Tricholomas of North America: A Mushroom Field Guide 9780292742345

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Tricholomas of North America

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NUMBER 68

The Corrie Herring Hooks Series

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TRICHOLOMAS of North America A MUSHROOM FIELD GUIDE

Alan E. Bessette Arleen R. Bessette William C. Roody UNIVERSIT Y OF TEX AS PRESS

Austin

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Steven A. Trudell

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Copyright © 2013 by Alan E. Bessette, Arleen R. Bessette, William C. Roody, and Steven A. Trudell All rights reserved Printed in China First edition, 2013 Requests for permission to reproduce material from this work should be sent to: Permissions University of Texas Press P.O. Box 7819 Austin, TX 78713-7819 www.utexas.edu/utpress/about/bpermission.html

Publisher’s Note: Although this book includes information regarding the edibility of the mushrooms described, it is not intended as a guide for the 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.

The paper used in this book meets the minimum requirements of ANSI/NISO Z39.48-1992 (R1997) (Permanence of Paper).

Library of Congress Cataloging-in-Publication Data Tricholomas of North America: A Mushroom Field Guide / by Alan E. Bessette . . . [et al.]. — 1st ed. p.  cm. — (The Corrie Herring Hooks series ; no. 68) Includes bibliographical references and index. ISBN 978-0-292-74233-8 (pbk. : alk. paper) — 1. Tricholoma—North America—Identification. 2. Mushrooms—North America—Identification. I. Bessette, Alan. II. Series: Corrie Herring Hooks series ; no. 68. QK629.T73N67 2012 579.6—dc23 2011048575

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Contents

Preface vii Acknowledgments ix Introduction to Tricholoma (Fries) Staude xii The Genus Tricholoma 1 Ecology of Tricholomas  4 Edibility of Tricholomas  4 Macroscopic Features Used for Identifying Tricholomas  6 Microscopic Features Used for Identifying Tricholomas  7 Notes on the Descriptions of Species  7 A Note about Names  8 Species Not Included in This Book  9 Keys to the Described Species 10 Keys for Eastern North America  12 Keys for Western North America  19 Descriptions of Species 29 Recognized Species  29 Some Additional Tricholomas  174 Excluded Species 184 Glossary 189 References 197 Photo Credits 201 Index to Common Names 203 Index to Scientific Names 205

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Preface

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ost tricholomas ar e r elatively large showy mushrooms that draw attention from mushroom hunters, nature lovers, and casual forest walkers alike. Although only a few of them are popular edibles, tricholomas have great ecological importance due to their mycorrhizal associations with many of the dominant temperate forest tree species. Unfortunately, when it comes to naming them, probably every North American field mycologist has struggled with the identification of Tricholoma species. By and large, the greatest obstacle has been a lack of readily available reference materials, especially those having illustrations in color, which are particularly valuable for distinguishing many tricholomas that are otherwise similar in size, stature, and other morphological characters. In addition, until now there has been no reasonably comprehensive treatment of the North American species. Popular mushroom guidebooks typically feature only a small number of the more common tricholomas that are encountered, and the technical literature is scattered, often difficult to obtain, and can be daunting to those not formally trained in mycology. Currently, it is not possible to provide complete or entirely definitive coverage of Tricholoma for a number of reasons. For one, compared to the situation in Europe, North American mushrooms are poorly known and nearly all groups are in need of much additional study. Unfortunately few studies are being conducted, as almost no funding is

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available to support them. Instead, the small number of North American systematic mycologists are more likely to be funded to study the mushrooms of other places. For another, mushroom taxonomy in general is in a state of rapid flux, due primarily to the impact of DNA sequencing and computerized phylogenetic analysis, and this is likely to continue for the foreseeable future. Thus, we recognize that this book represents only a first effort to describe and illustrate the tricholomas of North America and that it unavoidably will contain information that will require revision in the future. However, we look at it in much the same way that U.S. soil scientists viewed their initial efforts to classify our nation’s soils. The first published classification was called an “approximation” and, as information and understanding grew, it was replaced by six more “approximations.” Over 25 years passed before Soil Taxonomy was deemed ready to be issued without the tag of “approximation.” Hopefully, publication of this book will spur increased study of this interesting and important group of mushrooms and represent a first approximation to a future monograph of North American tricholomas. Our goal is to provide a means to identify the majority of North American tricholomas within the context of the current system of names and classification. Because of the disparity of knowledge of the mushrooms that inhabit different parts of the continent, our coverage focuses primarily on the United States and adjacent parts of Canada and much less on Mexico and northern Canada, where much less

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information is available. Nonetheless, we believe this book provides a valuable compilation of information and fills the gap between technical publications and the limited representation of tricholomas in North American mushroom field guides. Even if some of the information presented here fails to survive future revisions of Tricholoma classification and nomenclature, we feel the descriptions and especially the color images will have lasting value. We have selected the best documentary photos available from our own extensive collections, and augmented them with images graciously contributed by many of our fellow mushroom photographers. Whenever possible, we have included more than one image of a species to illustrate the wide variation exhibited by many tricholomas. It is our hope that, even with some unavoidable limitations, this book will be an important addition to the mycological literature and that it will serve its users well.

pr e fac e

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viii

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Acknowledgments

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ever a l individua ls have m a de important contributions to the production of this book, and we gratefully acknowledge them as follows:

• Dr. Clark Ovrebo for his Tricholoma research, contributions to the mycological literature, and assistance with technical issues • Dr. Joseph Ammirati for providing access to key literature • Harley Barnhart, the late Kit Scates Barnhart, Dr. Michael Beug, Dr. Bruce Horn, Sava Krstic, Renée Lebeuf, David Lewis, Dr. Clark Ovrebo, Drew Parker, John Plischke III, Christian Schwarz, Noah Siegel, Fred Stevens, Walt Sturgeon, and Michael Wood for helping us fill gaps with their excellent photographs • Walt Sturgeon for providing field collections for study, and for valuable taxonomic input • Drew Parker and two anonymous readers for reviewing the draft manuscript and providing valuable comments • Donna Mitchell for field collections for study and other valuable input • Sally Graupman for her tolerance and for providing support of many kinds, including review of parts of the manuscript • Bill Bishel, Casey Kittrell, Leslie Tingle, Renata Brunner Jass, Lindsay Starr, and the staff at University of Texas Press for making publication of the book possible and being wonderful to work with and producing a great-looking book.

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Tricholomas of North America

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Introduction to Tricholoma (Fries) Staude

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or e th a n 100 species of Tricholoma have been reported from North America. They are often referred to as trichs (pronounced either “tricks” or “trikes”), although some field guides call them cavaliers or knights. Most are relatively large and fleshy mushrooms, and some are rather colorful. Thus, they represent a conspicuous element of our mycota. Tricholomas grow on the ground near certain types of trees and typically fruit from late summer through early winter or even into spring in warmer areas. A few of them are fine edibles, while others are inedible or even poisonous. However, the edibility of the majority of tricholomas is not well known. Thus, great care should be taken by anyone inclined to consume any of them. As with any wild mushrooms gathered for food, positive identification is essential.

The Genus Tricholoma H istory

What we know as the genus Tricholoma originated with Elias Magnus Fries in his 1821–1828 publication, Systema Mycologicum. In this early version of Fries’s classification system, Tricholoma constituted one of 36 tribes (Tribe V) within the huge genus, Agaricus, which included nearly all of the gilled mushrooms. Fries defined Tricholoma as including fungi that produce white-spored terrestrial mushrooms that are fleshy and relatively robust. They lack a universal veil (so there is no volva on the stalk base or warts/ patches on the cap) and either lack a partial veil or have one that is fibrillose or floccose and disappears early, sometimes leaving remnants on the margin of the cap. The cap is hemispherical or somewhat bell shaped with a thin incurved (at least when young) margin. The gills are of unequal length and emarginate or rounded where they approach the stalk. The stalk is not smooth, but rather is fibrillose, scaly,

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or has coarse longitudinal striations formed by aggregated fibrils, and its flesh is confluent with that of the cap. The mushrooms exhibit diverse colors. Including additions in his later publications, Fries’s Tricholoma eventually included over 100 species. The name Tricholoma is derived from two Greek roots—tricho = hair and loma = fringe—referring to the fibrillose partial veil remnants found on the cap margin in several species. Interestingly, however, most tricholomas do not exhibit this feature. Another of Fries’s tribes—Tribe III, Armillaria— is also important in the history of Tricholoma. The principal difference between Fries’s armillarias and tricholomas is the existence in the former of a membranous partial veil that leaves a ring on the stalk. Not surprisingly, over time, taxonomists have done a lot of shuttling of species between Armillaria and Tricholoma, such that now a number of the Friesian armillarias reside in Tricholoma. Since 1972, Tricholoma has been considered to have become a genus in 1857 with the publication of Friedrich Staude’s Die Schwämme Mitteldeutschlands insbesondere des Herzogthurms. The decision to designate Staude as the authority for the genus was not without controversy, as some mycologists felt that Paul Kummer (Der Führer in die Pilzkunde), who had been designated the authority in 1953, was more appropriate. Still others favored the selection of Lucién Quélet (Champignons du Jura et des Vosges). Thus, each of those names appears as the authority in various publications. The type species for the genus is Tricholoma flavovirens (Persoon) S. Lundell, which currently is thought to represent the same species as T. equestre (Linnaeus) P. Kummer. A recent analysis concluded that if the two names are in fact synonyms, then T. equestre should be used because it is the earlier of the two (Deng and Yao 2005). Following Fries’s initial definition of Tricholoma, mushrooms continued to be collected and examined, and new technologies, such as microscopes, were employed, which allowed previously

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The current concept of Tricholoma differs little from Fries’s view of it—fleshy terrestrial mushrooms with a white spore-print, smooth, inamyloid spores, parallel gill trama, and (mostly) emarginate or sinuate gill attachment. However, because the concept now is based in part on microscopic features (or their absence), it is not always possible to be certain that you have a tricholoma without using a microscope. Many white-spored genera include species that could be mistaken for a tricholoma. Although it isn’t possible to list every genus that someone might consider to be a tricholoma “look-alike,” the following are the more likely ones. The descriptions include only the key features for distinguishing them from tricholomas. Amanita—universal veil present, leaving a volva on the stalk base, and often warts or patches on the cap; skirt-like ring often present; gills free or nearly so, often seceding. Cap margin often striate. Most have an elegant gestalt, not easily described, but readily appreciated with a bit of experience. Armillaria (the honey mushrooms, formerly Armillariella)—usually found on wood, often in clusters, tough black rhizomorphs often present in substrate, the typical honey colors of their fruitbodies are rare or absent in tricholomas, gills adnate to decurrent.

Calocybe—white or brightly colored fruitbodies, basidia contain siderophilous granules, spores sometimes roughened, cap cuticle sometimes cellular. Catathelasma—large, hard-textured mushrooms, adnate to decurrent gills, stalk tapered downward and with double ring, amyloid spores. Clitocybe—gills usually decurrent and fruitbodies often funnel-shaped, at least at maturity. Collybia sensu lato (including Gymnopus and Rhodocollybia)—cap margin incurved to inrolled (at least when young), stalk with a cartilaginous rind, gills adnexed to adnate, spores usually pinkish in Rhodocollybia. Dermoloma—smallish fruitbodies, cap cuticle hymeniform (filamentous in tricholomas), spores amyloid in some species. Floccularia—spores amyloid, stalks typically floccose, usually with a membranous ring. Hygrophorus (sensu stricto—the medium to large, mostly dull-colored species)—gills thick and waxylooking, adnate to decurrent, gill trama divergent. Laccaria—fruitbodies with distinctive orangebrown, pinkish brown, or purplish colors that are rare or absent in tricholomas, gills thick and waxylooking, usually pinkish brown, lilac, or purple, stalk rather tough and fibrous, spores roughened to spiny and often nearly spherical. Lepista—spore-print with pinkish tones, spores roughened. Leucopaxillus—abundant mycelial cords or mat often present at base of stalk, fruitbodies often very firm and slow to rot, spores amyloid, sometimes roughened. Lyophyllum (including Tephrocybe)—basidia contain siderophilous granules, fruitbodies usually dull colored and often with greasy appearance, many species stain black. Megacollybia—usually found on wood, gills rather broad, their edges with abundant cystidia. Melanoleuca—cap hygrophanous and often broad relative to stalk length, spores amyloid, roughened, gills usually with abundant cystidia.

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unrecognized features of mushrooms to be observed. This resulted in some Friesian tricholomas being transferred to other existing genera and others providing the basis for defining new segregate genera. Thus, some former tricholomas became members of Calocybe, Dermoloma, Lepista/Rhodopaxillus, Leucopaxillus, Lyophyllum, Melanoleuca, Porpoloma, Rhodocybe, Tephrocybe, Tricholomopsis, and Tricholosporum. Because many of the mushrooms in these genera share the essential macroscopic features of Tricholoma, they can cause confusion and produce identification challenges for mushroom hunters. C urrent Concept of T r i c h o lo m a

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Porpoloma—spores amyloid, gill edges typically with cystidia. Tricholomopsis—usually found on wood, often bright yellowish, gill edges often with abundant cystidia. Tricholosporum—spores angular, or shaped like crosses. In addition to these pale-spored genera, many species of Entoloma (sensu stricto) and Hebeloma have the same stature as a typical tricholoma. However, they are easily recognized by their salmon / pinkish brown and dull brown spores, respectively. Small brownish tricholomas can appear very much like species of Inocybe but, again, spore-color (brown in inocybes) quickly separates them. S ubdivision of T r i c h o lo m a

The genus, Tricholoma, has been divided into subgroups since its creation. In Systema Mycologicum (1821), Fries recognized four groups based on the nature of the cap surface. In Epicrisis Systematis Mycologici (1836–1838), he recognized seven groups, mostly based on the nature of the cap surface, but also including consideration of such things as season of occurrence and the fleshiness, fragility, and shape of the cap. Each of these groups was then divided further based on color, width of the gills, and discoloration of the gills. For the most part, Fries retained this latter classification in his later works, although he did make minor revisions, changed some names, and moved some species around. Most mycologists of the late 1800s and early 1900s, including Charles Horton Peck and C. H. Kauffman in the United States, divided the genus in ways very similar to Fries’s scheme. Additional macroscopic features such as odor, taste, size, and color were utilized in many of these groupings. Of course, as genus definitions evolved and species were moved (mostly) out of Tricholoma, the changing species composition of the genus was reflected in its subdivision. In parallel with the new genus alignments

based on macroscopic features, the subdivision of Tricholoma also began to reflect microscopic features, such as the presence of clamp connections, anatomy of the cap cuticle, and location of pigments in the colored species (within the hyphae versus encrusting them). Perhaps the two most widely used subdivisions of the genus are those by Marcel Bon (1984) and Rolf Singer (1986). However, we chose not to utilize formal subdivisions of the genus for this book because: there is no one widely accepted scheme for doing so; many of the features used to define subgroups are gradational and not always easy to apply, hence Kauffman’s (1918) observation that, “The grouping of this large genus is fraught with considerable difficulties.”; the existing schemes are likely to be revised substantially, or even replaced, as the results of molecular studies and morphologicalmolecular integration lead to increased understanding of the evolutionary relationships within the group; and, most of all, we don’t feel including one of the existing subdivision schemes would add significantly to readers’ ability to identify North American tricholomas. For those who are interested in this level of detail, we suggest reviewing the publications by Bon (1984, 1991), Kauffman (1918), Ovrebo (1980), Shanks (1997), and Singer (1986). H istory of the S tudy of T r i c h o lo m a in N orth A merica

Despite the fact that tricholomas are among the larger and more conspicuous of our woodland fungi, the genus historically has received very little attention in North America. Charles Horton Peck described over 60 Tricholoma species in the late 1800s and early 1900s, although many of them later were transferred to other genera. In what still is the only comprehensive treatment of the genus in North America, William Alphonso Murrill prepared the “Tricholoma” section of the North American Flora (1914a), under the

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genus names Melanoleuca and Cortinellus. Thus, many non-tricholomas were included in Murrill’s compilation. C. H. Kauffman’s Agaricaceae of Michigan (1918) included an extensive treatment of the tricholomas of the Great Lakes region. During his prolific career, Alexander Smith of course studied the genus and described a number of species. However, he did not deal with it in a comprehensive fashion. Probably the most active North American student of Tricholoma over the past few decades has been Clark Ovrebo. His master’s thesis (1973) dealt with the tricholomas of the Pacific Northwest and his PhD dissertation (1980) covered the tricholomas of the Great Lakes region. Since completing his graduate studies, Dr. Ovrebo has continued work on the genus and has published a number of articles, describing new species and clarifying some of Peck’s species concepts (see References for citations). Kris Shanks surveyed the tricholomas of California for her master’s degree and later published the bulk of her thesis as a fascicle of the Agaricales of California series (1997). A number of other mycologists, including Tim Baroni, Howard Bigelow, Roy Halling, and Scott Redhead, have made contributions to our knowledge of Tricholoma in North America. In addition, amateur mycologists have helped expand our understanding in areas such as the northeastern United States (Ed Bosman), southeastern Canada (Yves Lamoureux and Jean Després), and the Pacific Northwest (Charles Volz, Coleman Leuthy, and Drew Parker). It is our hope that publication of this book will encourage further studies of Tricholoma in North America and elsewhere, as there still is much to learn.

certain trees, shrubs, and possibly even some herbaceous plants. Thus, they are almost always found where trees are present—most often in forests and parks. Many trees, both hardwoods and conifers, are known to partner with tricholomas, including oak, beech, birch, willow, aspen, cottonwood/poplar, pine, spruce, fir, hemlock, and Douglas-fir. When found beneath isolated trees or in more or less pure stands, the association is fairly obvious, while in mixed woodlands containing a variety of ectomycorrhiza-forming trees, it can be more difficult to determine a specific relationship. However, although much remains to be learned about the specificity of mycorrhizal associations between tricholomas and host trees, knowing which species of tree the subject mushrooms were growing with can be critical for identifying them. At a minimum, it is important to know whether the nearby trees were conifers or hardwoods, and so this information should always be included in your field notes.

Edibility of Tricholomas

Tricholomas all are believed to be ectomycorrhizal, forming a nutritionally reciprocal association with

The genus Tricholoma consists mostly of relatively large fleshy mushrooms, some of which are common and often abundant. Because of this, they are likely to attract the attention of those who collect wild mushrooms for food. However tempting they may appear, relatively few members of the genus are known to be particularly good edibles. The edibility of many species is unknown, some are decidedly toxic, and one has proven deadly under certain circumstances. Add to this that edible tricholomas can be difficult to distinguish from those that are suspect or poisonous, and it becomes clear why they are not widely gathered for the table. There are exceptions such as the highly prized American matsutake (T. magnivelare), which some consider to be among the best of all edible wild mushrooms. Our matsutake is a close relative of the Japanese species (T. matsutake). Both are highly esteemed and can command steep prices in commercial markets. However, not everyone rates matsutake

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Ecology of Tricholomas

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at the top of his/her list of edible wild mushrooms, especially on first tasting it. For many, its unique flavor may be an acquired taste. In Japan, matsutake has considerable cultural significance and its almost reverent adoration stems partly from centuries of tradition. As availability of native matsutake has been declining in Japan, demand there for the American version has created a lucrative market that supports widespread commercial harvesting of the species in British Columbia, other parts of the Pacific Northwest, and, more recently, Mexico. Tricholoma portentosum is a widely distributed edible that, particularly in the East, can be quite common beneath pines from late fall to early winter, or later in the deep South. It is appreciated both for its flavor and availability well after most other edible wild mushrooms have finished fruiting for the season. Tricholoma terreum is a popular edible mushroom in Europe where it is commonly available in markets, both fresh and canned. Although the name “T. terreum” appears in many North American field guides and we have used it in this book, it is not clear whether that species actually occurs here, or whether we are misapplying the name to one or more different species. In any event, what has been referred to as T. terreum is not popular in North America, in large part because it is similar to several possibly poisonous species and one must learn to confidently recognize it. Although small for a tricholoma, it often grows in large groups and then can be collected in quantity. The gastronomic quality of Tricholoma caligatum has received mixed reviews, being described as everything from very good to disgustingly inedible. We have sampled this species a number of times in New York, West Virginia, North Carolina, and Florida and, until recently, had always found it to be intensely bitter and/or acrid and unpalatable. However, a collection made in Florida while this book was in preparation proved mild and palatable. Bitterness and inedibility also has been the typical

experience in the Pacific Northwest. However, there are other reports, from the southern Rocky Mountains for instance, of it being an excellent edible. Tricholoma caligatum is considered a highly variable species, so perhaps there is more than one species going by that name, or it may be that different populations differ in their palatability. It also is possible that at least some of the edible collections of T. caligatum actually are matsutake, T. magnivelare. Until more is known about the reason(s) for the inconsistent edibility of this mushroom, we recommend caution when trying it. There are a number of other tricholomas that are reported to be edible either in North America or Europe, but with which we have little or no first-hand experience. Examples include T. atrosquamosum, T. orirubens, T. populinum, T. scalpturatum, and T. subresplendens / T. columbetta. Fortunately, no tricholoma of which we are aware is dangerously poisonous when sampled in small amounts. However, no discussion of their edibility would be complete without addressing T. equestre / flavovirens. At one time, both in North America and Europe, this seemingly widespread mushroom was regarded by many as the best edible in the genus (especially when matsutake was classified in Armillaria), and it has a long history of use in many parts of the world. However, in 2001, the New England Journal of Medicine reported a dozen cases of poisoning, including three fatalities, that occurred in France, all following consumption of several consecutive meals that included large amounts of T. equestre / flavovirens. The victims experienced severe rhabdomyolysis, a disease that destroys muscle tissue. Similar poisonings also have been reported from Poland. For many years, we have enjoyed eating this mushroom with no ill effects and this has been the experience of many other mushroom eaters throughout North America. Nothing resembling the European incidents has been reported here. Despite this clean record, we no longer eat T. equestre, and we cannot recommend its use until further research either clarifies the reasons

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behind the puzzling occurrence of the European poisonings, or demonstrates that our North American species is different from the toxic European one. At a minimum, one should not eat large amounts of this or any other mushroom repeatedly over a short time (on the order of a few days).

Macroscopic Features Used for Identifying Tricholomas Identification of tricholomas relies heavily on size of the fruitbody and macroscopic features of the cap such as color, the presence or absence of scales or radiating fibrils on the surface, and whether it is dry, moist, or viscid. Odor and/or taste of the flesh is sometimes a defining character. The gills are white to off-white in most species but can also be grayish, buff, or yellow. Spotting, staining, or discoloring (often reddish brown) of the gills in age or after damage is a characteristic feature of many species. Although gill attachment is rarely, if ever, used to help differentiate species within Tricholoma, it is an important character for recognizing the genus. Usually it is described as being emarginate, notched, or sinuate. Unfortunately, gill attachment can be a rather variable feature and these terms have been interpreted differently from mycologist to mycologist (Leonard 2000). C. H. Kauffman (1918: 676) described the variability nearly a century ago: Theoretically, they [the gills] are always emarginate behind, but this condition varies considerably. It is true that, in the mature plant, when the pileus is fully expanded, they become either sinuate or emarginate in most cases, although a single specimen may not always be normal in this respect. When young, however, they often do not show this character clearly, but are then adnexed, roundedadnate, or adnate in such a way that they are merely a little less broad at the attached portion than they are a few millimeters from the stem, and this short distance is often marked by a straight edge rather

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than a rounded edge. . . . In old stages the gills may even become spuriously decurrent.

Thus, when attempting to determine the gill attachment of a putative tricholoma, it is best to examine multiple specimens of different ages and to allow for some latitude in interpretation. Size of the fruitbodies can vary within a species and so exact measurements are not always useful. Thus in the keys, we have used small, medium, and large as follows, based on the diameter of the mature caps: small refers to caps with diameter usually less than 5 cm; medium to caps usually between 5 and 10 cm; and large to caps usually greater than 10 cm. The stalk is usually more or less equal but may be swollen in the middle (ventricose), tapered downward, or enlarged to slightly bulbous at the base. It is fleshy or fibrous like the cap and can be solid, stuffed, or hollow. Its surface usually is dry and longitudinally striate, but also can be nearly smooth, appressedfibrillose, scaly, or slightly scurfy (at the apex only). Stalk color often is white, but it may be tinged with the cap color, or, more rarely, fully concolorous with the cap. Staining from bruising, handling, or with age is sometimes important in distinguishing between species. A number of tricholomas have a cortina. In these species, the veil often is delicate and fleeting, and may be visible only on very young specimens, leaving just a trace of fibrils as a faint ring-zone on the upper portion of the stalk or a fringed margin on the cap. A few species have a more substantial membranous veil that leaves a more or less prominent ring on the stalk. Common odors of tricholomas include farinaceous, cucumber, coal tar, and, of course, the matsutake’s cinnamon-Red-Hots-with-old-gym-socks aroma. In many cases, the odor is pleasant, although not easily describable, and we have referred to these as being fragrant. The more common tastes are mild, bitter, and farinaceous. Color-change reactions following the application of chemicals to the fruitbody (macrochemical

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reactions) are not used widely in identifying tricholomas, but they can be important in some cases. Potassium hydroxide (KOH), ammonium hydroxide (NH4OH), and paradimethylaminobenzaldehyde (PDAB) are of most use.

Microscopic Features Used for Identifying Tricholomas Microscopic features are important for distinguishing the genus Tricholoma from other genera that are similar macroscopically (see Current Concept of Tricholoma [p. 2]), and they also are of use in separating species within Tricholoma. However, although the shape and size of spores and cystidia, the presence of clamp connections, and the structure of the cap cuticle sometimes are of value, microscopic features generally are less distinctive, less variable, and less helpful for distinguishing species in Tricholoma than they are in many other genera. Consequently, our species descriptions include only those microscopic features that seem critical for identifying the species in question. Spore size and shape are given for every species. Cheilocystidia, pleurocystidia, clamp connections, and pseudoparenchymatous hypodermium are mentioned when they are present. For those seeking more complete information about microscopic characters, we recommend consulting the works listed in the References section.

