in small groups or clusters attached at the base of Scleroderma citrinum, parasitic; summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Mississippi and Minnesota; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 12–18.5 × 3.5–5 µm, ellipsoidal, smooth, pale brown. Edibility: edible or at least nonpoisonous. Lookalikes: none. Gen us P u lv e rob ol e t u s Mu r r i ll The genus Pulveroboletus was erected in 1909 to accommodate species formerly classified in several genera, including Boletus, Ceriomyces, and Suillus. At present, only four species are known to occur in eastern North America. Although united by DNA analysis, this odd assemblage of species has a variety of macroscopic features that are erratically expressed. Most of the species in this genus have a dry, often powdery, coated cap, but one has a viscid to glutinous cap. They have white to pale-yellow flesh that slowly stains blue in one of the four species and produce an olive-brown to ocher-brown or grayish-brown spore print.

Pulveroboletus   ·  289

Pulveroboletus atkinsonianus

Pulveroboletus atkinsonianus (Murrill) L. D. Gomez = Boletus atkinsonianus (Murrill) Sacc. & Trotter = Ceriomyces atkinsonianus Murrill Common name(s): none. Overview: This bolete was named in honor of American mycologist George F. Atkinson (1854–1918). The reddish-brown to reddish-cinnamon cap with a sterile margin, pale-yellow pores, and cartilaginous stalk that is enlarged downward are distinctive features.

290  ·  Boletes of Eastern North America

Cap: 9–14 cm wide, convex, becoming broadly convex at maturity, margin inrolled when young, with a thin band of sterile tissue; surface dry, viscid when wet, smooth, glabrous, sometimes becoming cracked at maturity, reddish brown to reddish cinnamon, becoming much darker brown in age, often with a pinkish tinge; flesh white, becoming pale pinkish red and sometimes staining blue when exposed; odor not distinctive; taste sweet or not distinctive. Pores and tubes: pale yellow at first, becoming olivaceous brown at maturity, sometimes slightly depressed at the stalk in age, unchanging when bruised; tubes up to 1 cm deep. Stalk: enlarged downward to somewhat bulbous at the base, solid, cartilaginous; surface dry, becoming viscid when wet, glabrous, often longitudinally ridged, concolorous with the cap but paler, pale yellow at the apex, sometimes staining reddish; flesh white, unchanging when exposed. Habitat and season: solitary, scattered, or in groups in broadleaf or mixed woods usually associated with oak or hemlock; summer. Distribution and frequency: reported from Georgia, New York, North Carolina, and West Virginia, distribution limits yet to be established; uncommon. Spore print: dark olivaceous to dull brownish. Microscopic features: spores 11–13 × 4–5 µm, fusiform, smooth, yellowish. Edibility: unknown. Lookalikes: Boletus nobilissimus (p. 123) has a similarly colored cap with a distinct sterile incurved margin. However, its cap is strongly pitted to corrugated or rarely only slightly pitted, and it has a reticulate stalk.

Pulveroboletus auriflammeus (A)

Pulveroboletus   ·  291

Pulveroboletus auriflammeus (B)

Pulveroboletus auriflammeus (Berk. and M. A. Curtis) Singer = Boletus auriflammeus Berk. & M. A. Curtis Common name(s): Flaming Gold Bolete Overview: The term auriflammeus means “flaming gold,” a reference to the overall colors of this bolete’s fruitbody. This small- to medium-size bolete is striking for its overall deep, rich yellow-orange color. Handling this beautiful bolete will stain fingers yellow. The cap surface shows a faint blue flash and then stains vinaceous brown or darker orange with NH4OH, stains dull brown or dark amber after the yellow is dissolved with KOH, and is negative with FeSO4. Cap: 4–9 cm wide, convex to broadly convex, becoming nearly plane in age, margin even; surface velvety to powdery at first, sometimes finely cracked, bright orange yellow to golden yellow or brownish orange; flesh white to cream or pale yellow, not staining blue and not darkening when exposed; odor not distinctive; taste somewhat acidic or not distinctive. Pores and tubes: yellow to yellow orange, sometimes tinged with red, not staining blue when bruised, radially elongated and often depressed at the stalk at maturity; tubes 6–15 mm deep. Stalk: nearly equal, tapered in either direction or sometimes clavate with a pinched base, solid; surface dry, somewhat powdery at first, concolorous

292  ·  Boletes of Eastern North America

with the cap or more intensely yellow orange, coarsely reticulate overall or at least on the upper portion of mature specimens, reticulation often absent or indistinct on young specimens but possibly having longitudinal ribs, with white basal mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary or in small groups or caespitose clusters in broadleaf or mixed woods, especially with oak; summer and fall. Distribution and frequency: New York south to Florida, west to Ohio and Tennessee; occasional. Spore print: olive brown to ocher brown. Microscopic features: spores 8–12 × 3–5 µm, subellipsoid to subfusiform, smooth, nearly hyaline. Edibility: unknown. Lookalikes: Boletus aurantiosplendens (p. 90) is similar but has an orange to brownish-orange or brownish-yellow cap, a yellow to apricot or orange stalk with tawny to reddish-brown streaks that do not stain fingers when handled, and brighter-yellow pores that do not develop crimson tints. Retiboletus ornatipes (p. 300) is a somewhat similar but more robust bolete with coarse, raised reticulation on the stalk and variable cap colors that range from yellow to olive, brown, or gray.

Pulveroboletus curtisii (A)

Pulveroboletus curtisii (B)

Pulveroboletus curtisii (C)

294  ·  Boletes of Eastern North America

Pulveroboletus curtisii (Berk.) Singer = Boletus curtisii Berk. Common name(s): none. Overview: Other than from a few locations, this small, relatively uncommon bolete is not well known. It was named in honor of the nineteenth-century American clergyman turned mycologist Moses Ashley Curtis (1808–1872). This brightly colored bolete bears a superficial resemblance to the genus Suillus, but it lacks the resinous dots on the stalk that characterize many in that genus. The gluten on its cap has an acidic taste and stains fingers yellow. Cap: 3–9.5 cm wide, obtuse to convex, becoming broadly convex to nearly plane in age, margin with a narrow band of sterile tissue, incurved when young; surface bright yellow to orange yellow, initially dusted with an orange powdery coating, viscid and glutinous when fresh, becoming glabrous as the orange coating disappears, sometimes with brownish tints or whitish areas in age; flesh whitish, unchanging when exposed; odor and taste not distinctive. Pores and tubes: whitish to buff, pale yellow, or bright yellow orange at first, duller and brownish at maturity, unchanging when bruised, often depressed at the stalk in age; tubes 6–12 mm deep. Stalk: nearly equal, solid or hollow; surface viscid to glutinous when fresh, somewhat scurfy near the apex, nearly smooth below, pale yellow to yellow down to a base sheathed with cottony white mycelium; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in conifer or mixed woods, often with pines; summer and fall. Distribution and frequency: New England south to Florida, west to Texas; occasional. Spore print: olive brown. Microscopic features: spores 9.5–17 × 4–6 µm, ellipsoidal to subventricose, smooth, yellowish. Edibility: unknown. Lookalikes: Boletus aureissimus var. aureissimus (p. 92) has a yellow cap, yellow unchanging flesh, yellow pores, and a yellow stalk that typically has delicate yellow reticulation on the upper portion.

Pulveroboletus   ·  295

Pulveroboletus ravenelii

Pulveroboletus ravenelii (Berk. & M. A. Curtis) Murrill = Boletus ravenelii Berk. & M. A. Curtis = Suillus ravenelii (Berk. & M. A. Curtis) Kuntze Common name(s): Ravenel’s Bolete, Powdery Sulphur Bolete Overview: The specific epithet honors American botanist-mycologist Henry William Ravenel (1814–1887). This bolete is easy to recognize when young by the floccose-powdery coating on the fruitbody and the delicate cottony ring that usually remains on the upper stalk, at least initially, after the partial veil ruptures. Cap: 2.5–10 cm wide, convex, becoming broadly convex to nearly plane, margin incurved at first, usually fringed with partial-veil remnants; surface dry and powdery to matted and tacky or glabrous when moist, bright yellow, soon becoming reddish orange to reddish brown on the disc; flesh thick, white to yellow, staining pale blue and then brownish when cut or injured; odor not distinctive or somewhat like hickory leaves; taste acidic. Pores and tubes: bright yellow becoming greenish yellow to greenish olive, bruising greenish blue, covered at first with a bright-yellow, cottony partial veil; tubes 5–8 mm deep. Stalk: nearly equal, solid; surface dry, floccose to fibrillose-matted from the base up to a delicate cottony ring near the apex, smooth above the ring, bright yellow; ring fragile, sometimes integrated with the floccose surface of the stalk and inconspicuous; flesh concolorous with and staining like the cap flesh.

296  ·  Boletes of Eastern North America

Habitat and season: solitary, scattered, or in groups in conifer woods, especially under pine or hemlock and rhododendron; summer and fall. Distribution and frequency: widely distributed from eastern Canada south to the Gulf of Mexico, west to Minnesota and Texas; occasional to fairly common locally. Spore print: olive brown to olive gray. Microscopic features: spores 8–10.5 × 4–5 µm, ellipsoidal to oval, smooth, pale brown. Edibility: edible. Lookalikes: Boletus melleoluteus (Snell, E. A. Dick & Hesler) T. J. Baroni & Halling (not illustrated) is a similar but uncommon bolete that has been recorded only from Georgia, North Carolina, and Tennessee. It grows in broadleaf woods, especially beneath oak. It has a smaller, bright-yellow to yellowish-olive or honey-yellow cap that does not develop reddish-orange to reddish-brownish tones at maturity, lacks a partial veil and ring, and has slightly longer spores, 7–12 × 3.5–4.5 µm. Gen us R e t i b ol e t u s M a n f r . Bi n der & Br e s. The genus Retiboletus was erected in 2002 to accommodate about a half-dozen boletes that produce unique pigment compounds called retipolides. The transfer of these species, which were formerly classified in the genus Boletus, is based on the combination of retipolide production, morphological characteristics, and molecular sequencing. Species in this genus have dry caps and prominently reticulate stalks. The spore-print colors are olive brown to dark yellow brown.

Retiboletus griseus

Retiboletus   ·  297

Retiboletus griseus var. fuscus

Retiboletus griseus (Frost) Manfr. Binder & Bres. = Boletus griseus ssp. pinicaribaeae Singer = Boletus griseus var. griseus Frost = Xerocomus griseus (Frost) Singer Common name(s): Gray Bolete Overview: The term griseus means “grayish,” a reference to this bolete’s cap color. The key identification features include the grayish cap, unchanging whitish or slowly staining brownish flesh, whitish to brownish pores, and whitish to grayish stalk that develops yellow tones from the base upward as the bolete matures. Cap: 5–14 cm wide, convex, becoming broadly convex to nearly plane, sometimes slightly depressed, margin even; surface dry, appressed-fibrillose, fibrils grayish, often scaly in age, pale to dark gray or brownish gray when young, sometimes developing ocher tints in age; flesh whitish with dark yellow brown around larval tunnels, unchanging or slowly staining brownish when cut or bruised; odor and taste not distinctive. Pores and tubes: whitish to grayish or dingy gray brown, becoming pinkish brown in age, not yellow at any stage, unchanging or staining brownish or gray when bruised; tubes 8–20 mm deep.

298  ·  Boletes of Eastern North America

Stalk: nearly equal or tapered downward, often curved near the base, solid; surface whitish or grayish when young, developing yellow tones from the base upward with maturation, sometimes with reddish stains, covered overall with a coarse pale to yellowish reticulum that becomes brownish to blackish in age; flesh white with deep yellow at the base. Habitat and season: solitary or scattered in mixed broadleaf woods, especially under oak; summer and fall. Distribution and frequency: eastern North America west to Minnesota; occasional. Spore print: olive brown. Microscopic features: spores 9–13 × 3–5 µm, fusoid to oblong, smooth, pale brown. Edibility: edible and very good, although often damaged by insects; blemish-free specimens seldom found. Lookalikes: Retiboletus griseus var. fuscus (see photo, p. 297) is a variety of Retiboletus griseus that is known from Massachusetts, Florida, North Carolina, Virginia, and Japan. It has a darker-gray cap and whitish to yellowish flesh that slowly stains red when cut or bruised, grows under pines or in mixed pine and oak woods, and has larger spores, 11–17 × 4–6 µm.

Retiboletus ornatipes (A)

Retiboletus ornatipes (B)

300  ·  Boletes of Eastern North America

Retiboletus ornatipes (C)

Retiboletus ornatipes (Peck) Manfr. Binder & Bres. = Boletus ornatipes Peck Common name(s): Ornate-stalked Bolete Overview: This common and attractive bolete with a strongly reticulate yellow stalk is easily recognized and always a pleasure to find. It seldom becomes damaged by insect larvae, but the flesh is too bitter to enjoy as an edible. The term ornatipes means “ornate foot,” a reference to this bolete’s coarsely reticulate stalk. The cap surface stains orangish to pale orange brown with KOH or NH 4OH and is negative with FeSO4. Handling this showy bolete will stain fingers yellow. Cap: 4–20 cm wide, convex, becoming broadly convex to nearly plane or slightly depressed in the center in age; surface dull to somewhat powdery or slightly velvety, smooth, glabrous and shiny when dry, color variable from yellow, mustard yellow, and olive yellow to yellow brown or gray; flesh thick, yellow, unchanging when exposed; odor not distinctive; taste usually very bitter.

Rubroboletus   ·  301

Pores and tubes: bright lemon yellow to deep rich yellow, becoming dingy brownish yellow in age, staining yellow orange to orange brown when bruised; tubes 4–15 mm deep. Stalk: variable from nearly equal or swollen in the middle to tapered toward the base or less often with a swollen or club-shaped base, solid; surface dry, prominently and coarsely reticulate usually over the entire length, yellow to somewhat brownish, staining darker when bruised or handled; flesh concolorous with the cap flesh, rarely staining bluish at the base. Habitat and season: solitary or more often in groups, often in caespitose clusters in broadleaf or mixed woods, especially with oak or beech, sometimes under pine; late spring, summer, and fall. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: olive brown to dark yellow brown. Microscopic features: spores 9–13 × 3–4 µm, oblong to slightly ventricose with an obtuse apex, smooth, pale brown. Edibility: Some collections in the Northeast are mild tasting and edible, but the flesh of most collections is intensely bitter. Lookalikes: Gray-capped forms of Retiboletus ornatipes bear a strong resemblance to Retiboletus griseus (p. 297), which has mild-tasting flesh and is typically yellow only on the lower portion of the stalk. Both species often appear in the same habitat at the same time. Some authors recognize Retiboletus retipes (Berk. & M. A. Curtis) Manfr. Binder & Bres. (not illustrated) as a distinct species having a smaller cap, up to 5 cm in diameter; mild- to somewhat bitter-tasting flesh; and slightly larger spores, 11–15 × 3.2–4.2 µm. Boletus aureissimus var. aureissimus (p. 92) has a bright-yellow to honey-yellow or yellow-ocher cap, a yellow to bright-yellow stalk with or without delicate reticulation on the upper portion, and mild-tasting flesh. Also compare with Pulveroboletus auriflammeus (p. 291), which is similar but generally smaller and has a brownish-orange to golden-yellow or chrome-yellow cap and stalk. Gen us Ru b rob ol e t u s Kua n Zh ao & Zh u L . Ya ng The genus Rubroboletus was erected in 2014 to accommodate ten species distributed worldwide and by means of phylogenetic analysis shown to be related. The name Rubroboletus means “red bolete.” Species in this genus have a pinkish to reddish or sometimes grayish cap surface and orange-red to blood-red pores. They typically have yellow tubes, pink to red punctae or reticulation on the stalk, and a bluish color change when injured. To date, only two species have been reported from eastern North America.

302  ·  Boletes of Eastern North America

Rubroboletus dupainii

Rubroboletus dupainii (Boud.) Kuan Zhao & Zhu L. Yang = Boletus dupainii Boud. = Suillellus dupainii (Boud.) Blanco-Dios Common name(s): none. Overview: The combination of reddish pores and a slimy-viscid cap is most unusual. Rubroboletus dupainii was described in 1902 from central France. The first specimens of this bolete recorded from outside of Europe were based on collections made by American mycologist Owen McConnell in North Carolina. Cap: 2.5–11 cm wide, rounded at first, becoming convex and finally broadly convex to nearly plane in age, margin incurved at first, becoming decurved and extending as a thin band of sterile tissue on mature specimens; surface smooth, slimy-viscid when fresh, becoming shiny when dry, purplish red to pinkish red or bright red, sometimes with yellowish spots; flesh whitish to pale yellow, quickly staining blue and then blackish blue when exposed; odor pleasant and musky or not distinctive; taste not distinctive. Pores and tubes: dark red at first, becoming carmine red to orange red and finally fading to yellow toward the margin in age, staining blue when bruised; tubes 4–10 mm deep, yellow to greenish yellow, bruising blue to greenish blue.

Rubroboletus   ·  303

Stalk: enlarged downward and typically clavate with a somewhat pointed base or nearly equal, solid; surface dry, coated with fine reddish punctae that are more abundant toward the base, ground color yellow, typically with white basal mycelium, staining blue when bruised; flesh yellowish, becoming reddish toward the base, staining blue when exposed. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak and hickory, and in mixed woods; summer and early fall. Distribution and frequency: reported from North Carolina and Iowa; rare. Spore print: olive brown. Microscopic features: spores 10–17 × 3.5–6 µm, fusiform-ellipsoid to ellipsoidal, with prominent oil drops, smooth, light yellow. Edibility: unknown. Lookalikes: Boletus carminiporus (p. 97) is similar, but its cap is dry to subviscid; its flesh does not stain blue when exposed; and it has a distinctly reticulate stalk.

Rubroboletus rhodosanguineus (A)

304  ·  Boletes of Eastern North America

Rubroboletus rhodosanguineus (B)

Rubroboletus rhodosanguineus (Both) Kuan Zhao & Zhu L. Yang = Boletus rhodosanguineus Both Common name(s): none. Overview: The term rhodosanguineus means “red to purplish-red blood,” a reference to this beautiful bolete’s red pores and stalk reticulation. Pink areas of the cap surface produce a bluish-slate flash with NH4OH. Cap: 6.5–14 cm wide, strongly rounded at first, becoming broadly convex to nearly plane in age, margin strongly incurved at first, becoming decurved, with a narrow band of sterile tissue; surface dry, somewhat velvety to nearly glabrous, becoming cracked in age, at first dark reddish pink to purplish pink or brick red with a silvery-white bloom, usually developing brown to olive tones, staining dark red to dark purplish (most noticeable on young specimens) and eventually gray when bruised; flesh yellow, instantly staining blue and then fading to dull grayish blue when exposed; odor of overripe fruit, perfume-like on drying; taste sweet to nauseatingly sweet. Pores and tubes: typically bright blood red to dark red or garnet red to moderate reddish brown at first, becoming dull orange red to dull coral red

Strobilomyces   ·  305

with yellowish-olive tints, sometimes yellow at first when very young, margin with a conspicuous golden-yellow zone, slightly depressed with a short decurrent tooth in age; tubes 7–14 mm deep, yellow to brownish olive, rapidly staining blue when cut. Stalk: strongly club-shaped to somewhat bulbous, solid; surface dry, golden yellow near the apex, pale yellow to pinkish below, typically red on the lower portion and white at the very base, sometimes with brownish spots and streaks, covered nearly overall by a delicate blood-red reticulum, staining blue to greenish on the red areas when bruised; flesh yellow with burgundy-red and golden-yellow areas in the base. Habitat and season: scattered or in groups, sometimes fused in clusters, under red oak; summer and fall. Distribution and frequency: known from eastern Canada, western New York, Ohio, and West Virginia, distribution limits yet to be established; rare to occasional. Spore print: olive brown. Microscopic features: spores 11–15 × 4.5–5.5 µm, subfusiform, smooth, pale yellow in water, yellow to pale amber in Melzer’s. Edibility: unknown. Lookalikes: Boletus flammans (p. 107) has a dark-red to brick-red, rosy-red, or brownish-red cap that lacks olive tones and does not stain dark red to purplish when bruised; the odor of its flesh is not distinctive; it has a less prominently reticulate stalk; and it grows under conifers. Gen us S t robi l om y c e s Ber k . The genus Strobilomyces was erected in 1851 to accommodate species formerly classified in the genus Boletus. The name Strobilomyces means “pinecone fungus,” a reference to the scales on the cap surface of boletes in this genus. Only three species of Strobilomyces are known to occur in eastern North America. They are easy to recognize as a group, although their macroscopic differences are subtle. In general, they are medium to large terrestrial boletes that have a dry, coarsely fibrillose to conspicuously scaly cap and a dry, woolly to shaggy stalk with a distinct ring zone. The pores, which are at first covered by a partial veil, are white but soon become gray and then black. These boletes have whitish flesh that quickly stains orange to orange red and then eventually blackish when exposed. Their spore-print color is brownish black to black.

Strobilomyces dryophilus

Strobilomyces confusus

Strobilomyces   ·  307

Strobilomyces dryophilus Cibula & N. S. Weber Common name(s): none. Overview: The term dryophilus means “oak loving,” a reference to an association with oak trees. The cap surface stains brown with NH4OH or FeSO4 and dark reddish brown with KOH. The key identification features include the grayish-pink to pinkish-brown scales over a whitish ground color, whitish flesh that quickly stains orange to orange red and then slowly blackish, white pores that become black in age, and a pinkish-tan to brownish stalk that is ridged or reticulate above a ring and shaggy below. Cap: 3–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin fringed with cottony pieces of a whitish to tan torn partial veil; surface dry, with a whitish ground color, covered with coarse, woolly or cottony, appressed or erect, grayish-pink to pinkish-tan or pinkish-brown scales; flesh whitish, quickly staining orange to orange red and then slowly blackish when cut or bruised; odor and taste not distinctive. Pores and tubes: white when young, soon becoming gray and eventually black, staining reddish orange or brick red and then black when bruised, covered by a whitish to pale pinkish-tan, cottony to woolly partial veil when young; tubes 1–1.7 cm deep. Stalk: nearly equal or sometimes enlarged at the base, solid, with a ring or shaggy zone; surface dry, pinkish tan to brownish, ridged or reticulate above the ring, shaggy to woolly below; flesh whitish, quickly staining orange to reddish orange and then slowly blackish when exposed. Habitat and season: solitary, scattered, or in groups under oaks or in mixed woods; summer, fall, and early winter. Distribution and frequency: Cape Cod, Massachusetts, south to Florida, west to Texas; occasional to frequent. Spore print: blackish brown to black. Microscopic features: spores 9–12 × 7–9 µm, subglobose to short-ellipsoidal, covered by a distinct and complete reticulum, grayish. Edibility: edible. Lookalikes: Strobilomyces strobilaceus (p. 309) is very similar, but it has a darker purplish-gray to blackish cap and stalk. Strobilomyces confusus (see photo, p. 306) is also similar, but it has a darker purplish-gray to blackish cap with smaller; more erect, stiff, and pointed scales; a darker stalk; and spores with irregular projections and short ridges that lack reticulation.

Strobilomyces strobilaceus (A)

Strobilomyces strobilaceus (B)

Suillellus   ·  309

Strobilomyces strobilaceus (Scop.) Berk. = Strobilomyces floccopus (Vahl) P. Karst. Common name(s): Old Man of the Woods, Pine Cone Bolete Overview: The term strobilaceus means “like a pine cone,” a reference to this bolete’s overlapping cap scales. The cap surface stains brown with NH4OH or FeSO4 and dark reddish brown with KOH. The gray to purplish-gray or blackish scales, whitish flesh that quickly stains orange to orange red and then black, white pores that become black in age, and distinct annular zone on the stalk are the distinctive features. Cap: 3–15.5 cm wide, convex, becoming broadly convex to nearly plane in age, margin fringed with cottony pieces of grayish partial veil; surface dry, with a whitish to grayish ground color, covered with coarse woolly or cottony, flattened or erect, sometimes overlapping, gray to purplish-gray or blackish scales; flesh whitish, quickly staining orange to orange red and then black when cut or bruised; odor and taste not distinctive. Pores and tubes: white when young, soon gray, and then finally black in age, staining reddish to brownish or reddish brown and then black when bruised, covered by a cottony or woolly partial veil when young; tubes 1–1.5 cm deep. Stalk: nearly equal or sometimes enlarged at the base, solid, usually with a distinct shaggy annular zone; surface dry, grayish or concolorous with the cap, reticulate at the apex above the ring zone, shaggy to woolly below; flesh whitish, staining orange to reddish orange and then black when exposed. Habitat and season: solitary, scattered, or in groups under pines or oaks, sometimes with beech; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: blackish brown to black. Microscopic features: spores 9.5–15 × 8.5–12 µm, short-ellipsoidal to globose, covered by a distinct reticulum, grayish. Edibility: edible. Lookalikes: Strobilomyces confusus (see photo, p. 306) is nearly identical but has erect and pointier cap scales as well as spores with irregular projections and short ridges that sometimes resemble a partial reticulum. It also is widely distributed throughout eastern North America. Strobilomyces dryophilus (p. 307) has a dull grayish-pink to pinkish-tan cap that becomes darker brown in age, and its spores have a complete reticulum. Gen us S u i l l e l lu s Mu r r i ll The genus Suillellus was erected in 1909 to accommodate species formerly classified in the genera Boletus, Dictyopus, Leccinum, and Tubiporus. Currently, there are four known members of this genus from eastern North America. Species

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in this genus are terrestrial, have a dry or slightly viscid cap that is glabrous or nearly so, a central stalk that is usually reticulate or pruinose to punctate, white to yellow flesh, and orange to red pores. The species that are currently included in this genus are closely related, as demonstrated by molecular analysis.

Suillellus hypocarycinus

Suillellus hypocarycinus (Singer) Murrill = Boletus hypocarycinus Singer Common name(s): none. Overview: The term hypocarycinus means “under hickory,” a reference to the type of trees in the habitat where this bolete grows. The yellow flesh that quickly stains blue when exposed, red to orange-red or dull-orange pores that quickly stain blue, and whitish to yellowish stalk with carmine-red punctae are its distinctive features. Cap: 4–12 cm wide, rounded to convex, becoming broadly convex in age, margin incurved when young, typically with a narrow band of sterile tissue; surface dry, somewhat velvety, brown to yellow brown, sometimes with olive or cinnamon tones; flesh yellow, quickly staining blue when exposed; odor and taste not distinctive. Pores and tubes: red to orange red or dull orange, quickly staining blue when bruised, usually somewhat depressed at the stalk at maturity; tubes 8–16 mm deep.

Suillellus   ·  311

Stalk: nearly equal or slightly enlarged downward, solid; surface dry, whitish to yellowish with a yellow apex, carmine-red punctae at least on the lower portion, with a white basal mycelium; flesh yellow, rapidly staining blue when exposed. Habitat and season: solitary, scattered, in groups on the ground or in decaying leaf litter under oak and hickory, especially in river bottomlands; summer and fall. Distribution and frequency: North Carolina south to Florida, west to Texas, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: spores 8–12 × 3–4 µm, subfusoid to ellipsoid, smooth, brownish yellow. Edibility: unknown. Lookalikes: Neoboletus luridiformis (p. 277) is very similar, but it has a more northern distribution, different macrochemical test reactions, and larger spores, 12–16 × 4.5–6 µm.

Suillellus luridus (A)

312  ·  Boletes of Eastern North America

Suillellus luridus (B)

Suillellus luridus (Schaeff.) Murrill = Boletus luridus Schaeff. Common name(s): Lurid Bolete Overview: The term luridus means “appearing dull dirty brown or smoky yellow,” a reference to the color of this bolete’s cap. The distinctive features include the variable yellowish to orange or brown cap, yellowish to reddish flesh that quickly stains blue, dark-red to orange-red or brownish-orange pores that instantly stain blackish blue when bruised, and yellow reticulate stalk with red tinges. Cap: 4–12.5 cm wide, convex, becoming broadly convex to nearly plane in age, margin with a narrow band of sterile tissue; surface dry, dull or shiny, somewhat velvety, color variable, yellowish to olive yellow, brownish orange, or olive brown, sometimes with pinkish or reddish tints, staining dark greenish blue to nearly blackish when bruised; flesh yellowish to reddish, sometimes with a red line above the tubes of freshly sectioned specimens, quickly staining blue when bruised; odor and taste not distinctive. Pores and tubes: dark red to orange red or brownish orange, instantly staining blackish blue when bruised, depressed at the stalk at maturity; tubes 1–2 cm deep.

Suillellus   ·  313

Stalk: nearly equal or enlarged downward, solid; surface dry, pruinose to punctate, yellow on the upper portion with a red base or yellow streaked with red, sometimes red at the apex, with red to reddish-yellow reticulation nearly overall or at least on the upper portion; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in broadleaf or mixed woods or landscaped areas, usually with oak or hickory, sometimes under conifers; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: olive brown. Microscopic features: spores 11–17 × 5–7 µm, ovoid to subellipsoid, smooth, yellow. Edibility: reportedly poisonous, causing gastrointestinal distress. Lookalikes: This bolete is sometimes confused with Suillellus subvelutipes (p. 315), which is similar but has an orange-brown cap, lacks reticulation on its stalk, and usually has short reddish or yellow hairs on its stalk base. Suillellus subluridus (p. 314) has a variably colored cap, yellow to orange with purple stains or entirely purplish red; yellow flesh that stains blue; and deep purple-red to dark-red pores that bruise blue. Its stalk is concolorous with the cap or yellow near the apex and red below, bruises blue, and lacks reticulation. Other similar species such as Suillellus hypocarycinus (p. 310) and Neoboletus luridiformis (p. 277) also lack reticulation on their stalks.

Suillellus subluridus

314  ·  Boletes of Eastern North America

Suillellus subluridus Murrill = Boletus miniato-olivaceus var. subluridus (Murrill) Singer = Boletus subluridus (Murrill) Murrill Common name(s): none. Overview: This southern bolete has not been recorded north of North Carolina. The term subluridus means “somewhat like Suillellus luridus” (p. 312), which is a typically drab-colored bolete. The key identification features include the variable orange-pink to orange-yellow or red to purplish-red cap, dark-red pores that stain blue, and the yellow stalk that is minutely punctate and reddish below. Cap: 4–11.5 cm wide, pulvinate to convex, becoming broadly convex to nearly plane in age, margin incurved at first, with a narrow band of sterile tissue when young, becoming even; surface dry, glabrous to velvety subtomentose, sometimes finely cracked in age, color variable, orange pink to orange yellow with vinaceous tints or red to purplish red, sometimes rusty red to cinnamon red, developing more brownish tones in age, staining dark blue when bruised; flesh pale yellow to yellow, staining blue when exposed, sometimes unchanging, especially in older specimens; odor weakly fetid or not distinctive; taste not distinctive. Pores and tubes: often partly yellow at first, especially near the margin, becoming purplish red to dark red overall and then fading to reddish orange to orange yellow in age, staining blue when bruised; tubes 6–12 mm deep. Stalk: tapered in either direction, sometimes ventricose, or nearly equal, with a pinched base, solid; surface dry, glabrous and yellow near the apex, minutely punctate and reddish below, often longitudinally striate, staining blue when bruised; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees or in mixed woods, usually with oaks and pines; summer and fall. Distribution and frequency: North Carolina south to Florida, west to Mississippi, distribution limits yet to be established; uncommon. Spore print: olive brown. Microscopic features: spores 9–14 × 4–6 µm, fusoid to subfusoid, smooth, yellow. Edibility: unknown. Lookalikes: Similar species include Boletus flammans (p. 107), which has a reticulate stalk, and Boletus carminiporus (p. 97), which also has a reticulate stalk and flesh that does not stain blue when exposed.

Suillellus   ·  315

Suillellus subvelutipes

Suillellus subvelutipes (Peck) Murrill = Boletus subvelutipes Peck Common name(s): Red-mouth Bolete Overview: The term subvelutipes means “somewhat velvety foot,” a reference to the yellow or dark-red hairs often present on this bolete’s stalk base. Suillellus subvelutipes is one of the first boletes to appear in early spring, often under planted Norway spruce in lawns, parks, and cemeteries. All parts instantly stain dark blue to bluish black when handled. Cap: 6–13 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety to nearly glabrous, occasionally cracked in age, color variable, cinnamon brown to yellow brown, reddish brown, or reddish orange to orange yellow, quickly staining dark bluish black when bruised; flesh bright yellow, quickly staining dark blue when exposed and then becoming whitish; odor not distinctive; taste slightly acidic or not distinctive. Pores and tubes: variable, red, brownish red, dark maroon red, or red orange to orange when fresh, often with a yellow rim, duller in age, quickly staining dark blue to blackish when bruised; tubes 8–26 mm deep.

316  ·  Boletes of Eastern North America

Stalk: nearly equal, solid; surface dry, scurfy, flushed red and yellow, typically yellow at the apex, quickly staining dark blue to blackish when handled, often with short, stiff, dark-red hairs at the base of mature specimens and yellow hairs on young specimens that become dark red with age; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak, and sometimes under conifers, especially hemlock and Norway spruce; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: dark olive brown. Microscopic features: spores 13–18 × 5–6.5 µm, fusoid-subventricose, smooth, pale brown. Edibility: traditionally considered to be poisonous. However, we have had several recent reports that some individuals have eaten this bolete without ill effects. We have not consumed it. Those who choose to experiment eating it should do so with great caution. Lookalikes: Neoboletus luridiformis (p. 277) has a dark-brown, velvety cap and lacks hairs on the base of its stalk. Boletus subluridellus (p. 163) has a bright-red to pinkish-red, orange-red, or brick-red cap and a yellow stalk with much less red, and the stalk base lacks yellow or dark-red hairs. Gen us S u i l lu s P. M ich eli In 1729, Pier Antonio Micheli  (1679–1737) first erected the genus Suillus. Because this genus predated the accepted start date of Linnean taxonomy in 1753, it was not considered valid until 1821, when British botanist Samuel Frederick Gray (1766–1828) used it to describe a group of mushrooms in his work A Natural Arrangement of British Plants (1821). The name Suillus is derived from the Latin word sus, which means “small pig.” Suillus is a large genus that includes more than thirty species known from eastern North America. They are medium to large terrestrial boletes that almost exclusively grow in association with conifer trees and are often the last boletes to fruit in fall and early winter. Their caps may be dry to moist or viscid to glutinous. Spore prints are olive yellow, yellow brown, olive brown, cinnamon brown, or dark brown. These mushrooms have white or yellow or sometimes brown pores that may be rounded, angular, or elongated and radially arranged. The stalks are solid, and most are covered with conspicuous resinous dots or smears composed of bundles of pigmented caulocystidia. Some species have a partial veil that leaves a membranous or glutinous ring on the stalk. A very few species have reticulation on the stalk, especially above the ring. Scabers are lacking.

Suillus acidus (A)

Suillus acidus (B)

318  ·  Boletes of Eastern North America

Suillus acidus (Peck) Singer = Suillus acidus var. intermedius A. H. Sm. & Thiers = Suillus intermedius (A. H. Sm. & Thiers) A. H. Sm. & Thiers = Suillus subalutaceus (A. H. Sm. & Thiers) A. H. Sm. & Thiers Common name(s): Sour-cap Suillus Overview: The term acidus means “sour or acidic,” a reference to the taste of the gluten on this bolete’s cap. This small- to medium-size bolete is recognized by a persistent ring on the stalk and sour-tasting gluten on its cap cuticle. Cap: 5–16 cm wide, rounded to convex at first, becoming broadly convex to nearly plane in age, margin strongly incurved and remaining so well into maturity, often fringed with soft yellowish remnants of a partial veil; surface smooth, viscid when fresh, shiny when dry, coated with gluten that typically but not always has an acidic taste, yellowish when young, becoming tan, ocher yellow, or pinkish cinnamon to yellow brown and often streaked or spotted in age; flesh whitish to pale yellow or orange yellow, unchanging or slowly staining pale reddish brown when exposed; odor and taste not distinctive. Pores and tubes: pale yellow, with beads of fluid when fresh, dingy yellow in age, slowly staining pale reddish brown when bruised or sometimes unchanging, covered at first with a soft, cottony, yellow partial veil that is coated with gluten; tubes 4–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface pale yellow to ocher yellow, sometimes pinkish brown on the lower portion, with reddish to brownish resinous dots and smears that darken with age and a gelatinous band-like ring near the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups on the ground or among mosses under red or white pine; summer and fall. Distribution and frequency: eastern Canada south at least to North Carolina and Tennessee, west to Minnesota; occasional to fairly common. Spore print: tawny brown to dull cinnamon brown. Microscopic features: spores 8–11 × 3–5 µm, ellipsoidal to subfusoid, smooth, pale brown. Edibility: edible, with a lemony flavor. Lookalikes: Compare with Suillus hirtellus (p. 340), which lacks a ring. Also compare with Suillus tomentosus (p. 360), which lacks a ring and whose pore surface bruises blue.

Suillus   ·  319

Suillus americanus

Suillus americanus (Peck) Snell = Boletus americanus Peck = Boletus americanus var. americanus Peck = Boletus americanus var. reticulipes Coker & Beers = Suillus americanus var. reticulipes (Coker & Beers) Grand Common name(s): American Slippery Jack, Chicken-fat Suillus Overview: This bolete always occurs under white pine, with which it forms a host-specific mycorrhizal relationship. The term americanus means “of America.” The cap surface displays a pink flash and then stains red and finally black with NH4OH. The tissue that hangs from the cap margin of young specimens is not a remnant of a true partial veil because it is never attached to the stalk. When handled, this bolete stains fingers brownish. Although negative reactions are uncommon, some individuals develop contact dermatitis consisting of swelling and an itchy rash or develop irritation of the eyes if rubbed after this bolete is handled. The synonym Suillus americanus var. reticulipes, reported from North Carolina, was originally described as a distinct variety because it has coarse reticulation on its stalk and strongly decurrent pores. Cap: 3–13 cm wide, rounded with an incurved margin when young, becoming broadly convex in age, occasionally with an umbo, margin hung with white

320  ·  Boletes of Eastern North America

to yellow or pale-brown, cottony false-veil tissue; surface viscid to glutinous when moist, covered beneath the gluten by appressed fibrils and flattened scales, or sometimes nearly glabrous on young specimens, color bright yellow to ocher yellow with cinnamon to reddish patches or streaks; flesh yellow, staining pinkish gray to purplish brown when exposed; odor and taste not distinctive. Pores and tubes: yellow when young, slightly browner in age, slowly staining reddish brown when cut or bruised; pores angular, occasionally elongated and radially arranged, often somewhat decurrent; tubes 7–10 mm deep. Stalk: nearly equal, often crooked, solid at first, developing cavities or becoming hollow with age, not reticulate or rarely so at the apex, up to 10 mm thick, lacking a ring; surface dry, yellow, speckled with reddish to darkbrown resinous dots and smears, often developing wine-red or wine-brown stains when bruised or in age; flesh concolorous with and staining like the cap flesh. Habitat and season: sometimes solitary but usually in groups or caespitose clusters under white pine, often common in grassy areas in parks; summer and fall. Distribution and frequency: eastern Canada south to North Carolina and Tennessee, west to Minnesota; fairly common. Spore print: brown. Microscopic features: spores 8–11 × 3–4 µm, nearly fusiform, smooth, pale brown. Edibility: edible though not highly regarded. Lookalikes: Suillus subaureus (p. 358) is somewhat similar, but it lacks fragments of cottony veil tissue on its margin, grows in mixed woods with oak and pine, and occurs in broadleaf woods with oak where pine is absent. Suillus sibiricus (Singer) Singer (not illustrated), which is nearly identical, if not the same species, has been reported in eastern North America from Michigan and Ontario. It is described as having a thicker stalk, up to 15 mm wide, which is bright yellow and often reticulate at the apex. Its veil occasionally leaves a superior ring on the stalk but typically leaves only remnants on the cap margin. It also grows in association with pines.

Suillus   ·  321

Suillus bovinus

Suillus bovinus (L.) Roussel = Boletus bovinus L. = Suillus bovinus var. viridocaerulescens (A. Pearson) Singer Common name(s): none. Overview: The term bovinus means “ox-like,” perhaps a reference to the color of this bolete’s cap. The cap surface and flesh stain vinaceous to burgundy with KOH or NH 4OH and pale olive gray with FeSO4. This European bolete was likely introduced into North America with plantings of Scots pine. Cap: 3–10 cm wide, obtuse to convex at first, becoming broadly convex to nearly plane in age, margin incurved to inrolled at first; surface viscid, glabrous, ochraceous to rusty orange, sometimes with brownish tints, margin whitish; flesh pale yellow to saffron, at least in some areas slowly becoming dull orange pink when exposed, not staining blue when exposed; odor fruity and pleasant; taste somewhat sweet or not distinctive. Pores and tubes: grayish olive at first, becoming ochraceous to dull yellow and then dull olive brown, not staining when bruised, somewhat decurrent; pores angular, compound, highly variable, up to 2 mm wide on mature specimens; tubes 3–6 mm deep. Stalk: nearly equal, solid; surface dry, glabrous, dull yellow to ochraceous at first, developing reddish-brown stains in age or when handled, lacking

322  ·  Boletes of Eastern North America

resinous dots and smears; flesh pale yellow and then dull ochraceous from the base upward in age, not staining when exposed. Habitat and season: solitary, scattered, or in groups among mosses and needle litter under Scots pine; late summer and fall. Distribution and frequency: Adirondack Mountains of New York, also reported from North Carolina, Pennsylvania, and Vermont; rare. Spore print: olive brown. Microscopic features: spores 8–10 × 3–4 µm, ellipsoid to subfusiform, smooth, hyaline to pale yellow in KOH, ochraceous in Melzer’s. Edibility: edible. Lookalikes: Suillus granulatus (p. 333) is similar, but its odor and the taste of its flesh are not distinctive; its stalk has conspicuous pinkish-tan to brownish resinous dots and smears; and it grows under white pine or very rarely with other pines, spruce, or hemlock. Suillus neoalbidipes (p. 344) is also similar. It has a paler cap with a wide band of whitish sterile tissue on the margin and a white stalk that at first lacks conspicuous resinous dots and smears but with age may exhibit some that become brownish.

Suillus bresadolae var. flavogriseus

Suillus   ·  323

Suillus bresadolae var. flavogriseus Cazzoli & Consiglio = Suillus nueschii Singer Common name(s): Bearded Bolete Overview: This bolete has only recently been recorded from Newfoundland and is rare in eastern North America. The term bresadolae honors Italian mycologist Giacomo Bresadola (1847–1929). The term flavogriseus means “yellow and gray,” in reference to the colors of the pores. Cap: 2.5–10 cm wide, convex to broadly umbonate, margin incurved at first, becoming uplifted at maturity, with conspicuous floccose and white to yellow veil tissue especially on immature specimens; surface covered with yellow gluten, pale yellow to dull ocher with reddish-brown tones especially toward the margin, mottled and appearing scaly when drying; flesh whitish to pale yellow, staining slightly pinkish vinaceous when bruised; odor and taste not distinctive. Pore and tubes: whitish to very pale yellow at first, soon developing a grayish tint overall, becoming grayish brown at maturity, not staining when bruised, attached to slightly decurrent; pores angular to irregular. Stalk: nearly equal to slightly club-shaped, stout, solid; surface viscid, colored like the cap on the upper portion, darker reddish brown on the lower portion, sometimes with a deep-yellow base, with a superior to median, sheathing, whitish to yellow membranous ring, basal mycelium white or slightly pinkish; flesh pale yellowish green at first and then becoming olivaceous in age, staining like the cap flesh, especially near the base. Habitat and season: solitary, scattered, or in groups among mosses and needle litter under larch trees; fall. Distribution and frequency: in North America currently known only from Newfoundland; rare. Spore print: dull cinnamon. Microscopic features: spores 7.8–10.9 × 3–4.7 µm, subfusoid-ellipsoid, smooth, pale straw. Edibility: unknown. Lookalikes: Suillus grevillei (p. 335) is similar, but it has yellow pores and a much darker orange-yellow to reddish-brown cap. Suillus bresadolae var. bresadolae (Quél.) Gerhold, which has not been reported from North America, has a dark chestnut-brown cap.

Suillus brevipes (A)

Suillus brevipes (B)

Suillus   ·  325

Suillus brevipes (Peck) Kuntze = Boletus brevipes Peck = Rostkovites brevipes (Peck) Murrill = Suillus brevipes var. subgracilis A. H. Sm. & Thiers = Suillus pseudogranulatus (Murrill) Murrill Common name(s): Short-stalked Suillus, Short-stemmed Slippery Jack Overview: This bolete is aptly named. The term brevipes means “short foot,” a reference to its typically short stalk, which is often so short that the cap appears to rest directly on the ground. The resinous dots exhibited on the stalk of most members of the genus Suillus are absent or colorless and inconspicuous (use hand lens) on this species. Cap: 4–10 cm wide, hemispheric to obtuse at first, becoming broadly convex to nearly plane in age, margin incurved at first, even; surface viscid to glutinous, smooth and glabrous, usually with adhering pine needles and debris, light brown to dark brown, vinaceous brown to cinnamon brown, or grayish brown, fading to cinnamon or tan in age, at times mottled with pallid areas or nearly cream-colored overall, usually with debris and needles adhering; flesh white at first, often becoming yellowish, at least near the stalk apex in age, unchanging when cut; odor and taste not distinctive. Pores and tubes: whitish to pale yellow at first, becoming yellow to dingy yellow in age, unchanging when bruised; tubes honey yellow to olivaceous yellow or sometimes bright yellow, 3–10 mm deep. Stalk: typically short, enlarged at the base or nearly equal, base often pinched, solid; surface dry, nearly glabrous, white to pale yellow, sometimes with brown stains near the base, occasionally with inconspicuous resinous dots that become visible only in age; flesh concolorous with the cap flesh. Habitat and season: scattered, in groups, or in caespitose clusters under pines, often with only the caps visible above the needles or duff; summer and fall, also winter in the southern part of its range, most commonly occurring in early to late fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: cinnamon brown. Microscopic features: spores 7–10 × 2.5–3.5 µm, narrowly ellipsoidal, smooth, yellowish. Edibility: edible. The flesh is soft and mild with good flavor. It can produce a laxative effect if the cuticle and tube layer are not removed and discarded before cooking. Also as for many species in the genus, the texture and flavor are improved on drying. Lookalikes: The Short-stalked Suillus is most likely to be confused with Suillus granulatus (p. 333), which has a paler-brown cap and a longer

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stalk covered with conspicuous pinkish-tan to brownish resinous dots and smears. Suillus neoalbidipes (p. 344) has a paler cap with a wide band of whitish sterile tissue on the margin and a longer stalk.

Suillus cavipes

Suillus cavipes (Opat.) A. H. Sm. & Thiers = Boletinus cavipes (Opat.) Kalchbr. = Boletus cavipes Opat. = Boletus porosus (Berk.) Sacc. Common name(s): Hollow-stalked Larch Suillus, Hollow-stemmed Tamarack Jack Overview: The term cavipes means “hollow foot,” a reference to this handsome bolete’s hollow stalk. The cap surface stains red with NH 4OH and blackish with KOH and is negative with FeSO4. The dry, fibrous to scaly brown cap with whitish veil remnants on the margin, hollow stalk typically with a slight ring, and association with larch are distinctive features. Cap: 3–10 cm wide, rounded to convex, becoming nearly plane in age, occasionally with a rounded umbo, margin often with hanging whitish veil remnants; surface dry, nearly suede-like, densely covered with darkbrown to yellow-brown fibers and small scales; flesh yellow, not staining

Suillus   ·  327

blue when exposed; odor somewhat pungent or not distinctive; taste not distinctive. Pores and tubes: pale yellow to olive yellow, unchanging when cut or bruised; pores large, angular, typically but not always elongated and radially arranged, covered with a whitish partial veil when young; tubes 2–6 mm deep. Stalk: nearly equal or enlarged downward, soon hollow, more or less reticulate at the apex, lacking resinous dots and smears, not staining blue or green when bruised, often with an inconspicuous whitish to ochraceous or brownish ring or a distinct ring zone, pale yellow on the upper portion, concolorous with the cap on the lower portion, basal mycelium white; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups on the ground or among sphagnum mosses as well as in bogs, fens, and other wet areas under larch; fall. Distribution and frequency: eastern Canada south to New England and New York, west to Minnesota; occasional to fairly common. Spore print: brown. Microscopic features: spores 7–10 × 3.5–4 µm, narrowly oval to ventricose, smooth, pale brown. Edibility: edible but tastes unpleasant. Lookalikes: Boletus paluster (p. 132) is similar, but it has a pale pinkish-purple to reddish-purple cap and a concolorous solid stalk.

Suillus cothurnatus

328  ·  Boletes of Eastern North America

Suillus cothurnatus Singer Common name(s): Baggy-veiled Suillus, Booted Suillus Overview: The term cothurnatus means “high boot,” a reference to the baggy partial veil that covers the pores and part of this bolete’s stalk. The cap cuticle stains brown with KOH and a quickly fading pinkish with NH 4OH. Cap: 1.5–6 cm wide, obtuse to convex, becoming broadly convex, sometimes with a low umbo, margin incurved at first, typically with hanging whitish veil fragments; surface smooth, glabrous, viscid, color variable, ochraceous to yellow brown, olive brown to grayish brown or dark brown; flesh marbled orange buff and pale yellow, not staining when exposed; odor fragrant or not distinctive; taste not distinctive. Pores and tubes: pale yellow to orange yellow, becoming brownish yellow in age, unchanging when bruised; pores irregular to radially elongated, when young covered by a thick, baggy, whitish to grayish partial veil that is rubbery and glutinous on the outer layer and that tears, leaving a ring on the stalk at maturity; tubes up to 5 mm deep. Stalk: 2–6 cm long, 5–10 mm thick, nearly equal or tapered downward, dry, solid, whitish to yellowish when young, brownish at maturity, with brownish resinous dots on the upper portion or sometimes overall; ring median to superior, collapsed, membranous, band-like, whitish to grayish, sometimes with a flaring lower edge; flesh colored like the cap flesh. Habitat and season: scattered or in groups in sandy soil under pines, sometimes on moss-covered trunks or stumps; spring through fall, year-round in the southern part of its range. Distribution and frequency: North Carolina south to Florida, west to Oklahoma and Texas; occasional to fairly common. Spore print: brown. Microscopic features: spores 8–10 × 2.5–3.5 µm, ellipsoid-oblong to subcylindric, smooth, honey brown. Edibility: edible and very good, with a lemony flavor, but the cuticle and partial veil must be removed. Lookalikes: Suillus salmonicolor (p. 348) is larger overall and has a more northern distribution, a more orange cap, a thicker veil, often salmon-orange flesh in the stalk base, and different macrochemical reactions.

Suillus decipiens (A)

Suillus decipiens (B)

330  ·  Boletes of Eastern North America

Suillus decipiens (Peck) Kuntze = Boletinus decipiens Berk. & M. A. Curtis = Boletus decipiens Peck Common name(s): none. Overview: The term decipiens means “deceiving,” referring to this species’ close resemblance to other species. The cap surface stains pale gray to greenish gray with KOH, pinkish gray with NH4OH, and slowly pale gray with FeSO4; the flesh stains blue gray with KOH, pinkish to pale raspberry with NH4OH, and slowly pale gray with FeSO4. Cap: 4–9 cm wide, convex to broadly convex when young, becoming nearly plane in age, margin strongly incurved, usually fringed with whitish to yellowish or grayish veil tissue; surface dry, distinctly fibrillose, sometimes with small flattened scales, almost glabrous in places, orangish to dull yellow, tan, or pale reddish brown, staining grayish to blackish, sometimes slowly when bruised; flesh pale yellow to pinkish buff, unchanging or sometimes slowly reddening or darkening when exposed; odor and taste not distinctive. Pores and tubes: orange yellow to yellow, becoming brownish yellow in age, staining brownish when bruised, covered at first by a fibrillose, whitish to yellowish or grayish partial veil that disappears or leaves a thin, fragile ring zone on the upper portion of the stalk; pores angular to irregular, usually elongated and radially arranged; tubes up to 8 mm deep. Stalk: usually enlarged downward and often curved at the base, solid; surface dry, cottony to somewhat velvety or fibrillose, orangish to dull yellow, often bright yellow orange above the ring zone, lacking resinous dots and smears; flesh bright yellow, orange in the base, slowly darkening and developing reddish tints. Habitat and season: scattered or in groups under pines or in mixed pine/oak woods, sometimes common among sphagnum mosses; summer and fall. Distribution and frequency: New England south to Florida, west to Minnesota; occasional to fairly common. Spore print: pale brown. Microscopic features: spores 9–12 × 3.5–4 µm, cylindrical to subellipsoidal, smooth, hyaline to pale ochraceous. Edibility: edible. Lookalikes: Suillus hirtellus (p. 340) has tiny appressed fibrils and scales on its cap, lacks a ring or ring zone, and has a glabrous stalk with prominent resinous dots and smears. Suillus spraguei (p. 356) has much redder cap colors when fresh, but faded specimens strongly resemble Suillus decipiens and can at times be very difficult to differentiate.

Suillus glandulosus (A)

Suillus glandulosus (B)

332  ·  Boletes of Eastern North America

Suillus glandulosus (Peck) Singer = Boletinus glandulosus Peck = Fuscoboletinus glandulosus (Peck) Pomerl. & A. H. Sm. Common name(s): Boreal Bolete Overview: The term glandulosus means “resembling glands,” a reference to the dense blackish dots on the tube walls on this bolete’s elongated pores. The cap surface stains red with KOH or NH4OH and bluish with FeSO4. Cap: 3–12 cm wide, convex, becoming nearly plane and sometimes shallowly depressed, margin with a narrow band of sterile tissue, incurved at first, becoming decurved in age, hanging fragments of the partial veil sometimes present; surface glabrous, glutinous when fresh, red to mahogany red, orange chestnut, or reddish brown, often developing black spots and streaks in age; flesh yellow, reddish around larval tunnels, not staining blue when exposed, separated from the tube layer by a brilliant yellow line; odor slightly pungent and unpleasant or not distinctive; taste not distinctive. Pores and tubes: yellow at first, becoming dull ocher to dull olive yellow or brownish yellow, not staining when bruised but often showing dark-brown spots in age; pores angular to irregular, elongated and radially arranged, sometimes decurrent, covered at first by a gelatinous partial veil that may leave remnants on the cap margin; tubes 4–10mm deep. Stalk: nearly equal or tapered downward, sheathed with gluten up to the ring, yellow with reddish dots above the ring, somewhat reticulate at the apex, dark red to chestnut brown or sometimes yellow below, especially near the base, often spotted and streaked with black at maturity, with a thick membranous ring on the upper portion, stalk dark red when young, becoming blackish brown in age; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under conifers, especially balsam fir and hemlock; fall. Distribution and frequency: eastern Canada south to New England and New York, west to Minnesota; occasional. Spore print: dark reddish brown to purple brown. Microscopic features: spores 8–12 × 3–5 µm, narrowly oblong, smooth, pale brownish ocher. Edibility: edible. Lookalikes: Suillus sinuspaulianus (p. 351) is nearly identical but lacks a gelatinous veil and has a dry stalk.

Suillus   ·  333

Suillus granulatus

Suillus granulatus (L.) Roussel = Boletus granulatus L. = Suillus granulatus subsp. leptopus Singer = Suillus granulatus subsp. snellii Singer = Suillus lactifluus (With.) A. H. Sm. & Thiers Common name(s): Butterball, Granulated Slippery Jack Overview: The term granulatus means “covered with granules,” a reference to the resinous dots and smears on this bolete’s stalk. The cap surface stains dark olive to olive gray with KOH and bluish gray to olive with NH4OH or FeSO4. Cap: 5–12 cm wide, broadly convex, margin even at all stages; surface viscid to glutinous and often streaked or mottled when fresh, color variable, pale yellow or some shade of tan, brown, cinnamon, or orangish cinnamon; flesh white to pale yellow, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: whitish to pinkish buff at first, soon becoming yellowish, often with pinkish-cinnamon moisture beads on young specimens, staining dull cinnamon in age or when bruised or sometimes unchanging; tubes 4–10 mm deep.

334  ·  Boletes of Eastern North America

Stalk: nearly equal, solid; surface dry, whitish when young, becoming yellowish in age, especially at the apex, with conspicuous pinkish-tan to brownish resinous dots and smears; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under pines; summer and fall. Distribution and frequency: widely distributed in eastern North America; fairly common. Spore print: brown. Microscopic features: spores 7–10 × 2.5–3.5 µm, oblong or tapered slightly to the apex, smooth, pale brown. Edibility: edible and regarded as good by some; the cap cuticle and tube layer of older specimens should be peeled and discarded. Lookalikes: Suillus neoalbidipes (p. 344) is similar, but it has a whitish band of sterile tissue on the cap margin and a white stalk that at first lacks conspicuous resinous dots but shows them more obviously with age. Suillus bovinus (p. 321) is also similar but has an ochraceous to rusty-orange cap with a whitish margin, flesh that has a pleasant, fruity odor, and a stalk that lacks resinous dots and smears at any stage. It grows with Scots pine.

Suillus grevillei (A)

Suillus   ·  335

Suillus grevillei (B)

Suillus grevillei (Klotzsch) Singer = Suillus clintonianus (Peck) Kuntze = Suillus grevillei var. clintonianus (Peck) Singer = Suillus proximus A. H. Sm. & Thiers Common name(s): Larch Suillus, Tamarack Jack Overview: The specific epithet honors Scottish botanist Robert K. Greville (1794–1866). The cap surface stains dark green to olive black with NH4OH and greenish black with KOH. This distinctive bolete forms a specific mycorrhizal relationship with native American larch as well as with introduced European larch. Cap: 3.5–14 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, sometimes with hanging remnants of partial-veil tissue; surface glabrous, shiny, viscid to glutinous, color variable, orange yellow, dull red, red brown, reddish brown to dark reddish brown or dark chestnut brown; flesh pale orange yellow, staining pinkish brown when bruised, odor acidic metallic or not distinctive; taste not distinctive. Pores and tubes: yellow when young, darkening to olive yellow or olive brown in age, staining brownish when bruised, attached or depressed at the

336  ·  Boletes of Eastern North America

stalk, covered at first by a yellowish and brownish, cottony partial veil with a gelatinous outer coating; tubes 4–12 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface yellow, reticulate and dry above the ring, viscid to glutinous and reddish brown to brown below, often whitish near the base, with a floccose ring on the upper portion that is covered by a gelatinous layer, lacking resinous dots and smears; flesh yellowish, sometimes staining green near the base when exposed. Habitat and season: scattered or in groups under larch trees; fall. Distribution and frequency: eastern Canada south to West Virginia, west to Minnesota; fairly common. Spore print: dull cinnamon brown. Microscopic features: spores 8–10 × 2.5–4.5 µm, ellipsoid, smooth, pale straw to nearly hyaline. Edibility: edible with a soft texture; young firm buttons are best. Lookalikes: The Slim Jack, Suillus flavidus (Fr.) Presl = Suillus umbonatus E. A. Dick & Snell (not illustrated), reported from eastern Canada and New England west to Minnesota, is a rare species in eastern North America that grows under introduced European larch. It has a viscid to glutinous, dark to pale olive-buff, yellowish to tan or olive-tan cap that is usually umbonate and pale-yellow flesh that stains dingy cinnamon. It has yellowish to olive-yellow pores that stain dingy pinkish cinnamon when bruised and are covered at first by a whitish to brownish, gelatinous partial veil. Its stalk is whitish to pale yellow, with whitish to yellowish resinous dots that darken to pinkish or brownish in age or when handled, and it has a dingy-cinnamon, median to superior, collapsed, band-like ring. Suillus luteus (p. 342) has purple tones on the underside of its ring and grows under pines.

Suillus grisellus (A)

338  ·  Boletes of Eastern North America

Suillus grisellus (B)

Suillus grisellus (Peck) Kretzer & T. D. Bruns = Boletinus grisellus Peck = Fuscoboletinus grisellus (Peck) Pomerl. & A. H. Sm. Common name(s): none. Overview: The term grisellus means “grayish,” a reference to the colors of the cap, pores, and stalk of this bolete. The cap surface stains pale reddish brown with KOH, pale pink with NH4OH, and bluish gray with FeSO4. The dry or slightly viscid grayish cap with an inrolled margin when young, the whitish to grayish or grayish-brown decurrent pores, and growth under larch trees in wet areas are distinctive features. Cap: 3–8 cm wide, obtuse at first, becoming broadly convex to nearly plane, margin typically with hanging remnants of partial-veil tissue, inrolled when young; surface dry or slightly viscid, covered with tiny flattened hairs or scales, whitish to pale olive or olive gray, sometimes with yellow tints; flesh white to whitish, not staining blue when exposed; odor and taste not distinctive. Pores and tubes: pores circular to angular at first, becoming elongated and radially arranged to somewhat gill-like at maturity; whitish to grayish or grayish brown, decurrent to strongly decurrent, not staining blue when bruised, covered when young by a whitish to grayish and somewhat membranous veil that typically does not form a ring on the stalk; tubes 3–6 mm deep.