Notes on the Descriptions of Species Each description consists of the following components. The Latin binomial is provided for each species. We have tried to use the currently most widely accepted names, usually following the listing in Index Fungorum (www.indexfungorum. org). Thus, the name used here might not be the same one found in other guides. In cases where this involves a shift from a widely recognized name, we Scientific name:

have included the former name in our comments. The scientific name also includes the author citation, for the convenience of readers who are interested in the origin of the species concept and name. In some instances, the author citation is simple. For example, the entry “Tricholoma muricatum Shanks” indicates that this species was described and named by Kris Shanks. Sometimes, however, the original author’s name is enclosed in parentheses and followed by the name of the person who later reclassified the mushroom. For example, “Tricholoma magnivelare (Peck) Redhead” was originally described by Charles Peck as Agaricus magnivelaris and later transferred to Tricholoma by Scott Redhead. Common name: One or more common names are provided in cases where we felt their usage was reasonably widespread and well established, or otherwise desirable. To reduce confusion, we used trich consistently as the base name, rather than cavalier or knight. In addition to common names that have appeared in other guides, we coined a few new common names where we thought this would be useful to some readers. Cap: Information about the size and shape of the cap, its disc and margin, surface texture, and color is provided. Flesh: The thickness, texture, color, and staining reaction when exposed are described in this section. Some tricholomas have distinctive odors and tastes (although often these are not easily describable in terms on which everyone would agree), and this information is noted whenever possible. It is important to remember that some mushrooms, including some tricholomas, taste hot and peppery and may irritate, burn, or numb your mouth if chewed for an extended period. However, there is negligible risk in properly tasting mushrooms. To taste mushrooms safely, place a small piece (no greater than the size of a small garden pea) on the tip of your tongue, chew it with your front teeth for a few seconds, and spit it out. If the taste initially is mild (not bitter, peppery, astringent, etc.), wait a minute and then chew a

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second small piece for 15 to 30 seconds before spitting it out. The tastes of some mushrooms are subtle or take time to develop. Gills: Attachment, width, spacing, color, bruising reactions, and additional features of the faces or edges are noted. Stalk: Information about the length, thickness, shape, texture, color, staining reactions, whether it is solid, stuffed, or hollow, and nature of the ring, when present, is provided. Microscopic features: Information about spore size and shape is presented here. Additional microscopic features, such as cap cuticle structure and cystidia, are described when present and distinctive. Occurrence: The habit (e.g., solitary, scattered, in groups or clusters), substrate, habitat, known or likely distribution, and commonness are described here. The usual fruiting period for many tricholomas is mid to late fall. However, their season extends later in areas such as California and the Gulf Coast, so typical “fall” species may appear there well into late winter or early spring. Commonness usually is described in terms of the overall distribution, and the descriptor will not be equally accurate in all parts of the species’s range. For instance, taxa listed as “occasional” may be quite common or locally abundant in some areas or years and rare or absent in others. At this time it is not possible to provide definitive information on distribution, frequency, and abundance of tricholomas because of the lack of data, historic confusion over the application of species concepts, and difficulty in identification. The state/province where each photo was taken is included in its caption. Edibility: Species known to be edible are listed as such. We recommend that none of the other species be eaten. In addition, it is important to remember that even the best “edible” mushrooms are not equally edible for everyone—some persons are sickened by species that are tolerated by the vast majority of mushroom eaters. Thus, the precautions

included in most mushroom guides should be observed, especially proceeding with moderation and caution until you know that a given species is edible for you. Comments: This section includes synonyms, comparisons with similar species or varieties, the meaning of the species epithet, and other useful or interesting information about the species described.

I n t roduct ion to T r icholom a

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A Note about Names Mushroom systematics (naming, classifying, and study of their evolutionary relationships) is in a state of rapid flux. Analysis of the large amount of DNA-sequence and other molecular data being produced is leading to many changes in how we view the evolutionary relationships among mushroomforming fungi and, as a result, in their classification and names. Fortunately, Tricholoma has been affected much less by these developments (so far, at least) than many other mushroom groups, and so the impact on this book has not been great. Nonetheless, in our effort to use the most current names, some that we’ve chosen might be unfamiliar to some North American mushroom hunters. Because name changes represent probably the biggest source of frustration for many mushroomers, where we have used a recent name we have also included the older, probably more familiar name in our comments and the index. A bigger issue for this book comes from not knowing whether the North American fungi to which European names have been applied really do belong to the same species. Few, if any, mycologists have spent enough time on both continents to have first-hand comparative knowledge of large numbers of their respective fungi, and few critical studies have been done to evaluate our use of European names. The more comprehensive studies that have been done, which unfortunately do not include Tricholoma, suggest that many of our species are essentially the same as their European counterparts. However,

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for many others, our fungi do not quite fit the European concepts and thus probably should be given new names or have existing, but unused, North American names resurrected. Although this can be confusing, it results from the fact that mycologists are developing better understandings of the evolutionary relationships among the mushroom-forming fungi.

Species Not Included in This Book In reviewing the literature on North American tricholomas, we encountered a large number of names of species that have been used rarely, if at all, and others whose application here could not be readily confirmed. In cases where the concept of the species could be ascertained clearly from the original description and seemingly represented a valid species, we included it. However, for most rarely used names, the available information was not sufficient to clearly define the species concept and, in those cases, we decided to not include the species. We have provided a list of these and other excluded species following the species descriptions. The list includes European species reported to occur in North America, but whose occurrence we were unable to substantiate.

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Keys to the Described Species

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B

ecause cur r ent species concepts in Tricholoma are based to a large extent on macroscopic characters and ecology, we chose to provide relatively simple “field” keys to the species that are described in this book, rather than more elaborate technical ones emphasizing microscopic features. However, in some cases, consideration of microscopic features is useful, or even necessary, for pinning down an identification. So for those readers with access to a compound microscope, we have included a table (p. 28) listing those species that have important distinctive features—clamp connections, cheilocystidia, and a pseudoparenchymatous hypodermium. There are two sets of keys: one for eastern North America and one for the western part of the continent. The Eastern Region extends from the Atlantic Coast to the eastern edge of the Rocky Mountains and includes most of Texas. The Western Region includes westernmost Texas, all of the Rocky Mountains, and everything from there to the Pacific Coast. Since the distributions of most tricholomas have not been firmly established, it is possible that a species currently known from only one region of the continent could occur in the other. Thus, if a collection fails to key out in one region, it may be worthwhile to try the corresponding key for the other region. The keys are mostly dichotomous, with occasional trichotomous sets of leads (in the latter case, down-arrows indicate the existence of a third lead). When keying mushrooms, selecting the appropriate lead to follow is not always easy. In such cases, a useful strategy is to take note of the place where you are uncertain so you can return and try the other lead if your original path doesn’t yield a convincing result. Unfortunately, this is likely to occur more often than usual with tricholomas, for several reasons. The fruitbodies of many species of Tricholoma exhibit a wide range of color and other features, and the appearance often changes in specimens as they age. Environmental conditions also can affect features such as surface colors and textures. For instance, a cap that is viscid when young and fresh,

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or in wet weather, may appear dry when collected. In this case, however, soil or plant debris stuck to the cap surface or touching the cap surface with a moistened fingertip may allow the feature to be accurately interpreted. In addition, many species of Tricholoma are macroscopically similar to one another, many of their features are gradational and can be difficult to interpret, and characters such as color, odor, and taste can be subjective and hard to describe with words. Finally, many species are believed to occur mainly, or even solely, with particular types of trees. However, when specimens are found in mixed woodlands, it is usually not possible to determine with which tree(s) the fungus is associated. Thus, where the key requires decisions about such features, it may be necessary to follow first one lead, then the other. To facilitate backtracking, we have included in each set of leads the number of the couplet that led to the current one. To reduce the need for backtracking, we have included several species in more than one place in the keys. No key is perfect, so it is important to remember that even when the key leads to a seemingly clear choice, comparison with the full description and photographs should always be made before accepting the resulting name. The first step in using the keys is to determine in which of the two regions the mushrooms were found—East (eastward of the Rocky Mountains) or West (the Rockies and all points to the west). Then decide in which of the following four groups your collection best fits, based on the color of the cap: 1. White, whitish, silvery or pallid. 2. Yellow, greenish yellow to olivaceous, or colored otherwise but having yellow areas, especially near the margin. 3. Ground color and/or surface features predominantly gray, dark gray, brownish gray, or blackish (at least on the disc). 4. Predominantly some shade of brown, orange, ochre, or tan.

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Then proceed to the appropriate key based on the combination of region and color-group. Keep in mind that the wide variation in cap color in many tricholomas may make it necessary to try more than one of the keys to reach a satisfactory determination. Key E-1—Eastern White-Caps (p. 12) Key E-2—Eastern Yellow-Caps (p. 13) Key E-3—Eastern Gray-Caps (p. 15) Key E-4—Eastern Brown-Caps (p. 17) Key W-1—Western White-Caps (p. 19) Key W-2—Western Yellow-Caps (p. 21) Key W-3—Western Gray-Caps (p. 23) Key W-4—Western Brown-Caps (p. 25)

Although the size of tricholoma fruitbodies can be quite variable, the general range in, for instance, the cap diameter is usually an important character. Thus rather than use exact dimensions in the keys, we have used small, medium, and large as follows, based on the diameter of the mature caps: small refers to caps with diameter usually less than 5 cm; medium refers to caps usually between 5 and 10 cm; and large refers to caps usually greater than 10 cm.

Keys for Eastern North America Key E-1—E astern W hite- C aps

fibrils or scales, ring thick, stalk with sheathing veil remnants from the base up, occurring with conifers or hardwoods, but rarely if ever willows .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 3. (2) Fruitbody usually with squatty or stocky stature, stalk base often pointed, cap and stalk scales pale to medium brown, white color usually dominant, odor reminiscent of cinnamon Red Hots candy with a variably intense component of old gym socks, occurring with conifers .. . . . . . . . . . . . . . . . . . . . . . . . .  magnivelare (p. 94) 3. Fruitbody may be stocky, but often with a relatively slender stalk that is rounded at base, cap and stalk scales medium to dark brown, brown color often dominant, odor varying from mild or slightly fragrant to highly unpleasant or, in some collections, virtually the same as that of T. magnivelare, occurring with conifers or hardwoods, especially oak  . . . . . . . . . . . . . . .  caligatum (p. 50) 4. (1) Fruitbody odor strong and perceived as unpleasant by most persons, resembling coal tar (sometimes with a fruity component) or a heavy floral scent like hyacinths or paperwhites .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  5 4. Fruitbody odor mild to strong but, if strong, then farinaceous, spicy, or otherwise, and not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  6

Cap white, whitish, silvery or pallid.

2. (1) Fruitbody small to medium-sized, cap pale gray, with radially appressed fibrils or scales, ring cottony-membranous to sub-membranous, oc curring with willows  . . . . . . .  cingulatum (p. 52) 2. Fruitbody medium-sized to large, cap white, covered to a greater or lesser extent with brown

5. (4) Occurring with hardwoods, especially oak and beech (although found with conifers in the West), fruitbody medium-sized to large, cap, gills, and stalk whitish, bruising or aging yellow to dull yellow or brown, gills close .. . . . . . . . . . . . . . . . . . . . . .  sulphurescens (p. 154) 5. Occurring with conifers, fruitbody small to medium-sized, cap whitish overall or with pale yellowish buff tones in age, not bruising yellow or brown, gills subdistant to distant .. . . . . . . . . . . . . . . . . . . . . . . . .  inamoenum (p. 86)

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1. Stalk with persistent cottony or membranous ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  2 1. Stalk lacking ring . . . . . . . . . . . . . . . . . . . . . . . . . .  4

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6. (4) Cap evenly whitish, often with yellow, tan, or gray tones, particularly when bruised or in age .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  7 6. ( ) Cap with a white background overlaid with darker-colored scales or tomentum . . . . . . . .  10 6. Cap silvery or some shade of pale gray . . . . . .  13 7. (6) Fruitbody bright silky or glistening white, at least when young and fresh . . . . . . . . . . . . . . . .  8 7. Fruitbody duller whitish, or with more obvious grayish, tan, or yellowish tones . . . . . . . . . . . . . .  9 8. (7) Fruitbody usually small, cap silky-white, lubricous, cap, gills, and stalk discoloring yellowish to ochraceous, apparently rare or not often recognized, recorded only from Michigan and New York . . . . . . . . . . . . . . . . . . .  albidum (p. 34) 8. Fruitbody small to fairly large, cap glistening white, cap and stalk base often with bluish green stains, widely distributed in the East .. . . . . . . . . . . . . . . . . . . .  subresplendens (p. 151) 9. (7) Fruitbody small to medium-sized, cap varies from entirely white to having grayish, tan, or yellow-brown tones, dry and densely fibrillose, gills discoloring dingy yellowish with blackish highly eroded edges in age, stalk at times dingy buff or pinkish at the base, widespread in the Northeast . . . . . . . . . . . . . .  serratifolium (p. 143) 9. Fruitbody medium-sized, cap shining white to creamy white or buff, at times with grayish or yellowish tints, stalk white above and orangish brown on lower half, known from Florida, but likely with a broader southern distribution .. . . . . . . . . . . . . . . . . . . . . . . .  silvaticoides (p. 145) 10. (6) Cap surface tomentose, in age usually cracking irregularly like dry mud, odor of celery or curry . . . . . . . . . . . . . . . . . . . . . . . . . .  apium (p. 36) 10. Cap surface scaly, odor farinaceous . . . . . . . . .  11

11. (10) Cap scales tan  . . . . . . . .  venenatum (p. 168) 11. Cap scales dark brownish to dark grayish or blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12 12. (11) Fruitbody usually medium-sized, cap surface wooly over the disc, breaking up into radiating fibrils and squamules toward the margin, fibrils and scales initially black over a very pale gray background, then becoming brownish black . . . . . . . . . . . . . . . .  atrosquamosum (p. 43) 12. Fruitbody usually large, surface with a whitish background overlaid with small, brownish gray to blackish, fibrillose scales that give an overall spotted appearance, typically darker over the disc where the scales are more concentrated .. . . . . . . . . . . . . . . . . . . . . . . . . .  pardinum (p. 119) 13. (6) Cap viscid, odor and taste farinaceous, cap broadly convex, often with a low umbo, frequently rugulose between the disc and margin, light medium gray with a white margin, sometimes with irregular white areas or rarely white overall, sometimes with gray streaks .. . . . . . . . . . . . . . . . . . . . . .  marquettense (p. 100) 13. Cap dry to moist, odor not distinctive, taste acrid to bitter . . . . . . . . . . . . . . . . . . . . . . . . . . . .  14 14. (13) Cap conic to obtuse with a ± prominent acute umbo, silky gray, darker on the disc, conspicuously virgate, taste very acrid or slowly bitter . . . . . . . . . . . . . . . . . . . . . .  virgatum (p. 172) 14. Cap obtuse becoming broadly convex, with or without a low umbo, evenly silvery to light gray over the disc, whitish buff near the margin, may be faintly virgate in places, taste bitter .. . . . . . . . . . . . . . . . . . . . . . . . . .  argenteum (p. 38) Key E-2—E astern Y ellow- C aps

Cap yellow, greenish yellow, to olivaceous, or colored otherwise but having yellow areas, especially near the margin.

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1. Fruitbody odor strong and perceived as unpleasant by most persons, resembling coal tar (sometimes with a fruity component) or a heavy floral scent like hyacinths or paperwhites . . . . . . . . . .  2 1. Fruitbody odor mild to strong but, if strong, then farinaceous, soap-like, or otherwise, and not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 2. (1) Cap greenish yellow when young, becoming yellowish buff to yellowish tan, at times with a brownish disc, flesh whitish .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  odorum (p. 114) 2. ( ) Cap whitish to cream or pale yellow, cap, gills, and stalk bruising or discoloring yellowish to buff, flesh whitish .. . . . . . . . . . . . . . . . . . . . . .  sulphurescens (p. 154) 2. Cap and stalk pale yellow to sulfur-yellow with reddish brown streaks, fibrils, or spots, flesh yellow . . . . . . . . . . . . . . . . . .  sulphureum (p. 157) 3. (1) Base of the stalk often pinkish orange, odor mild or like non-perfumed soap, cap color highly variable, pale greenish yellow, light green to grayish green, grayish olive, lead-gray, bluish gray, greenish gray, or grayish brown to brown, cap surface slippery when moist, occurring mainly with conifers  . . . . .  saponaceum (p. 134) 3. Base of the stalk usually not pinkish orange, odor mild to farinaceous, cap less variable in color, usually with yellow tones clearly dominant, cap surface not slippery when moist, occurring with conifers or hardwoods . . . . . . .  4 4. (3) Cap viscid, at least when young and fresh .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  5 4. Cap dry to moist but not viscid . . . . . . . . . . . . . .  6

5. Gills and stalk white to whitish with yellowish flushes, especially in age, cap distinctly coated with dark brownish to blackish radiating fibrils .. . . . . . . . . . . . . . . . . . . . . . . . .  sejunctum (p. 140) 6. (4) Flesh with acrid taste (may be slow to develop) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  7 6. Flesh with mild to farinaceous taste . . . . . . . . . .  8 7. (6) Occurs mainly with conifers, cap often develops a grayish cast  . . . . . . . .  palustre (p. 117) 7. Occurs with beech and oak, cap less likely to develop a grayish cast  . . . . . . . . .  aestuans (p. 32) 8. (6) Cap smooth or, at most, slightly tomentose or scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  9 8. Cap radially streaked with brownish, grayish, or blackish fibrils  . . . . . . . . . . . . . . . . . . . . . . . . . . . .  10 9. (8) Occurring with conifers in the Northeast .. . . . . . . . . . . . . . . . . . . . . .  fumosoluteum (p. 76) 9. Occurring with hardwoods, especially oak, in Florida or other parts of the Southeast .. . . . . . . . . . . . . . . . . . . . . . . . .  floridanum (p. 64) 10. (8) Gills white to whitish, at least when young .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  11 10. Gills buff to yellowish, yellowish green, green ish, olive, or grayish . . . . . . . . . . . . . . . . . . . . . . .  12 11. (10) Fibrils on cap dark brownish to blackish, radiating from the dark brownish to blackish matted fibrillose disc, gills developing yellowish flushes . . . . . . . . . . . . . . . . . . .  sejunctum (p. 140) 11. Fibrils on cap yellowish, disc smooth, fibrils mostly near margin, gills remaining white .. . . . . . . . . . . . . . . . . . . . . . . . .  subluteum (p. 149)

5. (4) Gills and stalk uniformly pale yellow to sulfur-yellow, cap may be slightly scaly, but not distinctly fibrillose . . . . . . . . . . .  equestre (p. 60) 5. ( ) Gills and stalk white to whitish, cap with golden brown to reddish brown fibrils or small scales . . . . . . . . . . . . . . . . . . .  intermedium (p. 89)

12. (10) Cap predominantly dark brownish gray or bluish gray overall or sometimes dull yellowish brown over the disc, bruising yellow .. . . . . . . . . . . . . . . . . . . .  luteomaculosum (p. 92) 12. Cap predominantly yellowish . . . . . . . . . . . . . .  13

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13. (12) Occurring with conifers, especially pine, cap dark brownish gray to dark green on the umbo or overall at first, becoming yellowish green to pale yellowish green or grayish yellow with darker streaks, then fading to yellowish buff overall, cap margin, gills, and stalk often with pinkish orange flushes in age .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .  davisiae (p. 56) 13. Occurring with hardwoods, especially beech, cap dark buff-brown, light golden brown, or dark olive-brown in the center, dull yellow to yellowish buff on the margin, cap margin, gills, and stalk lack pinkish orange flushes .. . . . . . . . . . . . . . . . . . . . . . . . .  subaureum (p. 148) Key E-3—E astern G ray- C aps

Ground color and/or surface features of cap predominantly gray, dark gray, brownish gray, or blackish (at least on the disc). 1. Stalk with persistent cottony or membranous ring, fruitbody small to medium-sized, cap pale gray, with radially appressed fibrils or scales, ocurring with willows  . . . . . .  cingulatum (p. 52) 1. Stalk lacking ring, although slight remnants of a cortina may be present, fruitbody small to large, cap color and surface texture various, rarely if ever occurring with willows . . . . . . . . . . . . . . . .  2 2. (1) Cap viscid, often occurring with pines or other conifers, occasionally with beech . . . . . .  3 2. Cap dry, occurring with conifers or hardwoods .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  6 3. (2) Cap virgate . . . . . . . . . . . . . . . . . . . . . . . . . . . .  4 3. Cap may have scattered fibrils, but not virgate .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  5 4. (3) Cap margin, gills, and stipe usually with yellow flushes, occurring on a variety of soils .. . . . . . . . . . . . . . . . . . . . . . .  portentosum (p. 125)

4. Fruitbody without yellow flushes, usually occur ring on sandy soils  . . . . . . . . . . .  niveipes (p. 112) 5. (3) Cap light to medium gray with a whitish margin, often wrinkled, gills white, not discoloring . . . . . . . . . . . .  marquettense (p. 100) 5. Cap dark brown, olive brown, or brownish gray, gills whitish to buff, discoloring brown on edge .. . . . . . . . . . . . . . . . .  olivaceobrunneum (p. 116) 6. (2) Fruitbody odor not distinctive, taste acrid or bitter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  7 6. Fruitbody odor and taste farinaceous, soap-like, or not distinctive . . . . . . . . . . . . . . . . . . . . . . . . .  11 7. (6) Cap dark fuscous-gray to nearly black, densely matted-fibrillose over the disc and appressed-scaly toward the margin, gill edges blackish, occurring with conifers .. . . . . . . . . . . . . . . . . . . . . . . . .  atrodiscum (p. 42) 7. Cap usually paler in color, not scaly, gill edges concolorous with faces or darkened, occurring with conifers or hardwoods . . . . . . . . . . . . . . . . .  8 8. (7) Occurring with conifers . . . . . . . . . . . . . . . .  9 8. Occurring with hardwoods . . . . . . . . . . . . . . .  10 9. (8) Cap conic to obtuse with a ± prominent acute umbo, silky gray, darker on the disc, conspicuously virgate, taste very acrid or slowly bitter . . . . . . . . . . . . . . . . . . . . . .  virgatum (p. 172) 9. Cap obtuse becoming broadly convex, with or without a low umbo, evenly silvery to light gray over the disc, whitish buff near the margin, may be faintly virgate in places, taste bitter .. . . . . . . . . . . . . . . . . . . . . . . . . .  argenteum (p. 38) 10. (8) Cap pale medium gray, the disc occasionally fuscous-gray in age, virgate only in places .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  acre (p. 30) 10. ( ) Cap with dark gray fibrils over a paler gray background, paler near the edge, virgate .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  hordum (p. 80)

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10. Cap dark gray, paler toward the margin and between the fibrils, conspicuously virgate nearly overall, at times developing bronze tones .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  pullum (p. 130) 11. (6) Base of the stalk often pinkish orange, odor mild or like non-perfumed soap, cap color highly variable, pale greenish yellow, light green to grayish green, grayish olive, lead-gray, bluish gray, greenish gray, or grayish brown to brown, cap surface slippery when moist, occurring mainly with conifers . . . .  saponaceum (p. 134) 11. Base of the stalk usually not pinkish orange, odor mild to farinaceous, cap less variable in color, often with yellow tones, cap surface not slippery when moist, occurring with conifers or hardwoods . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12 12. (11) Stalk distinctly scaly . . . . . . . . . . . . . . . . .  13 12. Stalk not, or only slightly, scaly . . . . . . . . . . . . .  14 13. (12) Gills white, often beaded with pinkish drops or streaked with pink, cap pale to dark gray, developing spot-like scales near the disc, margin slightly fringed and cottony, often beaded with pinkish drops or streaks .. . . . . . . . . . . . . . . . . . . . . . . . . .  huronense (p. 81) 13. Gills pale gray, discoloring dark gray on the edges, often staining salmon-pink in some areas, cap dark brownish gray with lighter grayish margin, with appressed or recurved scales .. . . . . . . . . . . . . . . . . . . . . . .  quarrulosum (p. 146) 14. (12) Gills having yellowish tones or discoloring yellow, especially toward the cap margin .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  15 14. Gills lacking yellow color . . . . . . . . . . . . . . . . .  16

tones, edges often dark gray to blackish, partial veil lacking . . . . . . . . . .  luteomaculosum (p. 92) 15. Cap convex with a low, broad umbo when young, wooly, then breaking up into fibrillose scales, often unevenly colored, with brownish, grayish, or black fibrils scattered over a white to pale gray background, gill edges not dark gray to blackish, evanescent cortina present .. . . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 16. (14) Cap densely interwoven-fibrillose to appressed-fibrillose, often with scattered minute scales, light gray to reddish gray over a light pink, buff, or reddish salmon background, becoming brownish gray, blackish brown, or dark reddish brown over the disc, often with brownish pink to reddish tones, paler near the margin, gills white to light buff, discoloring salmon or orange-buff, odor and taste farinaceous, occurring with hardwoods, especially oak and beech . . . . . . . . . . . . . . . . . . . . . . . .  insigne (p. 88) 16. ( ) Cap with scattered brownish gray to blackish scales over a whitish background, gills whitish, sometimes discoloring brownish, grayish, or blackish, odor and taste farinaceous, occurring with conifers and hardwoods . . . . . . . . . . . . . .  17 16. Cap ± evenly colored gray to dark gray, grayish brown, dull brown, or blackish brown, densely fibrillose to fibrillose-scaly, gills whitish to pale grayish, not discoloring, odor and taste not distinctive or slightly farinaceous, occurring with conifers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  18

15. (14) Cap brownish gray to bluish gray, sometimes dull yellowish brown over the disc, often bruising yellow, appressed-fibrillose, squamulose near margin, gills often developing yellow

17. (16) Fruitbody usually small to medium-sized, cap surface wooly over the disc, breaking up into radiating fibrils and squamules toward the margin, fibrils initially black over a very pale gray background, then becoming brownish black .. . . . . . . . . . . . . . . . . . . . .  atrosquamosum (p. 43) 17. Fruitbody usually large, cap with a whitish background overlaid by small, brownish gray to blackish, fibrillose scales that give an overall

K e ys to t h e De scr i be d Speci e s

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spotted appearance, typically darker over the disc where the scales are more concentrated .. . . . . . . . . . . . . . . . . . . . . . . . . .  pardinum (p. 119)

collections, virtually the same as that of T. magnivelare, occurring with conifers or hardwoods, especially oak  . . . . . . . . . . . . . . .  caligatum (p. 50)