Suillus   ·  339

Stalk: nearly equal, solid, margin even; surface dry, glabrous, whitish to grayish or yellowish, with yellow basal mycelium, typically lacking a ring; flesh white near the apex, yellow to bright yellow downward to a reddish-orange base, not staining when exposed. Habitat and season: scattered, in groups, or in small clusters on needle litter or among sphagnum mosses in bogs, fens, and other wet areas under larch trees; summer and fall. Distribution and frequency: eastern Canada south to Pennsylvania, west to Minnesota; rare to uncommon. Spore print: grayish brown to dark brown. Microscopic features: spores 9–14(20) × 4–6(7.5) µm, broadly ellipsoidal to elongate-ellipsoidal, smooth, pale olive gray. Edibility: edible but too uncommon to collect for the table. Lookalikes: Compare with Suillus viscidus (p. 361), which has a variably colored grayish to brown cap and a whitish to grayish pore surface that stains bluish to greenish when bruised.

Suillus hirtellus (A)

340  ·  Boletes of Eastern North America

Suillus hirtellus (B)

Suillus hirtellus (Peck) Snell = Boletus hirtellus Peck Common name(s): none. Overview: The term hirtellus means “hairy or shaggy,” a reference to the tiny appressed fibers and scales on this bolete’s cap. The fibrils and scales on the cap as well as the prominent resinous dots and smears are good field identification characters. The cap surface stains reddish brown with KOH and slowly pale grayish with FeSO4 and is negative with NH4OH. Cap: 5–12 cm wide, convex, becoming nearly plane in age, margin incurved when young, fringed with tiny woolly tufts of sterile tissue that disappear at maturity; surface dry to slightly tacky, covered with scattered tufts of reddish, brownish, or grayish fibrils and scales on a yellow ground color, in age glabrous or nearly so, sometimes staining vinaceous brown when handled; flesh pale yellow, unchanging or sometimes staining weakly and erratically blue when exposed; odor and taste not distinctive. Pores and tubes: pale yellow, becoming dull yellow to olive yellow or dingy orange buff in age, unchanging or staining vinaceous brown or more rarely bluish green when bruised, sometimes exuding whitish to pinkish droplets when young; pores slightly elongated and radially arranged at maturity; tubes 3–8 mm deep.

Suillus   ·  341

Stalk: nearly equal or enlarged downward, often curved, solid; surface dry, glabrous overall, pale yellow, sometimes with reddish tints, especially toward the base, with prominent resinous dots and smears that are yellowish at first and brown or blackish brown in age, often with a white basal mycelium; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under conifers, especially pine, spruce, and balsam fir; summer and fall, also winter in the Deep South. Distribution and frequency: widely distributed throughout eastern North America, though more common in the southern part of its range; occasional to locally common. Spore print: ochraceous brown to dull cinnamon. Microscopic features: spores 7–11(13) × 3–3.5 µm, nearly oblong, smooth, pale ochraceous. Edibility: edible. Lookalikes: Suillus decipiens (p. 330), which is often found in the same habitat, has a distinctly fibrillose cap, lacks resinous dots on its stalk, and has a fragile ring zone on the upper portion of the stalk.

Suillus luteus (A)

342  ·  Boletes of Eastern North America

Suillus luteus (B)

Suillus luteus (L.) Roussel = Boletus luteus L. Common name(s): Slippery Jack Overview: Some consider this species to be one of the better edibles in the genus Suillus. It appears in large numbers under pines in the fall, and it is not difficult to recognize. The viscid to glutinous brown cap, yellow pores, and sheathing, purple-tinged partial veil that leaves a prominent membranous ring on the stalk are distinctive field features. The term luteus means “yellow.” Cap: 5–12 cm wide, rounded when young, convex and becoming broadly convex to nearly plane in age, often with a low umbo, margin sterile, often with hanging remnants of the partial veil; surface glabrous, viscid to glutinous when fresh, often sticky and shiny when dry, dark reddish brown to cinnamon brown, yellow brown, or ocher; flesh thick in the center, white to pale yellow, not staining blue when cut or bruised; odor and taste not distinctive. Pores and tubes: whitish to pale yellow when very young, yellow to dark yellow or olive yellow in age, unchanging when bruised, depressed at the

Suillus   ·  343

stalk at maturity, covered when young by a white partial veil with purple to dark-purple or grayish-purple tones on the underside; pores small; tubes concolorous with the pores, 4–15 mm deep. Stalk: nearly equal, solid; surface white when young, becoming pale yellow at the apex and often developing dingy-purplish or brownish tones toward the base in age, speckled with resinous dots and smears, at least above the ring; typically with a large and flaring or sleeve-like white ring with purple or dark-purple tones on the underside; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under or near pines or, rarely, with spruce; summer and fall, also winter in the southern part of its range. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: brown. Microscopic features: spores 7–9 × 2.5–3 µm, nearly oblong, smooth, pale brown. Edibility: edible, but cap cuticle and tube layer of mature specimens must be removed. Lookalikes: Suillus grevillei (p. 335) is similar, but it does not have purple tones on the underside of its partial veil, and it grows with larch. Suillus brevipes (p. 325) is similar and often found in the same habitat at the same time, but it lacks a ring or colored resinous dots on the stalk.

Suillus neoalbidipes

344  ·  Boletes of Eastern North America

Suillus neoalbidipes M. E. Palm & E. L. Stewart = Suillus albidipes Peck Common name(s): White-stemmed Bolete Overview: The cap surface stains olivaceous gray with the application of KOH, lilaceous to greenish gray with NH4OH, and slowly pale olivaceous with FeSO4. The term neoalbidipes means “new white foot,” a reference to this bolete’s being a newly described species that supersedes Suillus albidipes Peck. Cap: 4–10 cm wide, obtuse when young, becoming broadly convex in age, margin inrolled at first, with a wide band of sterile tissue; surface viscid to glutinous, glabrous or coated with tiny fibers, color variable, white, pale pinkish cinnamon, dull orange yellow or yellowish tan when young, becoming darker cinnamon to pale brown or yellowish brown with moderate orange-yellow tones, margin typically white on younger specimens; flesh white, becoming pale yellow at maturity; odor and taste not distinctive. Pores and tubes: whitish at first, becoming yellow at maturity, unchanging when bruised; tubes up to 1 cm deep. Stalk: nearly equal or tapered in either direction, solid; surface dry, white and lacking resinous dots at first, becoming yellowish near the apex and brownish near the base, developing brownish resinous dots and smears in age; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups on sandy soil or among mosses under pines; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional to locally fairly common. Spore print: yellow brown. Microscopic features: spores 6–10 × 2–3 µm, ellipsoidal to fusiform or somewhat ovate, yellowish. Edibility: edible and good once the gluten is removed. Lookalikes: Suillus bovinus (p. 321) is similar, but it has an ochraceous to rusty-orange cap and a stalk that lacks resinous dots, and it grows with Scots pine. Suillus brevipes (p. 325) has a much darker-brown cap without a white margin and a shorter stalk. Compare with Suillus granulatus (p. 333) also, which has conspicuous resinous dots on the stalk at all stages and is almost always found under white pine.

Suillus   ·  345

Suillus placidus

Suillus placidus (Bonord.) Singer = Boletus placidus Bonord. = Suillus placidus f. placidus (Bonord.) Singer Common name(s): White Suillus Overview: This species is the only white Suillus known to occur in eastern North America. It is always found under white pine, with which it forms a host-specific mycorrhizal relationship. The white cap, pinkish-tan resinous dots on the stalk, and association with white pine make this bolete easy to identify. The term placidus means “calm or gentle.” Cap: 3–9 cm wide, obtuse to broadly convex, sometimes nearly plane in age, margin even, inrolled when young; surface viscid, glabrous, white or whitish at first, becoming yellowish in age; flesh white to pale yellow, slowly staining pale vinaceous when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, soon yellowish to pale ocher yellow in age, often beaded with pinkish droplets when moist, unchanging when bruised, evenly attached to the stalk or slightly decurrent; tubes 3–8 mm deep. Stalk: nearly equal or tapered in either direction, solid or sometimes hollow in age; surface dry, white, becoming yellow at maturity, with conspicuous pinkish-tan resinous dots and smears that become reddish brown or darker in age; flesh concolorous with and staining like the cap flesh.

346  ·  Boletes of Eastern North America

Habitat and season: scattered or in groups under white pine; summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Minnesota and Mississippi; occasional. Spore print: dull cinnamon. Microscopic features: spores 7–9 × 2.5–3.5 µm, nearly oblong, smooth, yellowish. Edibility: edible. Lookalikes: none.

Suillus punctipes

Suillus punctipes (Peck) Singer = Boletus punctipes Peck Common name(s): Fragrant Bolete, Spicy Suillus Overview: The term punctipes means “dotted foot,” a reference to the resinous dots and smears on this bolete’s stalk. The cap surface stains dull purple with KOH and purplish to brown with NH4OH. Cap: 3–10 cm wide, convex, becoming broadly convex, margin even; surface viscid, covered with tufts of tiny gray or brown fibrils when young, soon becoming smooth and dull ocher orange; flesh pale yellow to yellow orange, not staining blue when exposed; odor fragrant, spicy or like almond extract; taste not distinctive.

Suillus   ·  347

Pores and tubes: brown when young, becoming orange brown to dark or dingy yellow in age, not staining blue when bruised; tubes 4–8 mm deep. Stalk: equal or enlarged downward, often curved, solid; surface dry, dull orangish brown when young, becoming dull ocher orange to ocher yellow, sometimes with reddish stains at the base, densely covered with brown to dark-brown resinous dots and smears; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under spruce, balsam fir, white pine, and other conifers in forests or bogs; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: brown. Microscopic features: spores 7.5–12 × 3–4 µm, ellipsoidal to subfusiform, smooth, pale brown. Edibility: edible. Lookalikes: Suillus subaureus (p. 358) has orangish-yellow pores and flesh that lacks a distinctive odor. Suillus tomentosus (p. 360) has a yellow to orange-yellow, fibrillose-scaly cap that becomes nearly glabrous with age, brownish pores that bruise blue, and flesh that irregularly and sometimes slowly stains blue when exposed.

Suillus salmonicolor

348  ·  Boletes of Eastern North America

Suillus salmonicolor (Frost) Halling = Suillus subluteus (Peck) Snell = Suillus pinorigidus Snell and Dick Common name(s): Slippery Jill Overview: The term salmonicolor means “salmon-colored.” The cap cuticle stains purplish red to dark purple with KOH or NH4OH. The flesh stains dark purplish red with KOH or NH4OH. Cap: 3–10 cm wide, convex to broadly convex, sometimes with a low umbo, margin incurved at first, typically with hanging whitish veil remnants; surface smooth, glabrous, viscid to glutinous when moist, shiny when dry, color variable, yellowish orange to ochraceous salmon or olive brown to cinnamon brown or yellow brown; flesh pale orange yellow to orange, not staining when exposed; odor and taste not distinctive. Pores and tubes: yellow to dingy yellow or yellowish orange to salmon, darkening to brownish in age, not staining when bruised; pores circular to angular, when young covered by a thick, baggy, whitish to grayish partial veil that is rubbery, glutinous on the outer layer, with a thick cottony roll on the base that tears and leaves a ring on the stalk when mature; tubes up to 10 mm deep. Stalk: 2.5–10 cm long, 6–16 mm thick, equal or enlarged downward, whitish to yellowish or orangish, with reddish-brown to dark-brown resinous dot and smears; ring median to superior, collapsed, membranous, band-like, whitish, with a conspicuous flaring lower edge; flesh ochraceous to yellowish, often salmon orange at the base. Habitat and season: scattered or in groups under pines; summer and fall. Distribution and frequency: eastern Canada south to Virginia, west to Minnesota and Missouri, specific distribution range yet to be determined; occasional to fairly common. Spore print: cinnamon brown to brown. Microscopic features: spores 6–11 × 2.5–4 µm, ellipsoidal, smooth, pale brown. Edibility: edible and very good, with a lemony flavor, but the cuticle and partial veil should be removed. Lookalikes: Suillus cothurnatus (p. 328) is smaller overall and has a more southern distribution, a more brownish cap, a thinner veil, and different macrochemical reactions.

Suillus   ·  349

Suillus serotinus

Suillus serotinus (Frost) Kretzer & T. D. Bruns = Boletus serotinus Frost = Fuscoboletinus serotinus (Frost) A. H. Sm. & Thiers Common name(s): none. Overview: The term serotinus means “late in the season,” a reference to the time of year when this bolete appears. The flesh quickly stains green with FeSO4. When this bolete is wrapped in waxed paper or collected in waxed-paper bags, it leaves conspicuous blue stains on the paper. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane, often with a low umbo; margin incurved at first, usually with attached fragments of the partial veil; surface glutinous to viscid, with a dark reddish-brown layer of gluten that covers the whitish ground color, fading to pale reddish brown or yellow brown in age; flesh white to pale yellow, slowly staining bluish and then purplish gray and finally reddish brown when exposed or sometimes unchanging; odor somewhat pungent or not distinctive; taste not distinctive. Pores and tubes: whitish, becoming grayish white and then pale reddish brown with age, staining purplish gray and then reddish brown when bruised, typically somewhat decurrent and not depressed at the stalk, covered at first with a cottony-membranous, grayish-white veil; pores angular,

350  ·  Boletes of Eastern North America

not elongated and radially arranged or only slightly so; tubes 9–16 mm deep. Stalk: nearly equal, solid; surface dry, dingy white and weakly reticulate above the ring, whitish with pinkish-brown to yellowish-brown streaks or blotches below, having a cottony-membranous, grayish-white ring on the upper portion; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups, often among sphagnum mosses in wet areas under larch trees; late summer and fall. Distribution and frequency: eastern Canada south to New England and New York, west to Minnesota; uncommon. Spore print: purplish brown. Microscopic features: spores 8–12 × 4–5 µm, oblong to subellipsoidal, smooth, brownish. Edibility: edible but too uncommon to gather for the table. Lookalikes: Suillus viscidus (p. 361) is very similar but lacks the dark reddish-brown gluten on the cap, and its flesh stains only bluish green, not purplish gray or reddish brown. Suillus weaverae (A. H. Sm. & Shaffer) Kretzer & T. D. Bruns (not illustrated) is known only from the Great Lakes region. It has a viscid, vinaceous-buff to pinkish-cinnamon cap with a pinkish-gray to vinaceous-brown margin that is often adorned with the hanging remnants of a partial veil. The yellow flesh is unchanging when exposed. The pores are yellowish to ochraceous as well as elongated and radially arranged, and they do not stain when bruised. The stalk is tan and bruises yellow at the base. This species occurs in mixed woods of pine, oak, beech, and poplar.

Suillus sinuspaulianus (A)

Suillus   ·  351

Suillus sinuspaulianus with Gomphidius (B)

Suillus sinuspaulianus (Pomerl. & A. H. Sm.) E. A. Dick & Snell = Fuscoboletinus sinuspaulianus Pomerl. and A. H. Sm. Common name(s): none. Overview: The term sinuspaulianus refers to Baie-Saint-Paul, Quebec, where the type collection of this bolete was found. The key identification features include the chestnut-brown to red-brown cap, angular and radially arranged pores, and association with conifers. Cap: 3–13 cm wide, acutely convex at first, becoming convex to broadly convex and then nearly plane in age, sometimes slightly umbonate, margin incurved at first, with a narrow band of sterile tissue, sometimes with hanging remnants of a partial veil; surface smooth or somewhat wrinkled, obscurely innately fibrillose to glabrous, viscid, chestnut brown or rusty brown to red brown, fading to orange brown with age; flesh soft and watery, dingy orange buff to yellow, unchanging when exposed; odor somewhat mealy; taste not distinctive. Pores and tubes: yellow, becoming dingy yellow to dull yellow brown at maturity, darkening when bruised, evenly attached to somewhat decurrent, covered by a felt-like, nongelatinous, tufted veil when young; pores large, angular, and radially arranged; tubes yellowish brown, 4–10 mm deep. Stalk: nearly equal or enlarged in the middle or below, solid; surface dry, lacking resinous dots and smears, reddish brown, dull yellow to yellowish brown and reticulate above the ring, yellow brown to grayish brown with red lines or spots and covered with tiny hairs or scales below the ring; flesh concolorous with the cap flesh, reddish around worm holes and becoming slowly reddish toward the base when exposed. Habitat and season: scattered or in groups under conifers, especially pine, spruce, and balsam fir; fall.

352  ·  Boletes of Eastern North America

Distribution and frequency: eastern Canada south to New England, west to Minnesota; uncommon. Spore print: chocolate brown to purple brown. Microscopic features: spores 8–10.5(13.2) × 4–4.8 µm, ellipsoidal or narrowly ovate, smooth, pale yellowish brown. Edibility: edible. Lookalikes: Suillus glandulosus (p. 332) is nearly identical but has a gelatinous veil covering the stalk that causes it to be viscid.

Suillus spectabilis (A)

Suillus   ·  353

Suillus spectabilis (B)

Suillus spectabilis (Peck) Kuntze = Boletinus spectabilis (Peck) Murrill = Boletus spectabilis Peck = Fuscoboletinus spectabilis (Peck) Pomerl. & A. H. Sm. Common name(s): none. Overview: The term spectabilis means “spectacular or showy.” The viscid reddish cap covered with flat-brown scales, the gelatinous ring on the stalk, and growth under larch trees are this bolete’s main identifying features. Cap: 4–10 cm wide, convex to nearly plane, with or without an umbo, margin often with attached remnants of the partial veil, at least when young; surface covered with flat, coarse, pinkish-gray to reddish-brown scales or patches, viscid to glutinous beneath the scales, pinkish red to orange red, darkening in age; flesh yellow, slowly staining pinkish and then brown when exposed; odor disagreeable or somewhat pungent; taste astringent or acidic. Pores and tubes: yellow, becoming dull yellowish brown in age, usually staining pinkish when injured, covered by a gelatinous pale-red to yellowish-red partial veil when young; pores angular, elongated, and radially arranged; tubes 6–12 mm deep.

354  ·  Boletes of Eastern North America

Stalk: enlarged downward or nearly equal, solid, with a gelatinous, reddish to reddish-brown ring on the upper portion; surface yellow and smooth above the ring, whitish directly beneath the ring, lower portion with viscid pinkish-red to pinkish-gray fibrils over a yellowish ground color, often whitish at the base; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups among sphagnum mosses in bogs, fens, or other wet areas under larch trees, sometimes on moss-covered tree stumps; summer and fall. Distribution and frequency: eastern Canada south to Pennsylvania and Ohio, west to Minnesota; occasional. Spore print: purplish brown. Microscopic features: spores 9–15 × 4–6.5 µm, ellipsoidal, smooth, pale yellowish brown. Edibility: edible but too uncommon to consider collecting for the table. Lookalikes: Compare with Suillus glandulosus (p. 332) and Suillus sinuspaulianus (p. 351), both of which are somewhat similar in color but have glabrous caps.

Suillus spraguei (A)

Suillus spraguei (B)

Suillus spraguei (C)

356  ·  Boletes of Eastern North America

Suillus spraguei (Berk. & M. A. Curtis) Kuntze = Boletinus pictus Peck = Boletus pictus Peck = Boletus spraguei Frost = Suillus pictus (Peck) A. H. Sm. & Thiers Common name(s): Painted Bolete, Painted Suillus Overview: This attractive bolete is restricted to white pine, with which it forms mycorrhiza. The scaly, reddish cap and stalk, the web-like partial veil, and specific association with white pine make it easy to identify. The cap surface stains blackish with KOH, FeSO4, or NH4OH. The flesh stains grayish green to greenish black with FeSO4 and olive to greenish black with KOH or NH4OH. The name Spraguei honors Charles James Sprague (1823–1903), who first collected this species in 1856. Cap: 4–12 cm wide, convex with an incurved margin at first, becoming nearly plane in age, with or without a low umbo, margin fringed with fragments or flaps of whitish partial veil when young; surface dry, cottony-velvety, soon breaking into fibrils and soft, more or less flattened scales to reveal yellow flesh within, red to purplish red or rose red, fading in age to buff or ochraceous; flesh yellow, sometimes staining slightly reddish; odor and taste not distinctive. Pores and tubes: yellow, becoming dingy yellow to ochraceous in age, staining reddish to brownish when bruised, at times slightly decurrent, covered at first by a whitish, cottony or web-like partial veil; pores large and angular to elongated, radially arranged; tubes 4–8 mm deep, not easily separated from the cap flesh. Stalk: more or less equal, sometimes with a swollen base, solid; surface dry, cottony-scaly, the ornamentation sometimes arranged in concentric bands from the base up to a whitish or grayish cottony, flaring ring near the apex, concolorous with the cap or paler, lacking resinous dots and smears; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered and often gregarious under white pine; spring, summer, and fall. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: olive brown. Microscopic features: spores 8–12 × 3.5–5 µm, ellipsoidal, smooth, pale brown. Edibility: Although the flesh of the Painted Bolete is soft and turns black when cooked, some consider it to be one of the better edibles in the genus. It is very good when dried, crumbled, and added to soups, gravies, casseroles, and meat dishes.

Suillus   ·  357

Lookalikes: Suillus decipiens (p. 330) has an orangish to dull-yellow, tan, or pale reddish-brown cap. Faded specimens of Suillus spraguei strongly resemble Suillus decipiens and can be difficult to differentiate in the field.

Suillus subaureus (A)

Suillus subaureus (B)

358  ·  Boletes of Eastern North America

Suillus subaureus (Peck) Snell = Boletus subaureus Peck Common name(s): none. Overview: This bolete is an unusual species of Suillus because it is not associated exclusively with conifers. The term subaureus means “nearly golden,” in reference to the color of this species’ pores. Cap: 3–14 cm wide, convex, becoming broadly convex to nearly plane in age, margin even, incurved to inrolled at first and often remaining so well into maturity; surface with scattered, appressed, scarlet to reddish-brown fibrils over an apricot-orange to yellow ground color, viscid beneath the appressed fibrils, somewhat shiny when dry; flesh yellow, often reddening slightly when cut or bruised, not staining blue or green; odor not distinctive; taste slightly acidic or not distinctive. Pores and tubes: radially arranged, pale yellowish orange, sometimes with clear droplets when young, becoming dull ocher to dingy yellow in age, unchanging when bruised; tubes 3–10 mm deep. Stalk: nearly equal, solid; surface viscid or dry, yellow or yellowish, usually with resinous dots and smears that are yellow when young but darken in age and are somewhat raised near the base, apex typically glabrous or pseudoreticulate; flesh yellow, slowly darkening olive brown to brown when exposed. Habitat and season: solitary, scattered, or in groups in broadleaf woods, especially with birch or oak, and sometimes in conifer or mixed woods under white pine; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: brown. Microscopic features: spores 7–10 × 2.7–3.5 µm, narrowly ellipsoidal to subfusiform, smooth, pale brown. Edibility: edible. Lookalikes: Suillus punctipes (p. 346) has a smooth, dull-yellow to ochraceous or tawny cap, grayish-brown or cinnamon-brown or dull-ocher pores, and pale-yellow flesh with a spicy odor. Suillus tomentosus (p. 360) has a yellow to orange-yellow fibrillose-scaly cap that becomes nearly glabrous with age, brownish pores that bruise blue, and flesh that irregularly and often slowly stains blue when exposed.

Suillus tomentosus (A)

Suillus tomentosus (B)

360  ·  Boletes of Eastern North America

Suillus tomentosus (C)

Suillus tomentosus (Kauffman) Singer = Boletus tomentosus Kauffman = Suillus tomentosus var. discolor A. H. Sm., Thiers, & O. K. Miller Common name(s): Blue-staining Slippery Jack, Poor Man’s Slippery Jack Overview: The fibrillose-scaly, yellowish-orange cap that becomes nearly glabrous in age and brownish pores that bruise blue are distinctive features. The term tomentosus means “covered with matted hairs,” a reference to the cap surface of young specimens. Handling this bolete can sometimes stain fingers dark brown. Cap: 5–12 cm wide, obtuse to convex, becoming broadly convex to nearly plane in age, margin even, becoming uplifted and wavy in age; surface dry, conspicuously fibrillose-scaly with grayish fibers and scales when young, becoming nearly glabrous in age, yellow ocher, dark olive yellow, or reddish brown on a yellow to orange-yellow ground color; flesh yellow, erratically and often slowly staining blue when exposed; odor and taste not distinctive. Pores and tubes: brown to vinaceous brown or dingy cinnamon, sometimes beaded with yellowish droplets when young, becoming dingy yellow to olive yellow in age, staining blue when bruised, evenly attached to slightly decurrent; tubes 8–15 mm deep.

Suillus   ·  361

Stalk: enlarged downward or nearly equal, solid; surface dry or moist, yellow to dull orange yellow with darker-orange to brownish resinous dots and smears; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under conifers; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; fairly common. Spore print: brown. Microscopic features: spores 7–12 × 3–5 µm, fusoid to elongate-ovoid, smooth, yellowish. Edibility: edible. Lookalikes: Suillus punctipes (p. 346) has a similarly colored cap and pores, but its flesh has a spicy, fragrant odor, and its pores do not bruise blue. Other similar species of Suillus lack the blue staining reaction of the pore surface.

Suillus viscidus

Suillus viscidus (L.) Roussel = Fuscoboletinus aeruginascens (Secr. ex Snell) Pomerl. & A. H. Sm. = Fuscoboletinus laricinus (Berk.) Bessette, Roody, & A. R. Bessette = Fuscoboletinus viscidus (L.) Grund & K. A. Harrison = Suillus aeruginascens Secr. ex Snell = Suillus laricinus (Berk.) Kuntze

362  ·  Boletes of Eastern North America

Common name(s): none. Overview: The term viscidus means “sticky or tacky.” The flesh stains reddish brown with NH4OH. This bolete stains waxed paper blue after being wrapped in it for several minutes. Cap: 3–12 cm wide, obtuse at first, becoming broadly convex to nearly plane, margin often with remnant fragments of a partial veil when young, inrolled at first, becoming decurved with a narrow band of sterile tissue; surface glabrous, glutinous to viscid when fresh, gluten colorless, ground color variable from grayish to greenish gray or pale grayish brown to yellow brown or reddish brown, sometimes cracked when dry; flesh white to pale yellow, staining bluish green when exposed, sometimes slowly and weakly; odor somewhat pungent or not distinctive; taste not distinctive. Pores and tubes: whitish to pale gray, staining bluish or greenish when bruised, sometimes slowly or weakly; pores angular to irregular, covered at first by a whitish to grayish partial veil; tubes 5–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface viscid, fibrillose to scurfy, with a membranous, collapsed ring on the upper portion, sometimes broadly reticulate near the apex, whitish to pale brown above the ring, pale grayish to grayish brown or brown below. Habitat and season: scattered or in groups under native larch trees, usually in wet areas; fall. Distribution and frequency: eastern Canada and the northeastern United States, west to Minnesota, also reported from as far south as Mississippi with planted larch; occasional. Spore print: vinaceous brown. Microscopic features: spores 7–12 × 3.5–5 µm, ellipsoidal to subfusiform, smooth, pale yellow brown. Edibility: edible. Lookalikes: Suillus serotinus (p. 349) is very similar but has dark reddish-brown gluten on the cap, and its exposed flesh sometimes slowly stains bluish, then purplish gray, and finally reddish brown. Gen us S u t or i u s H a l l i ng, Nu h n & N. A. Fech n er The genus Sutorius was erected in 2012 to accommodate Boletus robustus, originally and illegitimately named in 1874 by Vermont mycologist Charles C. Frost (1805–1880) and renamed Boletus eximius by Charles H. Peck (1833–1917) in 1887. Since then, it has been reclassified in several additional genera, including Leccinum and Tylopilus. Molecular sequencing determined that it does not belong in any of these genera and that a new genus name was needed. The name Sutorius is Latin for “shoemaker” and honors Frost, who worked as a cobbler. Sutorius is a small genus with presently only two species described worldwide,

Sutorius   ·  363

one of which occurs in North America. They have a purple-brown to grayish-brown cap; white flesh mottled with grayish, reddish, or brownish lilac; and a stalk that has a dense coating of purple-brown to grayish-brown fine scales.

Sutorius eximius (A)

Sutorius eximius (B)

364  ·  Boletes of Eastern North America

Sutorius eximius (Peck) Halling, Nuhn & Osmundson = Boletus eximius Peck = Leccinum eximium (Peck) Singer = Tylopilus eximius (Peck) Singer Common name(s): Lilac-brown Bolete Overview: The term eximius means “distinguished or excellent in size or beauty.” This bolete’s key identification features include the purplish-brown to grayish-brown cap, dark purple-brown to chocolate-brown pores that become reddish brown at maturity, and pale purplish-gray stalk that is densely covered with tiny darker purple-brown scales. Cap: 5–12 cm wide, hemispherical to convex at first, becoming broadly convex to nearly plane in age, margin incurved at first; surface dry, glabrous to finely velvety, purplish brown to grayish brown, often with a whitish bloom when young; flesh whitish to grayish, reddish, or brownish lilac; odor slightly pungent or not distinctive; taste not distinctive or slightly bitter. Pores and tubes: dark purple brown to chocolate brown when young, becoming reddish brown at maturity; tubes 9–22 mm deep, concolorous or paler than the pores. Stalk: stout, nearly equal or sometimes enlarged downward, solid; surface dry, pale purplish gray, densely covered with tiny, darker purple-brown scales; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under conifers, especially hemlock, or in mixed woods with oak; summer and fall. Distribution and frequency: eastern Canada south to Georgia, west to Minnesota; occasional to locally common. Spore print: pinkish brown. Microscopic features: spores 11–17 × 3.5–5 µm, narrowly subfusoid, smooth, hyaline to pale brown. Edibility: not recommended. Although this bolete has been regarded as a safe edible in the past, there are reports that some collections from the Northeast have caused poisonings. Lookalikes: Tylopilus plumbeoviolaceus (p. 388) is similar, but it has a smooth stalk and intensely bitter-tasting flesh. Because of the fine scales on its stalk, the Lilac-brown Bolete might be mistaken for a species of Leccinum, a genus that differs by having a yellow-brown spore print. Gen us Ty l opi lu s P. K a r st. The genus Tylopilus was erected in 1881 to accommodate species formerly classified in two genera, Boletus and Suillus. The name Tylopilus means “bumpy or swollen cap.” Tylopilus species are medium to large terrestrial or, rarely, lignicolous boletes with solid stalks that are often reticulate at least near the apex. Caps

Tylopilus   ·  365

are dry and glabrous to somewhat velvety. Pores are most often white at first and typically become pinkish to pinkish brown or blackish at maturity. Spore-print colors range from pinkish to vinaceous, pinkish brown, reddish brown, purplish brown, or rusty brown.

Tylopilus alboater (A)

Tylopilus alboater (B)

366  ·  Boletes of Eastern North America

Tylopilus alboater (Schwein.) Murrill = Boletus alboater Schwein. = Porphyrellus alboater (Schwein.) E. J. Gilbert Common name(s): Black Velvet Bolete Overview: The term alboater means “white and black.” The cap surface stains amber orange with KOH and is negative with FeSO4 or NH 4OH. This bolete is easily overlooked because of its dark color. Although most species in the genus Tylopilus have bitter-tasting flesh, the Black Velvet Bolete is an exception. It has mild-tasting flesh and is popular with mushroom hunters. Cap: 3–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin often with a narrow band of sterile tissue; surface dry, somewhat velvety, occasionally finely cracked in age, black to dark grayish brown, often covered with a thin whitish bloom when young; flesh dense and firm on young specimens, white or tinged gray, staining pinkish to reddish gray when exposed, eventually blackening; odor and taste not distinctive. Pores and tubes: white or with a gray tinge when young, becoming dull pinkish brown in age, usually staining reddish and then slowly black when bruised; tubes 5–10 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, concolorous with the cap or paler, especially near the apex, often covered with a thin whitish bloom, not reticulate or only slightly so at the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary to scattered under broadleaf trees, especially oak; late spring, summer, and fall. Distribution and frequency: New England south to Florida, west to Minnesota and Missouri; occasional to locally fairly common. Spore print: pinkish brown. Microscopic features: spores 7–11 × 3.5–5 µm, narrowly oval, smooth, hyaline. Edibility: edible and good, with mild flavor and firm texture when young. Lookalikes: Compare with Tylopilus atronicotianus (p. 369), which is very similar but has a smooth, olive-brown cap and flesh that typically has a musty or disagreeable odor. Tylopilus atratus Both (not illustrated) is known only from western New York. It has a gray-brown to grayish-black cap with a cocoa-brown tinge and a cracked surface even when young. Its flesh is white and unchanging when exposed, and it grows under hemlock.

Tylopilus   ·  367

Tylopilus appalachiensis

Tylopilus appalachiensis Singer Common name(s): Appalachian Yellow-brown Bolete Overview: The term appalachiensis means “Appalachian,” a reference to the mountain range in North Carolina where this bolete was first collected. It is a dense and heavy bolete despite its fairly small stature. The cap surface stains orange with KOH or NH 4OH and olive green with FeSO4. The flesh is negative with KOH or NH 4OH and stains bluish gray with FeSO4. Cap: 2–9 cm wide, convex, becoming broadly convex to nearly plane at maturity, margin often with a narrow band of sterile tissue, sometimes upturned in age; surface dry, slightly velvety, sometimes cracked on older specimens, yellow brown over the disc, fading to pale yellow brown to dull tan on the margin or darker brown overall, at times with an orange or grayish tinge; flesh whitish to pale yellow, staining pinkish and then pale brown when exposed; odor not distinctive; taste bitter to intensely bitter, sometimes tardily. Pores and tubes: creamy white at first, becoming pinkish at maturity, staining brown when bruised, depressed at the stalk in age; pores often elongated and radially arranged near the stalk; tubes 3–10 mm deep. Stalk: typically tapered downward or swollen on the lower portion, solid; surface dry, finely velvety to nearly glabrous, usually lacking reticulation but

368  ·  Boletes of Eastern North America

sometimes finely reticulate near the apex, colored like the cap or paler; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups on the ground, often in sandy soil, sometimes along road banks or on mossy bases of stumps, under broadleaf trees, especially oaks; summer and fall. Distribution and frequency: eastern Canada south along the Appalachians to Georgia, distribution limits yet to be established; uncommon to occasional. Spore print: vinaceous pink. Microscopic features: spores 9–12 × 3–3.5 µm, narrowly ellipsoidal, smooth, hyaline. Edibility: unknown. Lookalikes: Xanthoconium montaltoense Wolfe (not illustrated) is similarly colored but with flesh that is unchanging when exposed, white pores that become cream to pale tan at maturity, and a yellow to yellow-ocher or brownish-yellow spore print.

Tylopilus atronicotianus (A)

Tylopilus   ·  369

Tylopilus atronicotianus (B)

Tylopilus atronicotianus Both Common name(s): False Black Velvet Bolete Overview: The term atronicotianus means “dark, like nicotine.” This bolete is sometimes collected in very dry weather when few other boletes are found. The cap surface stains blackish brown to reddish black with NH4OH and reddish brown to brown with KOH. Cap: 7.5–20 cm wide, hemispherical at first, becoming broadly convex to nearly plane in age, margin strongly inrolled when young, becoming decurved, with a narrow band of sterile tissue; surface dry, glabrous or nearly so, somewhat shiny, olive brown, bronze brown, grayish brown, or pale brownish, at times with yellowish tones, darker in age; flesh white, slowly staining pink to pinkish red or reddish vinaceous and then blackish when exposed; odor musty or disagreeable; taste not distinctive or somewhat unpleasant. Pores and tubes: white, slowly becoming vinaceous cinnamon to pale reddish brown, cocoa brown to dark reddish brown and depressed at the stalk in age, bruising dark pinkish cinnamon and then becoming nearly black; tubes 8–22 mm deep, bright brown, staining blackish when cut. Stalk: tapered in either direction, sometimes nearly equal or somewhat bulbous, usually with a whitish pointed root-like base, solid; surface dry,

370  ·  Boletes of Eastern North America

pruinose and delicately coated with blackish hairs, whitish at the apex, vinaceous gray to fuscous or dark brown below, becoming blackish near the base, sometimes finely reticulate only at the apex; flesh grayish to blackish. Habitat and season: solitary, scattered, or in groups under hemlock and oak or in mixed woods with oak present; summer and fall. Distribution and frequency: New England and New York south to North Carolina, distribution limits yet to be established; uncommon. Spore print: reddish brown. Microscopic features: spores 7.5–10.5 × 4–5 µm, narrowly oval, smooth, hyaline in KOH, pale yellow brown in Melzer’s. Edibility: unknown. Lookalikes: Compare with Tylopilus alboater (p. 366), which is very similar but has a blackish cap that lacks brown tones and flesh that lacks a distinctive odor.

Tylopilus badiceps (A)

Tylopilus   ·  371

Tylopilus badiceps (B)

Tylopilus badiceps (Peck) A. H. Sm. & Thiers = Boletus badiceps Peck Common name(s): Beveled-cap Bolete Overview: The partially folded to beveled or obliquely truncated cap margin, dark-brown to maroon cap, and the violaceous tones on the stalk are distinctive features of this attractive bolete. The cap surface stains amber brown with NH4OH. The stalk and white stalk base instantly stain blackish brown with NH4OH. The term badiceps means “reddish brown to maroon cap.” Cap: 4–8 cm wide, convex, becoming broadly convex and sometimes depressed at the center in age, margin even, typically beveled to folded or obliquely truncate in age; surface velvety and maroon when young and fresh, soon purplish brown to dark reddish brown, becoming duller and smoother with age; flesh white, unchanging or slowly staining pinkish to brown when exposed; odor sweet like molasses or not distinctive; taste not distinctive. Pores and tubes: white, becoming dingy white to brownish, not pinkish, at maturity; often staining brownish when bruised; tubes 7–12 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, colored like the cap but with more pronounced violaceous tones or sometimes violaceous

372  ·  Boletes of Eastern North America

nearly overall, usually white at the apex and base, at times forming obscure narrow reticulation at the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups in oak woods; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota, distribution limits yet to be determined; occasional. Spore print: pinkish brown. Microscopic features: spores 8–11 × 3.5–4.5 µm, narrowly ellipsoidal, smooth, hyaline to yellowish. Edibility: edible. Lookalikes: Tylopilus ferrugineus (p. 377) is very similar, but it has a reddish-brown cap, its stalk lacks violaceous tones, and the cap margin is not beveled.

Tylopilus balloui (A)

Tylopilus balloui (B)

Tylopilus balloui (C)

374  ·  Boletes of Eastern North America

Tylopilus balloui (Peck) Singer = Boletus balloui Peck = Boletus balloui var. fuscatus Corner = Gyrodon ballouii (Peck) Snell and Dick Common name(s): Burnt Orange Bolete Overview: Charles H. Peck (1833–1917) named this species in honor of American mycologist W. H. Ballou (1857–1937), who made several collections of this beautiful bolete. The epithet balloui is incorrectly spelled ballouii in most field guides. Tylopilus balloui has recently been transferred to Rubinoboletus balloui (Peck) Heinem. & Rammeloo, as listed in Index Fungorum. We have not accepted this transfer because of the conclusions reached by Todd Osmundson and Roy Halling (2010), which clearly indicate that this taxon is more closely aligned with species of Tylopilus rather than with species of Rubinoboletus. Cap: 5–12 cm wide, convex, becoming nearly plane in age, often irregular, margin even, incurved at first; surface dry, bright orange to bright orange red, fading to dull orange, cinnamon, or tan in age; flesh white, staining pinkish tan to violet brown when exposed; odor not distinctive; taste mild or bitter. Pores and tubes: white to dingy white, becoming tan or slightly pinkish in age, staining brown when bruised; tubes up to 8 mm deep. Stalk: equal or swollen on the lower portion, solid; surface smooth or scurfy, sometimes finely reticulate only at the apex, whitish or tinged yellow to orange, staining brownish when cut or bruised or in age; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups on lawns under trees or in woods, especially near oak, beech, and pine; summer and fall. Distribution and frequency: New England south to Florida, west to Ohio and Texas; occasional. Spore print: pale brown to reddish brown. Microscopic features: spores 5–11 × 3–5 µm, ellipsoidal, smooth, hyaline to pale brown. Edibility: edible but sometimes bitter. Lookalikes: Some species of Leccinum, including Leccinum insigne (p. 251) and Leccinum pseudoinsigne (p. 257), also have orange caps and white stalks, but they have scabers on their stalks.

Tylopilus   ·  375

Tylopilus felleus

Tylopilus felleus (Bull.) P. Karst. = Boletus felleus Bull. = Boletus felleus var. minor Coker & Beers = Tylopilus felleus var. minor (Coker & Beers) Pilát & Dermek = Tylopilus felleus var. uliginosus A. H. Sm. & Thiers Common name(s): Bitter Bolete Overview: The term felleus is derived from the Latin word fel, meaning “gall” or “bile,” relating to this common bolete’s bitter-tasting flesh. Cap: 5–30 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, margin even; surface dry or sometimes viscid when moist, glabrous, pinkish to reddish purple when young, soon becoming some shade of brown with or without purplish tints and finally brown to tan in age; flesh white, unchanging or staining reddish when exposed; odor not distinctive; taste very bitter and astringent. Pores and tubes: white when young, becoming pinkish, vinaceous, or pinkish tan in age, often staining brown when bruised; tubes 1–2 cm deep. Stalk: enlarged downward and typically bulbous, solid; surface dry, brown nearly overall, typically with a white apex and base, often developing olive or olive-brown stains when bruised or with age, with prominent brown reticulation, at least over the upper third; flesh concolorous with and staining like the cap flesh.

376  ·  Boletes of Eastern North America

Habitat and season: solitary or in groups on the ground or on decaying wood under conifers or in mixed woods; summer through winter. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: pinkish brown to reddish brown. Microscopic features: spores 11–17 × 3–5 µm, subfusoid, smooth, pale brown. Edibility: nonpoisonous but unpalatable for most owing to its very bitter-tasting flesh. Lookalikes: The Bitter Bolete can be mistaken for the delicious and popular King Bolete, Boletus edulis (p. 101), which has a fine, white reticulation on its stalk. Tylopilus indecisus (p. 380) is similar, but it has mild-tasting flesh and less-prominent reticulation on its stalk. Tylopilus variobrunneus (p. 400) is also similar, but it has white reticulation on the upper portion of its stalk, brown reticulation on the lower portion, and mild to slightly bitter-tasting flesh.

Tylopilus ferrugineus

Tylopilus   ·  377

Tylopilus ferrugineus (Frost) Singer = Boletus ferrugineus Frost = Tylopilus ferrugineus subsp. vinaceogriseus Snell, E. A. Dick & Hesler Common name(s): none. Overview: The term ferrugineus means “rust-colored,” in reference to the color of this bolete’s cap and stalk. NH4OH applied to the cap produces a dark-violet to vinaceous flash that becomes reddish brown to blackish brown. The cap stains dark brown to blackish brown with KOH and dark olive green with FeSO4. Cap: 4–12 cm wide, pulvinate, becoming broadly convex to nearly plane in age, margin incurved at first, often wavy or lobed when mature, sometimes with a narrow band of sterile tissue; surface dry, somewhat velvety, dark brown to reddish brown; flesh white, slowly staining pink and then brownish when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, becoming pale pinkish buff, staining dull brown when bruised, depressed at the stalk in age; tubes 6–12 mm deep. Stalk: nearly equal, solid; surface dry, glabrous or reticulate near the apex or on the upper half, dull brown to reddish brown, often whitish near the apex, with white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under oaks; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: pinkish brown. Microscopic features: spores 8–13 × 3–5 µm, subfusoid to ellipsoid, smooth, yellowish. Edibility: edible, if not bitter or astringent. Lookalikes: Tylopilus badiceps (p. 371) is very similar but has a maroon to purplish-brown or dark reddish-brown cap with a beveled to folded margin and violaceous tones on the stalk.

Tylopilus griseocarneus (A)

Tylopilus griseocarneus (B)

Tylopilus   ·  379

Tylopilus griseocarneus Wolfe & Halling Common name(s): none. Overview: The term griseocarneus means “gray flesh.” This bolete’s distinctive features are the dark reddish-brown, dark-gray, or blackish cap; grayish flesh that stains orangish to orange red and finally black when exposed; black pores that becomes gray in age; and a blackish-brown to gray stalk with prominent reticulation. The cap surface stains vinaceous black with KOH. The cap flesh stains vinaceous pink with KOH and is negative with NH4OH or FeSO4. The stalk flesh stains reddish orange with KOH and gray blue with FeSO4 and is negative with NH4OH. Cap: 4–11 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety when young, occasionally cracked in age, color variable, dull reddish brown to brown, dark olive brown, dark gray, brownish gray, or blackish; flesh grayish, staining orangish to orange red and finally black when exposed; odor and taste not distinctive. Pores and tubes: black when young, becoming gray in age, staining grayish orange or darker gray when mature, typically depressed at the stalk; tubes 3–11 mm deep. Stalk: nearly equal or tapered in either direction, solid; typically surface dry, blackish brown to gray, sometimes paler at the apex, prominently reticulate nearly overall or at least on the upper one-third, often with a dark pruinosity; flesh grayish, staining like the cap flesh. Habitat and season: scattered or in groups near or under oak or pine; summer and fall, also winter in the southern part of its range. Distribution and frequency: New England south to Florida, west to Texas, distribution limits yet to be established; uncommon. Spore print: pinkish to pinkish gray. Microscopic features: spores 8–14 × 3–5 µm, fusiform-ellipsoidal, smooth, hyaline. Edibility: not recommended, possibly poisonous; reported to cause severe vomiting and profuse perspiration. Lookalikes: Tylopilus alboater (p. 366) is similar, but it has white or whitish pores and lacks prominent reticulation on the stalk.

380  ·  Boletes of Eastern North America

Tylopilus indecisus

Tylopilus indecisus (Peck) Murrill = Boletus indecisus Peck = Boletus subpunctipes Peck = Tylopilus indecisus var. subpunctipes (Peck) Wolfe = Tylopilus subpunctipes (Peck) A. H. Sm. & Thiers Common name(s): none. Overview: The term indecisus means “undecided.” The cap surface stains dark blackish brown with NH4OH, instantly dark rusty brown to blackish with KOH, and dark gray with FeSO4. The flesh is negative with KOH, NH4OH, or FeSO4. Cap: 5–17 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety, ochraceous brown to pale brown or rarely reddish brown; flesh white, unchanging or slowly staining brownish or pinkish when exposed; odor and taste not distinctive. Pores and tubes: white, becoming pinkish or brownish with age, staining brown when bruised; tubes 6–12 mm deep. Stalk: enlarged downward when young, nearly equal in age, solid; surface dry, whitish when young, becoming pale brown on the lower portion at maturity, staining brown when bruised, reticulate toward the apex; flesh white, staining like the cap flesh.

Tylopilus   ·  381

Habitat and season: scattered or in groups near oak or pine, also in mixed broadleaf woods; summer and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota; occasional. Spore print: pinkish brown to reddish brown. Microscopic features: spores 10–15 × 3–5 µm, narrowly subfusiform, smooth, pale brown. Edibility: edible. Lookalikes: Tylopilus badiceps (p. 371) is similar but has a dark-brown to maroon cap with a beveled to folded margin and a darker stalk with pale to distinctly violaceous tones. Tylopilus ferrugineus (p. 377) is also similar, but it has a darker reddish-brown cap and stalk. Also compare with Tylopilus tabacinus (p. 398), which typically has a larger cap, often grows in grassy areas, and has different macrochemical test reactions.

Tylopilus intermedius (A)

382  ·  Boletes of Eastern North America

Tylopilus intermedius (B)

Tylopilus intermedius A. H. Sm. & Thiers Common name(s): Parchment Bitter Bolete Overview: The term intermedius means “intermediate between two species,” in this case referring to Tylopilus peralbidus (p. 386) and Tylopilus rhoadsiae (p. 391), both of which occur in the southeastern United States. The Parchment Bitter Bolete often typically appears soiled or dirty because of the brown stains that develop on the stalk. Cap: 6–15 cm wide, broadly convex, becoming nearly plane in age, margin incurved when young, with a narrow band of sterile tissue; surface uneven, often wrinkled like parchment, occasionally pruinose, white or whitish when young, occasionally with a pinkish tinge or sometimes ochraceous to tan, developing brownish stains with age; flesh white, firm, very slowly staining dark brown when exposed or where there has been insect damage; odor somewhat pungent or not distinctive; taste bitter. Pores and tubes: white when young, becoming pale tan to pinkish in age, slowly staining brownish when bruised, depressed at the stalk when mature; pores minute, almost appearing like a solid surface on young specimens; tubes 1–1.5 cm deep. Stalk: enlarged to club-shaped or bulbous, solid; surface dry, white or whitish, developing dingy-brownish or yellow-brown stains in age or when handled, weakly to distinctly reticulate (use hand lens); flesh white, staining like the cap flesh.

Tylopilus   ·  383

Habitat and season: scattered or in groups under broadleaf trees, especially oak, also reported under pine; summer and fall. Distribution and frequency: New England south to North Carolina, west to Minnesota; infrequent. Spore print: pinkish to pinkish brown. Microscopic features: spores 10–15 × 3–5 µm, nearly oblong, smooth, hyaline to pale brown. Edibility: inedible because of the bitter taste. Lookalikes: Tylopilus peralbidus (p. 386) is similarly colored but lacks reticulation on the stalk. Tylopilus rhoadsiae (p. 391) is also similarly colored, but its flesh does not stain when exposed, and it has prominent, coarse reticulation on the stalk. Boletus pallidus (p. 131) is somewhat similar, but it has whitish to yellow pores that become greenish yellow in age, a faintly or distinctly reticulate stalk, and mild to slightly bitter-tasting flesh.

Tylopilus minor (A)

384  ·  Boletes of Eastern North America

Tylopilus minor (B)

Tylopilus minor Singer = Boletus minor (Singer) Murrill Common name(s): Little Bitter Bolete Overview: The term minor means “lesser or smaller,” a reference to this bolete’s stature. The typically small, brown cap, bitter-tasting flesh, and finely to somewhat prominently reticulate stalk with white basal mycelium are reliable field features. Cap: typically 3–8 cm wide, sometimes up to 15.5 cm wide, pulvinate, becoming broadly convex to nearly plane in age, margin even; surface dry, glabrous to finely velvety, pale brown to brown or sometimes dull whitish, often with pinkish tones; flesh white, unchanging when exposed; odor not distinctive; taste bitter. Pores and tubes: white at first, becoming pale brownish pink and often depressed at the stalk in age, not staining when bruised; tubes 4–7(18) mm deep. Stalk: nearly equal or slightly enlarged downward, usually with an abruptly narrowed base, solid; surface dry, with pale to brownish reticulation near the apex or extending to the midportion or sometimes lacking, white to whitish at first, becoming brownish to cinnamon, apex and base whitish, basal mycelium white; flesh white. Habitat and season: solitary, scattered, or in groups in mixed woods or broadleaf forests with oak; late spring, summer, and fall.

Tylopilus   ·  385

Distribution and frequency: New England and New York south to Florida, west to Minnesota and Texas; uncommon. Spore print: pinkish brown. Microscopic features: spores 9–15 × 3–5 µm, fusoid, smooth, yellowish to hyaline. Edibility: inedible because of the bitter taste. Lookalikes: Tylopilus felleus (p. 375) is typically a larger species with coarse, brown reticulation on the stalk.

Tylopilus peralbidus (A)

Tylopilus peralbidus (B)

386  ·  Boletes of Eastern North America

Tylopilus peralbidus (Snell & Beardslee) Murrill = Boletus peralbidus Snell & Beardslee Common name(s): none. Overview: The term peralbidus means “whitish throughout,” a reference to the color of all parts of this bolete, at least when young. The cap surface stains yellow with KOH and dark blue green with FeSO4 and is negative with NH4OH. The flesh immediately stains dark bluish gray with FeSO4 and pale yellow with KOH or NH4OH. Cap: 4.5–13 cm wide, convex, becoming broadly convex to nearly plane in age, often depressed over the disc, margin even; surface dry, glabrous to slightly velvety, finely cracked in age, white when young, soon becoming ochraceous tan to chamois and finally brownish, staining cinnamon to brown when bruised; flesh white, sometimes staining pale pinkish brown or buff when exposed, often slowly; odor bleach-like or unpleasant; taste bitter. Pores and tubes: whitish when young, becoming buff or tinged pinkish in age, staining brown when bruised, depressed at the stalk in age; tubes 6–10 mm deep. Stalk: equal or tapered in either direction, often with a pointed base, solid; surface dry, smooth, white to brownish or concolorous with the cap, staining brown when bruised or with age; flesh white, staining like the cap flesh. Habitat and season: solitary, scattered, or in groups on shaded lawns, along roads, or in woodlands with oaks or sometimes pines; late spring, summer, and fall. Distribution and frequency: Maryland south to Florida, west to Texas; occasional to locally fairly common. Spore print: pinkish brown. Microscopic features: spores 7–12 × 2.3–3.5 µm, cylindric or cylindric-subclavate, smooth, pale honey yellow. Edibility: unknown. Lookalikes: Tylopilus intermedius (p. 382) is similar, but it has a reticulate stalk. Tylopilus rhoadsiae (p. 391) is also similar, but its pores do not stain when bruised, and its stalk has prominent, coarse reticulation at least over the upper half. Tylopilus rhodoconius (p. 392) has a brownish cap and stalk that stain darker brown when handled and whitish flesh with hyaline marbling.

Tylopilus plumbeoviolaceus (A)

Tylopilus plumbeoviolaceus (B)

388  ·  Boletes of Eastern North America

Tylopilus plumbeoviolaceus (Snell & Dick) Singer = Boletus plumbeoviolaceus Snell & E. A. Dick Common name(s): Violet-gray Bolete Overview: The term plumbeoviolaceus means “grayish violaceous,” in reference to this handsome bolete’s overall color. The cap surface stains orange to dull amber with KOH and pale brownish amber with NH4OH or FeSO4. This medium to large bolete is one of several in the genus Tylopilus that have flesh that tastes exceedingly bitter. The combination of a brownish to purplish-brown cap, a purple stalk, and bitter flesh makes this bolete easy to recognize. Cap: 4–15 cm wide, convex to broadly convex with a narrow sterile margin, becoming nearly plane in age; surface glabrous, smooth, shiny, at times becoming cracked, brownish to grayish brown or dull cinnamon, at times tinged with purplish areas, especially toward the margin, or entirely purple when young; flesh firm when fresh, white, not staining when cut or bruised, insect-damaged areas becoming brown; odor slightly pungent or not distinctive; taste very bitter. Pores and tubes: dull white when young, becoming very pale brownish to pinkish tan with age, not staining when bruised or just barely bruising very lightly pinkish brown, depressed at the stalk at maturity; pores minute; tubes 4–18 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, purple with white mycelium at the base when young, at times becoming grayish purple or purplish brown, smooth and glabrous except for a narrow zone of reticulation at the apex; flesh concolorous with the cap flesh, insect-damaged areas especially becoming brown. Habitat and season: solitary or in small groups in broadleaf woods with oak; early summer and early fall. Distribution and frequency: widely distributed; fairly common. Spore print: purplish brown. Microscopic features: spores 10–13 × 3–4 µm, ellipsoidal, smooth, pale brown. Edibility: inedible due to strongly bitter flesh. Lookalikes: Tylopilus plumbeoviolaceus is not likely to be confused with any bolete other than the equally bitter Tylopilus violatinctus (p. 402), which differs by having a paler grayish-lilac to pinkish-lilac cap, a pale-lilac stalk that is white at the apex, and smaller spores.

Tylopilus   ·  389

Tylopilus porphyrosporus

Tylopilus porphyrosporus (Fr. & Hök) A. H. Sm. & Thiers = Boletus pseudoscaber Secr. ex Singer = Porphyrellus porphyrosporus (Fr. & Hök) E. J. Gilbert = Porphyrellus pseudoscaber (Secr.) Singer = Tylopilus nebulosus (Peck) Wolfe = Tylopilus pseudoscaber (Secr.) A. H. Sm. & Thiers Common name(s): Dark Bolete Overview: The term porphyrosporus means “purple-red spores.” The cap surface stains mahogany red with KOH or NH4OH and is negative with FeSO4. The flesh stains dull orange with KOH and green with FeSO4 and is negative with NH4OH. Much confusion and contradiction occur in the literature concerning this species. Some authors have used the presence or absence of a blue staining reaction of the flesh and pores or growth with conifers rather than with broadleaf trees as a basis for naming different species. Specimens wrapped in waxed paper for an hour or so sometimes stain the paper blue. Cap: 5–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety, sometimes finely to conspicuously cracked in age, dark brown to olive brown or dark vinaceous brown, sometimes staining darker brown when bruised, usually paler at the margin; flesh white, sometimes rapidly or slowly staining blue when exposed

390  ·  Boletes of Eastern North America

and then staining reddish brown and finally grayish brown to grayish black; odor pungent, clove-like or resembling coal tar or burned rubber or unpleasant; taste weakly bitter to unpleasant or not distinctive. Pores and tubes: pinkish brown to dark reddish brown or dark brown, depressed at the stalk in age, bruising dark reddish brown, sometimes with dark greenish-blue stains; tubes 1.3–2 cm deep, concolorous with the pores or paler, typically staining blue when bruised. Stalk: enlarged downward or nearly equal; surface dry, scurfy-punctate, concolorous with the cap, often with a whitish base, with or without reticulation at the very apex; flesh concolorous with and staining like the cap flesh, sometimes with a strong odor of chlorine in the base. Habitat and season: solitary, scattered, or rarely in groups in broadleaf woods, conifer forests, or mixed woods, sometimes on well-decayed logs and stumps; summer and fall, also winter in the southern part of its range. Distribution and frequency: widely distributed throughout eastern North America; uncommon to occasional. Spore print: dark reddish brown. Microscopic features: spores 12–18 × 6–7.5 µm, ellipsoidal, smooth, pale brown. Edibility: unknown. Lookalikes: Tylopilus sordidus (p. 396) has a similarly colored cap and stalk, but its pores are whitish to grayish buff when young, and it has smaller spores and different macrochemical test reactions.

Tylopilus rhoadsiae (A)

Tylopilus   ·  391

Tylopilus rhoadsiae (B)

Tylopilus rhoadsiae (Murrill) Murrill = Boletus rhoadsiae (Murrill) Murrill = Gyroporus rhoadsiae Murrill Common name(s): Pale Bitter Bolete Overview: The white to whitish cap; white, unchanging, and bitter flesh; white pores that become dull pinkish in age; and a white to whitish, prominently reticulate stalk are this distinctive bolete’s key identifying features. The cap surface stains pale yellow with KOH or NH4OH and gray with FeSO4. Cap: 6–15 cm wide, convex, becoming broadly convex to nearly plane in age, margin even; surface dry, somewhat velvety to glabrous, sometimes shiny, white to whitish, often with buff, grayish-buff, pinkish, or pinkish-tan tinges or sometimes becoming tan, pinkish brown, or golden brown overall in age; flesh white, unchanging when exposed; odor not distinctive; taste bitter. Pores and tubes: white when young, becoming dull pinkish in age, unchanging when bruised, depressed at the stalk in age; tubes 9–20 mm deep. Stalk: nearly equal or enlarged above a pinched base, solid, basal mycelium white; surface dry, white or concolorous with the cap, prominently reticulate nearly overall or at least on the upper half; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups near or under pines or oaks; summer and fall, also winter in the southern part of its range. Distribution and frequency: New England south along the coastal plain to Florida, west to Texas; fairly common along the Gulf Coast and increasingly uncommon northward.

392  ·  Boletes of Eastern North America

Spore print: pinkish to brownish vinaceous. Microscopic features: spores 11–13.5 × 3.5–4.5 µm, oblong-ellipsoidal, smooth, hyaline to pale yellow. Edibility: unpalatable because of the bitter taste. Lookalikes: Tylopilus intermedius (p. 382) is similarly colored, but its pores slowly stain brownish when bruised, and it has finer reticulation on its stalk. Tylopilus peralbidus (p. 386) is also similarly colored, but its stalk lacks reticulation. Tylopilus rhodoconius (p. 392) has a darker-brown cap and a stalk that stains brown when handled, and its flesh is white to creamy white with hyaline marbling.

Tylopilus rhodoconius

Tylopilus rhodoconius T. J. Baroni, Both & Bessette = Tylopilus peralbidus var. rhodoconius Singer Common name(s): none. Overview: The white to creamy-white, soft flesh with hyaline marbling is distinctive. The cap surface slowly stains pale yellow brown with KOH or NH4OH and bluish green to bluish gray with FeSO4. Cap: 4–14 cm wide, hemispheric, becoming broadly convex to nearly plane in age, margin incurved at first and remaining so well into maturity, with a narrow band of sterile tissue; surface dry, finely velvety, delicately cracked at maturity, pale ochraceous to brownish orange at first, becoming pale brown and then darker brown with a paler margin, staining dark brown when handled or bruised; flesh firm when young, soft at maturity, white

Tylopilus   ·  393

to pale creamy white with hyaline marbling, slowly staining pale pinkish and then dingy pinkish red when exposed; odor faintly of chlorine or not distinctive; taste mild at first and then very bitter. Pores and tubes: white to whitish when young, becoming pale pinkish brown in age, staining dull brown and then darker brown when bruised; tubes 6–10 mm deep. Stalk: nearly equal or enlarged downward, tapered to a point at the base, often with a short, root-like projection, solid; surface dry, nearly glabrous, white near the apex, gradually yellow brown to reddish brown downward, or concolorous with the cap, staining brown to dark brown when handled, with white basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under oak, pine, or in mixed oak, beech, and pine woods; summer and fall. Distribution and frequency: Cape Cod, Massachusetts, and Florida, distribution limits yet to be established; rare to uncommon. Spore print: pink to reddish gray. Microscopic features: spores 8.4–13 × 2–3.5 µm, cylindric, smooth, hyaline. Edibility: unknown. Lookalikes: Tylopilus intermedius (p. 382) has a paler cap and stalk, white flesh that lacks hyaline marbling, and a reticulate stalk. Tylopilus peralbidus (p. 386) has a paler cap and stalk and white flesh that lacks hyaline marbling. Tylopilus rhoadsiae (p. 391) has a paler cap and stalk and white flesh that lacks hyaline marbling, and its stalk has prominent reticulation.