18. (16) Evanescent cortina present .. . . . . . . . . . . . . . . . . . . . . . . . .  myomyces (p. 108) 18. Cortina lacking  . . . . . . . . . . . . .  terreum (p. 159)

5. (2) Fruitbody small to medium-sized, cap dull orange, orange-yellow to orange-brown, stalk below ring-zone with horizontal bands of fibrils or scales colored like cap and often in a snakeskin pattern, flesh not staining upon exposure .. . . . . . . . . . . . . . . . . . . . . . . . . .  aurantium (p. 48) 5. Fruitbody usually large and bulky, cap pale beige-brown to reddish brown, sometimes with yellowish patches, stalk with reddish brown fibrils below ring-zone, often grayish green at the base, flesh staining reddish pink when exposed .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  colossus (p. 54)

Key E-4—E astern B rown- C aps

Cap predominantly some shade of brown, orange, ochre or tan. 1. Stalk with a persistent membranous ring or hav ing a distinct ring-zone . . . . . . . . . . . . . . . . . . . .  2 1. Stalk lacking a ring or distinct ring-zone  . . . . . . .  6 2. (1) Stalk with sheathing, flaring membranous ring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 2. Stalk merely with ring-zone . . . . . . . . . . . . . . . .  5 3. (2) Cap orange-brown, often with olivaceous tones, viscid, often breaking into scales when dry, odor strongly farinaceous .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  focale (p. 66) 3. Cap with brown to cinnamon-brown fibrils or scales over a whitish background, lacking orange and olivaceous tones, dry, odor spicy, fragrant, or somewhat unpleasant . . . . . . . . . . . . . . . . . . . .  4 4. (3) Fruitbody usually with squatty or stocky stature, stalk base often pointed, cap and stalk scales pale to medium brown, white color usually dominant, odor reminiscent of cinnamon Red Hots candy with a variably intense component of old gym socks, occurring with conifers .. . . . . . . . . . . . . . . . . . . . . . . .  magnivelare (p. 94) 4. Fruitbody may be stocky, but often with a relatively slender stalk that is rounded at base, cap and stalk scales medium to dark brown, brown color often dominant, odor varying from mild or slightly fragrant to highly unpleasant or, in some

6. (1) Odor distinctive, reminiscent of soap, celery, curry, corn silks, or green corn, but not farinaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  7 6. Odor farinaceous or not distinctive . . . . . . . . . .  9 7. (6) Odor strongly of celery or curry, cap with a white background overlaid with brownish tomentum, in age usually cracking irregularly like dried mud . . . . . . . . . . . . . . . . .  apium (p. 36) 7. Odor and cap characteristics different from above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  8 8. (7) Odor mild or like non-perfumed soap, cap dry to moist, cap color highly variable, often with olivaceous or yellowish tones, flesh in stalk base pinkish orange, taste somewhat like nonperfumed soap or mild, usually occurring with conifers . . . . . . . . . . . . . . . .  saponaceum (p. 134) 8. Odor having a green-corn or corn-silks component, cap somewhat viscid at first, orange-brown to reddish brown, becoming paler in age, flesh in stalk base whitish, taste strongly farinaceous or bitter, in sandy soil under cottonwood .. . . . . . . . . . . . . . . . . . . .  fulvimarginatum (p. 72)

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9. (6) Cap viscid to some extent, at least when young . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  10 9. Cap dry to moist but not viscid at any stage .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  17 10. (9) Occurring with conifers . . . . . . . . . . . . . . .  11 10. Occurring with hardwoods . . . . . . . . . . . . . . . .  14 11. (10) Gills developing reddish brown spots or stains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12 11. Gills not developing reddish brown spots or stains, although edges may be somewhat brownish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  13 12. (11) Gills whitish to buff, cap tan to light yellowish brown, dry in age with densely mattedfibrillose disc, at times with scattered scales, occurring with pine or spruce .. . . . . . . . . . . . . . . . . . . . . . .  muskokense (p. 106) 12. ( ) Gills whitish to buff, cap chestnut-brown, paler golden brown near the margin, smooth, occurring with a variety of tree species .. . . . . . . . . . . . . . . . . . . . . .  pessundatum (p. 121) 12. Gills yellowish buff to yellowish, cap dark brown to reddish brown with paler margin and densely matted fibrils over the disc, occurring with a variety of tree species .. . . . . . . . . . . . . . . . . . . . . . .  transmutans (p. 161) 13. (11) Cap dark brown, olivaceous brown, olivegray, or brownish gray, not virgate, gills whitish to pale buff, discoloring brownish or brownish gray on the edges, occurring with pine or hard woods . . . . . . . . . . .  olivaceobrunneum (p. 116) 13. Cap dark brown to grayish brown, virgate, gills snow-white, unchanging, typically occurring with pine, especially on sandy soil .. . . . . . . . . . . . . . . . . . . . . . . . . . .  niveipes (p. 112) 14. (10) Cap smooth, with pinkish to reddish brown tones and whitish to pale yellowish margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  15

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14. Cap smooth or otherwise, lacking pinkish tones .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  16 15. (14) Cap viscid, gills white, usually developing reddish brown spots or stains, odor farinaceous, occurring on sandy soil with poplar, cotton wood, and aspen . . . . . . . . .  populinum (p. 123) 15. ( ) Cap viscid, gills pinkish buff, discoloring reddish brown to brown, odor farinaceous, occurring with beech and oak .. . . . . . . . . . . . . . . . . . . . . . . .  pudorinum (p. 128) 15. Cap dry to moist, gills white to pale yellowish, occasionally developing brownish spots or stains, odor not distinctive, occurring with conifers . . . . . . . . . . . . . .  roseoacerbum (p. 132) 16. (14) Cap reddish brown over the disc and paler yellow-brown to dull yellow on the margin, radially streaked, gills yellowish buff to buff, staining brownish, usually occurring with birch .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .  fulvum (p. 73) 16. ( ) Cap dark brown to reddish brown, gills and stalk pinkish buff, discoloring reddish brown to brown, occurring with beech and oak .. . . . . . . . . . . . . . . . . . . . . . . .  pudorinum (p. 128) 16. Cap grayish orange to brownish orange with dark brown disc, gills pale buff or yellowish white, discoloring reddish brown, usually occur ring with oak . . . . . . . . . . . . . . . . .  ustale (p. 164) 17. (9) Occurring with conifers . . . . . . . . . . . . . . .  18 17. Occurring with hardwoods . . . . . . . . . . . . . . . .  24 18. (17) Cap smooth, fibrillose, or virgate, but not wooly-felty or distinctly scaly . . . . . . . . . . . . . .  19 18. Cap wooly-felty or scaly . . . . . . . . . . . . . . . . . .  20 19. (18) Cap dark brownish to blackish green becoming yellowish green to yellowish buff in age, typically with distinct umbo, appressedfibrillose, gills and stalk whitish or greenish yellow, often developing pinkish orange stains .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .  davisiae (p. 56)

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19. Cap dark brown to grayish brown, plane or with low rounded umbo, virgate, gills snow-white, stalk snow-white, sometimes discoloring grayish or brownish when handled .. . . . . . . . . . . . . . . . . . . . . . . . . . .  niveipes (p. 112) 20. (18) Cap wooly-felty, becoming scaly . . . . . . .  21 20. Cap scaly, or fibrillose-scaly, but not wooly-felty .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  22 21. (20) Cap and gills often with yellow staining, evanescent cortina present .. . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 21. Cap and gills lack yellow staining, partial veil lacking . . . . . . . . . . . . . . . . . . . . .  terreum (p. 159) 22. (20) Cap whitish with scattered to dense tan scales that become less distinct and lighter in color toward the whitish margin, gills whitish, not staining . . . . . . . . . . . . . .  venenatum (p. 168) 22. Cap more highly colored, gills staining reddish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  23 23. (22) Cap convex to obtuse when young, dull brown, fibrillose to fibrillose-scaly, often becoming coarsely cracked with overlapping scales on the disc, stalk normally solid, often oc curring with pine . . . . . . . . .  imbricatum (p. 82) 23. Cap often conical when young, covered with bright rusty brown scales over a whitish background, cap margin cottony with slight remains of a partial veil, stalk soon hollow, often occur ring with spruce  . . . . . . . . . . . .  vaccinum (p. 165) 24. (17) Fruitbody small to medium-sized, cap wooly, then breaking up into fibrillose scales, cap and gills staining yellowish .. . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 24. Fruitbody small to large, cap fibrillose or scaly, but not wooly, cap and gills generally lack yellow stains, although gills may occasionally stain dingy golden yellow . . . . . . . . . . . . . . . . . . . . . .  25

25. (24) Odor not distinctive, taste mild or sometimes bitter, cap whitish but typically having light gray, tan, brownish gray, or yellowish brown tones, gills white, becoming dingy golden yellow near the cap margin, gill edges usually highly eroded in age .. . . . . . . . . . . . . . . . . . . . . .  serratifolium (p. 143) 25. Odor and taste farinaceous, cap more highly colored, gills not discoloring golden yellow, gill edges entire or only slightly eroded . . . . . . . . .  26 26. (25) Cap whitish with scattered to dense tan scales that become less distinct and lighter in color toward the whitish margin, gills white to buff, not discoloring . . . . . .  venenatum (p. 168) 26. Cap more highly colored, fibrillose to somewhat scaly, gills discoloring . . . . . . . . . . . . . . . . . . . . .  27 27. (26) Cap light golden brown to dark buff-brown or dark olive-brown over the center, yellowish to buff at the margin, gills light buff to pale yellow, discoloring brown on the edges, stalk light buff, sometimes with dingy buff at the base .. . . . . . . . . . . . . . . . . . . . . . . .  subaureum (p. 148) 27. Cap at first light gray to reddish gray over a pale pink to buff or reddish salmon background, becoming brownish gray, blackish brown, or dark reddish brown over the disc, gills white to light buff, discoloring salmon or orange-buff, stalk white above, bruising or discoloring salmon, yellowish salmon, or dark reddish salmon at the base . . . . . . . . . . . . . . . . . . . . . .  insigne (p. 88)

Keys for Western North America Key W-1—W estern W hite- C aps

Cap white, whitish, silvery or pallid. 1. Stalk with distinct cottony or membranous ring .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  2 1. Stalk lacking ring . . . . . . . . . . . . . . . . . . . . . . . . . .  5

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2. (1) Fruitbody small to medium-sized, cap pale gray, with radially appressed fibrils or scales, ring cottony-membranous to sub-membranous, oc curring with willows . . . . . .  cingulatum (p. 52) 2. Fruitbody medium-sized to large, cap white, at least at first, occurring mainly with conifers or sometimes with hardwoods, but rarely if ever willows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 3. (2) Cap white when young, often with a whitish bloom, soon fuscous to brown, often with olive, grayish, or pale tan areas, ring thin, not sheathing, at times faint, odor strongly farinaceous, occurring mainly in montane habitats in late spring and early summer . . . . . . . .  vernaticum (p. 170) 3. Cap white and covered to a greater or lesser extent with brown fibrils or scales, ring thick, sheathing from the stalk base up, odor spicy, fragrant, or unpleasant but not strongly farinaceous, occurring mainly in fall in a variety of forest habitats . . . . . . . . . . . . . . . . . . . . . . . . . . .  4 4. (3) Fruitbody usually with squatty or stocky stature, stalk base often pointed, cap and stalk scales pale to medium brown, white color usually dominant, odor reminiscent of cinnamon Red Hots candy with a variably intense component of old gym socks, occurring with conifers . . . . . . . . . . . . . . . . .  magnivelare (p. 94) 4. Fruitbody may be stocky, but often with a relatively slender stalk that is rounded at base, cap and stalk scales medium to dark brown, brown color often dominant, odor varying from mild or slightly fragrant to highly unpleasant or, in some collections, virtually the same as that of T. magnivelare, occurring with conifers or hardwoods, especially oak . . . . . . . . . . . . . .  caligatum (p. 50) 5. (1) Fruitbody odor strong and perceived as unpleasant by most persons, resembling coal tar (sometimes with a fruity component) or a heavy floral scent like hyacinths or paperwhites . . . . .  6

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5. Fruitbody odor mild to strong but, if strong, then farinaceous to spicy or otherwise, and not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  7 6. (5) Fruitbody medium-sized to large, cap and stalk whitish, bruising or aging yellow to dull yellow or brown, gills close .. . . . . . . . . . . . . . . . . . . . . .  sulphurescens (p. 154) 6. Fruitbody small to medium-sized, cap whitish overall or with pale yellowish buff tones in age, not bruising yellow or brown, gills subdistant to distant . . . . . . . . . . . . . . . . . . .  inamoenum (p. 86) 7. (5) Cap evenly whitish, at least when young and/or covered with duff, often with yellow, tan, brown, or gray tones, particularly when bruised or in age . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  8 7. ( ) Cap with a white background overlaid with darker-colored scales or tomentum . . . .  11 7. Cap silvery or some shade of pale gray . . . . . .  14 8. (7) Cap white, unchanging or developing pale brown tints in age, dry or moist, odor strongly farinaceous, described from the Pacific Northwest with conifers, apparently rare .. . . . . . . . . . . . . . . . . . . . . . . . .  farinaceum (p. 63) 8. Cap whitish when young and/or covered with duff, viscid, odor farinaceous or not distinctive, occurring with oak or manzanita, fairly common in California  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  9 9. (8) Cap whitish to pale violet-gray or darker when young and/or covered with duff, then becoming medium to dark violet-gray, gills sometimes developing pinkish or orangish tints .. . . . . . . . . . . . . . . . . . . .  griseoviolaceum (p. 77) 9. Cap whitish to pale orange when young and/ or covered with duff, then becoming brownish, gills often spotting or staining reddish brown .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  10

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10. (9) Occurring with oak, cap pale orange to white at first, darkening to brownish orange in streaks and patches, finally uniformly brown overall, odor and taste farinaceous .. . . . . . . . . . . . . . . . . . . . . . . . .  dryophilum (p.58) 10. Occurring with manzanita, cap white overall when young, soon pale orange and eventually brownish orange or brown over the disc, odor not distinctive, taste mild or somewhat bitter .. . . . . . . . . . . . . . . . . . . . . . . . .  manzanitae (p. 98) 11. (7) Cap surface tomentose, in age usually cracking irregularly like dry mud, odor of celery or curry . . . . . . . . . . . . . . . . . . . . . . . . . .  apium (p. 36) 11. Cap surface scaly, odor farinaceous . . . . . . . . .  12 12. (11) Cap scales tan . . . . . . .  venenatum (p. 168) 12. Cap scales dark brownish to dark grayish or blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  13 13. (12) Fruitbody usually medium-sized, cap surface wooly over the disc, breaking up into radiating fibrils and squamules toward the margin, fibrils initially black over a very pale gray background, then becoming brownish black .. . . . . . . . . . . . . . . . . . . . .  atrosquamosum (p. 43) 13. Fruitbody usually large, surface with a whitish background overlaid by small, brownish gray to blackish, fibrillose scales that give an overall spotted appearance, typically darker over the disc where the scales are more concentrated .. . . . . . . . . . . . . . . . . . . . . . . . . .  pardinum (p. 119) 14. (7) Cap conic to obtuse with a ± prominent acute umbo, silky gray, darker on the disc, conspicuously virgate, taste very acrid or slowly bitter . . . . . . . . . . . . . . . . . . . . . .  virgatum (p. 172) 14. Cap obtuse becoming broadly convex, with or without a low umbo, evenly silvery to light gray over the disc, whitish buff near the margin, may be faintly virgate in places, taste bitter .. . . . . . . . . . . . . . . . . . . . . . . . . .  argenteum (p. 38)

Key W-2—W estern Y ellow- C aps

Cap yellow, greenish yellow, to olivaceous, or colored otherwise but having yellow areas, especially near the margin. 1. Fruitbody odor strong and perceived as unpleasant by most persons, resembling coal tar (sometimes with a fruity component) or a heavy floral scent like hyacinths or paperwhites . . . . . . . . .  2 1. Fruitbody odor mild to strong but, if strong, then farinaceous, soap-like, or otherwise not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 2. (1) Cap greenish yellow when young, becoming yellowish buff to yellowish tan, at times with a brownish disc, flesh whitish .. . . . . . . . . . . . . . . . . . . . . . . . . . .  odorum (p. 114) 2. ( ) Cap whitish to cream or pale yellow, cap, gills, and stalk bruising or discoloring yellowish to buff, flesh whitish .. . . . . . . . . . . . . . . . . . . . .  sulphurescens (p. 154) 2. Cap and stalk pale yellow to sulfur-yellow with reddish brown streaks, fibrils, or spots, flesh yellow . . . . . . . . . . . . . . . . . .  sulphureum (p. 157) 3. (1) Cap viscid, at least when young, yellow to golden yellow, usually overlaid with brownish to blackish fibrils or small scales . . . . . . . . . . . . . .  4 3. Cap moist or dry but not viscid at any stage, often dull greenish yellow, yellow mixed with other colors, or yellow present only as stains .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  6 4. (3) Gills and stalk uniformly pale yellow to sulfur-yellow, cap reddish brown on the disc, may be slightly scaly, but not distinctly fibrillose .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  equestre (p. 60) 4. Gills and stalk white to whitish with yellowish flushes, especially in age, cap yellowish brown or brownish yellow and innately fibrillose overall, or yellowish with golden brown, reddish brown, or dark brownish to blackish fibrils or small scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  5 21

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5. (4) Gills and stalk white to whitish, cap with tiny golden brown to reddish brown fibrils or small scales . . . . . . . . . . . . .  intermedium (p. 89) 5. ( ) Gills and stalk white to whitish with yellowish flushes, especially in age, cap distinctly coated with dark brownish to blackish fibrils radiating from the dark disc .. . . . . . . . . . . . . . . . . . . . . . . . .  sejunctum (p. 140) 5. Gills and stalk white to whitish, often with yellowish flushes, especially in age, cap yellowish brown or brownish yellow and innately fibrillose overall, may fade to almost white in age, but retaining vestiges of yellow .. . . . . . . . . . . . . . . . . . . . . . . . . .  arvernense (p. 40) 6. (3) Base of the stalk often pinkish orange, odor mild or like non-perfumed soap, cap color highly variable, pale greenish yellow, light green to grayish green, grayish olive, lead-gray, bluish gray, greenish gray, or grayish brown to brown, cap surface slippery when moist, occurring mainly with conifers .. . . . . . . . . . . . . . . . . . . . . . .  saponaceum (p. 134) 6. Base of the stalk usually not pinkish orange, odor mild to farinaceous, cap less variable in color, usually with yellow tones clearly dominant, cap surface not slippery when moist, oc curring with conifers or hardwoods . . . . . . . . .  7 7. (6) Fruitbody small, cap golden orange-yellow with rusty orange stains over the disc and yellowish tan on the margin at first, then duller orangish or yellowish brown to light brown with darker spots and stains, taste slightly acrid, gills white or buff, becoming brownish spotted or stained, stalk slender, often somewhat rooting, cream to buff, covered with orangish fibrils or granules . . . . . . . . .  aurantio-olivaceum (p. 46) 7. Fruitbody occasionally small, but usually medium-sized to large, stalk not particularly slender, taste acrid or not, coloration different from above  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  8

K e ys to t h e De scr i be d Speci e s

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8. (7) Gills honey-yellow or greenish yellow to greenish buff, sometimes becoming buff to whitish at maturity, older specimens may develop pinkish orange stains on the edge . . . . . . . . . .  9 8. Gills whitish to pale buff or pale gray to drab, may be stained yellow near the cap margin and on the edge or have dark brownish gray to black ish edge  . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  10 9. (8) Cap yellowish green to grayish yellow, brownish on the disc, radially streaked with grayish fibrils, gills honey-yellow to greenish yellow, sometimes bruising pale brownish gray, taste slowly but strongly acrid .. . . . . . . . . . . . . . . . . . . . . . . . . . . .  aestuans (p. 32) 9. Cap dark brownish gray to dark green on the umbo or overall at first, becoming yellowish green to pale yellowish green or grayish yellow with darker streaks, then fading to yellowish buff overall, gills yellowish to greenish buff, becoming whitish to buff at maturity, cap margin, gills, and stalk often with pinkish orange flushes in age, taste farinaceous . . . . . . . . .  davisiae (p. 56) 10. (8) Cap predominantly dark brownish gray or bluish gray overall or sometimes dull yellowish brown over the disc, bruising yellow, appressedfibrillose, gills pale gray to drab, often dark brownish gray to blackish on the edge, sometimes with yellow tones near the cap margin, stalk equal or with a tapered base, white to pale gray, occasionally darkening to pale yellowish brown, occurring with hardwoods, especially oak and beech . . . . . . .  luteomaculosum (p. 92) 10. ( ) Cap with a mixture of yellow-ochre, greenish to grayish yellow, and bright yellow, often whitish on the margin, with a dark brown to yellowish brown disc, virgate, gills whitish to pale buff, staining yellow on the edge, stalk ventricose, white with pinkish or pale orange base and longitudinal fibrils that darken to pale brown, reported from coastal evergreen forest in California . . . . . . . . . . . . . . . . .  tumidum (p. 163)

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10. Cap lemon yellow, brownish yellow, or with orange-brown or olivaceous tones over the disc, margin pale yellowish white when young, may become whitish overall with irregular yellowish patches in age, appressed-fibrillose, gills white or pale gray, sometimes with yellowish tones near margin of cap, stalk nearly equal or somewhat ventricose with a pointed and sometimes bulbous base, white to cream, sometimes with yellowish, brownish, or pinkish tones in lower portion, reported with conifers in Washington and Idaho  . . . . . . . . . . . . . . . . .  arvernense (p. 40) Key W–3—W estern G ray- C aps

Ground color and/or surface features of cap predominantly gray, dark gray, brownish gray, or blackish (at least on the disc). 1. Stalk with persistent cottony or membranous ring, fruitbody small to medium-sized, cap pale gray, with radially appressed fibrils or scales, oc curring with willows . . . . . .  cingulatum (p. 52) 1. Stalk lacking ring, although slight remnants of a cortina may be present, fruitbody small to large, cap color and surface various, rarely if ever oc curring with willows . . . . . . . . . . . . . . . . . . . . . . .  2 2. (1) Cap viscid and more or less radially streaked, odor and taste not distinctive or farinaceous, sometimes with a sweet component . . . . . . . . .  3 2. Cap dry to moist but not viscid, odor and taste various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  4 3. (2) Cap white at first or when covered with duff, becoming violet-gray with darker violet-gray to blackish radiating streaks, fibrillose, odor and taste of flesh not distinctive or somewhat farinaceous, gills white at first, discoloring pinkish brown to grayish orange in patches, occurring with oak and tanoak in coastal and low-elevation forests . . . . . . . . . . . . . . .  griseoviolaceum (p. 77)

3. ( ) Cap pale gray or violet-gray on the disc, becoming dark gray with violet tones, smooth, odor of flesh strongly sweet-farinaceous reminiscent of cucumber or grapefruit, gills and stalk white, discoloring pale golden brown, occurring with tanoak and conifers in mixed evergreen forests . . . . . . . . . . . . . . . . . . . . .  mutabile (p. 107) 3. Cap gray to grayish brown or nearly black, at times with faint yellow to purple tints, with dark gray radiating fibrils, odor and taste not distinctive or somewhat farinaceous, gills and stalk white, becoming flushed yellowish in age, occurring primarily with conifers .. . . . . . . . . . . . . . . . . . . . . . .  portentosum (p. 125) 4. (2) Flesh with acrid or bitter taste, odor not distinctive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  5 4. Taste and odor farinaceous, soap-like, or not distinctive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  6 5. (4) Cap convex to broadly convex, often with a low broad umbo, pale medium gray, the disc occasionally fuscous-gray in age, virgate in places, taste of flesh acrid, sometimes slowly so, or, at times, bitter, occurring with hardwoods in the East, presence in the West uncertain .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  acre (p. 30) 5. ( ) Cap obtuse becoming broadly convex, with or without a low umbo, evenly silvery to light gray over the disc, whitish buff near the margin, may be faintly virgate in places, taste bitter, occurring with conifers .. . . . . . . . . . . . . . . . . . . . . . . . . .  argenteum (p. 38) 5. Cap conic to obtuse with a ± prominent acute umbo, silky gray, darker on the disc, conspicuously virgate, taste very acrid or slowly bitter, occurring with conifers  . . . . . .  virgatum (p. 172) 6. (4) Base of the stalk often pinkish orange, odor mild or like non-perfumed soap, cap surface slippery when moist, cap color highly variable, pale greenish yellow, light green to grayish green,

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grayish olive, lead-gray, bluish gray, greenish gray, or grayish brown to brown .. . . . . . . . . . . . . . . . . . . . . . .  saponaceum (p. 134) 6. Stalk base not pinkish orange, odor farinaceous or not distinctive, cap appressed-fibrillose to scaly, less variable in color, usually with gray tones clearly dominant . . . . . . . . . . . . . . . . . . . . .  7 7. (6) Stalk distinctly scaly, gills pale gray, discoloring dark gray on the edges, often staining salmon-pink in some areas, cap dark brownish gray with lighter grayish margin, with appressed or recurved scales . . . . .  squarrulosum (p. 146) 7. Stalk not or only slightly scaly, gills variously colored, but not staining salmon-pink, cap color and texture various  . . . . . . . . . . . . . . . . . . . . . . . . .  8 8. (7) Gills having yellowish tones or discoloring yellow, especially toward the cap margin . . . . .  9 8. Gills lacking yellow color . . . . . . . . . . . . . . . . .  10 9. (8) Cap brownish gray to bluish gray, sometimes dull yellowish brown over the disc, often bruising yellow, appressed-fibrillose, squamulose near margin, gills often developing yellow tones, edges often dark gray to blackish, partial veil lacking . . . . . . . . . .  luteomaculosum (p. 92) 9. Cap convex with a low, broad umbo when young, wooly, then breaking up into fibrillose scales, often unevenly colored, with brownish, grayish, or black fibrils scattered over a white to pale gray background, gill edges not dark, evanescent cortina present .. . . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 10. (8) Cap with scattered brownish gray to blackish scales over a whitish background, odor and taste farinaceous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  11 10. Cap more evenly colored gray to dark gray, violet-gray, grayish brown, dull brown, or blackish brown, densely fibrillose to fibrillose-scaly, odor and taste slightly farinaceous or not distinctive .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12