Tylopilus rubrobrunneus (A)

Tylopilus rubrobrunneus (B)

Tylopilus rubrobrunneus (C)

Tylopilus   ·  395

Tylopilus rubrobrunneus Mazzer & A. H. Sm. Common name(s): Reddish-brown Bitter Bolete Overview: The term rubrobrunneus means “reddish brown.” During wet periods, this species is commonly attacked by a white mold. The purple areas of the cap surface bleach pinkish brown with KOH or NH4OH and are negative with FeSO4. Cap: 8–30 cm wide, broadly convex, becoming nearly plane and occasionally slightly depressed in age, margin even, incurved to inrolled at first; surface dry, glabrous or nearly so, sometimes becoming cracked in age, dark to bright purple when young, becoming purple brown, dark reddish brown, dull brown, or cinnamon in age; flesh white, unchanging or slowly staining brown when cut or bruised; odor not distinctive; taste very bitter. Pores and tubes: whitish to very pale brownish at first, becoming dingy pinkish brown at maturity, staining brown when bruised; tubes 8–20 mm deep. Stalk: nearly equal or enlarged downward, solid; surface dry, white to brown, developing olive or olive-brown stains from the base upward as specimens mature or overall when handled, glabrous or with fine reticulation at the apex; flesh white, staining like the cap flesh. Habitat and season: scattered, in groups, or caespitose in broadleaf woods with beech or oak, also in mixed oak and pine woods or with hemlock; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; fairly common. Spore print: reddish brown to dull pinkish brown. Microscopic features: spores 10–14 × 3–4.5 µm, suboblong to nearly fusoid, smooth, pale brown. Edibility: unpalatable because of the bitter taste. Lookalikes: The equally bitter-tasting Tylopilus violatinctus (p. 402) is similar but has a pale-purplish to grayish-violet or pale-brown cap, and it does not develop olive-brown stains on the stalk. Tylopilus felleus (p. 375) has less-pronounced purplish tones on the cap. Its stalk has more prominent reticulation and lacks olive stains.

396  ·  Boletes of Eastern North America

Tylopilus sordidus

Tylopilus sordidus (Frost) A. H. Sm. & Thiers = Boletus sordidus Frost = Porphyrellus cyaneotinctus (A. H. Sm. & Thiers) Singer = Porphyrellus fumosipes (Peck) Snell = Porphyrellus pseudoscaber ssp. cyaneotinctus Singer = Porphyrellus sordidus (Frost) Snell = Porphyrellus umbrosus (G. F. Atk.) Singer, J. Garcia & L. D. Gómez = Tylopilus cyaneotinctus A. H. Sm. & Thiers = Tylopilus fumosipes (Peck) A. H. Sm. & Thiers = Tylopilus umbrosus (G. F. Atk.) A. H. Sm. & Thiers Common name(s): none. Overview: This small- to medium-size dark bolete is recognizable by its typically cracked cap and dramatic bluish-green bruising of the pores and cap flesh. The term sordidus means “dirty or smoky,” a reference to this bolete’s overall colors. When its flesh is wrapped in waxed paper for several minutes, it stains the paper a dark blue-green color. Cap: 4.5–13 cm wide, convex, becoming broadly convex to nearly plane in age, margin with a band of sterile tissue when young, becoming even at maturity; surface dry, glabrous to somewhat velvety, becoming cracked in age, gray brown to olive brown or dark brown, often with dark-greenish

Tylopilus   ·  397

or bluish tints along the margin; flesh whitish, staining blue green when exposed, sometimes with reddish tints; odor slightly pungent or not distinctive; taste unpleasant or not distinctive. Pores and tubes: whitish to grayish buff when young, becoming pinkish brown and then reddish brown to yellow brown, staining dark blue to dark blue green and then dark brownish red when bruised; tubes 6–20 mm deep. Stalk: slightly enlarged downward or nearly equal, solid; surface dry, minutely scurfy, brownish with much darker longitudinal streaks, typically paler toward the upper portion and whitish at the base, with greenish or bluish-green tints near the apex; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees, especially oak, and under various conifers; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: reddish brown. Microscopic features: spores 10–14 × 4–6 µm, subellipsoidal, smooth, pale brown. Edibility: unknown. Lookalikes: Tylopilus porphyrosporus (p. 389) has a similarly colored cap and stalk, but its pores are dark reddish brown to dark brown or blackish brown in all stages, and it has larger spores.

Tylopilus tabacinus (A)

398  ·  Boletes of Eastern North America

Tylopilus tabacinus (B)

Tylopilus tabacinus (Peck) Singer = Boletus tabacinus Peck = Ceriomyces tabacinus (Peck) Murrill = Tylopilus tabacinus var. amarus Singer = Tylopilus tabacinus var. dubius Singer Common name(s): none. Overview: The term tabacinus means “color of tobacco.” This bolete’s cap surface stains rusty brown to dark maroon with KOH, vinaceous with NH4OH, and pale olive gray with FeSO4. Specimens with bitter-tasting flesh were previously identified as var. amarus. Cap: 4.5–18 cm wide, obtuse when young, becoming broadly convex to nearly plane in age, margin even and wavy; surface dry, smooth, slightly velvety, becoming finely cracked in age, yellow brown to orange brown or tobacco brown; flesh white, usually slowly staining purplish buff or pinkish buff when exposed, often brown at maturity; odor variously described as fruity, fishy, pungent, or not distinctive; taste slightly bitter or not distinctive. Pores and tubes: whitish or sometimes brown at first, becoming brown to yellow brown, with darker-brown patches and stains, usually depressed at the stalk in age; tubes 1–1.5 cm deep.

Tylopilus   ·  399

Stalk: enlarged to bulbous when young, becoming nearly equal in age, solid; surface dry, concolorous with the cap, prominently reticulate at least over the upper portion, usually glabrous on the lower portion; flesh concolorous with the cap flesh or sometimes burgundy brown near the base, staining like the cap flesh. Habitat and season: scattered or in groups in woods and at their edges or around trees in lawns, usually with oak and pine; summer and fall. Distribution and frequency: New England south along the coastal plain to Florida, west to Texas; occasional to locally fairly common. Spore print: pinkish brown to reddish brown. Microscopic features: spores (10)12–14(17) × 3.5–4.5 µm, fusoid to ellipsoidal, smooth, hyaline to pale honey yellow; pleurocystidia 45–60 × 6.5–12.5 µm, lanceolate to narrowly fusoid-ventricose; cheilocystidia absent. Edibility: unknown. Lookalikes: Tylopilus indecisus (p. 380) usually has a smaller cap, typically occurs in woodlands, and has different macrochemical test reactions. Tylopilus felleus (p. 375) is similar, but its pores are white to brown, not pinkish to vinaceous, and it has very bitter-tasting flesh. Boletus durhamensis (p. 99) has an olivaceous gray-brown spore print, finer reticulation on the upper portion of the stalk or at least near the apex, mild-tasting flesh, and cheilocystidia.

Tylopilus variobrunneus (A)

400  ·  Boletes of Eastern North America

Tylopilus variobrunneus (B)

Tylopilus variobrunneus Roody, A. R. Bessette & Bessette Common name(s): Variable Brown-net Bolete Overview: The term variobrunneus means “various shades of brown,” a reference to the color of this bolete’s cap, stalk, and staining reactions of the pores and flesh. When this handsome bolete is very young, it has a compact stature and firm flesh, and the stalk and reticulation can be totally white, although brown coloration is typical. The cap surface stains grayish green with FeSO4 and reddish brown to orange brown with KOH or NH4OH. The flesh stains grayish green with FeSO4 and is negative with KOH or NH4OH. Cap: 4–12 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, with a narrow band of sterile tissue when young, becoming decurved and even at maturity; surface dry, somewhat velvety and dark olive brown to greenish brown or blackish brown when young, becoming nearly glabrous and dull medium brown to chestnut brown at maturity, remaining darker over the disc and fading to pale chestnut brown in age; flesh dull white, slowly staining dull pinkish rose to pale brownish pink when exposed, insect-damaged areas staining dark brown; odor not distinctive; taste mild or slightly bitter. Pores and tubes: dull white to creamy white at first, becoming yellow brown to brownish pink, depressed at the stalk in age, staining brown to brownish rose or pinkish cinnamon, sometimes slowly, when bruised; tubes 5–13 mm deep.

Tylopilus   ·  401

Stalk: enlarged downward or nearly equal, usually with a pinched base, solid; surface dry, dull white on the upper portion, becoming pale brown on the lower portion, dark brown toward the base, basal mycelium white, with prominent reticulation over the upper two-thirds or at least over the upper half; reticulation white near the apex when young, brown below; flesh concolorous with and staining like the cap flesh, blackish brown around larval tunnels. Habitat and season: solitary, scattered, or in groups under oaks or in mixed oak–pine woods; summer and fall. Distribution and frequency: New England south to Florida, west to Minnesota and Mississippi; occasional to locally fairly common. Spore print: dull pinkish brown to cocoa powder brown. Microscopic features: spores 9–13 × 3–4.5 µm, subfusiform to subellipsoidal, smooth, pale ochraceous. Edibility: edible but with an unpleasant aftertaste. Lookalikes: Tylopilus variobrunneus is sometimes confused with Tylopilus felleus (p. 375), which has very bitter-tasting flesh. Tylopilus badiceps (p. 371) is similar but has a reddish-brown to maroon cap, typically with a beveled margin; a darker-brown stalk that usually lacks reticulation; and pores that remain white for a long time and then become dingy white to brownish, not pinkish, at maturity.

Tylopilus violatinctus (A)

402  ·  Boletes of Eastern North America

Tylopilus violatinctus (B)

Tylopilus violatinctus T. J. Baroni & Both Common name(s): Pale Violet Bitter Bolete Overview: The term violatinctus means “tinged violet.” The cap surface stains yellowish to brownish yellow with NH4OH and dingy pale yellowish to amber with KOH and is negative with FeSO4. The flesh stains yellow with FeSO4 and then quickly develops a blue ring around the yellow. Cap: 7.5–14 cm wide, hemispheric, becoming broadly convex to nearly plane in age, margin strongly incurved at first, with a narrow band of sterile tissue; surface dry, somewhat velvety when young, becoming nearly glabrous when mature, sometimes cracked in age, grayish violet to bluish violet when young, becoming pale purplish, purplish pink, or tan to dull brown in age, staining rusty violet to dark violet when bruised; flesh white, unchanging or becoming very pale slate-colored when exposed; odor pungent or not distinctive; taste intensely bitter. Pores and tubes: white to dull pink at first, becoming brownish, unchanging when bruised, typically depressed at the stalk in age; tubes up to 1.5 cm deep. Stalk: enlarged downward or nearly equal, sometimes with a bulbous base, solid; surface dry, nearly glabrous, concolorous with the cap or paler when young, becoming brown in age with a white apex and base, at times

Tylopilus   ·  403

staining dull yellow to yellow brown when bruised, sometimes with inconspicuous reticulation at the apex; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under oak and pine, also with beech, spruce, or hemlock; late spring, summer, and fall. Distribution and frequency: New England and New York south to Florida, west to Minnesota and Mississippi; occasional. Spore print: reddish brown. Microscopic features: spores 7–10 × 3–4 µm, subfusiform, smooth, pale yellow. Edibility: unknown but too bitter to be of interest. Lookalikes: Tylopilus plumbeoviolaceus (p. 388) is similar but has a dark-purple stalk. Tylopilus rubrobrunneus (p. 395) is typically more robust, has a purple-brown to vinaceous-brown cap that fades to pinkish cinnamon in age, and develops olive-brown stains on its stalk, especially when handled.

Tylopilus williamsii (A)

Tylopilus williamsii (B)

Tylopilus williamsii (C)

Veloporphyrellus   ·  405

Tylopilus williamsii Singer & J. Garcia Common name(s): none. Overview: This bolete was named in honor of Florida mycologist Robert S. Williams (1939–2014), who first collected and described it. The dark magenta to purple-violet or brownish-purple cap, bitter flesh, and white to pinkish pores that bruise yellowish are its reliable field features. The cap surface stains orange yellow with KOH and orange with NH4OH. The flesh stains greenish with FeSO4. Cap: 4–10 cm wide, hemispheric when young, becoming broadly convex to nearly plane in age, margin even, incurved at first; surface dry, somewhat velvety, often uneven in age, dark magenta to purple violet or brownish purple when young, fading to pinkish gray to pale yellow brown; flesh white, unchanging when exposed; odor not distinctive; taste bitter. Pores and tubes: white at first, becoming pale pinkish at maturity, staining yellowish to yellow brown when bruised; tubes 1–1.5 cm deep. Stalk: nearly equal overall, solid; surface dry, glabrous, pale yellow to grayish, often streaked with beige and grayish orange; flesh concolorous with the cap flesh. Habitat and season: scattered or in groups under oaks; summer and fall. Distribution and frequency: southeastern coastal Georgia and Florida, west to Mississippi, distribution limits yet to be established; rare to uncommon. Spore print: pink. Microscopic features: spores 7.5–11 × 3.5–4.5 µm, fusoid to ellipsoid, smooth, hyaline to pale yellow. Edibility: inedible because of the bitter taste. Lookalikes: Tylopilus plumbeoviolaceus (p. 388) is similar, but it has a violet or purple stalk, which is sometimes reticulate at the apex. Tylopilus violatinctus (p. 402) has a paler and more lilac cap that bruises violet, a concolorous stalk with a white apex, and a base that typically stains yellowish when bruised. Gen us Ve l op or ph y r e l lu s L . D. G óm ez & Si nger The genus Veloporphyrellus was erected in 1984 to accommodate species formerly classified in two genera, Boletus and Tylopilus. Veloporphyrellus is a genus known from North and Central America, Africa, and southeastern Asia. Only one species is currently known to occur in eastern North America. The distinguishing features of the genus are the pinkish to grayish-pink pores, the membranous veil remnants that hang on the cap margin, a trichodermium-like cuticle, and smooth spores. These boletes have caps that are typically dry and shaggy or scaly, with soft-textured white flesh. The pores are white at first and become grayish pink to pinkish brown at maturity.

Veloporphyrellus conicus (A)

Veloporphyrellus conicus (B)

Xanthoconium   ·  407

Veloporphyrellus conicus (Ravenel) B. Ortiz, Yan C. Li & Zhu L. Yang = Fistulinella conica var. conica (Ravenel) Pegler & T. W. K. Young = Mucilopilus conicus (Ravenel) Wolfe = Tylopilus conicus (Ravenel) Beardslee Common name(s): Conical Shaggy-capped Bolete Overview: The Conical Shaggy-capped Bolete is an easy-to-recognize southern species. It is distinctive for its bluntly conical yellowish floccose-shaggy cap and smooth, slender stalk. The term conicus means “cone-shaped,” a reference to the shape of the young cap. Cap: 2.5–9.5 cm wide, bluntly conical when young, becoming convex in age; margin fringed with thin flaps of membranous veil tissue, at least when young; surface dry, shaggy or scaly when young and often developing a network of ridges and small depressions in age, pinkish tan to golden yellow, yellow brown or salmon-tinged, with white flesh showing between the scales or beneath the ridges; flesh white, not staining when cut; odor fruity or not distinctive; taste not distinctive. Pores and tubes: white when young, becoming grayish pink to pinkish brown at maturity; tubes 8–14 mm deep. Stalk: slender, nearly equal, often curved, solid; surface dry, smooth or minutely wrinkled, rarely reticulate, white or yellow, sometimes with pinkish tones especially toward the midportion; flesh concolorous with the cap flesh. Habitat and season: solitary or scattered under pine and in mixed woods, often in bottomlands along streams; summer and fall. Distribution and frequency: North Carolina south to Florida, western limits yet to be established; occasional to fairly common. Spore print: pinkish brown to reddish brown. Microscopic features: spores 14–21 × 4–6 µm, elongate-fusoid, smooth, hyaline to honey yellow, pseudoamyloid in Melzer’s. Edibility: edible. Lookalikes: Fistulinella conica var. reticulata (Wolfe) Singer (not illustrated), described from Florida, is nearly identical but has prominent reticulation on its stalk. Gen us X a n t ho con i u m Si nger The genus Xanthoconium was erected in 1944 to accommodate species that produce a yellowish to ochraceous, bright yellow-brown, or bright rusty-brown spore deposit, formerly classified in the genus Boletus. The name Xanthoconium means “yellow cone,” a reference to the spore color and shape. It is a small genus, with only three species that occur in eastern North America. They have white to yellow pores that do not bruise blue and flesh that does not stain when exposed. Some species have reticulation on the stalk.

Xanthoconium affine (A)

Xanthoconium affine (B)

Xanthoconium   ·  409

Xanthoconium affine (C)

Xanthoconium affine (Peck) Singer = Boletus affinis Peck = Xanthoconium affine var. affine (Peck) Singer = Xanthoconium affine var. maculosus (Peck) Singer = Xanthoconium affine var. reticulatum (A. H. Sm.) Wolfe Common name(s): none. Overview: The term affine means “related,” referring to a similarity with Boletus edulis (p. 101). The cap surface stains reddish brown with NH4OH and is negative with KOH or NH4OH. The key identification features include

410  ·  Boletes of Eastern North America

the brown cap that sometimes has pale-yellow spots, white flesh that does not stain when exposed, white to yellow-brown pores, and a whitish stalk with brownish streaks and with or without reticulation. Cap: 4–11.5 cm wide, convex to broadly convex, becoming nearly plane in age, margin even; surface dry, somewhat velvety, smooth or wrinkled, dark brown to chestnut brown or ocher brown, sometimes with white to pale-yellow spots or patches; flesh white, unchanging when exposed; odor and taste not distinctive. Pores and tubes: white at first, becoming yellowish to dingy yellow brown, bruising dull yellow to brownish, depressed at the stalk in age; tubes 6–20 mm deep. Stalk: nearly equal or tapered in either direction, solid; surface smooth or with sparse to conspicuous brown reticulation, dry, whitish, often with brownish streaks on the middle portion or base; flesh concolorous with the cap flesh. Habitat and season: usually in groups, sometimes in caespitose clusters, or scattered in broadleaf woods, especially with oak or beech, also under pines; spring, summer, and fall. Distribution and frequency: widely distributed throughout eastern North America; common. Spore print: bright yellow brown to rusty ochraceous. Microscopic features: spores (9)12–16 × 3–5 µm, narrowly ventricose to nearly cylindrical, smooth, yellowish. Edibility: edible and good when young. Lookalikes: Xanthoconium purpureum (p. 412) is very similar, except it has a purplish-red to maroon cap. Xanthoconium chattoogaense Wolfe (not illustrated) is known only from the Chattooga River region in the Nantahala National Forest of North Carolina. This bolete grows in mixed woods with oaks, red maple, and rhododendron. It has a somewhat waxy, golden-brown cap; white flesh with a disagreeable odor that is reminiscent of rotting meat; and a thick, obclavate, yellowish-white stalk with a brownish apex.

Xanthoconium purpureum (A)

Xanthoconium purpureum (B)

412  ·  Boletes of Eastern North America

Xanthoconium purpureum Snell & E. A. Dick = Boletus purpureofuscus A. H. Smith Common name(s): none. Overview: The term purpureum means “purplish,” a reference to this bolete’s cap colors. The cap surface stains greenish blue with NH4OH and is negative with KOH or FeSO4. The flesh is negative with NH4OH, KOH, or FeSO4. Cap: 3–11.5 cm wide, rounded to convex, becoming broadly convex to nearly plane in age, margin even, incurved at first; surface dry, glabrous, sometimes wrinkled, dark red to purple red or maroon, sometimes with pale-brownish tints; flesh whitish, unchanging when exposed; odor and taste not distinctive. Pores and tubes: whitish at first, becoming ochraceous yellowish to rusty ochraceous with age, staining brownish when bruised, depressed at the stalk at maturity; tubes 8–16 mm deep. Stalk: nearly equal or slightly enlarged downward, solid; surface dry, smooth, pale yellow to brownish yellow, usually with reddish or brownish streaks, base white, weakly reticulate at the apex or at times more prominently reticulate, sometimes longitudinally striate on the upper portion; flesh concolorous with the cap flesh. Habitat and season: solitary or scattered in broadleaf woods under oak or in mixed oak and pine woods; spring, summer, and fall. Distribution and frequency: throughout northeastern North America, south to Georgia; fairly common. Spore print: rusty ochraceous to yellow ocher. Microscopic features: spores 8–14 × 3–4 µm, oblong-ellipsoid to subfusiform, smooth, pale yellow. Edibility: edible and good. Lookalikes: This bolete is sometimes confused with Xanthoconium affine (p. 409), but the latter lacks reddish to purplish tones on the cap and stains reddish brown when NH4OH is applied. Boletus separans (p. 153) has a paler cap and stalk with lilac tones, and its pores do not stain brownish when bruised.

Xanthoconium   ·  413

Xanthoconium stramineum

Xanthoconium stramineum (Murrill) Singer = Boletus stramineus (Murrill) Murrill Common name(s): none. Overview: The cap of Xanthoconium stramineum stains bluish gray with FeSO4, stains quickly salmon pink with KOH and then fades on dark areas, and is negative with NH4OH. The term stramineum means “straw-colored,” alluding to the cap color. Cap: 4–9 cm wide, convex, becoming broadly convex to nearly plane in age, margin incurved at first, with a narrow band of sterile tissue; surface dry, glabrous and smooth when young, sometimes becoming cracked and uneven with age, white at first, becoming whitish to pale straw-colored or tinged brownish; flesh white, unchanging or slowly staining pinkish brown when exposed; odor variously described as slightly fruity, unpleasant, or not distinctive; taste somewhat bitter or not distinctive. Pores and tubes: white to whitish at first, becoming buff to yellowish buff or pale yellow brown in age, unchanging when bruised; tubes 3–10 mm deep. Stalk: enlarged downward or nearly equal, usually with a swollen base, solid; surface dry, glabrous, white to whitish, not staining when bruised; flesh white, staining like the cap flesh.

414  ·  Boletes of Eastern North America

Habitat and season: solitary, scattered, or in groups on sandy soil or in grassy areas, under oak and pine; summer and fall. Distribution and frequency: coastal plain of North Carolina south to Florida, west to Texas; occasional to fairly common. Spore print: brownish yellow to yellowish rusty brown. Microscopic features: spores 10–15 × 2.5–4 µm, cylindrical, smooth, yellowish to hyaline. Edibility: reported to be edible but in our experience sour, bitter, and unpleasant. Lookalikes: This southern bolete can be confused with Boletus albisulphureus (p. 86), which has a reticulate stalk. Gyroporus subalbellus (p. 215) is similar but has a brittle stalk that becomes hollow at maturity. Xanthoconium montaltoense Wolfe (not illustrated) is known only from the Mixhaux State Forest in south-central Pennsylvania. It has a white to pale-cream cap that darkens to ocher or pale yellow gold in age, a stalk that is concolorous with the cap, and white flesh, and it grows under birch and hemlock. Gen us X e ro com e l lu s Šu ta r a The genus Xerocomellus was erected in 2008 to accommodate fairly small boletes that have short, broad, and encrusted cuticular cap hyphae. Species in this genus have soft flesh and caps that often become cracked. Microscopically, they have dense, parallel, nondivergent hyphal trama and short to elongate, ellipsoidal to oblong, or subfusiform spores. In some species, the spores are longitudinally striate or are smooth and sometimes truncate. The genus Xerocomus is very similar to Xerocomellus, but species in Xerocomus have long, smooth, nonencrusted cuticular cap hyphae and spores with a bacillate shape. A useful field character for recognizing species of Xerocomellus and Xerocomus is a tendency of the tubes to split lengthwise when cut or torn owing to their parallel, nondivergent hyphal trama. The tubes of species in other bolete genera, such as Boletus, remain intact when cut or torn because they have strongly divergent hyphal trama. There are no significant macroscopic characteristics that differentiate species of Xerocomellus from species of Xerocomus. Only a few species of Xerocomellus are known to occur in eastern North America.

Xerocomellus chrysenteron (A)

Xerocomellus chrysenteron (B)

416  ·  Boletes of Eastern North America

Xerocomellus truncatus

Xerocomellus chrysenteron (Bull.) Šutara = Boletus chrysenteron (Bulliard) Fries = Xerocomus chrysenteron (Bulliard) Quélet Common name(s): Red-cracked Bolete Overview: The cap surface stains dark brown to blackish with KOH, olive with FeSO4, and brownish or negative with NH4OH. During extended periods of wet weather, this common bolete is often infected with the parasitic mold Hypomyces chrysospermus, which envelops the fruitbody with a white to golden-yellow coating. The term chrysenteron means “gold intestine,” possibly an obscure reference to the meandering pattern of the cracks on the cap surface. Cap: 3–7.5 cm wide, convex, becoming broadly convex to nearly plane; surface dark olive to olive brown or grayish brown, often with a reddish zone at the margin, velvety at first, soon becoming cracked with red to pinkish color in the cracks; flesh soft, white to pale yellow, often with a pinkish-red zone beneath the cuticle, slowly staining blue when exposed; odor and taste not distinctive. Pores and tubes: yellow, becoming olivaceous with age, bruising greenish blue, sometimes slowly, typically depressed at the stalk with age; tubes 2–10 mm deep.

Xerocomellus   ·  417

Stalk: slender, more or less equal to slightly enlarged downward, solid; surface dry, finely granular to somewhat scurfy, sometimes with longitudinal streaks, yellow on the upper portion, usually reddish near the base, with white basal mycelium, at times staining bluish green when handled; flesh white to pale yellow, staining pinkish red and sometimes slowly and erratically staining blue. Habitat and season: solitary to scattered in broadleaf and conifer woods, also on roadside banks; late spring, summer, and fall. Distribution and frequency: eastern Canada south to North Carolina, west to Minnesota and Texas; fairly common but rarely found in large numbers. Spore print: olive brown. Microscopic features: spores 9–13 × 3.5–4.5 µm, oblong, smooth, pale brown. Edibility: edible but poor. Lookalikes: Xerocomellus truncatus (Singer, Snell & E. A. Dick) Klofac = Boletus truncatus (Singer, Snell & E. A. Dick) Pouzar (see photo, p. 416) is nearly identical, but many of its spores are distinctly truncated and measure 10–15 × 5–7 µm. Compare with Xerocomellus intermedius (p. 419), which has a dry, somewhat velvety, red to rose-red or olivaceous cap that becomes cracked in age. Like Xerocomellus chrysenteron, the cap of Xerocomellus intermedius typically shows pinkish tints in the fissures as it ages, and its flesh stains blue when exposed. Also compare with Boletellus chrysenteroides (p. 78), which often grows on decaying logs and stumps or at the base of standing trunks. It has a darker cap that becomes cracked but does not show reddish in the fissures. Boletus subtomentosus var. subtomentosus (p. 165) is somewhat similar but does not show reddish in the cap fissures. It has yellow pores that bruise slightly greenish blue, if at all. Boletus ferrugineus (p. 105) lacks reddish tinges in the cap fissures, and the cap surface displays a green flash with NH4OH and then stains reddish brown. Boletus subdepauperatus (not illustrated), reported from New England and New York west to Michigan, has a dingy olive-brownish to dingy yellow-brown cap that is finely cracked in age, whitish flesh that stains blue when exposed, yellow pores that stain blue when bruised, a yellowish-brown pruinose stalk, and spores that measure 9–12 × 4.5–5 µm. It grows in broadleaf woods.

Xerocomellus intermedius (A)

Xerocomellus intermedius (B)

Xerocomellus   ·  419

Xerocomellus intermedius (A. H. Sm. & Thiers) Svetasheva, Simonini & Vizzini = Boletellus intermedius A. H. Sm. & Thiers Common name(s): none. Overview: The term intermedius means “intermediate between two species,” referring to this bolete’s very similar appearance to both Boletellus pseudochrysenteroides (p. 80) and Xerocomellus chrysenteron (p. 416). The key identification features include the rose-red cap when young, pink tints showing in a cracked cap cuticle, blue-staining pores, and striate spores that measure 9–12 × 4–5 µm. Cap: 4–10 cm wide, convex, becoming broadly convex to nearly plane, margin even, incurved when young; surface dry, subtomentose, becoming cracked in age, red to rose red or soon grayish to olive brown or olive gray at maturity, typically showing pink tints in the cracks in age; flesh yellow, staining blue when exposed; odor and taste not distinctive. Pores and tubes: bright yellow to yellow when young, becoming brownish yellow to reddish brown in age, staining blue and then slowly brown when bruised; tubes 7–12 mm deep. Stalk: nearly equal or slightly enlarged in either direction, solid; surface dry, yellow at the apex, punctate with rhubarb-red dots and points over a yellow ground color below the apex, usually with longitudinal ribs, with a white to pale-yellow basal mycelium; flesh concolorous with and staining like the cap flesh. Habitat and season: scattered or in groups under broadleaf trees or in mixed woods with oak and pine; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: olive to olive brown. Microscopic features: spores 9–12 × 4–5 µm, subellipsoidal to fusoid, weakly longitudinally striate, pale brown. Edibility: unknown. Lookalikes: Xerocomellus chrysenteron (p. 416) has a dark-olive to olivebrown cap that becomes cracked in age, with reddish tinges in the cracks, and smooth spores. Boletellus pseudochrysenteroides (p. 80) grows on the ground in broadleaf woods, especially with beech or oak. When young, it has a dark rose-red to red-brown cap that typically becomes olivaceous and develops prominent cracks as it ages. It has whitish to pale-yellow flesh that quickly stains blue when exposed; its stalk base is sometimes coated with yellowish mycelium; and it has larger spores with longitudinal ridges that measure 11–18 × 5–8 µm.

420  ·  Boletes of Eastern North America

Gen us X e ro com u s Qu él. The genus Xerocomus was erected in 1887 to accommodate small- to medium-size boletes formerly placed in the genus Boletus that have a dry cap and stalk and tubes that split lengthwise when torn, forming split half-tubes. The tubes of Boletus species remain intact when torn apart. Many species that were at one time placed in Xerocomus have been reclassified, leaving less than a dozen species that occur in eastern North America. They have soft flesh and caps that often become cracked. Microscopically, species of Xerocomus have long, smooth, nonencrusted cuticular cap hyphae; dense, parallel, nondivergent hyphal trama; and short to elongate, ellipsoidal to oblong, or subfusiform spores with a bacillate shape. The genus Xerocomellus is very similar to Xerocomus, but species in Xerocomellus have short, broad, and encrusted cuticular cap hyphae and longitudinally striate or smooth spores. A useful macroscopic feature for recognizing species of Xerocomus and Xerocomellus is the tendency of their tubes to split lengthwise when cut or torn owing to the parallel, nondivergent hyphal trama. The tubes of boletes in other genera remain intact when cut or torn because they have strongly divergent hyphal trama. There are no significant macroscopic characteristics that differentiate species of Xerocomus from species of Xerocomellus.