K e ys to t h e De scr i be d Speci e s

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11. (10) Fruitbody usually medium-sized, cap surface wooly over the disc, breaking up into radiating fibrils and squamules toward the margin, fibrils initially black over a very pale gray background, then becoming brownish black .. . . . . . . . . . . . . . . . . . . .  atrosquamosum (p. 43) 11. Fruitbody usually large, surface with a whitish background overlaid with small, brownish gray to blackish, fibrillose scales that give an overall spotted appearance, typically darker over the disc where the scales are more concentrated .. . . . . . . . . . . . . . . . . . . . . . . . . .  pardinum (p. 119) 12. (10) Evanescent cortina present . . . . . . . . . . .  13 12. Cortina lacking  . . . . . . . . . . . . . . . . . . . . . . . . . .  14 13. (12) Odor and taste of flesh farinaceous, cap, gills, and stalk often developing yellowish stains .. . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 13. Odor and taste not distinctive, lacking yellow stains . . . . . . . . . . . . . . . . . . . .  myomyces (p. 108) 14. (12) Fruitbody medium-sized, cap densely fibrillose-scaly, with dark violaceous tones, becoming grayish brown overall in age, flesh whitish, unchanging or rapidly staining reddish gray when exposed . . . . .  atroviolaceum (p. 44) 14. Fruitbody small to medium-sized, cap fibrillose to somewhat scaly, violaceous tones lacking, flesh white to grayish, unchanging . . . . . . . . .  15 15. (14) Fruitbody small, occurring in montane conifer forests in late spring and early summer, often near melting snow  . . . . . . .  moseri (p. 101) 15. Fruitbody small to medium-sized, occurring with conifers in fall . . . . . . . . . . . . . . . . . . . . . . .  16 16. (15) Fruitbody medium-sized, cap moist to tacky, densely matted-fibrillose to slightly scaly in the center, smooth or with minute scales toward the margin, occasionally virgate in places, often wrinkled, dark slate-gray when young,

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becoming paler gray, gill edges sometimes blackening, odor and taste of flesh distinctly farinaceous . . . . . . . . . . . . . . . . . .  nigrum (p. 110) 16. Fruitbody small to medium-sized, cap dry, densely wooly-fibrillose to fibrillose-scaly, evenly grayish to blackish brown or dull brownish, gill edges not blackening, odor and taste slightly farinaceous or not distinctive .. . . . . . . . . . . . . . . . . . . . . . . . . . .  terreum (p. 159) Key W-4—W estern B rown- C aps

Cap predominantly some shade of brown, orange, ochre, or tan. 1. Stalk with a cottony-membranous ring or a clearly defined ring-zone . . . . . . . . . . . . . . . . . . .  2 1. Stalk lacking ring or ring-zone . . . . . . . . . . . . .  6 2. (1) Cap white, ± covered with brown fibrils or scales, dry, odor spicy, fragrant, or somewhat unpleasant, taste mild or somewhat bitter or unpleasant . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  3 2. Cap brown, orangish brown, or reddish brown, viscid, odor farinaceous, taste farinaceous or bitter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  4 3. (2) Fruitbody usually with squatty or stocky stature, stalk base often pointed, cap and stalk scales pale to medium brown, white color usually dominant, odor reminiscent of cinnamon Red Hots candy with a variably intense component of old gym socks, occurring with conifers .. . . . . . . . . . . . . . . . . . . . . . . .  magnivelare (p. 94) 3. Fruitbody may be stocky, but often with a relatively slender stalk that is rounded at base, cap and stalk scales medium to dark brown, brown color often dominant, odor varying from mild or slightly fragrant to highly unpleasant or, in some collections, virtually the same as that of T. magnivelare, occurring with conifers or hardwoods, especially oak . . . . . . . . . . . . . .  caligatum (p. 50)

4. (2) Stalk with persistent, thick, sheathing ring, cap color highly variable, often with a mix of orange and olivaceous tones, stalk white above, orange-brown and shaggy to scaly below the ring, stalk often tapered to a pointed base .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  focale (p. 66) 4. Stalk with clearly defined ring-zone near the apex, cap color similar to above or dark reddish brown, stalk fibrillose to finely scaly but not shaggy, stalk may be tapered but usually not as distinctly pointed as above . . . . . . . . . . . . . . . . .  5 5. (4) Cap dull orange or orange-yellow to orangebrown, stalk usually with horizontal bands of fibrils or fine scales colored like cap and often in a snakeskin pattern . . . . . . . .  aurantium (p. 48) 5. Cap reddish brown, sometimes darker over the disc, surface above the ring-zone silky-fibrillose and strongly pruinose, white to whitish, spotting brown in age, surface below the ring-zone with a coating of appressed orange-brown longitudinal fibrils that sometimes breaks up exposing the underlying white flesh . . . . . .  fracticum (p. 70) 6. (1) Odor of flesh distinctive, reminiscent of soap, celery, or curry, but not farinaceous, cap surface smooth or tomentose . . . . . . . . . . . . . . .  7 6. Odor farinaceous or not distinctive, cap surface viscid, fibrillose, or scaly . . . . . . . . . . . . . . . . . . .  8 7. (6) Odor strongly of celery or curry, cap with a white background overlaid with brownish tomentum, in age usually cracking irregularly like dried mud, flesh in stalk base whitish or darkening somewhat but not pinkish orange .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  apium (p. 36) 7. Odor mild or like non-perfumed soap, cap dry to moist, cap color highly variable, often with olivaceous or yellowish tones, flesh in stalk base pinkish orange  . . . . . . . . . .  saponaceum (p. 134)

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8. (6) Cap moist or dry, fibrillose, fibrillose-wooly, and/or distinctly scaly . . . . . . . . . . . . . . . . . . . . .  9 8. Cap viscid, at least when young, smooth, fibril lose, or slightly scaly . . . . . . . . . . . . . . . . . . . . . .  17 9. (8) Occurring in late spring and early summer in montane conifer habitats, often near melting snow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  10 9. Occurring in fall in a variety of habitats, rarely if ever near melting snow . . . . . . . . . . . . . . . . . . .  11 10. (9) Fruitbody small and relatively slender, cap grayish brown to blackish with a paler margin, margin inrolled and heavily bearded when young, but lacking partial veil, odor and taste farinaceous or not distinctive .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .  moseri (p. 101) 10. Fruitbody medium-sized to large and stocky, cap white when young, often with a whitish bloom, soon fuscous to brown, often with olive, grayish or pale tan areas, thin ring present, but at times faint, odor strongly farinaceous .. . . . . . . . . . . . . . . . . . . . . . . .  vernaticum (p. 170) 11. (9) Cortina present, at least in young specimens .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  12 11. Cortina lacking  . . . . . . . . . . . . . . . . . . . . . . . . . . .  13 12. (11) Cap convex with a low, broad umbo when young, wooly, then breaking up into fibrillose scales, often unevenly colored, with brownish, grayish, or black fibrils scattered over a white to pale gray background, cap and gills staining yellowish, stalk solid or hollow, not common, occurring with a variety of trees .. . . . . . . . . . . . . . . . . . . . . .  scalpturatum (p. 138) 12. Cap often conical when young, covered with bright rusty brown scales over a whitish background, cap margin cottony with slight remains of a partial veil, gills staining reddish brown, stalk typically hollow at maturity, fairly common and often abundant under spruce .. . . . . . . . . . . . . . . . . . . . . . . . . .  vaccinum (p. 165)

K e ys to t h e De scr i be d Speci e s

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13. (11) Fruitbody small, cap appressed-fibrillose, occasionally minutely squamulose or wooly on the disc when young, golden orange-yellow with rusty orange stains, taste mild or slowly acrid .. . . . . . . . . . . . . . . . .  aurantio-olivaceum (p. 46) 13. Fruitbody small to (more often) medium-sized or large, cap fibrillose to scaly, lacking orange stains although color may include orangish tones, taste farinaceous or mild . . . . . . . . . . . .  14 14. (13) Fruitbody medium-sized to large, cap with a whitish background overlaid with small, tan scales that give a spotted appearance, typically darker over the disc where the scales are more concentrated and whitish near the margin, gills whitish, not staining . . . . . .  venenatum (p. 168) 14. Fruitbody small to large, cap fibrillose to scaly, but not colored as above or appearing spotted, gills whitish or colored, unchanging, staining yellow or reddish brown, or flushed pinkish orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  15 15. (14) Fruitbody medium-sized to large, cap fibrillose, dark brownish to blackish green becoming yellowish green to yellowish buff, gills yellowish to greenish buff, becoming whitish to buff at maturity, cap margin, gills, and stalk often flushed pinkish orange . . . . . . . .  davisiae (p. 56) 15. ( ) Fruitbody medium-sized to large, cap fibrillose, lemon yellow, brownish yellow, or with orange-brown or olivaceous tones over the disc, margin pale yellowish white when young, may become whitish overall with irregular yellowish patches in age, gills white or pale gray, sometimes with yellowish tones near margin of cap .. . . . . . . . . . . . . . . . . . . . . . . . . .  arvernense (p. 40) 15. Fruitbody small to large, cap wooly-fibrillose to scaly, dull brown, grayish brown, or blackish brown, gills white to pale gray, unchanging or staining reddish brown . . . . . . . . . . . . . . . . . . . .  16

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16. (15) Fruitbody medium-sized to large, cap fibrillose to fibrillose-scaly initially, becoming coarsely cracked and scaly on the disc with whitish flesh showing through the cracks, gills white, then spotted or flushed reddish brown .. . . . . . . . . . . . . . . . . . . . . . . . .  imbricatum (p. 82) 16. Fruitbody small to medium-sized, cap densely wooly to radially fibrillose-scaly, grayish brown to blackish brown or dull brownish, gills white to grayish, not staining . . . . . . .  terreum (p. 159) 17. (8) Occurring with oak, cottonwood, aspen, or manzanita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  18 17. Occurring with conifers . . . . . . . . . . . . . . . . . . .  20

20. (17) Cap yellowish brown, brownish yellow, or with orange-brown or olivaceous tones over the disc . . . . . . . . . . . . . . . . . . . . . .  arvernense (p. 40) 20. Cap dark brown, reddish brown, or chestnutbrown, paler reddish brown, pinkish brown, or golden brown towards the margin . . . . . . . . .  21 21. (20) Gills and ground color of stalk buff to orangish buff, base of stalk usually not rooting, occurring primarily with pine . . . . .  muricatum (p. 104) and pessundatum (p. 121) 21. Gills and ground color of stalk yellowish buff to yellow, base of stalk often rooting, occurring with a variety of conifers .. . . . . . . . . . . . . . . . . . . . . . .  transmutans (p. 161)

18. (17) Occurring with manzanita, cap smooth or with brownish agglutinated appressed-fibrillose scales over the disc, white overall when young, soon pale orange and eventually brownish orange or brown, often developing reddish stains, upper portion of stalk pale yellow over a white background . . . . . . . . . . . . . .  manzanitae (p. 98) 18. ( ) Occurring, often in dense groups, with cottonwood or aspen, cap smooth, pinkish brown to reddish brown on the disc, paler or whitish toward the margin, stalk white at first, colored like the cap at maturity . . . .  populinum (p. 123) 18. Occurring with oak, cap smooth to fibrillose, cream to various shades of brown, stalk whitish to buff at first, often colored ± like the cap at maturity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .  19 19. (18) Cap smooth to fibrillose, cream to pale orange-white initially, darkening in irregular streaks and patches to brownish orange, and finally more uniformly brown overall, odor and taste farinaceous . . . . . . . . .  dryophilum (p. 58) 19. Cap smooth except for scattered appressed radial fibrils on the disc, grayish orange to brownish orange with a dark brown disc, darkening in age, odor slightly farinaceous or not distinctive, taste mild to bitter . . . . . . . . . . . . . . . . . .  ustale (p. 164)

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Table 1. Species of Tricholoma with Distinctive Microscopic Features. Epithet

Clamp Connections

acre aestuans argenteum arvernense atrodiscum atroviolaceum colossus davisiae dryophilum griseoviolaceum hordum huronense inamoenum insigne intermedium luteomaculosum marquettense moseri mutabile myomyces nigrum niveipes odorum olivaceobrunneum palustre pardinum pullum roseoacerbum saponaceum sejunctum serratifolium subaureum subresplendens sulphurescens sulphureum terreum tumidum venenatum vernaticum virgatum

Cheilocystidia

Pseudoparenchymatous hypodermium

   

    



   



   







 

 





  



 

  



   



    



  

 = feature usually well developed  = feature present but often not well developed  = feature present only occasionally

K e ys to t h e De scr i be d Speci e s

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Descriptions of Species Recognized Species

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Tricholoma acre Peck 

Hot Gray Trich

Cap: 2.5–12 cm wide, convex to broadly convex, often with a low, broad umbo, occasionally concave in age; surface dry, densely interwoven-fibrillose over the center and on the edge, elsewhere with interwoven-radiating fibrils, often with scattered squamules; pale medium gray overall, the disc occasionally fuscous-gray in age, occasionally virgate in places; margin incurved at first, expanded and sometimes wavy or eroded in age. Flesh: moderately thick; pale gray; odor not distinctive; taste hot and peppery, sometimes slowly so, or at times bitter. Gills: sinuate, close, broad; pale grayish overall or with fuscous-black edge, sometimes spotted fuscous-black; edges entire or sometimes eroded; lamellulae numerous. Stalk: 2.5–7.5 cm long, 1–2 cm thick, nearly equal, with a rounded or slightly bulbous base, solid, silkyfibrillose, often pruinose at the apex; dull white. Microscopic features: spores 6.5–8 × 4.5–6 µm, elliptic to broadly elliptic; cylindric to clavate cheilocystidia present. Occurrence: scattered or in groups under hardwoods, especially oak and hickory; northeastern North America, and possibly extending to the Pacific Northwest; not common.

unknown, however the taste suggests it is not edible and might be poisonous. Comments: Application of PDAB to the cap surface or flesh produces a bright pink reaction. Tricholoma serratifolium (p. 143) is similar and also grows with hardwoods, but its cap typically has extensive white areas or is entirely white, and its gills are often highly eroded. Tricholoma virgatum (p. 172) also is gray and has a peppery taste, but its cap has a pointed umbo and is conspicuously virgate nearly overall and it grows with conifers. The gray or gray-brown species such as T. terreum (p. 159) differ by their smaller size, often scaly caps, often farinaceous odor and taste, and, in many, the presence of a pseudoparenchymatous hypodermium. Tricholoma acre has been said to be similar to the European species T. impolitum (Lasch) P. Kummer, T. bresadolanum Clémençon, and T. murinaceum (Bulliard) Gillet, but comparisons are difficult because there seems to be no widespread agreement over the species concepts and taxonomy of these fungi. The epithet, acre, means sharp and here refers to the hot peppery taste.

De scr ip t ions of Speci e s

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Edibility:

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T. acre (A) NY

T. acre (B) NY

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Tricholoma aestuans (Fries) Gillet Cap: 3.5–9 cm wide, hemispheric or convex when young, becoming broadly convex or nearly flat in age, often with a low, broad umbo; surface dry, yellowish green to grayish yellow, brownish on the disc, radially streaked with grayish fibrils; margin incurved when young, expanded to slightly uplifted in age, sometimes lobed or wavy. Flesh: whitish, grayish white, to grayish green; odor not distinctive; taste bitter or mild initially, then becoming strongly acrid. Gills: emarginate to adnate, close, broad; honey-yellow to greenish yellow, bruising pale brownish gray. Stalk: 3–7 cm long, 1–2 cm thick, nearly equal or tapering slightly downward toward the base, hollow, silky; pale yellowish white to whitish green. Microscopic features: spores 6.5–8 × 4–5.2 µm, elliptic; cylindric to ventricose or clavate cheilocystidia present. Occurrence: solitary, scattered, or in groups under pine or sometimes oak and beech; widely distributed in northeastern North America and perhaps beyond; not common.

the acrid taste suggests it is inedible and possibly poisonous. Comments: Application of PDAB produces a bright pink color on all parts of the mushroom. Tricholoma palustre (p. 117) is similar (and possibly could represent the same species) but develops a grayish to brownish or blackish cast over the disc or in patches on the cap, is found under hardwoods, especially beech and oak, and has more broadly elliptic spores. Tricholoma equestre (p. 60) is similar in color, but is less likely to exhibit greenish tones. In addition, it has a somewhat viscid cap, and lacks the acrid taste, pink PDAB reaction, and cheilocystidia. Tricholoma sejunctum (p. 140) also is similar, but has dark brown to blackish radial fibrils on the cap, and gills that are usually whitish or white with yellowish areas rather than evenly yellowish green. The epithet, aestuans, means hot or burning, in reference to the taste.

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Edibility:

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T. aestuans NY

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Tricholoma albidum Bon Cap: 3–7 cm wide, convex or obtusely conical when young, becoming broadly convex or nearly plane; surface lubricous but not viscid, silky-fibrillose; white overall at first, dull white at maturity, then discoloring yellowish to ochraceous, sometimes watery spotted, especially on the disc; margin inrolled at first, becoming incurved to somewhat uplifted, sometimes rimose. Flesh: firm but brittle; white, unchanging or yellowing slightly; odor slightly farinaceous, sometimes with a fruity component; taste mild, weakly farinaceous, or nauseating at length. Gills: sinuate, sometimes with a decurrent tooth; white, slowly discoloring yellow; edges finely toothed; no cortina present. Stalk: 4–7 cm long, 5–10 mm thick, slightly tapered downward, solid; surface dry, fibrillose-silky; white, discoloring yellow when handled, sometimes ochraceous at the base. Microscopic features: spores 5–7 × 3–5 µm, elliptic to nearly ovoid. Occurrence: scattered, in groups or clusters on soil under hardwoods according to Marcel Bon’s original description, but see comments below; reported from Michigan by Alexander Smith (as T. albatum) and New York where we photographed it; seemingly rare.

unknown. The overall white color, small size, and tendency to bruise yellow characterize T. albidum. Tricholoma albatum (Quélet) Maublanc and d’Astis is considered a synonym, whereas T. albatum Velenovsky is thought most likely to be a species of Melanoleuca. Tricholoma columbetta (Fries) P. Kummer differs in its larger size and supposed lack of yellow staining. Although Bon described the odor and taste of the flesh as weakly farinaceous and then nauseating, Alexander Smith (as reported by Ovrebo 1980) stated that the odor of the flesh is slight and that it has a weakly acrid and disagreeable taste. Smith also reported that his collection of T. albatum possessed a fleeting cortina and that it was growing in association with pine and spruce, rather than hardwoods. These differences, together with differences between Smith’s and Bon’s spore sizes (Smith reported 4–5 × 3–4 µm), lead us to question whether the same taxon is involved. It seems quite possible that North American collections that have been called either T. albatum or T. albidum actually represent an undescribed North American species or, possibly, T. subresplendens (p. 151). Also see the comments under the latter species. The epithet, albidum, means whitish.

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Edibility:

Comments:

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T. albidum NY

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Tricholoma apium Jul. Schäffer Cap: 5–10+ cm wide, convex expanding to plane with low, broad umbo, margin strongly inrolled when young and long remaining so, later even or undulating; surface dry, dull, finely tomentose or granulose, becoming areolate; almost white at first, on exposure to light quickly becoming ochraceous, honey-yellow, or yellow-brown, then reddish brown with pale background showing in the cracks. Flesh: thick over the disc; white or dingy whitish; odor strong, like celery or somewhat curry-like (similar to the odor of Lactarius helvus or L. camphoratus); taste mild or a bit farinaceous or spicy. Gills: emarginate to arcuate-adnate, sometimes with a sub-decurrent tooth, close to rather crowded; creamy white to yellowish white, sometimes with ochre-yellow spots on the faces; edges entire to eroded; lamellulae numerous. Stalk: 4–15 cm long, 1–3 cm thick, nearly equal or with a tapered base, solid or stuffed, becoming hollow; surface dry, apex floccose or with transverse, somewhat fibrillose-scaly bands, strongly fibrillose or finely scaly below, with abundant white mycelial mat in the surrounding soil; white at first, then darkening somewhat to sordid gray, cinnamon buff, or yellow-ochre. Microscopic features: spores 3.5–6 × 3–5 µm, subglobose to broadly elliptic.

solitary or in small groups in mixed conifer forests; reported from the Pacific Northwest (British Columbia and Washington), Michigan, and Massachusetts, so probably widely distributed in northern North America; not common, possibly rare. Edibility: the single report we are aware of suggests it is inedible because of a bitter taste. Comments: Tricholoma apium is a little-known (in North America) but distinctive species on account of its odor and the yellowish brown tomentose cap surface that cracks like dry mud. The closest lookalike probably is T. imbricatum (p. 82) but its cap is dull brownish and less deeply cracked, and it lacks a strong odor. Tricholoma sulphurescens (p. 154) is similar in its stature, pale coloration, and occurrence with conifers, but stains yellow, lacks the highly cracked cap surface (although it can be somewhat cracked), and has a different odor. Tricholoma apium became an environmental poster child when it was discovered at Mt. Elphinstone, an old-growth forest site in British Columbia slated for logging. In part because of the presence of this seemingly rare fungus, a portion of the site was preserved as a Provincial Park. The epithet, apium, is Latin for celery.

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Occurrence:

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T. apium (A) MA

T. apium (B) WA

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Tricholoma argenteum Ovrebo Cap: 4.5–6.5 cm wide, obtuse and often with a low umbo when young, becoming broadly convex to nearly plane, with or without a low rounded or somewhat acute umbo at maturity; surface dry, densely matted-fibrillose over the disc and with innate radial fibrils extending toward the margin; light gray to silvery gray, generally whitish buff near the margin or occasionally elsewhere, occasionally with dark gray streaks but not distinctly virgate; margin incurved at first, becoming uplifted and wavy at maturity. Flesh: white to light gray; odor not distinctive; taste bitter. Gills: arcuate when young, sinuate when mature, close; white or pale gray, discoloring brownish on the edges; entire; lamellulae numerous. Stalk: 4–9.5 cm long, 1–1.5 cm thick, equal or slightly clavate with a rounded base, solid; surface silky-fibrillose with surface fibrils projecting, often pruinose at the apex; white, typically becoming sordid buff in places.

Microscopic features: spores 7.6–8.6 × 4.8– 5.7 µm, elliptic; cylindric or clavate cheilocystidia present. Occurrence: scattered or in groups under pine or spruce; northern North America. Edibility: unknown. Comments: Application of PDAB produces a bright pink reaction on all parts of the mushroom. The light gray to silvery gray cap with innate radial fibrils and the bitter-tasting flesh are the distinguishing features. Tricholoma virgatum (p. 172) is very similar but has a more acute umbo and a darker cap surface that is distinctly virgate. The epithet, argenteum, means silvery.

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T. argenteum (A) NS

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T. argenteum (B) QC

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Tricholoma arvernense Bon Cap: 3–14+ cm wide, more or less conical with acute umbo when young, expanding to flattened-convex with broad umbo; surface dry, smooth-satiny, radially fibrillose, sometimes slightly squamulose over disc, disc slightly lubricous when moist; lemon yellow, brownish yellow, or with orange-brown or olivaceous tones over the disc, margin pale yellowish white when young, may become whitish overall with irregular yellowish patches in age; margin downcurved, then becoming somewhat wavy and irregular. Flesh: solid; white overall or with yellowish zones near the cuticle, unchanging; odor and taste mildly farinaceous-fruity or like cucumber. Gills: sinuate, sometimes with decurrent tooth, close, broad; white or pale gray, often with yellowish tones near margin of cap. Stalk: length about the same as or slightly shorter than the cap diameter, nearly equal or somewhat ventricose with a pointed and sometimes bulbous base; white to cream overall, sometimes with yellowish, brownish, or pinkish tones in lower portion. Microscopic features: spores 4–7 × 3–5.5 µm; more or less clavate cheilocystidia present, some with narrowed apices or irregular shapes; clamp connections rare to occasional in gill trama and cap cuticle. Occurrence: in Europe under pine, fir, and spruce on nutrient-poor soils; in North America, reported scattered under hemlock and other conifers near Priest Lake, Idaho, from the Gifford Pinchot National Forest in south-central Washington, and from southern Oregon. Edibility: not well established. Comments: When the draft manuscript for this book was submitted to the publisher, the Idaho collection made by Drew Parker (Photo A) was included as an unknown species. During review of the manuscript, we received word from Dr. Brandon

Matheny that DNA sequencing of Drew’s collection indicated a very close (99%) match with a French collection of T. arvernense. To our knowledge, this collection represents the first North American report of this species. Shortly thereafter, we were alerted to another collection made by Sava Krstic (Photos B and C) and displayed at the Oregon Mycological Society’s annual mushroom show. Later, we learned of a third collection, this one from southern Oregon. Originally regarded in Europe as a subspecies of T. sejunctum (p. 140), T. arvernense differs from that species mainly by its lack of cap-surface green tones, lack of dark brown to black radial fibrils, and growth with conifers rather than hardwoods (although note that, in North America, the epithet sejunctum has been applied to conifer collections as well as those made with hardwoods). Tricholoma equestre (p. 60) is similar in color and also occurs with conifers, but its stipe and gills are evenly yellow-colored, rather than white with yellowish flushes. In age, fruitbodies can fade to almost whitish and then may appear similar to T. subresplendens (p. 151) and the other yellow-staining white species, but the highly pigmented caps of the young fruitbodies distinguishes T. arvernense from them. Tricholoma intermedium (p. 89) is very similar in having yellow tones in the cap and occurring with conifers; however, it differs in having gills and stipe that do not develop yellowish stains. The description of T. tumidum (p. 163) given by Kris Shanks (1997) and followed in our account seems very similar to T. arvernense and could possibly represent the latter species. The epithet arvernense refers to the species’s occurrence in the Auvergne region of south-central France, named after a Gallic tribe, Arverni, that once inhabited the area.