Xerocomus hortonii (A)

Xerocomus   ·  421

Xerocomus hortonii (B)

Xerocomus hortonii (A. H. Sm. & Thiers) Manfr. Binder & Besl = Boletus hortonii A. H. Sm. & Thiers Common name(s): Corrugated Bolete, Horton’s Bolete Overview: The species name honors American mycologist Charles Horton Peck (1833–1917). The deeply pitted to corrugated cap, yellow pores, and smooth to lightly pruinose stalk are this unusual bolete’s distinctive features. The cap surface develops a blue-green flash and then stains olive brown with NH4OH. Cap: 4–10 cm wide, convex to broadly convex, margin even; surface dry to somewhat viscid, deeply pitted and corrugated, tan to reddish tan or ocher brown to reddish brown; flesh whitish to pale lemon yellow, unchanging when exposed; odor and taste not distinctive. Pores and tubes: yellow at first, becoming olive yellow with age, unchanging or rarely and slowly staining blue when bruised; tubes 5–10 mm deep. Stalk: enlarged downward or nearly equal, solid; surface dry, smooth to lightly pruinose or occasionally with delicate reticulation on the upper half, pale yellow to tan, sometimes reddish at the base; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups under broadleaf trees or in mixed woods, especially with oak, hickory, beech, and hemlock; late spring through fall.

422  ·  Boletes of Eastern North America

Distribution and frequency: eastern Canada south to Georgia, west to Minnesota and Arkansas; occasional to fairly common. Spore print: olive brown. Microscopic features: spores 12–15 × 3.5–4.5 µm, somewhat boat-shaped, smooth, yellow. Edibility: edible and very good. Lookalikes: Hemileccinum subglabripes (p. 220) is nearly identical. It differs by having a relatively smooth cap and a scurfier stalk. Leccinum rugosiceps (p. 262) has a pitted to cracked cap, flesh that stains dull reddish when cut, and brownish scabers on its stalk.

Xerocomus hypoxanthus

Xerocomus hypoxanthus Singer = Boletus hypoxanthus (Singer) Murrill Common name(s): none. Overview: The term hypoxanthus means “yellow beneath,” referring to the color of this bolete’s pores and stalk. The cap surface stains blue to green blue with NH 4OH or KOH. The flesh stains blue to bluish with NH 4OH. Additional identification features include a brown cap that often becomes cracked with age and yellow flesh that sometimes stains blue when exposed.

Xerocomus   ·  423

Cap: 2.5–7 cm wide, convex, becoming broadly convex to nearly plane, margin even or nearly so; surface dry, somewhat velvety to granular, often finely cracked in age, tawny brown to pale red brown or pale yellow brown to orange brown; flesh yellowish to yellow, staining slightly blue or not at all when exposed; odor and taste not distinctive. Pores and tubes: yellow, staining bluish or greenish when bruised or sometimes unchanging, often depressed at the stalk in age; pores usually elongated near the stalk; tubes 6–12 mm deep. Stalk: nearly equal, solid; surface dry, pale yellow to pale tawny with a yellow apex and often with a yellow base, usually coated with tiny brownish powdery flakes over the midportion, basal mycelium yellow; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary, scattered, or in groups on sandy soil or humus in broadleaf woods, especially with oak, in mixed oak and pine woods, or on decaying trunks of broadleaf trees or saw palmetto; summer and fall. Distribution and frequency: South Carolina and Florida, west along the Gulf Coast to Mississippi, distribution limits yet to be established; occasional. Spore print: olive brown. Microscopic features: 8–14 × 4–5 µm, fusoid, smooth, yellowish. Edibility: unknown. Lookalikes: Xerocomus hemixanthus (Singer) Murrill (not illustrated) is similar but has a dark-brown to reddish-brown or chestnut-brown cap, reddish-brown raised ribs near the stalk apex, and a white to yellowish-white basal mycelium. Its cap surface stains dark reddish brown with KOH and tan with NH4OH. Boletus ochraceoluteus (p. 125) has flesh that stains reddish and then bluish green and a cap surface that stains reddish brown with KOH or NH4OH. Xerocomus pseudoboletinus (Murrill) Singer (not illustrated) occurs from Florida west to Texas, has a large (up to 18 cm wide), reddish-brown to ochraceous-tawny cracked cap, with yellow showing in the cracks of mature specimens, and yellow flesh that is unchanging or sometimes staining weakly green when exposed. The pores are yellow and stain blue when bruised. The pores are elongated near the stalk, which is pale yellow on the upper portion and brownish toward the base, with coarse, brownish longitudinal ribs.

Xerocomus illudens / tenax complex (A)

Xerocomus illudens / tenax complex (B)

Xerocomus illudens / tenax complex (C)

Xerocomus illudens / tenax complex (D)

426  ·  Boletes of Eastern North America

Xerocomus illudens (Peck) Singer / Xerocomus tenax (A. H. Sm. & Thiers) Halling & Nuhn complex Common name(s): none. Overview: In 1898, Charles H. Peck (1833–1917) described Boletus illudens. When Alexander Smith and Harry Thiers examined Peck’s type collection, they discovered that it was a mixture of two species, and they designated a lectotype for Boletus illudens Peck that represented Peck’s original description of it. In 1971, they described Boletus tenax, which is currently considered an illegitimate name, having been previously assigned to a different species in 1777. Smith and Thiers described the stalk of Boletus tenax as having a distinct, wide-meshed, fine reticulum overall or only over the upper half. In addition, they wrote in their description of Boletus illudens that its stalk was usually marked by coarse ridges and anastomosing lines but was not finely reticulate. The descriptions of Boletus illudens by other mycologists, including Ernst E. Both, William Coker and Alma Beers, Rolf Singer, and Walter Snell and Esther Dick, report its stalk to be strongly reticulated. For many years, mycologists have struggled with this conflicting information and the problem of how to differentiate Xerocomus illudens from Boletus tenax. Some authors have attempted to use macroscopic or microscopic features or macrochemical testing, but the results have always been inconclusive owing to taxonomic uncertainty of the specimens tested or examined. Recent phylogenetic analysis places Boletus tenax in the Xerocomus clade, therefore supporting the new combination Xerocomus tenax Nuhn & Halling (2015). It is possible that Xerocomus illudens and X. tenax are the same species or that two or more species are collectively represented by these two taxa. Until additional molecular analysis has been done, we have elected to consider the two a species complex. The term illudens means “to deceive,” a reference to this bolete’s similarity with other species. The term tenax means “tenacious, tough, or firm,” in reference to the texture of its stalk or its tenacious hold on the substrate. Cap: 3–10 cm wide, convex, becoming nearly plane, sometimes slightly depressed, margin even, incurved at first, sometimes uplifted, lobed, and irregular at maturity; surface dry, somewhat velvety, yellow brown to pinkish cinnamon or dull brick red to reddish brown, sometimes with an olivaceous tint; flesh whitish to pale yellow, unchanging or slowly staining pinkish red when exposed; odor not distinctive; taste acidic or not distinctive. Pores and tubes: bright yellow at first, becoming dingy yellow and depressed at the stalk when mature, usually slowly staining cinnamon red or sometimes slowly staining bluish and then cinnamon red, rarely unchanging

Xerocomus   ·  427

when bruised; pores large, angular to somewhat rounded, larger near the stalk; tubes 8–16 mm deep. Stalk: tapered downward or sometimes nearly equal, solid, firm or sometimes almost woody at the base; surface dry, reticulate overall or at least on the upper portion or with raised longitudinal lines that may form a partial reticulum at the apex or nearly overall, whitish with pale-rose to pinkish-cinnamon or yellowish tints; flesh concolorous with the cap flesh. Habitat and season: solitary, scattered, or in groups in broadleaf woods with oak or under pines; summer and fall. Distribution and frequency: widely distributed throughout eastern North America; occasional. Spore print: olive brown. Microscopic features: spores 9–14 × 4–6 µm, ellipsoidal to subfusoid, smooth, dull yellow to pale brown. Edibility: unknown. Lookalikes: none.

Xerocomus morrisii

428  ·  Boletes of Eastern North America

Xerocomus morrisii (Peck) M. Zang = Boletus morrisii Peck Common name(s): Red-speckled Bolete Overview: This species was named for Boston mycologist and artist George Edward Morris (1853–1916), the collector of the type specimen. The red-dotted stalk ornamentation on this unusual and distinctive bolete is similar to scabers on the stalks of species in the genus Leccinum, which are brown or black. Cap: 3–9 cm wide, convex to broadly convex with a narrow, overlapping band of sterile tissue at the margin; surface dry, somewhat velvety at first and becoming glabrous in age, olive brown to reddish brown, often yellow to olive gold on the margin; flesh pale yellow, slowly staining reddish when exposed; odor and taste not distinctive. Pores and tubes: yellow to orange yellow or tinged reddish near the stalk, becoming orange to brownish orange in age, deeply depressed at the stalk of mature specimens, partial veil absent; tubes 6–12 mm deep. Stalk: nearly equal or enlarged downward, solid; surface yellow beneath a covering of reddish or reddish-brown punctae, basal mycelium yellow; flesh concolorous with and staining like the cap flesh. Habitat and season: solitary or sometimes in small groups, with pine, often in sandy soil; summer and fall. Distribution and frequency: New England south to Georgia, west to Tennessee; generally uncommon to rare but may be somewhat common locally. Spore print: olive brown Microscopic features: spores 10–16 × 4–6 µm, ellipsoidal to spindle-shaped, smooth, yellowish. Edibility: reported to be edible but has an unpleasant taste. Lookalikes: Leccinum rubropunctum (p. 261) is similar, but its yellowish flesh does not stain when exposed, and it has an unpleasant odor reminiscent of stale cigarette butts in an ashtray. Its pores become dull yellow in age.

Xerocomus sclerotiorum (A)

430  ·  Boletes of Eastern North America

Xerocomus sclerotiorum (B)

Xerocomus sclerotiorum A. E. Bessette, M. E. Smith, A. R. Bessette & H. Hitchcock, nom. prov. Common name(s): Whitey’s Bolete Overview: The term sclerotiorum refers to the hidden, irregularly shaped orange sclerotia that this colorful bolete forms in leaf litter and soil. The common name honors Dr. Harold “Whitey” Hitchcock, who first collected it. All parts of this mushroom stain orange with KOH. Cap: 4–10 cm wide, hemispheric at first, becoming convex to broadly convex at maturity, margin bright yellow, often persistent, incurved at first, with a narrow band of sterile tissue; surface dry, dull or somewhat shiny, smooth or nearly so, pinkish red to rose red or purplish red, sometimes with olive tints, becoming dull rose pink to brownish pink with age, slowly staining blackish blue when bruised; flesh pale yellow, sometimes with a pinkish tinge under the cuticle, staining blue when exposed, sometimes weakly and erratically; odor not distinctive; taste acidic. Pores and tubes: bright yellow at first, becoming dull yellow then brownish yellow at maturity, staining blue then slowly dull brown when bruised, slightly depressed near the stalk in age; tubes 6–15 mm deep. Stalk: enlarged downward or nearly equal, solid; surface dry, yellow at the apex, red on the lower portion, with conspicuous red to reddish-brown punctae over a yellow ground color, staining dark blue and then dull brown

Xerocomus   ·  431

when handled or bruised, often with white basal mycelium and yellowish rhizomorphs, longitudinally striate, sometimes reticulate only at the very apex; flesh brighter and deeper yellow than the cap flesh, reddish brown around larval tunnels, staining blue green, sometimes slowly and erratically. Habitat and season: scattered or in groups on sandy soil or on pincushion moss in woods with mixed scarlet, chestnut, and white oaks; summer and fall. Distribution and frequency: reported from New England south to Florida and west to Texas, distribution limits yet to be determined; occasional. Spore print: olive brown. Microscopic features: spores (12)14–16(18) × 4–6 µm, subfusoid to fusiform, smooth, hyaline to pale brownish yellow. Edibility: unknown. Lookalikes: Boletus rubissimus A. H. Sm. (not illustrated) also has a pinkish-red to rose-red cap with a bright-yellow margin and a similarly colored stalk, but it has much smaller spores, 9–11 × 3–4.5 µm. Hortiboletus rubellus (p. 225) has reddish-orange flesh in the lower portion of its stalk, tubes that split lengthwise when torn, and smaller spores, 10–13 × 4–5 µm. Boletus rubricitrinus (p. 145) has a similarly colored cap without a bright-yellow margin, a more yellow longitudinally striate stalk streaked with red, and different macrochemical test reactions.

Undescribed Bolete Species Pending Publication

The photographs here illustrate species of boletes that at the time of publication of this book had not yet been fully described and published. Some are very distinctive and may be recognized from a photograph alone; however, a few brief comments are included, providing additional information.

UB-1

UB-1: Boletus sp. nov. Collected in North Carolina; growing on a lawn under oaks; cap brick red to brownish; stalk yellowish at the apex, red below; all parts immediately stain blue when bruised; odor and taste are not distinctive. 433

434  ·  Boletes of Eastern North America

UB-2

UB-2: Boletus sp. nov. Collected in northeastern Georgia; growing in a roadside grassy area with mixed oaks and pines; similar to Caloboletus peckii (p. 182) but differing by having a more rose-pink to rose-red cap that does not fade to brownish rose or dull brown from the disc outward and much finer reticulation; collection number ARB1268.

UB-3

Undescribed Bolete Species  ·  435

UB-3: Boletus sp. nov. Collected in southeastern Georgia; growing on the ground under pines or in mixed oak and pine woods; note the long stalk and the copious white basal mycelium; collection number ARB1297.

UB-4

UB-4: Boletus sp. nov. Collected in a mixed broadleaf parkland in West Virginia in July; the olivaceous yellow-brown caps are distinctive.

436  ·  Boletes of Eastern North America

UB-5

UB-5: Boletus sp. nov. Collected in Ohio, Pennsylvania, and West Virginia; growing on the ground in broadleaf or mixed woods; closely resembles Hemileccinum subglabripes (p. 220) but differs in having a more velvety reddish-purple cap with a whitish bloom; may be either uncommon or perhaps mistaken as a form of H. subglabripes.

UB-6

Undescribed Bolete Species  ·  437

UB-6: Boletus sp. nov. Collected in West Virginia at multiple sites; growing on the ground with hickory; cap cinnamon brown to golden brown, somewhat velvety, at times developing cracks on the disc; pores yellow, unchanging when bruised; flesh pale yellow, unchanging when exposed; stalk yellow, with or without a dull reddish zone on the midportion of mature specimens, finely reticulate on upper half or more, reticulation yellow or red in the area where there is red on the stalk.

UB-7

UB-7: Boletus sp. nov. Collected in mixed oak and hickory woods in West Virginia in August; distinctive blue staining on all parts when cut or bruised.

438  ·  Boletes of Eastern North America

UB-8

UB-8: Suillus sp. nov. Collected under pines or in mixed oak and pine woods in southeastern Georgia and Florida; very similar to Suillus cothurnatus (p. 328) and Suillus salmonicolor (p. 348), but much more robust and with flesh that tastes unpleasant to rather disgusting; collection number ARB1378.

Appendixes Glossary Recommended Resources References Index to Common Names Index to Scientific Names Photograph Credits

A PPE N DI X A

Chemical Reagents and Bolete Identification Chemical reagents are used as tools in the identification and classification of boletes. Two macroscopically similar species may be differentiated by their reactions to a chemical reagent. For example, Boletus ferrugineus (p.  105) and Boletus subtomentosus var. subtomentosus (p. 165) can be difficult to separate macroscopically. However, you can quickly distinguish the two species from each other by adding a drop of ammonium hydroxide, or ordinary household ammonia, to their cap surfaces. The cap surface of Boletus ferrugineus displays a fleeting but vivid blue or blue-green reaction that turns reddish brown, whereas the cap surface of Boletus subtomentosus var. subtomentosus immediately stains reddish brown when ammonia is applied. We have experimented using a variety of objects and techniques for applying macrochemical reagents to mushrooms, such as toothpicks, paper towels, glass and plastic droppers, metal probes, and forceps. Regarding technique, some individuals apply a very small amount of reagent, whereas others apply a copious amount. All of these methods can produce accurate results some of the time, but they are inconsistent. Using a cotton-tipped swab is easy and provides consistently accurate results. This is especially important when trying to observe a “flash” or fleeting chemical reaction. The application of too much reagent can obscure this response, sometimes obliterating rapidly changing color sequences. To use a cotton-tipped swab, dip it into the reagent and apply it to the mushroom tissue with minimum pressure. Use a new, clean swab every time you are testing tissue to prevent contamination of your reagent. Reagents should be fresh, not outdated. Although they will keep for several weeks or sometimes for months, periodic testing of the reagents using bolete tissues with known macrochemical reactions is strongly recommended. Reagents should be stored at room temperature, preferably in plastic bottles. They may also be kept in clear glass bottles for several weeks or more, but they should not be stored longer than this. Some reagents, especially KOH, can react with glass to form silicates and may no longer produce accurate macrochemical test results. Several factors can affect macrochemical test results: color variations on a single specimen, the age of the mushroom, whether the tissue is parasitized or not, if the specimen has been refrigerated, and moisture content. Best results are obtained when using recently collected, fresh, unrefrigerated mushrooms with dry surfaces. If macrochemical testing is done on refrigerated boletes, allow the specimens to warm to room temperature before testing them. In addition, some chemicals are used in microscopic work for bolete identification. They are used to rehydrate dried specimens; as a mounting medium in slide preparation; for identifying some species based on the appearance of their spores, hyphae, and cystidia; and for noting the specific staining reactions of other microscopic structures.

441

442  ·  Appendix A

Boletus huronensis with chemical reagents The chemical reagents listed here are used for both macrochemical testing of bolete tissues and microscopic study. NH 4OH = ammonium hydroxide: 3–14 percent aqueous solution or household ammonia without added soap. Ammonium hydroxide is also used as a mounting medium for microscopic work. It is ideal for use with dried material that has been rehydrated in a 70–95 percent ethyl alcohol solution. FeSO4 = iron sulfate: 10 percent aqueous solution KOH = potassium hydroxide: 5–14 percent aqueous solution. Potassium hydroxide is also used as a mounting medium for fresh and dried specimens. Unless otherwise stated, spore colors reported in the “Microscopic Features” section of each description are based on their appearance when mounted in potassium hydroxide. Melzer’s reagent: Melzer’s reagent is a special solution of iodine (0.5 g), potassium iodide (1.5 g), chloral hydrate (20 g), and water (20 ml). It is an important reagent used for identifying some mushroom species based on

Appendix B  ·  443 the appearance of their spores, hyphae, cystidia, and other microscopic structures. These structures sometimes stain specific colors when mounted in Melzer’s reagent. If they stain bluish gray to bluish black, they are described as amyloid; those that stain reddish brown are dextrinoid; and those structures that stain yellow or remain colorless are inamyloid. Lactophenol Cotton Blue: This blue dye is a mixture of lactic acid, phenol, and cotton blue. It is used as a mounting fluid and stain and is useful because it stains cytoplasm pale blue and hyphal wall structures and ornamentation a darker blue. Phloxine: Phloxine is a red-colored dye used for microscopic examination, especially of hyphae. It is useful because it stains cytoplasm red and thus makes other contrasting structures, such as cystidia, much easier to differentiate. Many other chemical reagents are useful for microscopic examination of bolete tissues but are beyond the scope of this work. We have decided not to include macrochemical reactions for most of the species in this book owing to striking inconsistencies in the literature, variable results in our own testing, and difficulties arising from the subjective nature of color interpretation. For some species, we have provided macrochemical test reactions that are reliable, valid, and specifically diagnostic.

A PPE N DI X B

Microscopic Examination of Boletes Microscopic structures are commonly used to differentiate genera and species when macroscopic characteristics are not enough for a positive identification. Many mycological papers and books, especially technical monographs, use identification keys based on microscopic features such as cystidia, basidia, clamp connections, and hyphae. An in-depth discussion of microscopic structures and evaluation techniques is beyond the scope of this book. We have, however, included the following information for those interested. A good-quality microscope is required to do microscopy. At minimum, you need to have a microscope that has a substage condenser, a built-in light source, and good-quality lenses: at least three objective lenses, 10×, 40×, and an oil-immersion 100× as well as one or two ocular lenses (eyepieces). An ocular micrometer is also required to make accurate measurements of microscopic features. Some ocular lenses have an ocular micrometer already inserted; others do not, and it must be bought separately. Because the magnification of each combination of lenses (ocular plus objective) is different, the ocular micrometer needs to be calibrated using a tiny glass ruler known as a stage

444  ·  Appendix B micrometer. Most microscope dealers as well as college and university biology departments have stage micrometers. The folks in these places are usually more than willing to help with this calibration process. Spores used in microscopic evaluation can be obtained from three sources: a spore print, fresh boletes, or dried boletes. • If you are using a spore print as your source, scrape off a minute sample

with a razor or knife blade. Mix it into a small drop of water or another mounting medium such as KOH, and carefully place a coverslip over the mixture. Gently apply pressure on the coverslip using a pencil eraser to distribute the spores and remove air bubbles and excess fluid. • If you are using fresh material, remove a small portion of the tubes, place it in your mounting medium of choice, add the coverslip, and gently apply pressure as noted earlier. • When you are using dried boletes as your source of spores, allow a small piece of the pore surface to soak and soften in the mounting medium for two or three minutes before adding the coverslip and applying pressure as directed. Before placing pieces of dried material into a mounting medium, you can soak them in wetting agents, such as 70–95 percent ethyl alcohol, so that hyphae and other structures absorb water and regain their original appearance. Also, some microscopic structures, such as cystidia and hyphae, may be easier to see if dyes such as phloxine are used (see appendix A, “Chemical Reagents and Bolete Identification,” p. 441). Once the coverslip is in place, your slide is ready for use. Place a drop of immersion oil on the coverslip and carefully lower the 100× objective lens into the oil. Fine-focus the image, adjust the lighting, and examine the spores, cystidia, hyphae, and other structures you are interested in. It is helpful to have a basic understanding of the organization of the fruitbody for doing microscopic study. Boletes are composed of specific tissues that provide critical information and clues that help lead to a positive identification. The following features are commonly examined, beginning with the cap and working downward to the stalk. To start, the inner portion of a bolete (cap and stalk) is the trama, commonly referred to as the context or flesh. • Cap: The pileipellis (cuticle or skin) is the outermost region of the cap and

can consist of a single layer or as many as three layers. The layer directly beneath the cap cuticle is the pileus trama. Beneath that layer is the tube tissue, known as the hymenophore or hymenophoral trama. It supports a thin layer composed of the reproductive cells (the basidia and basidiospores). This layer is called the hymenium. • Stalk: The outer layer of the stalk is called the cuticle, or the stipitipellis. Like the cap’s cuticle, it can consist of a single layer or as many as three layers. The inner portion of the stalk, its context or flesh, is called the stipe trama.

Appendix C  ·  445 Additional microscopic structures mentioned in some of the descriptions in this book are defined in the glossary. For a more in-depth explanation about microscopic features, refer to How to Identify Mushrooms to Genus III: Microscopic Features by David Largent, David Johnson, and Roy Watling (1977). This valuable reference describes laboratory techniques, equipment and materials, calibration of the microscope, planes of sectioning, rehydration of dried materials, microscopic structures, and much more.

A PPE N DI X C

Collecting, Cooking, and Preserving Boletes Gold and silver and dresses may be trusted to a messenger, but not a boletus, because he will eat it on the way. —Marcus Valerius Martialis, Epigrams

Edi bi l i t y of Bol et e s There are several outstanding edible boletes, including the King Bolete (Boletus edulis, p. 101). It is the most sought-after wild mushroom anywhere it occurs and has a host of regional names that are commonly used, including “Cep” in France, “Porcini” in Italy, “Steinpilz” in Germany, and “Penny Bun” in England. Although the King Bolete is the best-known and most-prized bolete, several other lesser-known species are also excellent. However, not all boletes are good edibles. Many are of mediocre quality or bitter tasting and unpalatable. A few are somewhat poisonous, and the edibility of others is still unknown. In the collection of any wild mushrooms for the table, it is well to adhere to the following rules. 1. Be certain of correct identification, and know that a species collected is an established edible. 2. Collect edible mushrooms only in places that are uncontaminated from industrial waste or agricultural chemicals, including lawn pesticides. Mushrooms are known to accumulate soil toxins such as heavy metals, so do not gather any along busy roadways if you plan to eat them. 3. Consume only specimens that are fresh, and avoid any that have mold spots or severe damage from insect infestation. 4. When trying a species for the first time, test your personal tolerance by eating only a small amount. Mushroom allergies are not uncommon. 5. Thorough cooking is recommended. It makes mushrooms more digestible and improves the flavor. Heat also destroys toxins that are present in some mushrooms— morels, for instance—and has the added benefit of killing any “overlooked” insects/ larvae. Although this statement is true, it is also important to emphasize that many mushroom toxins, including the most deadly ones, cannot be destroyed by heat.

446  ·  Appendix C Bol et e s i n t h e K i tch e n Preparation of boletes for consumption begins when they are collected in the field. Trim away the soil on the stalk base, and remove any needles or debris on the cap with a small brush such as a small paint brush. More refined cleaning can be done in the kitchen by removing stubborn dirt with a moist paper towel. Cut away blemishes and localized insect damage. Larval tunnels that are typically most evident in the stalk base become less concentrated toward the apex. The degree of minor insect-damage removal is largely a matter of personal choice and aesthetics. Some individuals experience an adverse dermatological reaction from handling certain species of Suillus. Others experience gastric distress from ingesting mature tubes or the cuticle slime layer of viscid or glutinous caps. However, species of Suillus are generally considered safe for eating, and, although often scorned by connoisseurs, some are actually good when properly prepared. In preparing species that have a glutinous coating, peel and discard the cap cuticle. Also remove the tube layer on older specimens because it becomes slimy when cooked. The tubes can be dried and ground into a flavorful seasoning powder for use in soups and sauces. P r e se rv i ng Bol et e s At times, nature will provide an overabundance of edible boletes. Few things are more thrilling to a mushroom hunter than finding a large flush of pristine edible boletes. Depending on the age and moisture content, fresh specimens will keep in the refrigerator for two or three days. When you have more than can be consumed when fresh, any surplus can be preserved in several ways. By far the preferred method for keeping and storing boletes is to dry them. It is a simple and efficient way to enjoy boletes long after they are no longer available fresh. Drying also concentrates the flavors, and many aficionados actually prefer them dried and reconstituted. The best way to dry boletes is to slice them thinly and place the slices on trays in a commercial food dryer, preferably one that has a thermostat control. Drying at 105 degrees is about the right temperature, but this is not absolute. It is better to dry them more slowly over a longer period than to have the temperature too hot, which will cook the mushrooms. It is also possible, though less reliable, to dry them on screens or racks in an oven at a very low temperature. In arid conditions, it is even possible to dry boletes in the sun if they aren’t too water-logged to begin with. Once the boletes are dry and brittle, store them in air-tight glass jars. Once they are dried, it is a good idea to put them in the freezer for approximately two weeks to ensure that any remaining insect eggs or larvae or both are killed. Properly dried boletes will keep for months or even years. To reconstitute them, soak them in lukewarm water or wine for fifteen to twenty minutes and then drain and save the liquid for later use. Dried mushrooms can be directly added to soups and broth without reconstituting. Freezing is another suitable method for preserving boletes. Although they can be frozen raw, it is preferable to cook them first. Sliced or quartered boletes sautéed in butter or olive oil can be divided into serving-size portions and put into freezer containers or bags. Be sure to label and date the container or bag because frozen mushrooms are

Appendix C  ·  447 difficult to identify. These frozen boletes will keep for three or four months. Bolete duxelles also make handy flavor packets when frozen in ice-cube trays. To prepare duxelles, chop the boletes finely, sauté with garlic or other seasoning until little moisture remains, and then pack the mixture into ice-cube trays. Once frozen, the cubes can be transferred to freezer containers or bags. They are excellent for adding to soups and other dishes. Although it is possible to freeze raw boletes whole, this method of preserving them works best if the specimens are young and firm. Pickling or marinating boletes in oil and spices is popular, but, again, this method works best with young specimens. Numerous variations and recipes can be found on the Internet. Pickled boletes will keep for several weeks in the refrigerator, although they are so delicious that you will likely consume them much sooner than that. Bol et e Sch na pps Dried specimens of some flavorful boletes, such as Boletus edulis (p. 101) and Boletus separans (p. 153), can be added to vodka or other neutral spirits to enhance their flavor, although this recommendation is not intended as a method of preserving boletes. After a week or more of “curing,” the resulting infusion is ready to consume. Adding dried boletes to dry white wine is another possibility worth experimenting with. In the Dolomite region of northern Italy, a superb popular grape-based brandy called Grappa is often flavored to great advantage with Porcini (Boletus edulis).

Porcini Grappa

448  ·  Appendix D

A PPE N DI X D

Common Tree Associates of Eastern Boletes Ash, Fraxinus species Black, Fraxinus nigra Carolina, Fraxinus caroliniana Green, Fraxinus pennsylvanica White, Fraxinus americana Aspen, Populus species Big-toothed, Populus grandidentata Cottonwood, Populus deltoides Quakinq, Populus tremuloides Beech, Fagus species American, Fagus grandifolia Birch, Betula species Dwarf, Betula glandulosa Gray, Betula populifolia Paper, Betula papyrifera River, Betula nigra Yellow, Betula alleghaniensis Chestnut, Castanea and Aesculus species American, Castanea dentata Horsechestnut, Aaesculus hippocastanum Fir, Abies species Balsam, Abies balsamea Hemlock, Tsuga species Carolina, Tsuga caroliniana Eastern, Tsuga canadensis Hickory, Carya species Bitternut, Carya cordiformis Mockernut, Carya tomentosa Pecan, Carya illinoensis Pignut, Carya glabra Sand, Carya pallida Shagbark, Carya ovata Shellbark, Carya laciniosa Water, Carya aquatica Hornbeam, Carpinus species American, Carpinus caroliniana Larch, Larix species Eastern, Larix laricina European, Larix decidua

Oak, Quercus species Bear, Quercus ilicifolia Black, Quercus velutina Blackjack, Quercus marilandica Bluejack, Quercus incana Chestnut, Quercus prinus Chinkapin, Quercus muehlenbergii Laurel, Quercus laurifolia Live, Quercus virginiana Northern red, Quercus rubra Post, Quercus stellata Shingle, Quercus imbricaria Southern red, Quercus falcata Turkey, Quercus laevis White, Quercus alba Pine, Pinus species Eastern white, Pinus strobus Jack, Pinus banksiana Longleaf, Pinus palustris Pitch, Pinus rigida Pond, Pinus serotina Red, Pinus resinosa Sand, Pinus clausa Scots, Pinus sylvestris Shortleaf, Pinus echinata Slash, Pinus elliottii Table mountain, Pinus pungens Virginia, Pinus virginiana Spruce, Picea species Black, Picea mariana Norway, Picea abies Red, Picea rubens White, Picea glauca

Glossary

acrid: producing a burning sensation in the mouth amyloid: staining grayish to blue black in Melzer’s reagent anastomosing: fusing to form a network angular: having multiple angles, not rounded annular zone: a partially formed ring composed of partial-veil remnants, also called a ring zone apex: the uppermost portion of the stalk apical: pertaining to the apex apical pore: a small opening or thin area in the wall at the apex of a spore; also known as a germ pore appressed: flattened onto the surface appressed-fibrillose: having fibrils that are flattened onto the surface appressed-tomentose: having a thick coating of hairs that are flattened onto the surface aquamarine: pale greenish blue aqueous: containing water astringent: drying and puckering bacillate: rod-shaped basal: located at the base of the stalk base: the lowest portion of the stalk basidia: typically club-shaped cells on which basidiospores are formed basidiospores: spores formed on basidia bay: reddish brown or chestnut color beige: a light yellowish brown beveled: having a slanted edge bloom: a dull, thin coating that is typically whitish bolete: a fleshy-pored mushroom

boletinoid: having elongated and radially arranged pores brick red: brownish red to dull brownish orange red broadleaf: referring to any non-conebearing deciduous tree or shrub bulbous: having a bulb-like base buff: pale yellow brown to pale grayish brown burgundy: a dark reddish purple burgundy red: a dark purplish red button: immature and typically rounded stage caespitose: occurring in clusters with stalk bases fused or nearly so cap: the upper part of the bolete fruitbody that supports the tubes carmine: rose red to dull rose red cartilaginous: tough and breaking with a snap when bent far enough caulocystidia: cystidia found on the stalk central: located at the center chambered: divided into distinctly separate spaces chamois: medium to grayish yellow cheilocystidia: cystidia found on the outer edge of the tubes cinnamon: reddish brown clade: a group of organisms that includes an ancestor and all the descendants of that ancestor clamp connection: a microscopic semicircular bridge-like structure that connects two adjoining cells in some boletes

449

450  ·  Glossary clavate: club-shaped clavate-mucronate: club-shaped with a short, sharp point cleft: a split-like division concentric: having rings or zones within one another in a series concolorous: having the same colors, colored like conical: more or less cone-shaped conifer: a cone-bearing tree with needle-like leaves, such as spruce, fir, hemlock, or pine convex: curved or rounded coral red: a reddish or pinkish shade of orange corrugated: coarsely ridged or wrinkled cottony-membranous: having a soft, cotton-like membrane cottony-scaly: having soft, cotton-like scales coverslip: a thin, flat piece of glass used for microscopy crimson: deep purplish red cuticle: the outermost layers of a bolete’s cap cuticular: referring to the cuticle cutis: a type of cuticle with cells arranged more or less parallel to the surface cylindric(al): somewhat cylinder-like, with nearly parallel sides cylindric-subclavate: cylinder-like and slightly club-shaped cystidia: sterile cells that project between and usually beyond the basidia cystidioid: resembling cystidia decurrent: descending or running down the stalk decurved: bent downward depressed: sunken dextrinoid: staining orange to orange brown, pinkish red to dark red, or reddish brown in Melzer’s reagent dichotomous: divided into two parts

disc: the central area of the surface of the cap divergent: turning outward from a middle line drab: a dull light brown duff: decaying plant matter on the ground in a forest eccentric: away from the center ecological: pertaining to the relationship between an organism and its environment ectomycorrhiza: a type of mycorrhiza characterized by fungal cells growing around the roots and between the outer root cells but not penetrating the cells ectomycorrhizal: pertaining to an ectomycorrhiza ellipsoid(al): resembling an elongated oval with similarly curved ends ellipsoid-cylindric: resembling an elongated oval with somewhat parallel sides ellipsoid-fusoid: mostly spindle-shaped but somewhat ellipsoidal ellipsoid-oblong: resembling an elongated oval that is longer than wide and with somewhat flattened ends elongate-ellipsoidal: having an extended oval with similarly curved ends elongate-fusoid: having an extended spindle shape elongate-ovoid: egg-shaped and somewhat extended encrusted: covered with a hard layer that gives a roughened appearance epithet: the descriptive portion of a species name; the second part of a Latin binomial scientific name equal: having the same thickness over the entire length evanescent: slightly developed and soon disappearing even: stopping at the tube layer, not projecting as a band of sterile tissue

Glossary  ·  451 exudate: a discharged substance exude: to come out slowly in drops fascicle: a small bundle fenugreek: a Eurasian plant with aromatic seeds used as a seasoning, especially in curry FeSO4: iron sulfate, usually a 10 percent concentration in water fiber: a hair-like structure present on the cap or stalk of some boletes fibril: a tiny fiber fibrillose: composed of fibrils fibrillose-punctate: composed of fibrils and tiny points, dots, scales, or spots fibrillose-scaly: with tiny scales composed of appressed fibrils fibrous-membranous: having a membrane composed of fibers fissure: a deep crack or cleft flesh: the inner tissue of a fruitbody flexuous: bent alternately in opposite directions floccose: tufted like a cotton ball floccose-powdery: resembling a powder-covered cotton ball floccose-scaly: having scales that are tufted like a cotton ball fruitbody: the fleshy reproductive structure of a fungus commonly called a mushroom (or, in this work, a bolete) fulvous: reddish cinnamon, colored like a red fox fuscous: dark brownish gray to brownish black or purplish brownish black fusiform: spindle-shaped and narrowed at both ends fusiform-ellipsoid(al): more or less ellipsoidal and spindle-shaped fusoid: somewhat spindle-shaped fusoid-ampullaceous: swollen and somewhat spindle-shaped fusoid-subventricose: slightly enlarged in the middle and somewhat spindle-shaped

fusoid-ventricose: enlarged in the middle and somewhat spindle-shaped garnet red: a deep or dark red gelatinous: composed of a jelly-like substance genus (pl. genera): taxonomic grouping of closely related species geotropic: a directional growth pattern in response to gravity gill: blade-like structure located on the underside of some mushrooms’ caps glabrous: bald, lacking hairs or scales globose: more or less round gluten: a sticky, glue-like substance glutinous: having gluten granular: composed of tiny grains granular-scaly: having scales composed of tiny grains granulose: composed of tiny grains gregarious: closely scattered over a small area habitat: the environment in which a species occurs hazel: a light brown to strong yellowish brown hemispheric(al): shaped like half of a sphere host: the organism on or in which a parasitic fungus lives or a plant partner in a mycorrhizal relationship hyaline: transparent, clear and nearly colorless hymenial cystidia: cystidia that occur on the inner surface or edge of the tubes, typically very similar among most boletes hymenium: the spore-bearing layer hymenophoral trama: sterile tissue supporting the hymenium hymenophore: tube tissue, also known as “hymenophoral trama” hyphae: thread-like filaments of fungal cells hyphal: pertaining to hyphae

452  ·  Glossary hyphal trama: a type of supporting tissue between basidia inamyloid: unchanging or pale yellow in Melzer’s reagent; neither amyloid nor dextrinoid incurved: bent inward toward the stalk indigo: very dark blue to almost violet inequilateral: having unequal sides inrolled: bent inward toward the stalk and upward intervenose: having veins on the surface that extend various distances ITS: internal transcribed spacer, one of the most widely sequenced DNA regions of fungi KOH: potassium hydroxide, usually made up in a 3–5 percent concentration in water lacerate(d): appearing as if torn lacerate-reticulate: covered with a netlike pattern and appearing torn lacunose: covered with elongated pits or indentations lageniform: flask-shaped, having the shape of a Florence flask, or gourd-shaped lamellate: having gills lanceolate: longer than wide, tapered, and pointed lateral: attached to the margin of the cap lignicolous: growing on wood lilac: a pale shade of violet or light purple lilaceous: having lilac coloration lobed: having rounded divisions longitudinal(ly): oriented along the vertical axis of the stalk lubricous: smooth and slippery macrochemical: relating to the use of a chemical reagent to observe reactions of bolete tissues without the use of a microscope macroscopic: observable without the use of a microscope

magenta: purplish red mahogany: dark reddish brown margin: the edge of a cap maroon: dark brownish red median: located in the middle medicinal: unpalatable, disagreeable Melzer’s reagent: a solution used for testing color reactions of spores or tissues; see p. 442 in appendix A for the formula membranous: having a membrane mercaptan: also known as methanethiol, a harmless but pungent-smelling gas that has been described as having the stench of rotting cabbages or smelly socks microscopic: of a size requiring a microscope to be seen clearly mixed woods: a mixture of trees containing both broadleaf and conifers monotypic: pertaining to a single type morphological: pertaining to the shape of an organism mottled: colored with streaks or blotches of different shades mottling: pattern of different shades or colors mycelium: a mass of hyphae, typically hidden in the substrate mycorrhiza: a mutually beneficial relationship with a tree or other plant mycorrhizal: having a mycorrhiza or mutually beneficial relationship with a tree or other plant NH 4OH: ammonium hydroxide; used to test color reactions on bolete tissues and sometimes used as a mounting fluid nondivergent: remaining parallel, not turning outward from a middle line nonmycorrhizal: not forming a mycorrhizal relationship with a host plant obclavate: the reverse of club-shaped oblong: longer than wide and with somewhat flattened ends

Glossary  ·  453 oblong-ellipsoid(al): longer than wide and resembling an elongated oval with similarly curved ends obtuse: rounded or blunt ochraceous: pale brownish orange yellow ocher: brownish orange yellow olivaceous: having olive tints opaque: dull, not shiny ornamentation: spines, warts, ridges, grooves, or other markings oval: egg-shaped, with the larger end at the base ovate: shaped like an egg ovoid: somewhat egg-shaped pallid: having a pale or whitish appearance parasitic: living on or in another organism and deriving nourishment from it partial veil: a layer of fungal tissue that covers the pore surface of some immature boletes phylogenetic tree: a branching diagram showing the inferred evolutionary relationships among various biological species piedmont: an area of land formed or lying at the foot of a mountain range pileipellis: the outermost layer of cap tissue, also called a cuticle pileus trama: the supporting tissue of the cap located between the cuticle and the spore-bearing layer of cells pith: soft tissue that often disappears plane: more or less flat pleurocystidia: cystidia found along the inner surface of the tubes polypore(s): a common name for fibrous to woody fungi that typically grow on wood and have tubes with pores pores: the open ends of the tubes of a bolete or polypore poroid: resembling pores or composed of pores

pruina: powdery particles, flakes, or dots pruinose: appearing finely powdered pruinose-scurfy: appearing powdered and roughened by tiny particles or flakes pruinosity: a coating of powdery particles, flakes, or dots pseudoamyloid: reacting strongly and actively to the iodine in Melzer’s reagent by staining a deep purplish brown pseudoreticulate: having a false reticulum usually formed by anastomosing ridges pseudoreticulation: false reticulation pseudoreticulum: a false reticulum that is usually formed by anastomosing ridges pseudorhiza: a root-like extension of the stalk pulverulent: somewhat powdery pulvinate: cushion-shaped, slightly convex punctae: tiny points, dots, scales, or spots punctate: marked with tiny points, dots, scales, or spots pungent: sharp or irritating radial(ly): pointed away from a common central point, like the spokes of a wheel range: geographical distribution of a species resinous dots: sticky and varnish-like spots on the surface of the stalk resupinate: reclining on the substrate and facing outward reticulate(d): covered with a net-like pattern of ridges reticulation: a pattern or arrangement of interlacing lines resembling a net reticulum: see reticulation retipolide(s): chemical compound(s) unique to species of the genus Retiboletus

454  ·  Glossary rhubarb red: pinkish red to crimson red ring: a portion of the partial veil that remains attached to the stalk after the veil ruptures ring zone: remnants of a partial veil that resemble a ring; also called an annular zone riparian: relating to the banks of a natural course of water saprotroph: an organism that obtains nourishment from dead organic matter saprotrophic: obtaining nourishment from dead or decaying organic matter scabers: small, stiff, granular projections on the surface of the stalks of some boletes; a distinctive feature of the genera Leccinellum and Leccinum scale: an erect, flattened, or recurved projection or torn portion of the cap or stalk surface scaly-punctate: composed of scales and tiny points, dots, or spots scarlet: vivid red with an orange tinge sclerotia: hard, dormant structures of variable shape and size that give rise to fruitbodies; composed of hyphae, usually with a dark outer rind scurfy: roughened by tiny particles or flakes scurfy punctae: scales, dots, points, or spots roughened by tiny particles or flakes scurfy-punctate: having a combination of tiny particles or flakes and scales, dots, points, or spots septa: microscopic crosswalls of hyphae septate: having microscopic crosswalls sheathing: to enclose with a cover layer sinuously: of a wavy or winding form slimy-viscid: glue-like and sticky solitary: occurring singly

spore: a microscopic reproductive cell with the ability to germinate and form hyphae spore print: a deposit of spores from the bolete tubes that is made on a piece of paper, plastic, or glass stalk: the structure that arises from the substrate and supports a bolete’s cap sterile: not involved with reproduction sterile margin: tissue not directly involved with the reproductive process stipe trama: the inner tissue, or portion, of a stalk stipitipellis: the outer tissue layer of a stalk, also called the cuticle striate: having parallel or longitudinal lines striations: minute parallel or longitudinal lines or furrows stuffed: containing a soft tissue that usually disappears with age and leaves a hollow space subcylindrical: somewhat cylinder-like subdecurrent: extending slightly down the stalk subellipsoid(al): somewhat elliptic subfusiform: nearly spindle-shaped subfusoid: somewhat spindle-shaped, tapered slightly at both ends subfusoid-ellipsoid: somewhat spindle-shaped and resembling an elongated oval subglobose: nearly round sublongitudinal: oriented somewhat along the longest axis suboblong: slightly longer than wide and having somewhat flattened ends substrate: the material on or in which an organism grows subtomentose: coated with a more or less thick, matted covering of hairs subventricose: somewhat swollen in the middle and tapered in both directions

Glossary  ·  455 subviscid: slightly sticky or tacky suede-like: having a soft, downy surface superior: located near the top symbionts: organisms living together in a state of symbiosis tawny: dull yellowish brown taxon (pl. taxa): unit having position and rank, such as a species or genus terrestrial: growing on the ground tomentose: coated with a thick, matted covering of hairs trama: sterile supporting tissue trichodermium: a cuticle with the distal portion of the filiform elements of equal length and arranged perpendicularly to the surface truncate(d): appearing cut off at the end tubes: narrow, parallel, spore-producing cylinders on the undersurface of a bolete’s or polypore’s cap tube trama: the sterile supporting tissue of the tubes umbo: a pointed or rounded elevation at the center of the cap umbonate: having an umbo undulating: having a wavy or uneven surface universal veil: a layer of tissue that completely encloses immature stages of some fruitbodies veil: a layer of fungal tissue that covers all or part of some immature boletes; see also partial veil and universal veil ventricose: swollen in the middle and tapered in both directions vinaceous: pinkish red to pale purplish red violaceous: having a bright bluish-purple color viscid: sticky or tacky viscidity: the condition of being sticky or tacky

References and Recommended Resources

R e f e r e nce s Bessette, A. E., and O. K. Miller. 2000. Boletus rainisii. In North American Boletes: A Color Guide to the Fleshy Pored Mushrooms, 145. Syracuse Univ. Press, Syracuse, NY. Bessette, A. E., W. C. Roody, and A. R. Bessette. 2000. North American Boletes: A Color Guide to the Fleshy Pored Mushrooms. Syracuse Univ. Press, Syracuse, NY. 400 pp. Both, E. E. 1993. The Boletes of North America: A Compendium. Buffalo Museum of Science, Buffalo, NY. 436 pp. Brundrett, M. C., and B. Kendrick. 1987. The Relationship between the Ash-tree Bolete (Boletinellus merulioides) and an Aphid Parasitic on Ash Tree Roots. Symbiosis 3:315–320. Den Bakker, H. C., and M. E. Noordeloos. 2005. A Revision of European Species of Leccinum Gray and Notes on Extralimital Species. Persoonia 18:511–587. Gelardi, M., A. Vizzini, and G. Simonini. 2014. Cyanoboletus. Index Fungorum 176. At http://www.indexfungorum.org. Kendrick, B. 2000. The Fifth Kingdom. Focus Publishing, Newburyport, MA. 373 pp. Kuo, M., and A. S. Methven. 2014. Mushrooms of the Midwest. Univ. of Illinois Press, Champaign. 427 pp. Largent, D. L., D. Johnson, and R. Watling. 1977. How to Identify Mushrooms to Genus III: Microscopic Features. Mad River Press, Eureka, CA. 148 pp. Nuhn, M. E., M. Binder, A. Taylor, R. E. Halling, and D. S. Hibbett. 2013. Phylogenetic Overview of the Boletineae. Fungal Biology 117:479–511. Osmundson, T. W., and R. E. Halling. 2010. Tylopilus oradivensis sp. nov.: A Newly Described Member of the Tylopilus balloui Complex from Costa Rica. Mycotaxon 113:475–483. Siegel, N., C. F. Schwarz, and J. L. Frank. 2014. [As “rainisii.”] Index Fungorum 179 (1). At http://www.indexfungorum.org. Snell, W. H., and E. A. Dick. 1970. The Boleti of Northeastern North America. Cramer, Lehre, Germany. 115 pp. Stamets, P. 2005. Mycelium Running: How Mushrooms Can Help Save the World. Ten Speed Press, Berkeley, CA. 343 pp. Vizzini, A., G. Simonini, and M. Gelardi. 2014. Exsudoporus. Index Fungorum 183. At http://www.indexfungorum.org. Wu, G., B. Feng, J. Xu, X-T. Zhu, Y-C. Li, N-K. Zeng, Md. H. Hosen, and Z. L. Yang. 2014. Molecular Phylogenetic Analyses Redefine Seven Major Clades and Reveal 22 New Generic Clades in the Fungal Family Boletaceae. Fungal Diversity 67:1–23.

457

458  ·  References and Recommended Resources R ecom m en ded R e sou rce s Books and Articles

Arora, D., and J. L. Frank. 2014. Clarifying the Butter Boletes: A New Genus, Butyriboletus, Is Established to Accommodate Boletus Sect. Appendiculati, and Six New Species Are Described. Mycologia 106 (3): 464–480. Bessette, A. E., A. R. Bessette, M. E. Nuhn, and R. E. Halling. In press. Lanmaoa roseocrispans, a New Bolete Species from Florida. North American Fungi. Calzada Dominguez, A. 2007. Guia de los BOLETOS de ESPAÑA y PORTUGAL. Náyade Editorial, Medina del Campo (Valladolid), Spain. 407 pp. Coker, W. C., and A. H. Beers. 1943. The Boletaceae of North Carolina. Univ. of North Carolina Press, Chapel Hill. 218 pp. Cotter, H. V., and O. K. Miller Jr. 1985. Sclerotia of Boletinellus merulioides in Nature. Mycologia 77:927–931. Den Bakker, H. C., G. C. Zuccarello, Th. W. Kuyper, and M. E. Noordeloos. 2004. Evolution and Host Specificity in the Ectomycorrhizal Genus Leccinum. New Phytologist 163:201–215. Dentinger, B. T. M., J. F. Ammirati, E. E. Both, D. E. Desjardin, R. E. Halling, T. W. Henkel, P-A. Moreau, E. Nagasawa, K. Soytong, A. F. Taylor, R. Watling, J-M. Moncalvo, and D. J. McLaughlin. 2010. Molecular Phylogenetics of Porcini Mushrooms (Boletus section Boletus). Molecular Phylogenetics and Evolution 57:1276–1292. Gelardi, M., G. Simonini, E. Ercole, and A. Vizzini. 2014. Alessioporus and Pulchroboletus (Boletaceae, Boletineae), Two Novel Genera for Xerocomus ichnusanus and X. roseoalbidus from the European Mediterranean Basin: Molecular and Morphological Evidence. Mycologia 106 (6): 1168–1187. Grand, L. F., and A. H. Smith. 1971. A Previously Unrecognized Southern species of Boletus. Mycologia 63:114–117. Grund, D. W., and K. A. Harrison. 1976. Nova Scotian Boletes. J. Cramer, Vaduz, Germany. 283 pp. Halling, R. E. 1983. Boletes Described by Charles C. Frost. Mycologia 75:70–92. Halling, R. E., A. R. Bessette, and A. E. Bessette. 2015. Aureoboletus innixus. Index Fungorum 258. At http://www.indexfungorum.org. Halling, R. E., N. Fechner, M. Nuhn, T. Osmundson, K. Soytong, D. A. M. Binder, and D. Hibbett. 2015. Evolutionary Relationships of Heimioporus and Boletellus (Boletales), with an Emphasis on Australian Taxa Including New Species and New Combinations in Aureoboletus, Hemileccinum, and Xerocomus. Australian Systematic Botany 28:1–22. Halling, R. E., M. E. Nuhn, N. A. Fechner, T. W. Osmundson, K. Soytong, D. Arora, D. S. Hibbett, and M. Binder. 2012. Sutorius: A New Genus for Boletus eximius. Mycologia 104 (4): 951–961. Kibby, G. 2012. British Boletes with Keys to Species. The author, London. 79 pp. Klofac, W. 2010. The Genus Aureoboletus, a World-wide survey. A Contribution to a Monographic Treatment. Österreichische Mykologische Gesellschaft (Osterr. Z. Pilzk.) 19:133–174.

References and Recommended Resources  ·  459 Korhonen, M. 1995. New Boletoid Fungi in the Genus Leccinum from Fennoscandia. Karstenia 35:53–66. Korhonen, M., J. Hyvönen, and T. Ahti. 1993. Suillus grevillei and S. clintonianus (Gomphidiaceae), Two Boletoid Fungi Associated with Larix. Karstenia 33:1–9. Kretzer, A., Y. Li, T. Szaro, and T. D. Bruns. 1996. Internal Transcribed Spacer Sequences from 38 Recognized Species of Suillus sensu lato: Phylogenetic and Taxonomic Implications. Mycologia 88 (5): 776–785. Kuan, Z., W. Gang, and Z. L. Yang. 2014. A New Genus, Rubroboletus, to Accommodate Boletus sinicus and Its Allies. Phytotaxa 188 (2): 61–77. Kuo, M., A. S. Methven, A. M. Minnis, and R. E. Halling. 2013. Studies of American Macrofungi 1. Validation of Lactarius rubidus comb. nov. and Leccinellum quercophilum sp. nov. Mycotaxon 124:323–332. Lamoureux, Y., and J. Després. 1997. Champignons du Québec Tome 1 Les Bolets. Cercle des mycologues de Montréal, Montreal. 115 pp. Lannoy, G., and A. Estades. 1995. Monographie des Leccinum d’Europe. La Roche-surForon, France, Federation Mycologique Dauphine-Savoie, 229 pp. Li, Y-C., B. Ortiz-Santana, N-K. Zeng, B. Feng, and Z. L. Yang. 2014. Molecular Phylogeny and Taxonomy of the Genus Veloporphyrellus. Mycologia 106 (2): 291–306. Murrill, W. A. 1948. Florida Boletes. Lloydia 11:21–35. Ortiz-Santana, B., A. E. Bessette, and O. L. McConnell. In press. A New Bolete Species from North Carolina. Mycotaxon. Ortiz-Santana, B., and E. E. Both. 2011. A Preliminary Survey of the Genus Buchwaldoboletus. Bulletin of the Buffalo Society of Natural Sciences 40:1–14. Palm, M. E., and E. L. Stewart. 1984. Suillus neoalbidipes: A New Species for Suillus albidipes. Mycologia 76 (3): 433–438. Peck, C. H. 1889. The Boleti of the United States. Bulletin of the New York State Museum 2 (8): 73–166. Roody, W. C. 2003. Mushrooms of West Virginia and the Central Appalachians. Univ. Press of Kentucky, Lexington. 522 pp. Singer, R. 1945a. The Boletineae of Florida with Notes on Extralimital Species. I. The Strobilomycetaceae. Farlowia 2:97–141.    . 1945b. The Boletineae of Florida with Notes on Extralimital Species. II. The Boletaceae (Gyroporoideae). Farlowia 2:223–303.    . 1945c. New Boletaceae from Florida (a Preliminary Communication). Mycologia 37:797–799.    . 1947. The Boletineae of Florida with Notes on Extralimital Species. III. American Midland Naturalist 37:1–135. Singer, R., J. Garcia, and L. D. Gomez. 1990. The Boletineae of Mexico and Central America I and II. Cramer, Berlin. 73 pp.    . 1991. The Boletineae of Mexico and Central America III. Cramer, Berlin. 128 pp.    . 1992. The Boletineae of Mexico and Central America IV. Cramer, Berlin. 62 pp. Singer, R., and R. Williams. 1992. Some Boletes from Florida. Mycologia 84:724–728. Smith, A. H., and H. D. Thiers. 1964. A Contribution toward a Monograph of North American Species of Suillus. The authors, Ann Arbor, MI. 116 pp.

460  ·  References and Recommended Resources    . 1971. The Boletes of Michigan. Univ. of Michigan Press, Ann Arbor. 428 pp. Snell, W. H., and E. A. Dick. 1957. A Glossary of Mycology. Harvard Univ. Press, Cambridge, MA. 181 pp. Šutara, J. 2000. Xerocomus sensu lato in the Light of the Present State of Knowledge. Czech Mycology 60 (1): 29–62. Treu, R. 1993. Studies on Boletus Section Luridi. Mycotaxon 47:367–377. Vizzini, A. 2014. Imleria. Index Fungorum 147. At http://www.indexfungorum.org. Watling, R., and A. E. Hills. 2005. British Fungus Flora: Agrics and Boleti 1/Boletes and Their Allies. Royal Botanic Gardens, Edinburgh. 172 pp. Weber, N. S., and A. H. Smith. 1985. A Field Guide to Southern Mushrooms. Univ. of Michigan Press, Ann Arbor. 280 pp. Williams, R. S. 1980. A Key to Some Boletes of the Deep South. McIlvainea 4 (2): 40–46. Wolfe, C. B. 1979. Austroboletus and Tylopilus Subgenus Porphyrellus with Emphasis on North American Taxa. Cramer, Vaduz, Germany. 148 pp.    . 1987. Studies in the Genus Xanthoconium (Boletaceae). I. New Species and a New Combination. Canadian Journal of Botany 65:2142–2146. Zhao, K., G. Wu, R. E. Halling, and Z. L. Yang. 2015. Three New Combinations of Butyriboletus (Boletaceae). Phytotaxa 234 (1): 51–62. Magazines and Journals

FUNGI Magazine: www.fungimag.com. Mushroom: The Journal of Wild Mushrooming: www.mushroomthejournal.com. Field Mycology, a publication of the British Mycological Society: http://www.britmy colsoc.org.uk/society/publications/journals/field-mycology. McIlvainea: Journal of American Amateur Mycology: http://www.namyco.org/mcil vainea.php. Websites

MushroomExpert.com: www.mushroomexpert.com. Mushroom Observer: mushroomobserver.org. European Boletes: boletales.com. Index Fungorum: www.indexfungorum.org. Dr. Roy Halling, New York Botanical Garden Boletineae Surveys and Revisions: www .nybg.org/science/scientist_profile.php?id_scientist=6.