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T. arvernense (A) ID

T. arvernense (B) WA

T. arvernense (C) WA 41

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Tricholoma atrodiscum Ovrebo Cap: 3–6.5 cm wide, obtusely conic when young, becoming broadly convex to nearly plane, often with a low, broad umbo; surface dry, densely matted-fibrillose over the disc, appressed-fibrillose and appressed-squamulose over the remainder; dark fuscous-gray to dark sooty gray, paler near the margin; margin incurved at first, becoming uplifted and wavy. Flesh: gray or dark watery gray; odor not distinctive; taste quickly bitter. Gills: sinuate, close; light buff to light pinkish buff when young, buff to light gray at maturity, discoloring blackish on the edges and in small spots on the faces; entire; with numerous lamellulae. Stalk: 4–9 cm long, 6–14 mm thick, nearly equal to slightly clavate with a rounded base, hollow; surface

silky-fibrillose with some projecting surface fibrils; drab grayish buff. Microscopic features: spores 7.6–8.6 × 5.2– 5.7 µm, elliptic; cylindric to clavate cheilocystidia present. Occurrence: scattered or in groups under conifers or in mixed woodlands; eastern Canada. Edibility: unknown. Comments: Application of PDAB produces a bright pink reaction on all parts of the mushroom. Tricholoma hordum (p. 80) has similar cap coloration and its gills also have a pinkish cast when young, but it is larger, has fewer squamules on its virgate cap, and grows in association with hardwoods. The epithet, atrodiscum, means having a black disc.

T. atrodiscum QC

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Tricholoma atrosquamosum (Chevalier) Saccardo Black-scaled Trich (Not illustrated)

Cap: 4–5.5 cm wide, broadly convex with a low umbo; surface dry, wooly over the disc, breaking up into radiating fibrils and squamules toward the margin; fibrils initially black over a very pale gray background, then becoming brownish black; margin inrolled and distinctly bearded when young, remaining incurved well into maturity, finally becoming wavy and uplifted. Flesh: pale gray; odor and taste farinaceous. Gills: emarginate to adnexed, close, moderately broad; white to very pale gray, sometimes with black edges. Stalk: 4.5–7 cm long, 1–2 cm thick, nearly equal or enlarged toward the base, solid; surface dry, appressed-fibrillose to appressed-squamulose except at the apex, which is pruinose; medium gray overall; base usually carmine-red when dry. Microscopic features: spores 5–9 × 3.5–5 µm, elliptic to narrowly elliptic. Occurrence: scattered or in groups under hardwoods or in mixed woodlands, also with conifers (especially spruce) according to some European authors; reported from California, the Pacific Northwest, and Québec, but appears to be uncommon.

considered edible in Europe; we have no information for North America. Comments: Tricholoma nigromarginatum Bresadola, is a synonym, and T. squarrulosum (p. 146) is considered by many to be a variety of T. atrosquamosum, differing primarily by having more abundant squamules on the stalk. Others maintain them as separate species, differing in features such as stalk scaliness, odor, and habitat. Tricholoma atroviolaceum (p. 44) is similar but its cap margin is not bearded when young, its cap flesh may stain reddish gray when cut, and its gills are darker brownish gray at maturity. Tricholoma luteomaculosum (p. 92) has a cap that is more appressed-fibrillose and sometimes develops dingy yellow stains on the margin and gills. The latter two species also differ by having a pseudoparenchymatous hypodermium and T. luteomaculosum has cystidia on its gills. The European species, T. orirubens Quélet, occasionally said to occur in North America, has a white stalk, slowly develops reddish stains on its gills and stalk, and has smaller broader spores. The grayish species such as T. terreum (p. 152) are smaller and have white to paler gray stalks. The epithet, atrosquamosum, means having black scales. Edibility:

43

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Tricholoma atroviolaceum A.H. Smith Cap: 4–10 cm wide, broadly convex when young, nearly plane at maturity; surface dry, densely covered by small blackish violet to violaceous gray-brown fibrillose scales; nearly black over the disc, grayish violet-brown to violet-gray toward the margin, fading to grayish brown overall; margin incurved at first, becoming uplifted and wavy, often split radially. Flesh: thin; whitish and unchanging (according to the original description) or whitish at first, then rapidly staining reddish gray and finally sordid grayish brown when cut (according to Shanks 1997); odor and taste slightly to distinctly farinaceous. Gills: sinuate, moderately broad, close, thick, sometimes anastomosing; very pale gray when young, becoming pinkish gray to sordid brownish gray or cinnamon at maturity; edges sometimes discoloring fuscous or blackish. Stalk: 5–14 cm long, 1.5–4 cm thick, nearly equal or enlarged downward, sometimes abrupt or tapered at the base, solid, tough; surface dry, appressedfibrillose with some fibrils projecting, apex pruinose; whitish to pale violet-gray or brownish, discoloring yellowish tan towards the base or where handled.

Microscopic features: spores 7–10 × 4.8–7.2 µm, elliptic to broadly elliptic; pseudoparenchymatous hypodermium present. Occurrence: solitary to scattered under conifers, possibly a Douglas-fir associate; Pacific Northwest south into California; not common. Edibility: unknown. Comments: Tricholoma atrosquamosum (p. 43) is similar but usually is smaller, its cap margin is inrolled and distinctly bearded when young, it lacks violet tones, its flesh does not stain reddish gray when cut, and it lacks a pseudoparenchymatous hypodermium. Tricholoma luteomaculosum (p. 92) differs in its cap lacking violet tones and being more appressed-fibrillose, sometimes developing dingy yellow stains on the cap margin and gills, and having cystidia on its gills. The epithet, atroviolaceum, means black-violet.

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T. atroviolaceum (A) ID

T. atroviolaceum (B) ID

T. atroviolaceum (C) WA 45

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Tricholoma aurantio-olivaceum A.H. Smith Cap: 1–4 cm wide, obtusely conic when young, becoming broadly convex to nearly plane, with a low to prominent umbo; surface dry, appressed-fibrillose, occasionally minutely squamulose or wooly on the disc when young; golden orange-yellow with rusty orange stains over the disc and yellowish tan on the margin at first, soon becoming duller and yellowish to orangish brown to light brown with brown to dark brown spots and stains; margin inrolled at first, expanding slightly and then remaining down-turned well into maturity. Flesh: whitish when young, becoming buff and finally grayish brown; odor not distinctive; taste mild or slowly acrid. Gills: emarginate to adnexed, narrow, close to subdistant; white or buff when young, gradually spotting light brown or olive-brown to dark brown; with one or two tiers of lamellulae. Stalk: 4–8 cm long, 3–7 mm thick, nearly equal or tapered downward and sometimes with a rooting base, stuffed or hollow; covered with orange to rusty orange fibrils or granules over a cream to buff background, fibrils becoming brown in age; sometimes beaded with ochraceous orange droplets at the apex.

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Microscopic features:

spores 5–7.6 × 3.4–5 µm,

elliptic. scattered or in groups under conifers, especially Douglas-fir; Pacific Northwest, south into the coastal forests of northern California; not commonly reported. Edibility: unknown. Comments: The small stature, dry cap, and the presence of orange to rusty orange fibrils on the stalk are the distinctive features. It is similar to T. psammopus (Kalchbrenner) Quélet, a European species with yellow gills, acrid, mildly bitter, or bitter-astringent flesh, and association with larch. Tricholoma vaccinum (p. 165) is larger, has a rusty brown cap with a partial veil when young, and is associated with spruce (mostly) and pine. Tricholoma imbricatum (p. 82) also is larger, has a duller brown cap, lacks orange to rusty orange fibrils on the stalk, and is associated with pine (mostly) and spruce. The epithet, aurantio-olivaceum, refers to the golden orange and olive-green colors. Occurrence:

46

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T. aurantio-olivaceum CA

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Tricholoma aurantium (Schaeffer) Ricken 

Orange-sheathed Trich

Cap: 4–8 cm wide, convex when young, becoming broadly convex to nearly flat in age; surface sticky to slimy when fresh, shiny when dry, covered with tiny, flattened scales or fibrils; brownish gold to orangebrown or tawny ochre, often darker on the disc; margin incurved when young, becoming expanded. Flesh: white; odor and taste strongly farinaceous or unpleasant. Gills: emarginate to adnexed, close, narrow; white, staining brownish red to brownish orange. Stalk: 4–8 cm long, 1–2 cm thick, nearly equal or tapered downward, solid; white, dry, and smooth at the apex, sheathed from the base up to an obscure ring-zone with tiny, bright orange-brown to reddish orange, flattened scales; cortina thin and evanescent. Microscopic features: spores 4–6 × 3–4 µm, elliptic. Occurrence: solitary, scattered, or in groups under conifers and in mixed woodlands; widely distributed in North America and fairly common.

inedible. Tricholoma aurantium var. olivascens Mitchel and A.H. Smith was described from the Colorado Rockies, apparently based on a single collection. Its main distinguishing features are the bright orange coloration with olive-fuscous “overwash” and orange droplets on the incurved cap margin. We have no information regarding how widespread this form might be. Tricholoma focale (p. 66) can have similar cap and stalk colors and is also strongly farinaceous. However, it has a thick, membranous partial veil that typically forms a more or less conspicuous ring on the upper portion of the stalk. Tricholoma aurantio-olivaceum (p. 46) also is similar but has a dry cap, lacks the strong farinaceous odor, and usually is smaller. The epithet, aurantium, means orange.

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Edibility:

Comments:

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T. aurantium (A) CO

T. aurantium (B) WV

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Tricholoma caligatum (Viviani) Ricken 

Brown Matsutake

Cap: 6–12 cm wide, hemispheric when young, becoming broadly convex to nearly flat in age, sometimes shallowly depressed; surface dry, reddish brown to pale cinnamon-brown, grayish brown, or dark brown, breaking up into coarse flattened scales and fibers on a whitish to pinkish buff background; margin inrolled and cottony-membranous when young, becoming expanded to uplifted, sometimes rimmed with veil remnants. Flesh: white to cream; odor varies from mild to fragrant to often strongly unpleasant or, at times, virtually the same as the cinnamon-Red-Hots-plus-dirtysocks aroma of the matsutake; taste bitter (at times even when the odor is pleasant), nutty, or mild. Gills: emarginate to adnate, close, broad; white, often developing brownish spots or stains. Stalk: 4–10 cm long, 1.5–3 cm thick, nearly equal or ventricose, solid, dry, sheathed from the usually rounded base up to the ring by a brown veil (typically concolorous with the cap) that breaks into patches on a white background; white above the ring, sometimes pruinose; ring white, cottony-membranous, often flaring upward, median to superior; base and surrounding duff may develop a blue-green tint due to the growth of the mold, Penicillium simplicissimum (P. janthinellum). Microscopic features: spores 6–8 × 4.5–5.5 µm, elliptic.

solitary, scattered, or in groups, sometimes abundant, under conifers, especially hemlock, or hardwoods, especially oak; widely distributed in North America. Edibility: highly variable; edible when flesh is mild tasting, but often it is bitter. See Edibility section (p. 4). Comments: Like many tricholomas, what we are calling T. caligatum encompasses several forms that possibly represent different species, and some of these forms have been given formal variety names by Alexander Smith, under the genus name Armillaria. The existence of multiple taxa could explain the wide range of colors, odors, tastes, and host tree species reported. For instance, Smith distinguished var. glaucescens by its bluish gray to bluish green staining and disagreeable taste, and var. occidentalis by its occurrence in forests of various conifers, usually spruce and fir, its very dark colored pileus, the fragrant cinnamon odor resembling that of T. magnivelare (p. 94), and the disagreeable taste of the raw flesh. T. caligatum is most likely to be confused with T. magnivelare, which usually is larger and stockier, has paler cap colors, a more robust stipe that usually tapers to a point, a more consistently cinnamon-spicy odor, and mild tasting flesh. The epithet, caligatum, is from the Latin for heavily shod (as in wearing a heavy boot), and refers to the sheathing veil on the stalk.

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T. caligatum (A) FL

T. caligatum (B) FL

T. caligatum (C) NM 51

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Tricholoma cingulatum (Almfelt) Jacobashch Belted Trich, Girdled Trich

Cap: 2.5–6.5 cm wide, obtuse or soon obtusely campanulate, becoming expanded or umbonate with a spreading margin; surface dry, radially appressedfibrillose with the fibrils sometimes becoming arranged in patches forming appressed scales, caps of young specimens often with a fine whitish reticulum of fibrils; pale gray overall or pale gray with a bluish cast along the margin, sometimes becoming tinted yellowish or pinkish; margin incurved and cottony at first and long remaining incurved, often rimose when mature. Flesh: thick under the disc but abruptly thinner toward the margin, fragile; watery drab gray when moist, whitish when dry; odor and taste farinaceous. Gills: sinuate or occasionally adnate or adnexed, close; pale grayish to nearly white, often staining yellowish; edges even; lamellulae numerous. Stalk: 4–7 cm long, 3–10 mm thick, nearly equal or slightly clavate, rounded or abruptly tapered at the base, solid; surface silky at the apex and more or less glabrous at the base, with a persistent, flaring, cottony-membranous ring and appressed white fibrils or belts for some distance below the ring; whitish overall or sometimes with a pale bluish gray cast near the apex.

Microscopic features: spores 4.5–7 × 2.5–4.3 µm, elliptic to narrowly elliptic. Occurrence: scattered, in groups or clusters under willow; widely distributed in North America; not commonly collected, perhaps because its habitat seldom is visited. Edibility: in Europe, said to be edible by some, inedible by others; we have no firsthand knowledge for North America but, given the farinaceous odor and taste, it seems unlikely to be a good edible. Comments: Although similar to several other smallish gray tricholomas, the combination of a gray fibrillose cap, persistent cottony ring, and association with willow separates this species from the others. Recent phylogenetic analyses in Europe have supported this concept of a ringed willow-associate, distinct from similar species such as T. scalpturatum (p. 138), T. argyraceum (Bulliard) Gillet, and T. inocybeoides A. Pearson ( Jargeat et al. 2010). The epithet, cingulatum, means girdled, belted, or collared, in reference to the ring.

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T. cingulatum (A) QC

T. cingulatum (B) CA

T. cingulatum (C) CA 53

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Tricholoma colossus (Fries) Quélet 

Giant Trich

Cap: 10–25 cm wide, hemispherical to convex and remaining so well into maturity, sometimes with a depressed center; surface slightly to distinctly viscid when moist, fibrillose and somewhat shiny when dry; pale beige-brown when young, becoming darker reddish brown or brick-color with yellow-brown patches, often streaked; margin strongly involute, usually somewhat ribbed, only slightly expanding. Flesh: thick, firm; white, quickly turning pinkish red when cut; odor not distinctive; taste mild or astringent to slowly slightly bitter. Gills: emarginate, crowded, thick, narrow; white to buff at first, developing brownish or pinkish to reddish stains; edges irregular. Stalk: 5–20 cm long, 3–8 cm wide, nearly cylindrical or clavate to slightly bulbous, sometimes tapered towards the base, with a wooly-fibrillose annular zone that disappears in age; surface dry, appressedfibrillose, white above the annular zone, with reddish brown fibrils on a paler background below, sometimes with grayish or grayish green streaks and patches on the base.

Microscopic features: spores 7–10 × 5–6.5 µm, elliptic to oblong; filiform, clavate, or lageniform cheilocystidia present. Occurrence: solitary, scattered, or in groups under pines, often in sandy soil; widespread in the eastern United States, especially in coastal areas; not common. Edibility: reportedly edible and sold in markets in Italy, but we have no personal experience with it. Comments: This species also is known as Megatricholoma colossus (Fries) Kost. The robust caps and stalks with a veil that soon disappears are distinctive. Tricholoma aurantium (p. 48) is smaller, more brightly orange-colored, and has snakeskin-like bands on the stalk. Tricholoma focale (p. 66) can be fairly large and occur in similar habitats, but it often is olivaceous orange-colored and has a prominent ring and strongly farinaceous odor and taste. The epithet, colossus, means gigantic.

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T. colossus (A) FL

T. colossus (B) MA

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Tricholoma davisiae Peck Cap: 3–16 cm wide, broadly conic to bell-shaped when young, becoming broadly convex to plane, typically with a conspicuous blunt or acute umbo, somewhat thin and fragile; surface dry, appressedfibrillose overall when young, developing tiny scales near the margin; dark brownish gray to dark green on the umbo or nearly overall at first, becoming yellowish green to pale yellowish green or grayish yellow with darker streaks and umbo, usually then fading to yellowish buff nearly overall, sometimes with salmon tinges, especially near the edge; margin incurved at first, becoming wavy and uneven, often uplifted and split. Flesh: greenish, fading to buff at maturity; odor farinaceous; taste farinaceous or disagreeable. Gills: arcuate when young, becoming sinuate, moderately close to subdistant; yellowish to greenish buff, becoming whitish to buff at maturity, older specimens often developing pinkish orange stains on the edges, faces sometimes marked with transverse white stripes; lamellulae numerous. Stalk: 4.5–18 cm long, 1–3 cm thick, equal or sometimes enlarged downward with a rounded base, or ventricose, solid; surface dry, fibrillose; pale greenish yellow on the upper portion and whitish below, or sometimes whitish overall, often developing pinkish orange stains.

Microscopic features: spores 5.5–9.5 x 4–5.5 µm, narrowly elliptic to elliptic; cylindric or clavate cheilocystidia and a pseudoparenchymatous hypodermium present. Occurrence: scattered or in groups under conifers, especially pine; widely distributed in northeastern North America, but not common. Edibility: unknown. Comments: The stalk surface and cap flesh slowly stain pale pink with PDAB. Tricholoma cheilolaminum Ovrebo and Tylutki is a synonym. Charles Peck’s original description of young caps as bright yellow and often tinged with red or green differs somewhat from the one presented here (which is based primarily on Ovrebo 1980), perhaps because the collection he examined was quite mature. Because of its great variation in color, this species can be confusing and difficult to identify. The epithet, davisiae, honors Mrs. H. C. Davis, who collected the species near Falmouth, Maine, and sent material and notes to Peck.

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T. davisiae (A) NY

T. davisiae (B) NY

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Tricholoma dryophilum (Murrill) Murrill Cap: 4–10 cm wide, obtusely conic when young, becoming convex to broadly convex sometimes with a low, broad umbo; surface viscid, glabrous to innately radiating-fibrillose; cream to pale orange-white initially, darkening in irregular streaks and patches to brownish orange, and finally more uniformly brown overall; margin inrolled to decurved, lighter than the disc, sometimes with watery spots. Flesh: white; odor and taste farinaceous. Gills: deeply sinuate to adnexed, narrow, thin, close; white to pale cream, staining or spotting brownish orange to brown. Stalk: 6–17.5 cm long, 9–20 mm thick, equal or tapered slightly toward the base; surface dry, silkyfibrillose to scurfy at the apex; solid or hollow; whitish or brownish, apex white when young and often remaining so, with remainder darkening to brownish orange or yellowish brown to dark brown from the base upward. Microscopic features: spores 5.3–7.7 × 3.8– 5.8 µm, elliptic to narrowly elliptic (according to Shanks 1997, globose according to Murrill’s original description); scattered filiform cheilocystidia present.

scattered or in groups under coast live oak in California, also reported under Garry oak in southern Washington and so likely to occur with oaks in Oregon and possibly southwestern British Columbia as well; occasional. Edibility: unknown, but in light of the poisonous nature of some of the other viscid red-brown tricholomas, this is best avoided. Comments: Tricholoma dryophilum belongs to the confusing group of viscid-capped, brownish tricholomas. It is distinguished by the color changes of the cap from pale cream to brownish orange and finally brown, and its growth with oak. Tricholoma ustale (p. 164) also occurs under oaks, but it has a mild odor, bitter taste, and a more fibrillose cap that is reddish brown from the first. See comments under T. pessundatum (p. 121) for other species in this group. The epithet, dryophilum, means oak-loving.

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Occurrence:

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T. dryophilum (A) CA

T. dryophilum (B) CA

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Tricholoma equestre (Linnaeus) P. Kummer Man-on-horseback, Canary Trich, Yellow Trich

In North America, T. equestre is better known as T. flavovirens (Persoon) S. Lundell, which now is considered a synonym. However, by either name, this is a widespread and highly variable fungus and so it is quite possible that more than one species is involved. Based on his molecular studies, mycologist Tom Horton feels this is likely to be the case. If so, this might help explain the puzzling occurrence of fatal poisonings being caused by a mushroom widely thought to be a good edible species (see the Edibility section [p. 4]). Tricholoma sejunctum (p. 140) often is confused with T. equestre, however it has a golden yellow or greenish yellow cap with conspicuous brownish to blackish radial fibrils and white to pale yellow gills, or whitish gills and stalk flushed with yellow in places. The epithet, equestre, means horseman.

Cap: 5–10 cm wide, convex when young, becoming broadly convex to nearly flat in age; surface sticky when fresh or in wet weather, dry in age, smooth or often breaking into small scales on disc; pale yellow to sulfur-yellow, and usually golden brownish on the disc. Flesh: white; odor farinaceous; taste farinaceous or mild. Gills: sinuate to adnexed or seceding, close, broad; pale yellow to bright sulfur-yellow. Stalk: 3–15 cm long, 1–3 cm thick, nearly equal, occasionally enlarged at the base, solid, dry; whitish to pale yellow or sulfur-yellow. Microscopic features: spores 6–7 × 4–5 µm, elliptic. Occurrence: solitary to scattered or in groups under conifers, particularly pines, and in mixed woodlands; widely distributed in North America; fairly common. Edibility: traditionally considered a choice edible species, but we do not recommend it due to the occurrence of fatal poisonings in Europe when large quantities were eaten over a fairly short period. See the Edibility section (p. 4).

Comments:

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T. equestre (A) NY

T. equestre (B) FL

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T. equestre (C) CO

T. equestre (D) ID

T. equestre (E) OR De scr ip t ions of Speci e s

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Tricholoma farinaceum (Murrill) Murrill Cap: 3–8 cm wide, convex to expanded, umbonate; surface smooth, glabrous; white, often developing pale brown tints as the cap matures; margin somewhat incurved at first, then expanding, often wavy, entire or sometimes split. Flesh: thin; white; odor strongly farinaceous; taste not reported in the original description, but likely to be farinaceous. Gills: sinuate, broad, not crowded; white; with several tiers of lamellulae. Stalk: 5–6 cm long, 5–10 mm thick, nearly equal or enlarged downward, sometimes bulbous, with whitish mycelium at the base, stuffed or hollow; surface more or less glabrous, smooth, white. Microscopic features: spores globose, smooth, 4.5–6.5 µm in diameter. Occurrence: in leaf litter in woods; described from Washington.

unknown but, given the unpleasant odor, not likely to be a good edible. Comments: Murrill described T. farinaceum from collections made, in 1911, in forests near Seattle. The photograph, which represents the only modern collection of which we are aware, was taken by the late Kit Scates Barnhart under cottonwoods and mixed conifers at Priest Lake, Idaho. It fits Murrill’s description, and the spore length recorded by Kit (and checked by us) also fits, although the shape is somewhat elliptical rather than globose. The closest western look-alike is T. inamoenum (p. 86), which differs in having a strong coal-tar odor and thick distant gills. Other white tricholomas, such as T. albidum (p. 34) and T. subresplendens (p. 151), would be distinguished principally by odor and habitat. The epithet, farinaceum, refers to the odor. Edibility:

T. farinaceum ID 63

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Tricholoma floridanum (Murrill) Murrill Cap: 4–15 cm wide, conic-convex to nearly plane, sometimes broadly umbonate; surface dry, smooth to slightly tomentose; pale yellow overall or sometimes pale grayish yellow and weakly scurfy over the disc, paler along the edge; margin inrolled to strongly incurved well into maturity, even or sometimes wavy or lobed. Flesh: thick, firm; whitish to pale yellow; odor not distinctive; taste mild or slowly mildly bitter to astringent. Gills: adnate to sinuate, close, up to 1.5 cm broad; whitish with a pinkish tint; with several tiers of lamellulae. Stalk: 5–10 cm long, 2–4.5 cm thick, nearly equal or clavate to slightly bulbous, sometimes pinched at the base, solid at first, becoming hollow; surface dry, finely fibrillose, sometimes weakly striate; pale yellow, often with dull reddish brown stains near the base.

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Microscopic features:

spores 6–8 × 4–4.5 µm,

elliptic. solitary, scattered, or in groups under hardwoods, especially oak; described from Florida and we have seen it in Georgia, but likely to occur elsewhere in the southeastern United States. Edibility: unknown. Comments: This is a very robust, firm, and heavy mushroom. Tricholoma aestuans (p. 32) has an acrid taste and often occurs with pine. Tricholoma saponaceum (p. 134) has a soap-like odor and pinkish orange flesh in the base of the stalk. The epithet, floridanum, means from Florida; the type collection was made in Sanchez Hammock, 18 kilometers northwest of Gainesville, in 1938. Occurrence:

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T. floridanum (A) FL

T. floridanum (B) GA

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Tricholoma focale (Fries) Ricken Cap: 5–14 cm wide, convex when young, becoming broadly convex to nearly plane at maturity, occasionally broadly umbonate; surface viscid initially or when moist, usually becoming dry, with radiating fibrils or scales; color highly variable, typically a mixture of orange or yellow-orange, brown, and olivaceous green, or tans and browns with only a hint of orange and no green; margin inrolled at first, becoming uplifted and wavy. Flesh: thick, firm; white; odor farinaceous; taste very farinaceous-bitter. Gills: sinuate, close to crowded; whitish when young, becoming darker and developing rusty orange or brown stains. Stalk: 4.5–11 cm long, 1.5–3 cm thick, nearly equal or ventricose, usually tapered toward the often pointed base, solid; surface above the ring silkyfibrillose to pruinose, white, discoloring orangebrown; surface below the ring appressed-fibrillose with loose clumps of fibrils that appear shaggy, colored somewhat like the cap over a white background; ring superior to median, persistent, ragged, cottony, flaring or appressed, whitish on upper surface. Microscopic features: spores 5–6 × 3–4 µm, elliptic. Occurrence: scattered or in groups under conifers, especially pine; northeastern North America south to Florida and western North America; fairly common.

generally considered inedible because of the bitter taste, however Orson Miller reported that some people considered it an excellent edible while others found it worthless. Comments: Tricholoma zelleri (Stuntz and A.H. Smith) Ovrebo and Tylutki generally is considered to be a synonym, and the name T. robustum (Albertini and Schweinitz) Ricken, in the opinion of many, but not all, mycologists, has been misapplied to this species. British Columbia mycologist Paul Kroeger has observed what he believes are representatives of all three species (T. focale, T. zelleri, and T. robustum) growing in the same area in Manning Provincial Park, British Columbia, so it is quite possible that we have more than one species currently going by the epithet focale. Tricholoma aurantium (p. 48), has similar coloration but lacks a ring, usually is smaller, and often has well-developed snakeskin banding on its stalk. Tricholoma caligatum (p. 50) and T. magnivelare (p. 94) are similar in size, stature, and habitat, but are basically white mushrooms overlain with brownish cap and stalk scales, lack orange and green tones, and do not have a strong farinaceous odor. The epithet, focale, means scarf, in reference to the usually well-developed ring.