Index to Common Names

Abruptly Bulbous Sand Bolete, 84 Acidic Golden-pored Bolete, 60 American Slippery Jack, 319 Appalachian Yellow-brown Bolete, 367 Ashtray Bolete, 261 Ash Tree Bolete, 82 Aspen Bolete, 251 Aspen Scaber Stalk, 251 Baggy-veiled Suillus, 328 Bay Bolete, 227 Bearded Bolete, 323 Beveled-cap Bolete, 371 Billie’s Bolete, 95 Birch Bolete, 265 Bitter Beech Bolete, 186 Bitter Bolete, 375 Black Velvet Bolete, 366 Blue-staining Slippery Jack, 360 Booted Suillus, 328 Boreal Bolete, 332 Both’s Bolete, 172 Brick-red Conifer Scaber Stalk, 274 Brown and Yellow Bolete, 165 Brown Birch Bolete, 265 Burnt Orange Bolete, 374 Butterball, 333 Cep, 101 Chalk-white Bolete, 86 Chestnut Bolete, 209 Chicken-fat Suillus, 319 Chrome-footed Bolete, 217

Clustered Brown Bolete, 62 Common Scaber Stalk, 265 Conical Shaggy-capped Bolete, 407 Corrugated Bolete, 421 Dark Bolete, 389 Double-veiled Bolete, 282 False Black Velvet Bolete, 369 Flaming Gold Bolete, 291 Foxy Bolete, 274 Fragrant Bolete, 346 Frost’s Bolete, 205 Gertrude’s Bolete, 108 Ghost Bolete, 248 Gilled Bolete, 286 Golden-pored Bolete, 60 Graceful Bolete, 70 Granulated Slippery Jack, 333 Gray Bolete, 297 Half Yellow Powdery Bolete, 174 Hollow-stalked Larch Suillus, 326 Hollow-stemmed Tamarack Jack, 326 Horton’s Bolete, 421 King Bolete, 101 Larch Bolete, 132 Larch Suillus, 335 Lilac Bolete, 153 Lilac-brown Bolete, 364 Little Bitter Bolete, 384 Lurid Bolete, 312

Mahogany Bolete, 115 Noble Bolete, 121 Old Man of the Woods, 309 Ornate-stalked Bolete, 300 Painted Bolete, 356 Painted Suillus, 356 Pale Bitter Bolete, 391 Pale Rose Bolete, 129 Pale Violet Bitter Bolete, 402 Pallid Bolete, 131 Parasitic Bolete, 288 Parchment Bitter Bolete, 382 Patriotic Bolete, 133 Peck’s Bolete, 182 Peppery Bolete, 195 Piedmont Bolete, 189 Pineapple Bolete, 76 Pine Cone Bolete, 309 Pine-loving King Bolete, 137 Poor Man’s Slippery Jack, 360 Porcini, 101 Powdery Sulphur Bolete, 295 Purple-red Bolete, 198 Ravenel’s Bolete, 295 Red and Olive Velvet-cap, 117 Red and Yellow Bolete, 74 Red Bog Bolete, 132 Red-cracked Bolete, 416 Reddish-brown Bitter Bolete, 395 Red Gyroporus, 213 Red-mouth Bolete, 315 Red-speckled Bolete, 428 Roody’s Bolete, 141

461

462  ·  Index to Common Names Rosy Brick-red Bolete, 143 Rosy-footed Hemlock Bolete, 192 Rosy-footed Oak Bolete, 191 Roxanna’s Bolete, 67 Russell’s Bolete, 207 Rusty Iron Bolete, 105 Sand-loving Leccinum, 274 Scarlet-stemmed Bolete, 186 Sensitive Bolete, 150 Shaggy-stalked Bolete, 219 Short-stalked Suillus, 325 Short-stemmed Slippery Jack, 325 Showy Bolete, 155 Showy Orange Bolete, 90

Showy Reddish Brown Bolete, 181 Slender Red-pored Bolete, 277 Slim Jack, 336 Slippery Jack, 342 Slippery Jill, 348 Snell’s Bolete, 267 Snell’s Leccinum, 267 Sour-cap Suillus, 318 Spicy Suillus, 346 Staining Blue Bolete, 211 Steinpilz, 101 Suede Bolete, 165

Variable Brown-net Bolete, 400 Variable-stalk Bolete, 167 Violet-gray Bolete, 388

Tamarack Jack, 335 Two-toned Bolete, 74

Yellow-cracked Bolete, 165 Yellowfoot Bolete, 217

White Birch Bolete, 248 White Bog Bolete, 248 White Coarsely-ribbed Bolete, 71 White-stemmed Bolete, 344 White Suillus, 345 Whitey’s Bolete, 430 Wormy Bolete, 169 Wrinkled Leccinum, 262

Index to Scientific Names

Each general genus name is set in boldface small capital letters, with bold page numbers. Current binomial names for principal species described and illustrated in this book are set in regular (nonitalic) type face, with bold page numbers. Synonyms, partially described species, and cross-references are shown in italic type face, with lightface page numbers. abruptibulbus, Boletus, 84 acidus, Suillus, 318 acidus var. intermedius, Suillus, 318 aeruginascens, Fuscoboletinus, 361 aeruginascens, Suillus, 361 affine, Xanthoconium, 227, 409, 412 affine var. affine, Xanthoconium, 409 affine var. maculosus, Xanthoconium, 409 affine var. reticulatum, Xanthoconium, 409 affinis, Boletus, 409 albellum, Leccinellum, 238, 249 albellum, Leccinum, 238 albidipes, Suillus, 344 albisulphureus, Boletus, 86, 414 albisulphureus, Gyroporus, 86 alboater, Boletus, 366 alboater, Porphyrellus, 366 alboater, Tylopilus, 366, 370, 379 Alessioporus, 55 alutaceus, Boletus, 67, 87, 165 alutaceus, Xerocomus, 87 americanus, Boletus, 319 americanus, Suillus, 319

americanus var. americanus, Boletus, 319 americanus var. reticulipes, Boletus, 319 americanus var. reticulipes, Suillus, 319 ananas, Boletellus, 76 ananas, Boletus, 76 appalachiensis, Tylopilus, 367 arenicola, Leccinum, 274 atkinsonianus, Boletus, 289 atkinsonianus, Ceriomyces, 289 atkinsonianus, Pulveroboletus, 289 atkinsonii, Boletus, 89, 168 atratus, Tylopilus, 366 atronicotianus, Tylopilus, 366, 369 atrostipitatum, Leccinum, 271 aurantiacum, Leccinum, 274 aurantiacum var. vulpinum, Leccinum, 274 aurantiosplendens, Boletus, 90, 93, 292 aureissimus, Ceriomyces, 92 aureissimus var. aureissimus, Boletus, 91, 92, 95, 294, 301 aureissimus var. castaneus, Ceriomyces, 93, 95 Aureoboletus, 57

auriflammeus, Boletus, 291 auriflammeus, Pulveroboletus, 91, 291 301 auripes, Boletus, 93, 94 auripes var. aureissimus, Boletus, 92 auriporus, Aureoboletus, 60, 62, 85 auriporus, Boletus, 60 Austroboletus, 68 badia, Imleria, 65, 227 badiceps, Boletus, 371 badiceps, Tylopilus, 371, 377, 381, 401 badius, Boletus, 227 badius, Xerocomus, 227 balloui, Boletus, 374 balloui, Rubinoboletus, 374 balloui, Tylopilus, 374 balloui var. fuscatus, Boletus, 374 ballouii, Gyrodon, 374 Baorangia, 72 betula, Austroboletus, 219 betula, Frostiella, 219 betula, Heimiella, 208, 219 betula, Heimioporus, 219 bicolor, Baorangia, 74, 129, 142, 149, 151, 183, 231, 234, 235 bicolor, Boletus, 74, 229 bicoloroides, Boletus, 75

463

464  ·  Index to Scientific Names bicolor var. borealis, Boletus, 229 bicolor var. subreticulatus, Boletus, 74 billieae, Boletus, 95 Boletellus, 76 Boletinellus, 81 boletinoides, Phylloporus, 285 Boletus, 83 borealis, Lanmaoa, 97, 107, 229 Bothia, 171 bovinus, Boletus, 321 bovinus, Suillus, 321, 334, 344 bovinus var. viridocaerulescens, Suillus, 321 bresadolae var. bresadolae, Suillus, 323 bresadolae var. flavogriseus, Suillus, 323 brevipes, Boletus, 325 brevipes, Rostkovites, 325 brevipes, Suillus, 325, 343, 344 brevipes var. subgracilis, Suillus, 325 brunneus, Butyriboletus, 57, 156, 181 Buchwaldoboletus, 173 Butyriboletus, 180 caespitosus, Boletus, 62 Caloboletus, 185 calopus, Boletus, 186 calopus, Caloboletus, 186, 191, 193 calopus var. frustosus, Boletus, 186 campestris, Boletus, 223 campestris, Hortiboletus, 160, 223, 225 carminipes, Boletus, 231 carminipes, Lanmaoa, 231 carminiporus, Boletus, 97, 107, 303, 314 castanella, Bothia, 172

castanellus, Boletinus, 172 castanellus, Suillus, 172 castanellus, Xerocomus, 172 castaneus, Boletus, 209 castaneus, Gyroporus, 70, 209, 214 castaneus f. purpurinus, Boletus, 213 cavipes, Boletinus, 326 cavipes, Boletus, 326 cavipes, Suillus, 133, 326 Chalciporus, 193 chalybaeum, Leccinum, 245, 254, 266 chattoogaense, Xanthoconium, 410 chippewaensis, Boletus, 98, 102 chromapes, Boletus, 217 chromapes, Harrya, 217 chromapes, Leccinum, 217 chromapes, Tylopilus, 217 chrysenteroides, Boletellus, 78, 80, 81, 417 chrysenteroides, Boletus, 78 chrysenteron, Boletus, 416 chrysenteron, Xerocomellus, 79, 106, 165, 416, 419 chrysenteron, Xerocomus, 416 citrinovirens, Boletus, 105 clintonianus, Suillus, 335 confusus, Strobilomyces, 307, 309 conica var. conica, Fistulinella, 407 conica var. reticulata, Fistulinella, 407 conicus, Mucilopilus, 407 conicus, Tylopilus, 407 conicus, Veloporphyrellus, 407 cothurnatus, Suillus, 328, 348, 438 crocipodium, Leccinellum, 240 242, 244, 263 curtisii, Boletus, 294 curtisii, Pulveroboletus, 294

cyaneotinctus, Porphyrellus, 396 cyaneotinctus, Tylopilus, 396 cyanescens, Boletus, 211 cyanescens, Gyroporus, 211, 213 cyanescens var. cyanescens, Gyroporus, 211 cyanescens var. violaceotinctus, Gyroporus, 211 Cyanoboletus, 200 decipiens, Boletinus, 330 decipiens, Boletus, 330 decipiens, Suillus, 173, 330, 341, 357 discolor, Boletus, 277 dryophilus, Strobilomyces, 307, 309 dupainii, Boletus, 302 dupainii, Rubroboletus, 302 dupainii, Suillellus, 302 durhamensis, Boletus, 99, 399 edulis, Boletus, 98, 99, 111, 101, 122, 123, 138, 153, 158, 168, 376, 409 edulis f. pinicola, Boletus, 137 edulis ssp. subcaerulescens, Boletus, 157 erythropus f. michiganensis, Boletus, 277 erythropus ssp. discolor, Boletus, 277 eximium, Leccinum, 364 eximius, Boletus, 364 eximius, Sutorius, 364 eximius, Tylopilus, 364 Exsudoporus, 202 fagicola, Boletus, 161, 170 fairchildianus, Boletus, 104, 144, 146 felleus, Boletus, 375 felleus, Tylopilus, 102, 168, 375, 385, 395, 399, 401 felleus var. minor, Boletus, 375

Index to Scientific Names  ·  465 felleus var. minor, Tylopilus, 375 felleus var. uliginosus, Tylopilus, 375 ferrugineus, Boletus, 105, 165, 377, 417 ferrugineus, Tylopilus, 372, 377, 381 ferrugineus, Xerocomus, 105 ferrugineus subsp. vinaceogriseus, Tylopilus, 377 firmus, Boletus, 189 firmus, Caloboletus, 189 flammans, Boletus, 97, 107, 305, 314 flavidus, Suillus, 336 flavissimus, Boletus, 57 flavostipitatum, Leccinum, 270 floccopus, Strobilomyces, 309 flocculosipes, Boletellus, 78 floridanus, Boletus, 204 floridanus, Exsudoporus, 204, 206 floridanus, Suillellus, 204 foliiporus, Phylloporus, 286 fraternus, Boletus, 225 Frostiella, 206 frostii, Boletus, 205 frostii, Exsudoporus, 204, 205 frostii ssp. floridanus, Boletus, 204 frustosus, Boletus, 186 fumosipes, Porphyrellus, 396 fumosipes, Tylopilus, 396

gracilis, Tylopilus, 70 gracilis var. flavipes, Austroboletus, 70 gracilis var. gracilis, Austroboletus, 70 210, 214 gracilis var. pulcherripes, Austroboletus, 70 granulatus, Boletus, 333 granulatus, Suillus, 322, 325, 333, 344 granulatus subsp. leptopus, Suillus, 333 granulatus subsp. snellii, Suillus, 333 grevillei, Suillus, 323, 335, 343 grevillei var. clintonianus, Suillus, 335 grisellus, Boletinus, 338 grisellus, Fuscoboletinus, 338 grisellus, Suillus, 338 griseocarneus, Tylopilus, 379 griseonigrum, Leccinum, 270 griseum, Leccinellum, 240, 242, 244, 263 griseum, Leccinum, 242 griseus, Retiboletus, 297, 301 griseus, Xerocomus, 297 griseus ssp. pinicaribaeae, Boletus, 297 griseus var. fuscus, Retiboletus, 298 griseus var. griseus, Boletus, 297 Gyroporus, 208

gentilis, Aureoboletus, 60 gertrudiae, Boletus, 108, 122 glabellus, Boletus, 227 glandulosus, Boletinus, 332 glandulosus, Fuscoboletinus, 332 glandulosus, Suillus, 332, 352, 354 glutinopallens, Leccinum, 247, 249 gracilis, Porphyrellus, 70

harrisonii, Boletus, 223, 225 Harrya, 215 Heimioporus, 217 hemichrysus, Boletus, 174 hemichrysus, Buchwaldoboletus, 174, 178 hemichrysus, Pulveroboletus, 174 hemichrysus f. sphaerocephalus, Boletus, 179 Hemileccinum, 220

hemixanthus, Xerocomus, 125, 423 hirtellus, Boletus, 340 hirtellus, Suillus, 318, 330, 340 holopus, Boletus, 248 holopus, Leccinum, 238, 248, 248 holopus var. americanum, Leccinum, 248 holopus var. holopus, Leccinum, 248 Hortibolitus, 221 hortonii, Boletus, 421 hortonii, Xerocomus, 221, 421 huronensis, Boletus, 99, 102, 110, 421 hypocarycinus, Boletus, 310 hypocarycinus, Suillellus, 278, 310, 313 hypoxanthus, Boletus, 422 hypoxanthus, Xerocomus, 125, 422 illudens, Boletus, 426 illudens, Xerocomus, 426 Imleria, 226 indecisus, Boletus, 380 indecisus, Tylopilus, 376, 380, 399 indecisus var. subpunctipes, Tylopilus, 380 inedulis, Boletus, 191 inedulis, Caloboletus, 171, 187, 189, 191, 193 inedulis, Ceriomyces, 191 innixus, Aureoboletus, 60, 62, 67 innixus, Boletus, 62 innixus, Pulveroboletus, 62 insigne, Leccinum, 251, 257, 258, 272, 275, 374 insolens, Leccinum 253 insolens var. brunneo-maculatum, Leccinum, 253 insolens var. insolens, Leccinum, 253

466  ·  Index to Scientific Names intermedius, Boletellus, 419 intermedius, Suillus, 318 intermedius, Tylopilus, 131, 382, 386, 392, 393 intermedius, Xerocomellus, 79, 81, 417, 419

minor, Boletus, 384 minor, Tylopilus, 384 montaltoense, Xanthoconium, 368, 414 morrisii, Boletus, 428 morrisii, Xerocomus, 428

lactifluus, Suillus, 333 Lanmaoa, 228 laricinus, Fuscoboletinus, 361 laricinus, Suillus, 361 Leccinellum, 236 Leccinum, 245 leucomycelinus, Phylloporus, 286, 287 lignicola, Boletus, 177 lignicola, Buchwaldoboletus, 175, 177, 179 lignicola, Pulveroboletus, 177 longicurvipes, Boletus, 112 luridellus, Boletus, 114 luridiformis, Boletus, 277 luridiformis, Neoboletus, 277, 311, 313, 316 luridiformis ssp. discolor, Boletus, 277 luridus, Boletus, 312 luridus, Suillellus, 278, 312, 314 luteum, Leccinum, 256 luteus, Boletus, 342 luteus, Suillus, 336, 342

nebulosus, Tylopilus, 389 neoalbidipes, Suillus, 322, 326, 334, 344 Neoboletus, 275 nigrescens, Leccinum, 240 nobilis, Boletus, 89, 109, 121, 123, 168 nobilissimus, Boletus, 122, 123, 168, 290 nueschii, Suillus, 323

mahogonicolor, Boletus, 115, 127 melleoluteus, Boletus, 91, 296 merulioides, Boletinellus, 82, 280 merulioides, Gyrodon, 82 miniato-olivaceus, Boletus, 117, 134, 151 miniato-olivaceus var. sensibilis, Boletus, 150 miniato-olivaceus var. subluridus, Boletus, 314 miniatopallescens, Boletus, 119, 151

ochraceoluteus, Boletus, 125, 423 oliveisporus, Boletus, 116, 127, 202 oliveisporus, Ceriomyces, 127 ornatipes, Boletus, 300 ornatipes, Retiboletus, 93, 292, 300 oxydabile, Leccinum, 265 pallidoroseus, Boletus, 75, 129, 151 pallidus, Boletus, 131, 383 paluster, Boletinellus, 132 paluster, Boletinus, 132 paluster, Boletus, 132 327 paluster, Fuscoboletinus, 132 paluster, Suillus, 132 Paragyrodon, 280 parasiticus, Boletus, 288 parasiticus, Pseudoboletus, 288 parasiticus, Xerocomus, 288 patrioticus, Boletus, 133 peckii, Boletus, 182 peckii, Butyriboletus, 182 peckii, Caloboletus, 182, 434 peralbidus, Boletus, 386 peralbidus, Tylopilus, 215, 382, 383, 386, 392, 393

peralbidus var. rhodoconius, Tylopilus, 392 phaeocyanescens, Gyroporus, 211, 212 Phylloporus, 283 piceinum, Leccinum, 274 pictiformis, Boletellus, 78 pictiformis, Suillellus, 78 pictus, Boletinus, 356 pictus, Boletus, 356 pictus, Suillus, 356 piedmontensis, Boletus, 189 pinicola, Boletus, 137 pinophilus, Boletus, 137 pinorigidus, Suillus, 348 piperatoides, Chalciporus, 196 piperatus, Boletus, 195 piperatus, Chalciporus, 195 placidus, Boletus, 345 placidus, Suillus, 345 placidus f. placidus, Suillus, 345 plumbeoviolaceus, Boletus, 388 plumbeoviolaceus, Tylopilus, 364, 388, 403, 405 porosus, Boletus, 326 porphyrosporus, Porphyrellus, 389 porphyrosporus, Tylopilus, 389, 397 potteri, Leccinum, 252 projectellus, Aureoboletus, 64, 227 projectellus, Boletellus, 64 projectellus, Boletus, 64 projectellus, Ceriomyces, 64 proximus, Boletinellus, 82 proximus, Suillus, 335 pseudoboletinus, Xerocomus, 423 Pseudoboletus, 287 pseudochrysenteroides, Boletellus, 79, 80, 419 pseudogranulatus, Suillus, 325 pseudoinsigne, Leccinum, 257, 374

Index to Scientific Names  ·  467 pseudo-olivaceus, Boletus, 135 pseudopeckii, Boletus, 181 pseudopinophilus, Boletus, 102, 137, 158 pseudorubinellus, Boletus, 196 pseudorubinellus, Chalciporus, 196, 199, 200 pseudoscaber, Boletus, 389 pseudoscaber, Porphyrellus, 389 pseudoscaber, Tylopilus, 389 pseudoscaber ssp. cyaneotinctus, Porphyrellus, 396 pseudosensibilis, Boletus, 233 pseudosensibilis, Lanmaoa, 114, 233 pseudoseparans, Boletus, 153 pseudosulphureus, Boletus, 279 pseudosulphureus, Neoboletus, 57, 279 pseudosulphureus var. pallidus, Boletus, 279 Pulveroboletus, 288 pulverulentus, Boletus, 202 pulverulentus, Cyanoboletus, 127, 202 punctipes, Boletus, 346 punctipes, Suillus, 346, 358, 361 purpureofuscus, Boletus, 412 purpureorubellus, Boletus, 139 purpureum, Xanthoconium, 153, 410, 412 purpurinus, Gyroporus, 70, 210, 213 quercophilum, Leccinellum, 240, 242, 244, 263 rainisiae, Xerocomellus, xiv ravenelii, Boletus, 295 ravenelii, Pulveroboletus, 295 ravenelii, Suillus, 295 Retiboletus, 296 retipes, Retiboletus, 301

rhoadsiae, Boletus, 391 rhoadsiae, Gyroporus, 391 rhoadsiae, Tylopilus, 72, 86, 382, 383, 386, 391, 393 rhodoconius, Tylopilus, 386, 392, 392 rhodosanguineus, Boletus, 304 rhodosanguineus, Rubroboletus, 107, 304 rhodoxanthus, Phylloporus, 286, 286 rhodoxanthus, Xerocomus, 286 rhodoxanthus ssp. foliiporus, Phylloporus, 286 robustus, Boletus, 362 roodyi, Boletus, 141 roseipes, Boletus, 192 roseipes, Caloboletus, 187, 189, 191, 192 roseocrispans, Lanmaoa, 235 roseofractum, Leccinum, 265 roseolateritius, Boletus, 143 roseopurpureus, Boletus, 184 roseopurpureus, Butyriboletus, 155, 156, 184 roseoscabrum, Leccinum, 259 rotundifoliae, Leccinum, 265 roxanae, Aureoboletus, 67, 88 roxanae, Boletus, 67 roxanae, Ceriomyces, 67 roxanae, Xerocomus, 67 roxanae var. auricolor, Boletus, 67 rubelloides, Baorangia, 74 rubellus, Boletus, 225 rubellus, Hortiboletus, 140, 159, 160, 223, 225, 431 rubellus, Xerocomellus, 225 rubellus, Xerocomus, 225 rubellus subsp. bicolor, Boletus, 74 rubinellus, Boletus, 198 rubinellus, Chalciporus, 196, 197, 198, 200 rubinellus, Suillus, 198

Rubinoboletus, 374 rubissimus, Boletus, 142, 155, 156, 185, 431 rubricitrinus, Boletus, 104, 116, 145, 431 rubricitrinus, Ceriomyces, 145 rubricitrinus var. fairchildianus, Boletus, 104 rubriflavus, Alessioporus, 56, 182, 280 rubritubifer, Boletus, 199 rubritubifer, Chalciporus, 197, 199, 199 Rubroboletus, 301 rubrobrunneus, Tylopilus, 395, 403 rubroflammeus, Boletus, 107 rubropunctum, Leccinum, 261, 428 rubropunctus, Boletus, 261 rufocinnamomeus, Boletus, 146, 163 rufomaculatus, Boletus, 75, 148 rugosiceps, Boletus, 262 rugosiceps, Krombholzia, 262 rugosiceps, Krombholziella, 262 rugosiceps, Leccinum, 240, 242, 244, 257, 262, 422 russellii, Boletellus, 207 russellii, Boletus, 207 russellii, Frostiella, 65, 207, 219 salmonicolor, Suillus, 328, 348, 438 satanas var. americanus, Boletus, 189 scabrum, Leccinum, 245, 246, 260, 265 sclerotiorum, Xerocomus, 430 sensibilis, Boletus, 75, 118, 120, 129, 150, 233 sensibilis var. subviscidus, Boletus, 150

468  ·  Index to Scientific Names separans, Boletus, 109, 153 412 separans, Xanthoconium, 153 serotinus, Boletus, 349 serotinus, Fuscoboletinus, 349 serotinus, Suillus, 349, 362 sibiricus, Suillus, 320 sinuspaulianus, Fuscoboletinus, 351 sinuspaulianus, Suillus, 332, 351, 354 snellii, Krombholziella, 267 snellii, Leccinum, 267 270 sordidus, Boletus, 396 sordidus, Porphyrellus, 396 sordidus, Tylopilus, 390, 396 spadiceus, Boletus, 105 spadiceus, Xerocomus, 105 spadiceus var. furcatus, Boletus, 105 spadiceus var. gracilis, Boletus, 105 spadiceus var. rufobrunneus, Boletus, 105 speciosus, Boletus, 155, 182, 185 speciosus, Ceriomyces, 155 speciosus var. brunneus, Boletus, 181 spectabilis, Boletinus, 353 spectabilis, Boletus, 353 spectabilis, Fuscoboletinus, 353 spectabilis, Suillus, 353 sphaerocephalus, Boletus, 179 sphaerocephalus, Buchwaldoboletus, 175, 178, 179, 279 sphaerosporus, Boletus, 282 sphaerosporus, Gyrodon, 282 sphaerosporus, Paragyrodon, 282 sphaerosporus, Suillus, 282 spraguei, Boletus, 356 spraguei, Suillus, 330, 356 squarrosoides, Boletinus, 172 stramineum, Xanthoconium, 86, 413 stramineus, Boletus, 413

strobilaceus, Strobilomyces, 307, 309 Strobilomyces, 305 subalbellus, Gyroporus, 215, 414 subalbellus, Suillus, 215 subalutaceus, Suillus, 318 subaureus, Boletus, 358 subaureus, Suillus, 320, 347, 358 subcaerulescens, Boletus, 99, 102, 138, 157 subclavatipes subsp. inedulis, Boletus, 191 subdepauperatus, Boletus, 417 subflavidus, Austroboletus, 71, 219 subflavidus, Porphyrellus, 71 subflavidus, Tylopilus, 71 subfraternus, Boletellus, 159 subfraternus, Boletus, 159, 223, 225 subglabripes, Boletus, 220 subglabripes, Hemileccinum, 113, 220, 261, 422, 436 subglabripes, Leccinum, 220 subgranulosum, Leccinum, 263 subgraveolens, Boletus, 161, 170 subleucophaeum, Krombholziella, 269 subleucophaeum, Leccinum, 266, 267, 269 subluridellus, Boletus, 147, 163, 314, 316 subluridus, Boletus, 314 subluridus, Suillellus, 97, 107, 163, 313, 314 subluteus, Suillus, 348 subpunctipes, Boletus, 380 subpunctipes, Tylopilus, 380 subsensibilis, Boletus, 114 subsensibilis, Ceriomyces, 114 subtestaceum, Leccinum, 272 subtomentosus, Ceriomyces, 165 subtomentosus, Xerocomus, 165

subtomentosus var. perplexus, Boletus, 165 subtomentosus var. subtomentosus, Boletus, 106, 165, 417 subvelutipes, Boletus, 315 subvelutipes, Suillellus, 127, 163, 278, 313, 315 Suillellus, 309 Suillus, 316 sulphureus, Boletus, 279 sulphureus, Buchwaldoboletus, 179 sulphureus, Phlebopus, 179 Sutorius, 362 tabacinus, Boletus, 398 tabacinus, Ceriomyces, 398 tabacinus, Tylopilus, 100, 381, 398 tabacinus var. amarus, Tylopilus, 398 tabacinus var. dubius, Tylopilus, 398 tenax, Boletus, 426 tenax, Xerocomus, 426 testaceoscabrum, Leccinum, 271 tomentosus, Boletus, 360 tomentosus, Suillus, 318, 347, 358, 360 tomentosus var. discolor, Suillus, 360 truncatus, Boletus, 417 truncatus, Xerocomellus, 417 Tylopilus, 364 umbonatus, Suillus, 336 umbrinisquamosus, Gyroporus, 211, 213 umbrosus, Porphyrellus, 396 umbrosus, Tylopilus, 396 variipes, Boletus, 89, 122, 167 variipes var. fagicola, Boletus, 167 variipes var. variipes, Boletus, 167

Index to Scientific Names  ·  469 variobrunneus, Tylopilus, 376, 400 Veloporphyrellus, 405 vermiculosoides, Boletus, 100, 136, 161, 169 vermiculosus, Boletus, 100, 136, 161, 169, 170 versipelle, Leccinum, 252, 271

violatinctus, Tylopilus, 388, 395, 402, 405 viscidocorrugis, Boletus, 112 viscidus, Fuscoboletinus, 361 viscidus, Suillus, 339, 350, 361 vulpina, Krombholziella, 274 vulpinum, Leccinum, 252, 274

vulpinus, Boletus, 274 weaverae, Suillus, 350 weberi, Boletus, 171 williamsii, Tylopilus, 405 Xanthoconium, 407 Xerocomellus, 414 Xerocomus, 420

PHO T O C R E DI T S Except for the following, all photographs were taken by the authors. Bill Bakaitis Boletus huronensis with chemical reagents

New York Botanical Garden Dr. Roy E. Halling

Harley Barnhart Buchwaldoboletus sphaerocephalus, Chalciporus rubritubifer

John Plischke III Boletus huronensis (B), Boletus nobilissimus, Leccinum luteum (A), Suillus tomentosus (B), Tylopilus rubrobrunneus (A)

Dan Guravich Boletus aurantiosplendens, Boletus fairchildianus (C), Pulveroboletus curtisii (A), Tylopilus balloui (A), Tylopilus indecisus Roy Halling Boletus aureissimus var. aureissimus Mike Hopping Boletellus pseudochrysenteroides, Boletus harrisonii, Buchwaldoboletus lignicola (B) Josh Hutchins Frostiella russellii (A) Josh K ielsmeier-Cook Paragyrodon sphaerosporus (A, B) Michael Kuo Leccinellum quercophilum (B) Renée LeBeuf Boletus atkinsonii (A, B), Leccinum luteum (B), Suillus spectabilis (B), Suillus tomentosus (C), Owen McConnell Boletellus ananas (A), Boletus durhamensis, Butyriboletus peckii, Pulveroboletus curtisii (B), Rubroboletus dupainii, Veloporphyrellus conicus (B), Xerocomus illudens/tenax complex (C)

Noah Siegel Boletus pallidoroseus (B), Gyroporus purpurinus, Lanmaoa carminipes (A), Pulveroboletus curtisii (C) Eva Skif ic Chalciporus pseudorubinellus Mary Smiley Tylopilus williamsii (A, B, C) Walt Sturgeon Suillus grisellus (A), Suillus spectabilis (A), UB-1 Greg Thorn Suillus sinuspaulianus (A, B) The 3 Foragers Leccinum insigne (A) A ndrus Voitk Suillus bresadolae var. flavogriseus Robert Williams Xerocomus hypoxanthus Bill Yule Lanmaoa carminipes (B)

A BOU T T H E AU T HOR S

Alan E. Bessette is a professional mycologist and distinguished professor emeritus of biology at Utica College of Syracuse University. He has published numerous papers in the field of mycology and has authored or coauthored more than twenty books, including Edible Wild Mushrooms of North America (1992), Mushrooms of Northeastern North America (Syracuse Univ. Press, 1996), Mushrooms of the Southeastern United States (Syracuse Univ. Press, 2007), Milk Mushrooms of North America: A Field Identification Guide to the Genus Lactarius (Syracuse Univ. Press, 2009), and most recently Ascomycete Fungi of North America: A Mushroom Reference Guide (2014). Alan served as a consultant for the New York State Poison Control Center and was the scientific adviser to the Mid-York Mycological Society for more than twenty years. He has been the principal mycologist at national and regional forays and was the recipient of the Mycological Foray Service Award in 1987 and of the North American Mycological Association Award for Contributions to Amateur Mycology in 1992. William C. Roody has been studying and photographing fungi since 1975. He recently retired as a wildlife-diversity field biologist for the West Virginia Division of Natural Resources in Elkins, West Virginia. He has authored or coauthored several books, including North American Boletes: A Color Guide to the Fleshy Pored Mushrooms (Syracuse Univ. Press, 2000), Mushrooms of West Virginia and the Central Appalachians (2003), Mushrooms of the Southeastern United States (Syracuse Univ. Press, 2007), Macrofungi Associated with Oaks of Eastern North America (2008), Waxcap Mushrooms of Eastern North America (Syracuse Univ. Press, 2012), and most recently Tricholomas of North America: A Mushroom Field Guide (2013). Bill has taught workshops and lectured at numerous national and regional mycological forays, and he was the recipient of the North American Mycological Association Award for Contributions to Amateur Mycology in 2000. Arleen R. Bessette is a mycologist and botanical photographer. She has published several papers in the field of mycology and has authored or coauthored more than fifteen books, including North American Boletes: A Color Guide to the Fleshy Pored Mushrooms (Syracuse Univ. Press, 2000), Mushrooms of Cape Cod and the National Seashore (Syracuse Univ. Press, 2001), The Rainbow beneath My Feet: A Mushroom Dyer’s Field Guide (Syracuse Univ. Press, 2001), Waxcap Mushrooms of Eastern North America (Syracuse Univ. Press, 2012), Tricholomas of North America: A Mushroom Field Guide (2013), and most recently Ascomycete Fungi of North America: A Mushroom Reference Guide (2014). Arleen has won several awards in the North American Mycological Association’s annual photo competition, including top honors in both the documentary and the pictorial divisions. She presents programs and workshops on mycology, dyeing with mushrooms, and the culinary aspects of mycophagy both regionally and nationally.