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Edibility:

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T. focale (A) MA

T. focale (B) OR

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T. focale (C) WA

T. focale (D) WA

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T. focale (E) AK

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Tricholoma fracticum (Britzelmayr) Kreisel Cap: 2–10 cm wide, convex when young, becoming broadly convex to nearly plane; surface viscid then dry, with scattered, appressed, radiating fibrils, reddish brown, sometimes darker over the disc, often paler and with water spots or striations near the margin; margin inrolled at first, becoming decurved and remaining so well into maturity. Flesh: white or sometimes orange-brown near the surface of the cap; odor not distinctive or faintly farinaceous; taste strongly bitter. Gills: adnate at first, becoming sinuate with a subdecurrent tooth, close; orange-white, usually brown-spotted in age. Stalk: 2–7 cm long, 1–1.7 cm thick, equal or tapered somewhat toward the base, with a clearly defined ring-zone, solid; surface above the ringzone silky-fibrillose and strongly pruinose, white to whitish, spotting brown in age; surface below the ring-zone appressed-fibrillose with a coating of orange-brown fibrils that sometimes breaks up, exposing the underlying white flesh; cortina present when young, leaving a clearly defined superior to median ring-zone. Microscopic features: spores 5–7.2 × 3–5.5 µm, elliptic.

Occurrence:

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scattered, or in groups or clusters; California coastal forests; not common. Edibility: unknown, but several close relatives are poisonous. Comments: Application of the name, T. fracticum, like most aspects of the brownish tricholomas, is confusing. Tricholoma batschii Gulden and T. subannulatum (Batsch) Bresadola are considered synonyms and, of the three names, T. batschii seems to be preferred by most European mycologists. By whatever name, in Europe it is generally considered to occur primarily with pines. Tricholoma subannulatum (Peck) Zeller was described from a collection made under oaks in southern California and later reported from Corvallis, Oregon, but little is known about it. Most of the other tricholomas with viscid brownish caps, such as T. pessundatum (p. 121), T. muricatum (p. 104), and T. populinum (p. 123), lack the clearly defined ring-zone. Among the species with a fairly well defined ring-zone, T. colossus (p. 54) is larger and differently colored, T. aurantium (p. 48) is differently colored and has snakeskin-patterned bands on the stalk, and T. ustaloides Romagnesi (occasionally said to occur in North America) occurs in association with hardwoods such as oak.

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T. fracticum CA

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Tricholoma fulvimarginatum Ovrebo and Halling 

(Not illustrated)

Cap: 3–9.5 cm wide, obtusely convex when young, expanding to convex, and finally nearly plane; surface viscid, sometimes slightly so, drying in age, glabrous or sometimes slightly virgate or areolate; orange-brown to reddish brown when young, becoming pale reddish brown to orange-brown, sometimes paler yellowish tan near the margin; margin even, inrolled at first, becoming decurved or straight. Flesh: 10–15 mm thick; white, unchanging; odor strongly farinaceous and somewhat resembling that of green corn; taste strongly farinaceous or somewhat bitter. Gills: emarginate to nearly free, crowded to close; whitish, sometimes with small brownish spots on the faces; edges entire at first, becoming eroded, more or less concolorous with the cap. Stalk: 3–8 cm long, 1–2.5 cm thick, nearly equal to clavate, sometimes with an abruptly bent base, solid or hollow in age; surface dry, conspicuously fibrillose; white near the apex, more or less concolorous with the cap below but often blotchy and mixed with whitish areas, staining pale brown when handled.

Microscopic features: spores 4.6–6.1 × 3–4 µm, narrowly elliptic to elliptic. Occurrence: scattered, in groups, or somewhat clustered in sandy soil under cottonwood; reported occasionally from Massachusetts and New York but likely to occur elsewhere in New England and southeastern Canada. Edibility: unknown. Comments: The combination of an orange-brown cap, brown marginate gills, and the association with cottonwood is distinctive. Tricholoma populinum (p. 123) also grows in association with cottonwood but differs by usually having a darker reddish brown cap (at least in age) and non-marginate gills. Other viscid brownish tricholomas occur with other tree species (see comments under T. pessundatum [p. 121] ). The epithet, fulvimarginatum, means having a tawny (cap) margin.

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Tricholoma fulvum (Bulliard) Saccardo  Cap: 3–8 cm wide, somewhat conical to convex when young, becoming broadly convex to nearly flat, sometimes shallowly depressed, with or without a small umbo; surface smooth, slightly viscid when fresh or in wet weather, often shiny when dry, streaked with radial fibrils; reddish brown to yellowbrown, darker over the disc and paler yellow-brown to dull yellow on the margin. Flesh: white to yellowish; odor and taste farinaceous. Gills: emarginate to adnexed, close to crowded, broad; yellowish buff to buff, staining brownish. Stalk: 7–15 cm long, 1–2 cm thick, nearly equal or slightly enlarged toward the base, slippery to viscid when young and fresh, becoming dry, solid, hollow; lower portion concolorous with cap, pale yellow to yellowish at the apex. Microscopic features: spores 5.5–7 × 4–6 µm, elliptic.

Brown Birch Trich

solitary, scattered, or in groups under hardwoods, especially birch, or in mixed woodlands; most often reported from northeastern North America, but also in Alaska and probably in between where birch occurs; occasional. Edibility: not recommended; reported to be nonpoisonous, but poor. Comments: There is considerable confusion over the authority for this species, with at least five versions appearing in the principal European literature. We have chosen what seems to be the earliest name. Tricholoma flavobrunneum (Fries) P. Kummer is a synonym. Tricholoma pseudonictitans Bon, considered by some to be the same species, differs by its occurrence with conifers or in mixed forests that lack birch, and gills that are whitish to paler yellow than those of T. fulvum. The epithet, fulvum, means tawny or reddish yellow; flavobrunneum means yellow-brown. Occurrence:

T. fulvum (A) ME 73

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T. fulvum (B) QC

T. fulvum (C) QC

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T. fulvum (D) QC

T. fulvum (E) AK

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Tricholoma fumosoluteum Peck Cap: 4.5–9 cm wide, convex when young, becoming broadly convex to nearly plane, often with a low umbo; surface moist or dry, dull, glabrous or with distantly spaced appressed squamules, sometimes hygrophanous; dull yellow to brownish yellow with a darker smoky brown center; margin incurved when young, becoming uplifted and often wavy. Flesh: buff to pale yellow; odor not distinctive; taste bitter, astringent, or mild. Gills: sinuate, close to subdistant; whitish to light yellowish buff, occasionally discoloring brown on the edge and in small spots on the faces, becoming fuscous in places when dried; entire at first, then becoming eroded; lamellulae numerous. Stalk: 4.5–10 cm long, 1–1.8 cm thick, nearly equal with a rounded or abruptly tapered base, solid or stuffed; surface silky-fibrillose with projecting surface fibrils; whitish to light yellowish buff, not discoloring.

Microscopic features: spores 6.2–7.6 × 4.8– 5.7 µm, broadly elliptic to subglobose. Occurrence: scattered or in groups under conifers; northeastern North America; fairly common. Edibility: unknown. Comments: Yellowish forms of T. saponaceum (p. 134) can appear similar, especially when the caps have a smoky brown disc, but they usually have a soap-like odor and the gills and stalk often develop pinkish orange discolorations. Tricholoma subluteum (p. 149) lacks the smoky brown disc and has long, radiating fibrils on its cap. The epithet, fumosoluteum, means smoky yellow.

T. fumosoluteum QC De scr ip t ions of Speci e s

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Tricholoma griseoviolaceum Shanks Cap: 4–8 cm wide, campanulate with a prominent umbo when young, but typically lacking an umbo at maturity, becoming broadly convex and finally plane; surface viscid, innately radially fibrillose; white at first or when covered in duff, developing a pale violet to violet-gray ground color with dull violet-gray, dark violet-gray, or blackish radiating streaks, and finally becoming dark violet-gray over the disc; margin uplifted, wavy, and often splitting in age. Flesh: thin; white to watery gray; odor faintly farinaceous; taste farinaceous or sweet-farinaceous. Gills: sinuate, close, forking near the stalk; white, discoloring pinkish brown to grayish orange in patches with age. Stalk: 2–13 cm long, 1–2.2 cm thick, nearly equal or slightly tapered downward, with a pointed or abrupt base, solid, hollow, or stuffed; surface dry, dull, silky-fibrillose; white, occasionally pale orange at the base. Microscopic features: spores 4.8–7.2 × 3.4– 4.8 µm, elliptic; scattered clavate cheilocystidia sometimes present.

solitary to gregarious under oaks; coastal and low-elevation Sierra Nevada forests, California. Edibility: unknown. Comments: Tricholoma portentosum (p. 125) also has a viscid cap with radiating gray fibrils, but its cap is brownish gray and typically lacks the violet tones, its gills and stalk often have yellow rather than pinkish brown tints, its odor is not as strong, and it grows in association with conifers, especially pine. Tricholoma mutabile (p. 107) is similar but its cap is pale warm gray or pale violet-gray and often radially rugulose between the disc and margin, its stalk flesh is dull pink to pale orange at the base, and it has a pseudoparenchymatous hypodermium. Tricholoma vernaticum (p. 170) has a larger, dry to moist, whitish to gray cap without violet tones, a fibrillose ring, and grows with montane conifers in the late spring and early summer. The epithet, griseoviolaceum, means gray-violet. Occurrence:

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T. griseoviolaceum (A) CA

T. griseoviolaceum (B) CA

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T. griseoviolaceum (C) CA

T. griseoviolaceum (D) CA

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Tricholoma hordum (Fries) Quélet 

(Not illustrated)

Cap: 5–15 cm wide, obtusely conic or convex when young, becoming broadly convex to nearly plane or occasionally depressed; surface dry, densely interwoven-fibrillose over the disc, with long, innately radiating fibrils over the remainder of the cap, sometimes with scattered squamules near the margin; medium dark gray over a paler gray background, becoming paler near the margin, virgate; margin incurved when young, becoming uplifted. Flesh: light drab gray; odor not distinctive; taste bitter to bitter-acrid. Gills: sinuate or adnexed, occasionally seceding, close; pale vinaceous gray or grayish buff when young, then grayish buff, sometimes tinged dingy yellow, discoloring dark gray to fuscous on the edges and occasionally in small spots on the faces; entire; with numerous lamellulae. Stalk: 3.5–8.5 cm long, 1–2.5 cm thick, nearly equal with a rounded or slightly bulbous base, solid or hollow; surface silky-fibrillose, with projecting surface fibrils; whitish to very pale gray, occasionally tinged dingy yellow in age.

Microscopic features: spores 6.7–8.1 × 5.2– 6.2 µm, broadly elliptic to subglobose; cylindric to clavate cheilocystidia present. Occurrence: scattered or in groups under hardwoods; northeastern North America. Edibility: unknown. Comments: Application of PDAB produces a bright pink reaction on all parts of the mushroom. The faint vinaceous tinge on the young gills, which disappears with age, and the dark gray virgate cap are the distinguishing features. Tricholoma virgatum (p. 172) has a paler silvery gray to medium gray cap with an acute umbo, lacks the vinaceous tinge on the gills, and grows with conifers. Tricholoma argenteum (p. 38) also has a pale silvery gray cap, lacks the vinaceous tinge on the gills, and grows with conifers. Tricholoma atrodiscum (p. 42) has a fibrillose-scaly cap, smaller stature, and grows with conifers.

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Tricholoma huronense A.H. Smith  Cap: 6–12 cm wide, obtusely conic when young, becoming convex and finally plane, sometimes with a low, broad umbo; surface dry, fibrillose, pale to moderately dark gray at first, developing spot-like scales around the disc, with white flesh showing through along the margin; margin incurved when young, becoming uplifted in age, slightly fringed and somewhat cottony, often beaded with pink drops or streaked pinkish. Flesh: thick; white, unchanging; odor and taste distinctly farinaceous. Gills: sinuate, close to crowded, narrow; dull white, often with pinkish drops or streaks; edges slightly eroded; with two or three tiers of lamellulae. Stalk: 4–8 cm long, 1–2 cm thick, clavate, solid; surface whitish, covered over the lower portion with scales that are pale when young and then become gray; often with pinkish streaks or beaded with pinkish drops. Microscopic features: spores 7.5–9 × 4.5–6 µm, elliptic to slightly obovate; cheilocystidia clavate, rare; clamp connections typically common.

(Not illustrated)

in groups or clusters under hardwoods, especially oak; described from collections made near Ann Arbor, Michigan, likely to occur in similar habitat throughout the Great Lakes region and possibly beyond; rarely reported. Edibility: unknown, but the farinaceous odor and taste plus its similarity to poisonous species argue for avoiding it. Comments: This is a little-known, seemingly rare species. It could be confused with T. pardinum (p. 119), which lacks the pinkish drops and streaks on the gills, stalk, and cap margin as well as the gray scales on the stalk, and grows with conifers. Tricholoma venenatum (p. 168) is also similar but the cap scales are pale tan and it has white gills that become pale ivory buff at maturity. Both of these species have caused severe cases of gastric distress, suggesting that T. huronense should not be eaten. The epithet, huronense, means of or belonging to (Lake) Huron, near which the first collections were made. Occurrence:

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Tricholoma imbricatum (Fries) P. Kummer 

Shingled Trich

Cap: 2.5–9 cm wide, somewhat conical to convex when young, becoming broadly convex, usually with a low, broad umbo; surface dry, fibrillose to fibrillose-scaly initially, becoming coarsely cracked and scaly on the disc with whitish flesh showing through the cracks; dull brown on a whitish background; scales near the disc often overlapping and arranged like shingles; margin thin, incurved, and coated with soft, downy hairs when young, expanded in age. Flesh: whitish; odor and taste farinaceous or not distinctive. Gills: sinuate to nearly free in age, close, moderately broad; white when young, spotted or flushed reddish brown in age. Stalk: 5–9 cm long, 1–1.6 cm thick, nearly equal, tapered downward or slightly swollen at the base, solid, dry, fibrillose; white to whitish, slowly staining reddish brown from the base upward; apex often white-pruinose.

Microscopic features: spores 5.6–7.5 × 4.3– 5.5 µm, broadly elliptic. Occurrence: scattered or in groups under conifers, especially pines; widely distributed in North America, but not particularly common. Edibility: edible according to some sources; none of us has tried it. Comments: Tricholoma vaccinum (p. 165) is similar but it has a brighter rusty brown cap that is usually more scaly, but not coarsely cracked on the disc, a thin partial veil that often leaves remnants on the margin of the cap, and typically has a hollow stalk. Tricholoma vaccinum also is more common with spruces than with pines, at least in the West. The other brownish tricholomas, such as T. pessundatum (p. 121), differ primarily in having viscid caps and usually more reddish coloration. The epithet, imbricatum, means covered with overlapping tiles or scales, in reference to the texture of the cap surface.

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T. imbricatum (A) CO

T. imbricatum (B) CO

83

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T. imbricatum (C) NM

T. imbricatum (D) AZ

De scr ip t ions of Speci e s

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T. imbricatum (E) WA

85

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Tricholoma inamoenum (Fries) Quélet Cap: 3–6 cm wide, convex when young, becoming broadly convex to nearly flat in age, often with a low, broad umbo; surface smooth, dull, pale buff to whitish, often developing grayish to brownish tints in age. Flesh: white; odor pungent and disagreeable like coal tar or with a heavy floral scent like that of hyacinths or paperwhites; taste unpleasant. Gills: emarginate to adnexed, subdistant to distant, broad, often intervenose; white; with several tiers of lamellulae. Stalk: 4–9 cm long, 4–10 mm thick, nearly equal, or slightly enlarged downward to a swollen base that often tapers abruptly below, smooth, dry, pruinose at the apex; white to cream, brownish near the base. Microscopic features: spores 9.5–11 × 5.5– 6.5 µm, elliptic to almond-shaped; clamp connections rare. Occurrence: solitary or scattered under conifers; widely distributed in North America. Edibility: considered poisonous in Europe, the odor and taste suggest it would be inedible, even if not poisonous.

Comments: Tricholoma inamoenum is thought by many to be the same as T. platyphyllum (Murrill) Murrill, described from the vicinity of Seattle, Washington. However, Murrill’s description makes no mention of odor, so accepting the synonymy seems premature as does applying the latter name to collections exhibiting the coal tar odor. Tricholoma odorum (p. 114) also has a coal tar odor, but has a cap that is greenish yellow when young and then fades to yellowish buff to yellowish tan, pale yellow flesh that tastes farinaceous to disagreeable, a pale yellow stalk, and smaller spores. Tricholoma sulphureum (p. 157), which grows in association with both hardwoods and conifers, also has a coal tar odor, but its cap is dull sulfur-yellow with red-brown streaks and spots. Tricholoma sulphurescens (p. 154) has a whitish cap that bruises or ages dull yellow and a more variable and often not-unpleasant odor. The epithet, inamoenum, means disagreeable or not pleasing.

De scr ip t ions of Speci e s

86

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T. inamoenum (A) WA

T. inamoenum (B) WA

87

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Tricholoma insigne Ovrebo 

(Not illustrated)

Cap: 3.5–10 cm wide, obtusely conic to convex when young, becoming broadly convex to nearly plane, often with a low umbo; surface dry, densely interwoven-fibrillose over the disc, with radially appressed or innate fibrils toward the margin, often with scattered minute squamules; light gray to reddish gray over a light pink, light buff, or reddish salmon background when young, occasionally tinged green near the margin, becoming brownish gray, blackish brown, or dark reddish brown over the disc, and buff to light reddish buff along the margin; margin incurved when young, becoming uplifted and sometimes wavy. Flesh: buff or sometimes pinkish salmon; odor and taste farinaceous. Gills: sinuate or adnexed, close; white to light buff, discoloring salmon or orange-buff overall, and sometimes fuscous or reddish brown on the edges; entire or eroded; with many lamellulae. Stalk: 3–8 cm long, 6–17 mm thick, nearly equal or sometimes clavate, with a rounded or slightly bulbous base, solid when young, becoming hollow; surface silky-fibrillose with projecting surface fibrils; white over the upper half, bruising or becoming salmon, yellowish salmon, or dark reddish salmon below.

Microscopic features: spores 5.7–6.7 × 4.3– 4.8 µm, elliptic; cylindric or clavate cheilocystidia present. Occurrence: scattered, in groups, or clusters under hardwoods, especially oak and beech; described from southeastern Michigan but not a well-known species so its distribution, frequency, and abundance have not been determined. Edibility: unknown. Comments: Application of KOH to the surface of the cap, stalk, or flesh produces a yellowish green reaction. Tricholoma orirubens Quélet, a European species whose occurrence in North America has not been well established, has a grayish, fibrillosefelty to squamulose cap, white gills that become light pink to dull pinkish red, and white stalk flesh that becomes bluish to bluish green at the base. The epithet, insigne, means decorated, remarkable, or extraordinary.

De scr ip t ions of Speci e s

88

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Tricholoma intermedium Peck Cap: 4–10 cm wide, convex when young, becoming plane, sometimes with a broad umbo; surface viscid when fresh and moist, soon dry, coated with tiny golden brown to yellow-brown or reddish brown flattened scales and fibrils over a yellowish background; margin typically inrolled at first, then decurved well into maturity, smooth, yellow to brownish yellow. Flesh: white; odor farinaceous; taste mild to farinaceous. Gills: sinuate, close; white to whitish; with many tiers of lamellulae; edges often eroded at maturity. Stalk: 3–9 cm long, 1–2.5 cm thick, nearly equal, scurfy; white to whitish, often stained brownish, especially near the base. Microscopic features: spores 5–7 × 3.7–4.8 µm, elliptic, variously shaped cheilocystidia sometimes present and may be abundant.

scattered or in groups under conifers; widely distributed across northern North America, extending south along the West Coast to California; not common. Edibility: reportedly edible, but we haven’t tried it. Comments: Tricholoma leucophyllum Ovrebo and Tylutki is considered a synonym by some mycologists and we have followed that view here. However, others feel that two species are involved. Tricholoma subluteum (p. 149) differs primarily in having long radiating fibrils on the cap margin. The epithet, intermedium, means intermediate, in reference to T. equestre (p. 60) and T. sejunctum (p. 140); C. H. Kauffman (1918) felt that T. intermedium merely represents a variety of T. sejunctum, whereas, according to Michael Beug, Daniel Stuntz considered T. leucophyllum to be a variant of T. equestre, which would lend support for the existence of two species. Obviously more work is needed to better define the species concept(s) and its (or their) application. Occurrence:

T. intermedium (A) WV 89

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T. intermedium (B) WV

T. intermedium (C) WV

De scr ip t ions of Speci e s

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90

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T. intermedium (D) WV

T. intermedium (E) ID

91

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Tricholoma luteomaculosum A.H. Smith Cap:

4–11 cm wide, convex at first, becoming broadly convex to nearly plane at maturity, sometimes broadly umbonate; surface dry, appressed-fibrillose and typically squamulose near the margin; dark brownish gray or bluish gray overall or sometimes dull yellowish brown over the disc, staining yellow where bruised; margin incurved at first, becoming uplifted and wavy. Flesh: whitish to pale gray, bruising greenish yellow; odor and taste farinaceous. Gills: sinuate, moderately broad, close to subdistant; pale gray to drab, often stained dark brownish gray to blackish on the edges, sometimes developing yellow tones near the cap margin in age. Stalk: 5–12 cm long, 1–2.5 cm thick, nearly equal or with a tapered base, solid or hollow; surface dry, silky fibrillose, white to pale gray, occasionally discoloring pale yellowish brown. Microscopic features: spores 5.5–7 × 4–5 µm (original description), 6–10 × 4–6.5 µm (reported in the literature), elliptic to broadly elliptic; clavate to ventricose or flask-shaped cheilocystidia and pleurocystidia present, the pleurocystidia readily collapsing and sometimes hard to see; pseudoparenchymatous hypodermium present. Occurrence: solitary to gregarious under hardwoods or in mixed woodlands, usually with oak; apparently widely distributed across northern North America and south into California; not common, or not often recognized.

unknown, but the farinaceous odor and taste suggest it is inedible. Comments: When Smith described this species, he remarked on its similarity to T. scalpturatum (p. 138), which he felt differed by growing with fir, lacking well-developed pleurocystidia, and often having a cortina when young, cap scales that often turn reddish, and gills that turn yellow only in age. In contrast to Smith’s concept of the species, recent European descriptions of T. scalpturatum indicate a species that grows with oak or other hardwoods, often lacks a cortina, and does not necessarily have red-staining cap scales. Thus, T. luteomaculosum and T. scalpturatum could be closely related or even the same. Tricholoma atroviolaceum (p. 44) is similar but it has darker violet-gray cap scales, flesh that may stain reddish gray when cut, and lacks cystidia on its gills. Tricholoma atrosquamosum (p. 43) has darker blackish fibrils and squamules on its cap, the cap margin is inrolled and distinctly bearded when young, its gills are whitish, the stalk is pale gray with appressed black fibrils, and it lacks cystidia on its gills. The epithet, luteomaculosum, means yellow-spotted.

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Edibility:

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T. luteomaculosum TN

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Tricholoma magnivelare (Peck) Redhead Matsutake, Pine Mushroom, American Matsutake, White Matsutake

Cap: 5–20 cm wide, convex when young, becoming broadly convex to plane; margin cottony and inrolled initially; surface dry, white to whitish and nearly smooth at first, soon developing flattened fibers which aggregate into squamules; squamules becoming yellowish brown to reddish brown, especially on the disc. Flesh: white; very firm, dense; odor variously described as aromatic, spicy-sweet like cinnamon Red Hots candy, like old gym socks, or a mix of the above; taste mild. Gills: slightly sinuate to adnexed, crowded to close; white to whitish, often staining brownish in age; covered by a thick, white, cottony-membranous partial veil when young. Stalk: 4–15 cm long, 1.3–5 cm thick, nearly equal or often tapered downward, usually with a pointed base, dry, solid, sheathed by a white veil from the base up to a thick cottony ring, veil breaking into patches and becoming reddish brown as cap expands; ring white on the upper surface, reddish brown below. Microscopic features: spores 5–7 × 4.5–5.5 µm, broadly elliptic. Occurrence: solitary, scattered, or in groups under conifers, usually on nutrient-poor soils such as dune sands; widely distributed in North America but is most common on the West Coast where it occurs in quantities large enough to support commercial harvesting. Edibility: edible and considered choice by many, but mediocre by others. Comments: Tricholoma magnivelare is one of the more widely sought-after fungi in the West, both as a recreational and commercial commodity. Formerly known as Armillaria ponderosa Peck, it is recognized by the stocky stature, overall white color, often with brown fibrils or scales on cap and lower stalk, thick stalk that usually tapers to a pointed base and bears

a large cottony ring, and firm flesh with a characteristic spicy odor. The very similar T. matsutake (S. Ito and S. Imai) Singer is enormously popular in Japan and large quantities of matsutake collected in western North America are imported to satisfy the demand, creating an important conservation and sociological issue there. The volatile spicy odor (famously described by David Arora as a cross between cinnamon Red Hots candy and dirty socks) demands that the mushroom be used in simple dishes and not over-cooked. Results of molecular studies (Chapela and Garbelotto 2004) suggest that there are three main evolutionary lineages comprising the matsutakes: one from the western United States and Canada, one from Mexico (perhaps extending into Central America), and a third extending from the eastern United States and Canada across Europe and into Asia. Thus, it is possible that three or more species of matsutake exist in North America. Matsutake is most likely to be confused with T. caligatum (p. 50), which usually is smaller and more slender, has darker brown colors, usually lacks the spicy to fragrant odor, and often has a bitter taste; or with Catathelasma imperiale and C. ventricosum, both of which are pale colored and have white spores, prominent (double) rings, very firm texture, and pointed stalk bases. However, both of the latter species lack matsutake’s spicy odor and have an overall coloration with more gray than warm brown tones. Amanita smithiana is a dangerously poisonous western species that also is white with an often rooting stalk base. It too lacks the spicy odor, usually is more slender in stature, and has softer flesh than matsutake. If you squeeze the stipe of A. smithiana between your thumb and forefinger, it will crush or burst, whereas the flesh of T. magnivelare is so firm it will only give a little, unless riddled by fly larvae. Many other large whitish mushrooms have been mistaken for matsutake, including Russula brevipes

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and Tricholoma apium (p. 36). The mycelium of T. magnivelare often is parasitized by Allotropa virgata (candystick), a red and white plant that lacks chlorophyll and cannot make its own food as most plants do. The epithet, magnivelare, means having a large veil.

T. magnivelare (A) WA

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T. magnivelare (B) OR

T. magnivelare (C) OR

De scr ip t ions of Speci e s

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T. magnivelare (D) ID

97

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Tricholoma manzanitae Baroni and Ovrebo Cap: 4–10 cm wide, convex when young, becoming broadly convex to plane or depressed; surface viscid when fresh, often with leaf litter adhering to it, glabrous or with brownish agglutinated appressedfibrillose scales over the disc; white overall when young, soon orange-white to pale orange over the disc and nearly to the margin, eventually becoming brownish orange or brown over the disc, often developing scattered reddish stains; margin inrolled at first, soft-cottony, becoming decurved, entire, or irregularly wavy in age. Flesh: moderately thick, firm; white; odor not distinctive; taste mild or slightly bitter. Gills: sinuate, moderately broad, crowded, edges entire at first, becoming eroded; initially white, soon developing a pale salmon cast, becoming pale yellowish then light grayish salmon or darker, often with scattered reddish stains. Stalk: 3–4.5 cm long, 1.5–3 cm thick, equal, slightly tapered, or somewhat bulbous toward the base; scurfy on upper portion, glabrous and smooth to irregularly bumpy to the base, solid, dry; upper portion pale yellow over a white background, white below, often developing reddish stains over the base especially after handling.

Microscopic features: spores 5–7 × 4–5.5 µm, elliptic; very long, slender caulocystidia present. Occurrence: scattered, clustered, or in groups under manzanita or madrone, sometimes with nearby Douglas-fir and/or live oak; described from northern California and fairly common there; also likely to occur elsewhere along the Pacific Coast, following the distribution of manzanita and madrone. Edibility: unknown, but avoidance is recommended because of its likely close relationship to poisonous brownish tricholomas. Comments: The striking color changes of the cap and gills are the most distinctive features of this mushroom. Reddish stains can develop on all parts in any stage of its development. Tricholoma dryophilum (p. 58) also is white when young, but develops more reddish brown coloration, lacks any yellow coloration on the cap and stalk, and grows with oak. The many other brown-capped tricholomas differ by lacking the great change in color exhibited by T. manzanitae. See the comments under T. pessundatum (p. 121) for key features of these species. The epithet, manzanitae, refers to the fungus’s association with manzanita.

De scr ip t ions of Speci e s

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T. manzanitae CA

99

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Tricholoma marquettense Ovrebo 

(Not illustrated)

The distinctive features of T. marquettense are the viscid, gray cap that is frequently rugulose, the strongly farinaceous odor and taste, and the white gills and stalk that do not discolor. Tricholoma olivaceobrunneum (p. 116) has a similarly rugulose cap, but it is dark brown, olive-brown, or brownish gray, and its gills are whitish to buff and discolor brown on the edge. Tricholoma nigrum (p. 110) has a darker cap and its gills may discolor, but otherwise it is very similar to T. marquettense both macro- and microscopically. Tricholoma niveipes (p. 112) can occur in similar habitats and has a farinaceous odor, but differs by its darker, grayish black, virgate cap and lack of a pseudoparenchymatous hypodermium. Tricholoma virgatum (p. 172) has a non-viscid, silvery gray virgate cap with a sharp umbo, while T. argenteum (p. 38) has a non-viscid, pale silvery gray cap that can be virgate in places. The latter two species also lack a strong farinaceous odor and taste, and neither has a pseudoparenchymatous hypodermium. The epithet, marquettense, refers to Marquette, Michigan, where the type collection was made.

Cap: 2.8–10 cm wide, broadly convex to nearly plane at maturity, often with a low umbo; surface viscid when fresh, becoming dry, glabrous, or with scattered appressed fibrils, frequently rugulose between the disc and margin; light to medium gray with a white margin, sometimes with irregular white areas or gray streaks, rarely white overall; margin often wavy and sometimes uplifted. Flesh: white to watery gray; odor and taste strongly farinaceous. Gills: sinuate, close; white, becoming sordid yellowish when old; edges entire; lamellulae numerous. Stalk: 4–7 cm long, 8–14 mm thick, nearly equal with a rounded base, solid initially but becoming hollow; surface silky-fibrillose with occasional projecting fibers, typically pruinose at the apex; white, not discoloring. Microscopic features: spores 6.7–7.6 × 4.8– 5.7 µm, elliptic to broadly elliptic, often slightly pip-shaped; cylindric to clavate cheilocystidia and a pseudoparenchymatous hypodermium present. Occurrence: solitary, scattered, or in groups under pines, especially jack pine; eastern North America; fairly common. Edibility: unknown, but the strong farinaceous odor and taste suggest it is inedible.

Comments:

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Tricholoma moseri Singer Cap: 2–4.5 cm wide, campanulate when young, becoming broadly convex to nearly plane, occasionally with an umbo; surface dry, wooly overall when young, remaining so over the disc at maturity, radially appressed-fibrillose to minutely squamulose overall, often splitting radially to reveal the whitish flesh; grayish brown to blackish with a paler margin; margin inrolled and heavily bearded when young, usually with white fibrils remaining on the edge in age. Flesh: white to gray; odor and taste farinaceous or not distinctive. Gills: sinuate, narrow, close to subdistant; pale gray to gray, developing dark gray to blackish spots. Stalk: 2–6 cm long, 5–18 mm thick, nearly equal or occasionally with a slightly bulbous base; dry, solid, silky-fibrillose, pruinose at the apex; white to pale gray overall; cortina absent. Microscopic features: spores 7.2–10.6 × 3.8– 5.8 µm, elliptic to narrowly elliptic; cheilocystidia sometimes present; clavate, cylindric, to broadly clavate caulocystidia present.

solitary, scattered, or cespitose under conifers, including pines and, in California, red fir; described from a high-elevation pine forest in Mexico, quite common in the Sierra Nevada, and has been reported from the Washington Cascades and the Rockies, so it appears to be widespread in the western mountains; usually late spring to mid-summer in montane areas, encountered less frequently in fall. Edibility: unknown. Comments: The small stature, minutely squamulose dark brownish gray cap, pale gray gills, white stalk, lack of a cortina, lack of a pseudoparenchymatous hypodermium, and montane occurrence often near melting snowbanks are the distinctive features of this mushroom. Tricholoma scalpturatum (p. 138) is paler than T. moseri, has a farinaceous odor and taste, and often has a cortina when young. Tricholoma myomyces var. cystidiotum (see comments under T. myomyces, p. 108) also is similar and its flesh lacks a farinaceous odor and taste, but it has a cortina when young, a pseudoparenchymatous hypodermium, and shorter, broader spores. The epithet, moseri, honors the late Austrian mycologist Meinhard Moser. Occurrence:

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T. moseri (A) WA

T. moseri (B) WA

T. moseri (C) WA De scr ip t ions of Speci e s

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T. moseri (D) CA

T. moseri (E) CA

103

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Tricholoma muricatum Shanks Cap: 4–11 cm wide, convex to broadly convex, sometimes with a low, broad umbo when young, becoming plane and sometimes with a depressed center at maturity; surface viscid, becoming dry, often shiny, innately radially fibrillose and streaked; dull reddish brown on the disc, pinkish brown towards the margin; margin inrolled at first, becoming down-turned, and finally lobed and wavy, often with short radial ridges. Flesh: white; odor farinaceous or not distinctive; taste farinaceous or bitter-farinaceous. Gills: sinuate, moderately broad, close, thin; buff when young, rapidly becoming orange-white, discoloring brown, often with red-brown edges. Stalk: 3–8 cm long, 1–2.5 cm thick, nearly equal or tapered slightly toward the base, which may be somewhat root-like, solid or hollow; surface dry, silky-fibrillose with irregular belts of recurved fibrils; pruinose and concolorous with the gills at the apex; becoming brown to brownish orange, darkening from the base upwards or when bruised or handled. Microscopic features: spores 4.8–7.2 × 2.9– 4.3 µm, narrowly elliptic to elliptic.

solitary, scattered, or in groups or clusters under Bishop, Monterey, and shore pines in coastal woodlands of California, Oregon, and Washington, possibly extending into British Columbia and southern Alaska; fairly common. Edibility: unknown, but should be avoided because several similar brownish tricholomas are poisonous. Comments: Tricholoma muricatum is one of several very similar mushrooms with viscid brownish caps, a group that is in need of critical study. Tricholoma pessundatum (p. 121) is very similar and also grows in association with pines, but its cap is not radially fibrillose, it has slightly smaller spores, and the hypodermium of the cap is slightly different. Given the small magnitude of these differences, one might be forgiven for considering these two species to be the same. Most of the other species with viscid brownish caps occur with trees other than pine. See the comments under T. pessundatum (p. 121) for the key features of the species in this group. The epithet, muricatum, is derived from the specific epithet of Bishop pine, Pinus muricata, with which T. muricatum often occurs.

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Occurrence:

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T. muricatum (A) OR

T. muricatum (B) CA

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Tricholoma muskokense nom. prov. (Ovrebo) 

(Not illustrated)

Cap: 4.5–8 cm wide, convex when young, becoming broadly convex to nearly plane; surface viscid at first, later dry, densely matted-fibrillose over the disc with minute, appressed, radial fibrils, typically naked near the margin, occasionally with scattered squamules; tan or light yellowish brown nearly overall, paler cinnamon-buff to pinkish buff near the margin. Flesh: buff; odor and taste farinaceous. Gills: sinuate, close; light buff, staining or discoloring cinnamon-brown to darker brown on the edges and in patches on the faces; margins entire; lamellulae numerous. Stalk: 5–6.5 cm long, 7–15 mm thick, nearly equal, typically with a rounded base, solid; surface silkyfibrillose with projecting fibrils; buff at the apex, light tan to cinnamon-buff over the lower portion, discoloring cinnamon-brown.

Microscopic features: spores 5–6.2 × 3.5–4 µm, narrowly elliptic. Occurrence: in groups under pine and spruce; northeastern North America; not common. Edibility: unknown. Comments: The tan to light yellowish brown viscid cap and light buff gills and stalk that discolor cinnamon-brown are the distinctive features of T. muskokense. Tricholoma pessundatum (p. 121) and related species have darker brownish caps. The epithet, muskokense, refers to the Muskoka area of Ontario where the species was described. The species name was proposed by Clark Ovrebo in his doctoral dissertation (1980), but it has not yet been validly published.

De scr ip t ions of Speci e s

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Tricholoma mutabile Shanks 

(Not illustrated)

Cap: 3–9 cm wide, broadly convex to nearly plane, with a broad or prominent umbo, sometimes depressed over the center in age; surface viscid, glabrous, often radially rugulose between the disc and the margin; pale warm gray or pale violetgray on the disc, progressively paler with irregular darker gray streaks toward a white or silvery margin, darkening in age to medium or dark gray with violet tones, occasionally with yellow-brown areas; margin incurved when young, becoming uplifted and wavy or lobed, remaining white to silvery gray. Flesh: very thin; white, becoming grayish in age, dull pink or pale orange in base of stalk; odor strongly sweet-farinaceous or reminiscent of grapefruit; taste strongly farinaceous. Gills: sinuate, close; white, discoloring pale golden brown in age. Stalk: 5–10 cm long, 9–25 mm thick, nearly equal or clavate, base abrupt, solid or hollow; surface dry, silky fibrillose; white, discoloring pale golden brown when handled or bruised, base dull pink or pale orange. Microscopic features: spores 5.8–7.7 × 4.3– 5.8 µm, broadly elliptic to subglobose; cylindric to clavate cheilocystidia and a pseudoparenchymatous hypodermium present.

scattered to gregarious under tanbark oak or in mixed conifer forests; coastal mountains and foothills of the Sierra Nevada, California, and also reported from the Olympic Peninsula, Washington, so probably present in similar habitats in Oregon and southern British Columbia; a fairly recently described and largely unknown species so its distribution and frequency are not known. Edibility: very little information is available, but reportedly has caused gastrointestinal upset. Comments: Tricholoma griseoviolaceum (p. 77) has a similar cap color when young, but it becomes darker in age and is not radially rugulose, it lacks a pseudoparenchymatous hypodermium, and has narrower spores. Tricholoma nigrum (p. 110) has a dark slate-gray cap that becomes medium dark gray with a lighter gray edge at maturity, often has blackish squamules over the upper portion of its stalk, the flesh in its stalk base is white, not dull pink or pale orange, and it has longer spores. The epithet, mutabile, means changeable, presumably in reference to the color changes the cap undergoes with age. Occurrence:

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Tricholoma myomyces (Persoon) J. E. Lange Cap: 2–6 cm wide, bluntly conic when young, becoming convex to nearly plane, sometimes shallowly depressed or with a low, broad umbo; margin incurved when young; surface dry, covered with matted fibrils, often forming dense squamules at maturity; gray to dark gray, fuscous, or brownish black. Flesh: pale gray; odor and taste not distinctive. Gills: sinuate, close; pale gray when young, becoming nearly free and whitish or pale gray; covered by an evanescent cortina in the button stage. Stalk: 4–7 cm long, 5–10 mm thick, nearly equal; silky to slightly scurfy; whitish. Microscopic features: spores from specimens with 4-spored basidia 6.5–7.5 × 4–5 µm; spores from specimens with 2-spored basidia 8.5–10.5 × 4–6 µm; elliptic to ovate; pseudoparenchymatous hypodermium present. Occurrence: scattered or in small to large groups under conifers; widely distributed across northern North America; fairly common.

Edibility:

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edible, but use caution as some similar grayish tricholomas are poisonous. Comments: Tricholoma myomyces var. cystidiotum Shanks is virtually identical but its gills have scattered to abundant, broadly clavate cheilocystidia. Tricholoma terreum (p. 159) is very similar and also grows under conifers, especially pines, but is said by some mycologists to lack a cortina in the button stage. Increasingly, the two species are being synonymized by European mycologists. Tricholoma scalpturatum (p. 138) has a paler cap with fewer squamules, its flesh has a farinaceous odor and taste, the gills often develop yellow stains, and it lacks a pseudoparenchymatous hypodermium. Tricholoma moseri (p. 101) is small, has abundant tiny scales on a dark gray cap, pale gray gills, a white stalk, and relatively long narrow spores, lacks a cortina and pseudoparenchymatous hypodermium, and occurs primarily in the spring and early summer in the mountains of western North America. The epithet, myomyces, means mouse fungus, referring to the cap color.

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T. myomyces (A) NY

T. myomyces (B) QC

T. myomyces (C) CA 109

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Tricholoma nigrum Shanks and Ovrebo Cap: 3.5–10 cm wide, rounded-conic when young, becoming broadly convex to nearly plane at maturity; surface moist to tacky, densely matted-fibrillose to squamulose at the center, becoming glabrous or with scattered minute squamules toward the margin, occasionally virgate in places, often rugulose when mature; dark slate-gray overall when young, becoming medium dark gray with a lighter gray margin, finally dingy brownish gray; margin decurved to straight, frequently wavy, occasionally lobed. Flesh: white to very light gray; odor and taste strongly farinaceous. Gills: deeply adnexed at first, becoming sinuate at maturity, close to subdistant; white or occasionally light gray, sometimes discoloring blackish on the edges; entire or often eroded. Stalk: 7.5–15 cm long, 1–3 cm thick, equal to slightly clavate, often twisted or bent, with a rounded or abruptly tapered base, solid or hollow; surface silky-fibrillose; white, often with blackish squamules over the upper half or near the apex, not discoloring when handled, becoming dingy buff in age. Microscopic features: spores 6.5–9 × 4.5– 6 µm, elliptic; clavate cheilocystidia present, but not always conspicuous, pseudoparenchymatous hypodermium present, but not always well developed. Occurrence: scattered under coastal pines or in mixed conifer forests dominated by Douglas-fir and western hemlock; reported from Oregon and Washington, but it is a relatively recently described and not often recognized species, so little is known about its distribution, frequency, and abundance.

unknown, but the farinaceous odor and taste argue against it being edible. Comments: Tricholoma marquettense (p. 100) is very similar in most respects, including microscopic features, but its cap is paler gray, the gills don’t discolor, and the stalk lacks black scales. Tricholoma mutabile (p. 107) has a pale gray cap when young, lacks the blackish squamules on the stalk, the flesh at the base of its stalk discolors dull pink or pale orange, and its spores are shorter. Tricholoma atroviolaceum (p. 44) is similar but its cap has violet tones and the flesh may stain reddish gray when cut. Tricholoma atrosquamosum (p. 43) has darker blackish fibrils and squamules on its cap, the margin is inrolled and distinctly bearded when young, and it lacks a pseudoparenchymatous hypodermium. Tricholoma pardinum (p. 119) is somewhat similar but its cap has a distinctly spotted appearance from the blackish scales on a whitish background, its stalk lacks blackish squamules, and it usually is larger and bulkier, has clamp connections, and lacks a pseudoparenchymatous hypodermium. The epithet, nigrum, means black, in reference to the color of the young caps.

De scr ip t ions of Speci e s

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Edibility:

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T. nigrum (A) WA

T. nigrum (B) WA

T. nigrum (C) WA 111

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Tricholoma niveipes Peck Cap: 3–12 cm wide, hemispheric to convex, becoming nearly plane at maturity, occasionally with a low, broad umbo; surface viscid at first, then dry, matted fibrillose over the disc, elsewhere virgate with blackish fibrils; dark grayish brown or grayish black to purplish black, often paler toward the margin; margin entire, sometimes wavy or split. Flesh: fairly thin; white to pale gray; odor and taste farinaceous. Gills: arcuate to emarginate or sinuate, close, rather narrow; snow-white to pale gray; lamellulae numerous. Stalk: 3–8 cm long, 6–25 mm thick, equal or enlarged downward, solid or stuffed; surface silkyfibrillose, snow-white, sometimes with grayish or brownish discolorations, especially when handled. Microscopic features: spores 7–9 × 3–4 µm, nearly cylindric to elongate; cylindric to clavate cheilocystidia and pleurocystidia sometimes present.

solitary, scattered, or in groups, usually in sandy soil under conifers, especially pines such as Pinus rigida and P. banksiana; widely distributed in eastern North America from Ontario and Québec south to Florida and Texas; locally common. Edibility: reportedly edible, but we haven’t tried it. Comments: The lower portion of its stalk often is coated with sand. Tricholoma portentosum (p. 125) differs by having broader spores and yellowish tones in its cap, gills, and stalk. Tricholoma myomyces (p. 108) is similar but its cap is dry, gray to dark gray, fuscous, or blackish and covered with matted fibrils that often form dense squamules at maturity, its flesh lacks a distinctive odor and taste, and a cortina is present on young specimens. The epithet, niveipes, means white-footed, in reference to the snow-white stalk.

De scr ip t ions of Speci e s

112

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Occurrence:

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T. niveipes (A) FL

T. niveipes (B) FL

T. niveipes (C) TX 113

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Tricholoma odorum Peck Cap: 2–9 cm wide, convex with a broad umbo and incarved margin when young, becoming flat or shallowly depressed; surface dry, dull or shiny, smooth, greenish yellow at first then fading to yellowish buff to yellowish tan, at times with a brownish disc. Flesh: moderately thick; whitish; odor farinaceous or like coal tar; taste farinaceous or otherwise unpleasant. Gills: emarginate to adnate, close, broad; pale yellow to yellowish buff. Stalk: 4–12 cm long, 6–12 mm thick, nearly equal or enlarged toward the base, sometimes bulbous, dry, smooth, solid, at times longitudinally twisted; pale yellow to greenish yellow, especially toward the base.

Microscopic features: spores 7–11 × 5–7 µm, elliptic, spindle-shaped, or almond-shaped; clamp connections present but not abundant. Occurrence: solitary or in groups or clusters in broadleaf and mixed woodlands; widely distributed in North America; occasional. Edibility: inedible. Comments: Tricholoma odorum belongs to a small group of malodorous tricholomas, the members of which differ principally in their color. Tricholoma inamoenum (p. 86) is whitish to buff. Tricholoma sulphureum (p. 157) has a pale to bright sulfuryellow cap and stalk, subdistant gills, and somewhat smaller spores. Tricholoma sulphurescens (p. 154) has a whitish cap that bruises or ages dull yellow and has a more variable odor. The epithet, odorum, means scented.

T. odorum (A) WV

De scr ip t ions of Speci e s

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114

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T. odorum (B) GA

T. odorum (C) FL

115

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Tricholoma olivaceobrunneum Ovrebo 

(Not illustrated)

Cap: 3–8 cm wide, obtuse to convex when young, becoming broadly convex to nearly plane, occasionally slightly depressed over the disc, sometimes with a low umbo; surface viscid when fresh, becoming moist or dry, glabrous or with scattered minute appressed fibrils, frequently rugulose; dark brown, olivaceous brown, olive-gray, or brownish gray with a buff, olive-buff, or grayish buff margin; margin occasionally wavy or lobed. Flesh: pale buff to watery olive-gray; odor and taste strongly farinaceous. Gills: sinuate to adnexed, close; whitish to pale buff, discoloring brownish or brownish gray on the edges; entire; with numerous lamellulae. Stalk: 4–9 cm long, 4–24 mm thick, equal or tapered slightly toward the apex, with a rounded or abruptly tapered base, solid or sometimes hollow; surface silky-fibrillose with some projecting fibrils; white but often dingy at the base. Microscopic features: spores 6.2–8.6 × 4.3– 5.7 µm, elliptic; cylindric to clavate cheilocystidia and a pseudoparenchymatous hypodermium present.

solitary, scattered, or gregarious under oak and beech, as well as jack pine and possibly other conifers; northeastern North America; occasional. Edibility: unknown, but the strong farinaceous odor and taste suggest it is inedible. Comments: Tricholoma fucatum (Fries) P. Kummer is a very similar European species that differs by having a somewhat scaly cap and stalk that are similarly colored. Tricholoma marquettense (p. 100) is very similar in many respects, but its cap is light medium gray with a white margin, and its gills are white and do not discolor. Tricholoma luteomaculosum (p. 92) has a dry fuscous to grayish brown cap that is densely matted-fibrillose and has pleurocystidia on its gills. Tricholoma portentosum (p. 125), which occurs under conifers, is distinguished by the considerable yellow coloration of the gills and stalk, and by the lack of both cheilocystidia and a pseudoparenchymatous hypodermium. The epithet, olivaceobrunneum, means olive-brown, in reference to the cap.

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Occurrence:

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Tricholoma palustre A.H. Smith Cap: 3–9 cm wide, obtuse to convex when young, becoming broadly convex to nearly plane or slightly depressed; surface dry or slightly moist, mattedfibrillose or rarely areolate-squamulose over the disc, with long, innate, radial fibrils and often with scattered squamules between disc and margin; light yellowish green to pale green or olive and generally tinged brown or grayish to blackish over the disc or outward in irregular patches, occasionally virgate with darkened fibrils or streaked gray or black; margin incurved at first, becoming uplifted and wavy. Flesh: buff or pale green; odor not distinctive; taste slowly acrid. Gills: sinuate, close to crowded; light greenish yellow, yellowish green, or pale green, occasionally discoloring brownish on the edges; entire at first, then eroded, often splitting transversely; lamellulae numerous. Stalk: 2.5–8 cm long, 5–13 mm thick, nearly equal, with a rounded, abruptly tapered, or slightly bulbous base, solid or hollow; surface silky-fibrillose with projecting fibrils; whitish or with a very faint tinge of olive, occasionally discoloring brown or light pinkish, especially at the base. Microscopic features: spores 6.2–7.6 × 5.2– 5.7 µm, broadly elliptic to subglobose; filiform, cylindric, or clavate cheilocystidia often present and sometimes abundant.

scattered or in groups or clusters under beech and oak; northeastern North America; fairly common. Edibility: unknown, but the acrid taste suggests it is inedible. Comments: Application of PDAB anywhere on the mushroom produces a bright pink reaction. Tricholoma aestuans (p. 32) also has an acrid taste and is similar in color, but its cap has a less conspicuous brown or grayish to blackish cast over the disc, it grows in association with conifers, and its spores are slightly narrower. Given the seemingly minor nature of these differences, the two could represent one species, in which case the epithet aestuans would have priority. Tricholoma sejunctum (p. 140) has a viscid cap with dark radiating fibrils, a white stalk and gills that are tinged yellow, farinaceous odor and taste, and it lacks cheilocystidia. Tricholoma acre (p. 30) has the same stature and taste and can grow in the same habitats, but it differs in its grayish rather than greenish yellow coloration. The epithet, palustre, means marsh or swamp, in reference to the habitat where the type collection was made; however, it is not restricted to marshes and swamps. Occurrence:

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T. palustre (A) TX

T. palustre (B) VA

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118

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Tricholoma pardinum (Persoon) Quélet  Cap: 4–15+ cm wide, convex to hemispheric when young, becoming broadly convex to plane, sometimes with a low, broad umbo; surface dry, with a whitish background overlain by small, brownish gray to blackish, fibrillose scales that give an overall spotted appearance, typically darker over the disc where the scales are more concentrated; margin inrolled at first, becoming decurved or uplifted. Flesh: fairly thick, firm; white or pale gray; odor and taste farinaceous. Gills: sinuate to deeply adnexed, close, broad; white to creamy white, sometimes discoloring grayish or brownish when bruised; edges even at first, becoming eroded. Stalk: 8–15+ cm long, 1–3 cm thick, solid, clavate at first, becoming nearly equal at maturity; surface nearly smooth or somewhat appressed-fibrillose at the apex; white to whitish, sometimes tinged pale gray or stained pale yellowish or brownish at the base.

Dirty Trich, Poisonous Trich

Microscopic features: spores rather variable in length, 6–10 × 5–6.5 µm, elliptical to slightly ovate; clavate cheilocystidia present, but not always conspicuous; clamp connections abundant. Occurrence: scattered or in groups under conifers; widely distributed in northern North America, south into Colorado and California; occasional. Edibility: poisonous, can cause severe gastrointestinal upset. Comments: Other large grayish tricholomas, such as T. huronense (p. 81), T. nigrum (p. 110), and T. virgatum (p. 172), differ by having more evenly colored, smooth or streaked rather than scaly-spotted caps. The grayish to brownish species, typified by T. terreum (p. 159), are smaller and their often more densely wooly or scaly caps are more evenly colored and often have a felt-like appearance. Tricholoma venenatum (p. 168) is very similar in size and general appearance but is pale tan or buff overall, and the cap scales appear more brownish than blackish. The epithet, pardinum, refers to a leopard, from the spotted appearance of the cap.

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T. pardinum (A) WA

T. pardinum (B) WA

T. pardinum (C) WA De scr ip t ions of Speci e s

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120

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Tricholoma pessundatum (Fries) Quélet  Cap: 5–15 cm wide, convex when young, becoming broadly convex to flat; surface viscid when moist, smooth; chestnut-brown over the disc, paler golden brown toward the margin; margin incurved when young, uplifted and often faintly striate or with short ridges in age. Flesh: white, staining reddish; odor and taste farinaceous. Gills: emarginate to adnexed, close, broad; whitish to buff, spotting and staining reddish. Stalk: 7–10 cm long, 1–2 cm thick, nearly equal or enlarged toward the base, solid, smooth or slightly fibrillose or scurfy; whitish when young, becoming reddish brown. Microscopic features: Ovrebo (1980) observed two spore-size variants: 5.7–6.7 × 3.8–4.8 µm and 4.8–5.7 × 2.9–3.8 µm, with the conifer collections tending to fall in the latter range, similar to the size reported in most European publications (4–6 × 2.5–3.5 µm). Occurrence: scattered or in groups or clusters, sometimes among mosses, under conifers, hardwoods, or in mixed woodlands; across northern North America and extending south in the Appalachians; commonly reported but the potential for confusion with similar species makes it hard to assess the reliability of many reports. Edibility: poisonous, like many similar red-brown species, causing severe gastrointestinal upset.

Red-brown Trich

Comments: Tricholoma pessundatum is a member of the confusing group of viscid-capped brownish tricholomas. In Europe, the dominant concept is of a species associated principally with pines, or sometimes other conifers, but not hardwoods, so many North American reports probably represent misidentifications. Tricholoma tridentinum Singer is considered a synonym by some European mycologists, although not by others. Tricholoma terriferum Peck is a little-known North American species that, from its description at least, seems to not differ in any significant way from T. pessundatum. Similar species include T. populinum (p. 123), which grows in dense groups in association with cottonwood, poplar, and aspen; T. fulvimarginatum (p. 72), also with poplars; T. muricatum (p. 104) with West Coast, coastal, closed-cone pines; T. manzanitae (p. 98), another West Coast species, with manzanita; T. transmutans (p. 161) with conifers or in mixed woodlands; T. ustale (p. 164) with oaks; T. fracticum (p. 70) with pines (in Europe at least); and T. stans (Fries) Saccardo (considered by many to be a synonym of T. pessundatum) also with pines. This whole group is in desperate need of critical study, including molecular analyses and comparison of North American and European material, before we can have a hope of assigning names confidently.

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T. pessundatum (A) NY

T. pessundatum (B) NY

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122

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Tricholoma populinum J.E. Lange The Sandy, Sand Mushroom, Poplar Trich

Cap: 8–12 cm wide, convex when young, becoming broadly convex; surface viscid when moist, often with adhering debris and sand, glabrous, sometimes with water spots near the margin; pale pinkish brown to reddish brown, darker over the disc, paler or whitish toward the edge; margin inrolled at first, becoming uplifted and wavy. Flesh: thick; white; odor and taste farinaceous. Gills: sinuate or adnexed, close; white, staining reddish brown in age, especially along the edges. Stalk: 5–10 cm long, 1–3 cm thick, nearly equal, tapered toward the base, or sometimes clavate, solid; surface white at first, colored like the cap at maturity, sometimes yellowish at the base. Microscopic features: spores 5–6.5 × 3.5–4 µm, elliptic.

scattered or often in dense groups or clusters in sandy soil under poplar, cottonwood, and aspen; widely distributed in North America, but seemingly more common in the West. Edibility: edible. Comments: Tricholoma populinum is one of several very similar mushrooms with viscid brownish caps, a group that is in need of critical study in North America. Current species concepts often emphasize the type of tree with which they occur, and the association of T. populinum with Populus is distinctive. Tricholoma fulvimarginatum (p. 72) also occurs with poplars but differs by the orange-brown cap with yellowish tones on the margin and the brown-marginate gills. For other similar species, see the comments under T. pessundatum (p. 121). The epithet, populinum, refers to the tree genus Populus, with which this fungus forms ectomycorrhizas. Occurrence:

T. populinum (A) WA

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T. populinum (B) WA

T. populinum (C) AK

De scr ip t ions of Speci e s

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124

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Tricholoma portentosum (Fries) Quélet  Cap: 6–12 cm wide, broadly conic when young, becoming broadly convex to nearly flat; surface smooth, viscid to nearly slimy when fresh and moist; gray to grayish brown or olive-brown to nearly black overall, sometimes with faint yellow or purplish tints, streaked with innate dark gray fibrils; margin incurved at first, then expanding. Flesh: white; fragile; odor and taste farinaceous or not distinctive. Gills: emarginate, adnexed, or almost free, close, broad; white, becoming flushed with yellow, especially near the cap margin, occasionally tinted yellow or gray overall. Stalk: 5–10 cm long, 1–2 cm thick, nearly equal, often curved, solid, coated with tiny fibrils; whitish, sometimes with yellow tints in age, especially in the flesh at the base. Microscopic features: spores 5.5–7.6 × 3–5 µm, elliptic to oval. Occurrence: solitary, scattered, or often in groups, sometimes among mosses, typically under pines but also reported from other habitats; widespread in North America; common, at least in the East. Edibility: edible and choice.

Sticky Gray Trich

Comments: Tricholoma niveipes (p. 112) also has a viscid, dark-colored cap and white stalk, but differs in having narrower spores and lacking yellow coloration in its cap, gills, and stalk. Tricholoma virgatum (p. 172) is similar but has a sharply umbonate cap with a more conspicuously virgate surface and acrid or bitter flesh. Tricholoma sejunctum (p. 140) has a yellow to golden yellow cap covered with radiating brownish to blackish fibrils and occurs with either conifers or hardwoods. Tricholoma griseoviolaceum (p. 77), described from California, differs by the absence of yellow colors, having violet tones in the cap, gills that turn pinkish brown, a stronger farinaceous odor, and its association with oaks. Alexander Smith reduced T. avellaneifolium (Murrill) Murrill to a variety of T. portentosum. He felt it differs by lacking yellowish tints and streaks on the cap, and by having a longer stalk and more pronounced odor and taste. In many areas, T. portentosum is among the last mushrooms to fruit in the fall and, in the Northeast, sometimes can be collected under snow. The epithet, portentosum, is derived from the Latin portentos, meaning monstrous, hideous, unnatural, or marvelous. We’re guessing Fries had the last meaning in mind when he named it.

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T. portentosum (A) NY

T. portentosum (B) WV

De scr ip t ions of Speci e s

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T. portentosum (C) WV

T. portentosum (D) ID

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Tricholoma pudorinum nom. prov. (Ovrebo) Cap: 3–11 cm wide, hemispheric when young, becoming broadly convex to nearly plane; surface viscid then dry in age, matted-fibrillose on the disc, with scattered appressed fibrils toward the margin, occasionally striate on the margin; dark brown to dark reddish brown over the center, dark buff, yellowish cinnamon, light pinkish brown or light brown near the margin; margin incurved at first, becoming uplifted and somewhat wavy. Flesh: buff to light brown; odor and taste farinaceous. Gills: sinuate or adnexed, close; light pinkish buff, discoloring brownish on the edges or in patches on the faces, sometimes becoming reddish brown to brown nearly overall; entire; lamellulae numerous. Stalk: 2.5–11 cm long, 7–15 mm thick, nearly equal with a rounded or abruptly tapered base, often twisted and bent, hollow; surface silky-fibrillose with projecting fibrils; light pinkish buff at first, soon reddish brown to brown in large patches or nearly overall. Microscopic features: spores 5.7–6.7 × 4.3– 5.2 µm, elliptic to broadly elliptic.

scattered or in clusters under beech and oak; described from Michigan and known from Québec, but not often reported so its distribution, frequency, and abundance are not well established, however it is likely to occur in similar habitats elsewhere in the Great Lakes region and perhaps beyond. Edibility: unknown, but several brown-capped species are poisonous. Comments: The pinkish buff gills and stalk that both discolor brown, the pinkish tones of the cap, and the often twisted and bent stalk are the distinguishing features of T. pudorinum. Among the other brownish, viscid-capped tricholomas, T. ustale (p. 164) is the likeliest to occur with oak and beech. It differs by lacking the distinctive pink coloration. The epithet, pudorinum, means blushing (literally, full of shame), in reference to the pinkish tones of the gills and stalk. The species name was proposed by Clark Ovrebo in his doctoral dissertation (1980) but has not yet been validly published.

De scr ip t ions of Speci e s

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Occurrence:

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T. pudorinum (A) QC

T. pudorinum (B) QC

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Tricholoma pullum Ovrebo 

Dusky Trich

Cap: 4.5–15 cm wide, obtusely conic or convex with an incurved margin, becoming broadly convex to plane; surface dry, densely fibrillose over the disc with the fibrils extending toward a smooth to sometimes squamulose margin, conspicuously virgate nearly overall; dark gray, typically paler between the fibrils and toward the margin, at times developing bronze tones in age. Flesh: thick, firm; grayish white in the cap, whitish in the stalk; odor not distinctive; taste bitter or acrid. Gills: sinuate or narrowly attached, often seceding, moderately broad, close; whitish to pale pinkish gray or grayish buff, often developing dull yellow tints, dark gray on portions of the edges and faces; lamellulae numerous. Stalk: 3–8 cm long, 1–2.5 cm thick, nearly equal, smooth to somewhat silky; whitish, sometimes with yellowish tints in age. Microscopic features: spores 6.5–8 × 5–6 µm, broadly elliptic; cylindric to clavate cheilocystidia present.

scattered or in groups under hardwoods, especially beech and oak; northeastern North America south to the central Appalachians; not common. Edibility: unknown, although the bitter or acrid taste suggests it is not edible. Comments: The dark gray, virgate cap along with the bitter or acrid taste and dark gray tints on the edge of the gills are distinctive. Tricholoma sciodes (Secretan) Martin, described from Europe and reported from California, appears to be nearly identical and could be the same species. Tricholoma virgatum (p. 172), widely distributed and fairly common in North America, is similar but has a sharply pointed umbo, grows in conifer and mixed woodlands, and lacks the dark gray tints on the gill edge. Tricholoma acre (p. 30) is also similar but it is virgate over, at most, portions of the cap. This is another group of species that is in need of careful study, including integration of morphological, ecological, and molecular data. The epithet, pullum, means dusky or dark-colored.

De scr ip t ions of Speci e s

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Occurrence:

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T. pullum (A) MA

T. pullum (B) WV

T. pullum (C) WV 131

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Tricholoma roseoacerbum A. Riva 5–12 cm wide, convex with inrolled margin, expanding to plane, margin remaining inrolled or incurved well into maturity; smooth or minutely granulose without radial patterning; pinkish brown to pale reddish, margin pale yellowish or whitish, radially ridged, at least when young. Flesh: whitish in cap, pale yellowish in stalk, sometimes pinkish below cap cuticle or above the gills; firm; without distinctive odor or taste, or with slightly farinaceous or bitter taste. Gills: emarginate, crowded; whitish to pale yellowish, may become stained or spotted brownish; entire or eroded; lamellulae numerous. Stalk: 2–7 cm long, 1.5–3 cm thick, nearly equal or with a tapered base; surface dry, smooth or slightly fibrillose, apex floccose; white or whitish at first, yellowish at apex, and brownish at base, sometimes yellowing or browning overall in age. Microscopic features: spores 4–6 × 2.5–4 µm, elliptic; cylindric, clavate, or balloon-like cheilocystidia and a pseudoparenchymatous hypodermium present. Cap:

under conifers, often in moss beds; northeastern North America; not common. Edibility: unknown. Comments: Tricholoma roseoacerbum is essentially a pinkish-capped version of T. acerbum (Bulliard) Ventura, whose cap is cream, pale yellowish, or cinnamon buff, and which typically occurs with oak and beech. In fact, some mycologists suspect that T. roseoacerbum is merely a color form rather than a distinct species. Tricholoma acerbum has been said to occur in North America, but we are unaware of welldocumented reports. Other similar species include T. apium (p. 36) with a yellowish brown to reddish brown cap surface and areolate cracks exposing the underlying whitish flesh, and T. imbricatum (p. 82), often larger, with a brown, somewhat cracked cap. The epithet, roseoacerbum, refers to the species appearing to be a rosy-capped version of T. acerbum. Occurrence:

T. roseoacerbum (A) QC De scr ip t ions of Speci e s

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132

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T. roseoacerbum (B) QC

T. roseoacerbum (C-1) ON

T. roseoacerbum (C-2) ON 133

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Tricholoma saponaceum (Fries) P. Kummer 

Soap-scented Trich

Cap: 2.5–15+ cm wide, convex when young, becoming broadly convex to nearly flat, often with a broad umbo; surface initially smooth, often cracked over the disc at maturity, moist and slippery when fresh; cap color highly variable, pale greenish yellow, light green to grayish green, grayish olive, lead-gray, bluish gray, greenish gray, or grayish brown to brown; margin inrolled when young, becoming expanded, sometimes whitish or discoloring pinkish brown. Flesh: white to pale green, usually pinkish brown or orangish in the stalk base; odor similar to nonperfumed soap, farinaceous, or not distinctive; taste mild or slightly soapy. Gills: emarginate to nearly adnate or adnexed, subdistant, broad; white to yellowish or with a greenish tinge, sometimes discoloring pinkish brown or orangish. Stalk: 3.5–10+ cm long, 1–4 cm thick, nearly equal or sometimes ventricose or clavate with a rounded, tapered, or slightly bulbous base; solid, dry, smooth; white to pale green, often pinkish to orangish near the base. Microscopic features: spores 5–7.6 × 3.5–5 µm, elliptic; clamp connections scattered throughout. Occurrence: solitary, scattered, or in groups under conifers or sometimes hardwoods; widely distributed in North America and fairly common in many areas; often found in spring, unlike most tricholomas.

inedible. The highly variable appearance of T. saponaceum, including features such as stature, cap color, and the occasional presence of scales on the stalk, has led some, mostly European, mycologists to name several different varieties. For instance, T. saponaceum var. squamosum (Cooke) Rea (= var. ardosiacum Bresadola) has blackish scales over most of the stalk and its form ardosiacum has, in addition, a slate-gray to bluish gray cap. An informal variant mentioned by Ovrebo (1980) has a brown to olive brown cap. Whether these and the many other varieties and forms recognized in Europe represent truly discrete entities, as opposed to points on a continuum within a single variable taxon, and whether they can be widely applied to North American collections remains unknown. Thus, until T. saponaceum, in the broad sense, has been subjected to critical study combining morphological, ecological, and molecular data, we agree with Shanks (1997) that the variation in color with age combined with differences between populations produces too ill-defined a picture to warrant use of formal sub-specific names. The soap-like odor and the pinkish or orangish color of the flesh at the base of the stalk are the features that tie all of the various forms together. Unfortunately, neither feature is present in every collection. The epithet, saponaceum, means soapy.

De scr ip t ions of Speci e s

134

Bessette_4937_BK.indd 134

Edibility:

Comments:

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T. saponaceum (A) NY

T. saponaceum (B) NY

T. saponaceum (C) WV 135

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T. saponaceum (D) NS

T. saponaceum (E) NM

T. saponaceum (F) NM De scr ip t ions of Speci e s

Bessette_4937_BK.indd 136

136

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T. saponaceum (G) AZ

T. saponaceum (H) AK

137

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Tricholoma scalpturatum (Fries) Quélet Cap: 2–7 cm wide, convex with a low, broad umbo when young, becoming flattened-convex; surface dry and wooly becoming matted-fibrillose, appressed-fibrillose, or appressed-scaly; often unevenly colored due to brownish or pale gray to medium gray or black fibrils scattered over the white to pale gray background, sometimes with yellowish staining in age, especially near the margin; margin incurved when young, becoming uplifted and wavy. Flesh: whitish to pale gray; odor and taste farinaceous. Gills: sinuate, moderately broad, close, thin; pale gray when young, becoming white and sometimes discoloring yellow, especially near the cap margin; entire, but often eroded; lamellulae numerous; evanescent whitish cortina present. Stalk: 2–7 cm long, 6–15 mm thick, equal to slightly clavate with a rounded or abruptly tapered base, solid or hollow; surface dry, silky-fibrillose; white, sometimes discoloring yellow-brown or pale gray-brown or pale yellow when bruised or in age. Microscopic features: spores 4.8–6.7 × 2.9– 4.3 µm, narrowly elliptic to elliptic. Occurrence: solitary, scattered, or in groups under conifers or hardwoods; widely distributed across northern North America; locally common in areas such as southeastern Canada. Edibility: unknown; in Europe reported both as edible and inedible. Comments: This species has been interpreted variously for many years and this contributes to the difficulty in knowing whether the “real thing” occurs in North America. Currently, the dominant concept in Europe is of a hardwood species that occurs only occasionally with conifers. This is opposite of the prevailing concept in North America. Similar European species include T. cingulatum (p. 52), T. argyraceum (Bulliard) Gillet, T. inocybeoides A. Pearson, T. alboconicum ( J.E. Lange) Clémençon, and

T. ramentaceum (Bulliard: Fries) Ricken, and these have been the subject of recent phylogenetic analyses (Carriconde, Gardes, et al. 2008, Carriconde, Gryta, et al. 2008, Jargeat et al. 2010, Mouhamadou et al. 2008). Results of these studies suggest that four phylogenetic species can be recognized: T. scalpturatum, T. argyraceum, T. cingulatum, and T. inocybeoides. The first two are widespread in Europe, occur mostly with hardwoods but also conifers, and are difficult to separate morphologically. These studies shed little or no light on their occurrence in North America (a collection said to represent a North American occurrence of T. argyraceum [ Jargeat et al. 2010] actually came from Europe). Tricholoma cingulatum is widespread in Europe, also occurs in North America, and is found exclusively with willows, and T. inocybeoides is less common in Europe, found with poplars and birches, but not known to occur in North America. Given the difficulty in knowing the degree to which the situation in North America corresponds to these findings, we have chosen to include just T. cingulatum and T. scalpturatum pending future study of this group. Tricholoma myomyces (p. 108) is very similar to T. scalpturatum, with a cortina when young, but it lacks the yellow staining and has a darker colored cap, pseudoparenchymatous hypodermium, and longer broader spores. Tricholoma terreum (p. 159) differs in the same features, but lacks a cortina. Tricholoma moseri (p. 101) is smaller, lacks a cortina, has flesh with either farinaceous or non-distinctive taste, and occurs primarily in spring or early summer in the the mountains of western North America. The epithet, scalpturatum, means, among other things, carved or engraved, perhaps in reference to the manner in which the cap surface breaks up into patches.

De scr ip t ions of Speci e s

138

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T. scalpturatum (A) AZ

T. scalpturatum (B) QC

139

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Tricholoma sejunctum (Sowerby) Quélet Cap: 4–9 cm wide, broadly conical to bell-shaped or convex at first, becoming broadly convex with an umbo; surface viscid at first, then dry; covered with conspicuous brownish to blackish fibrils radiating from the dark-colored disc, over a yellow to golden yellow background, color and density of the fibrils highly variable. Flesh: white to yellowish, fragile; odor farinaceous or not distinctive; taste unpleasant, typically bitter or astringent. Gills: emarginate to adnexed, subdistant to close, broad; white or tinged yellowish in places or overall. Stalk: 5–9 cm long, 1–2 cm thick, nearly equal, sometimes enlarged at the base, solid, smooth, dry; whitish to yellowish. Microscopic features: spores 5–7 × 3–5 µm, broadly elliptic to subglobose; clamp connections abundant throughout. Occurrence: solitary, scattered, or in groups under conifers, especially pine and spruce, or in mixed woodlands; widely distributed and fairly common in North America. Edibility: inedible, generally considered unpalatable.

Comments: Tricholoma sejunctum often is confused with T. equestre (p. 60), which has warmer brown colors on its (basically) yellow cap, lacks dark radial fibrils, and has gills and stalk that usually are uniformly pale to bright yellow, rather than whitish flushed with yellow. Tricholoma subluteum (p. 149) is similar in background cap color, but lacks a dark disc, its cap fibrils occur mostly toward the margin, and it lacks clamp connections. Tricholoma portentosum (p. 125) also is similar but its cap is dark grayish. Tricholoma sejunctum is a European name and it has not yet been shown that we have the “real thing” in North America. If future study shows our fungus to be different, the name T. subsejunctum Peck seemingly would be available for it. Currently Peck’s name is considered a synonym for T. sejunctum by many mycologists. Of course, given the wide variation in cap color, degree of yellow coloration of stalk and gills, and habitat, it is quite possible that we have more than one species in North America. For instance, many of our collections from conifer forests are similar in appearance to T. viridilutescens Moser, a European conifer associate. The epithet, sejunctum, means separated or to part or sever, possibly referring to the fungus differing from similar species, or perhaps to the separate nature of the cap fibrils.

De scr ip t ions of Speci e s

140

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T. sejunctum (A) NY

T. sejunctum (B) NY

141

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T. sejunctum (C) WV

T. sejunctum (D) WA

T. sejunctum (E) AK De scr ip t ions of Speci e s

Bessette_4937_BK.indd 142

142

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Tricholoma serratifolium Peck 

Saw-tooth Trich

Cap: 3–8 cm wide, convex or obtusely conical when young, becoming broadly convex to nearly plane, occasionally depressed; surface dry, densely interwoven-fibrillose, sometimes with radiating fibrils between the disc and margin, often with scattered grayish or brownish squamules, occasionally striate at the margin; sometimes white overall but typically with areas of light gray, tan, brownish gray, or yellowish brown, especially over the disc, the margin usually remaining white, at times with scattered blackish brown streaks or patches; margin incurved at first, becoming uplifted and sometimes wavy or lobed. Flesh: white or watery gray; odor not distinctive; taste mild or sometimes bitter. Gills: sinuate, close to subdistant; white, becoming dingy golden yellow near the cap margin, discoloring fuscous-black on the edges where bruised; entire at first, then eroded; lamellulae numerous. Stalk: 2.5–7 cm long, 8–30 mm thick, nearly equal to slightly clavate with a rounded or slightly bulbous base, solid or hollow; surface silky-fibrillose with projecting surface fibrils, often pruinose at the apex; white, sometimes dingy buff or pinkish at the base.

Microscopic features: spores 6.2–7.6 × 4.8– 5.7 µm, broadly elliptic; filiform, cylindric, or clavate cheilocystidia may be present and sometimes are abundant. Occurrence: scattered or in groups or clusters under oak and beech; northeastern North America; occasional. Edibility: unknown. Comments: Application of PDAB to any part of the mushroom produces a bright pink reaction. Tricholoma serratifolium can be similar in color and habitat to T. subresplendens (p. 151), but the latter has a viscid cap that lacks grayish tones, its flesh has a farinaceous odor and taste, and it does not react with PDAB. Tricholoma acre (p. 30) also can occur in the same habitats but it differs by its overall medium gray cap and acrid taste. Tricholoma venenatum (p. 168) is another pale species that can occur with oak and beech, however it is predominantly tan, generally has a scalier cap and farinaceous odor and taste, lacks the pink PDAB reaction and cheilocystidia, and possesses clamp connections. The epithet, serratifolium, means having saw-toothed leaves, in reference to the prominently eroded gill edges.

143

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T. serratifolium (A) NS

T. serratifolium (B) NS

De scr ip t ions of Speci e s

Bessette_4937_BK.indd 144

144

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Tricholoma silvaticoides (Murrill) Murrill  Cap: 4–8 cm wide, convex, becoming nearly plane, with a rounded umbo; surface viscid, shining, sometimes hygrophanous, white to creamy white or buff, sometimes with grayish or darker yellowish tints; margin weakly translucent-striate when moist, even, sometimes uplifted in age. Flesh: about 2 cm thick, somewhat fragile; white to buff, sometimes staining pale orange-brown; odor farinaceous; taste farinaceous, then bitter or unpleasant. Gills: sinuate, sometimes seceding, close to subdistant, up to 2 cm broad, often forked near the cap margin; white; with several tiers of lamellulae. Stalk: 4.5–11.5 cm long, 1–2 cm thick, nearly equal or tapered downward, often curved and pinched at the base, hollow at maturity; surface dry, silkyfibrillose; white on the upper portion, pale orangebrown on the lower half or more.

(Not illustrated)

Microscopic features:

spores 8.5–11 × 5–6 µm,

ovoid. solitary, scattered, or in groups under hardwoods, especially oak; described from Florida, but likely to occur elsewhere in the southeastern United States. Edibility: unknown. Comments: The combination of a shiny viscid cap, sinuate to nearly free gills that are often forked, and a white stalk that is orange-brown on the lower half is most distinctive. The epithet, silvaticoides, means resembling T. silvaticum, a poorly known species described by Charles Peck, the epithet of which means belonging to the forest. Occurrence:

145

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Tricholoma squarrulosum Bresadola Cap: 2–10 cm wide, convex when young, becoming broadly convex to nearly plane, often depressed; surface dry, densely interwoven-fibrillose over the disc, appressed-squamulose or recurved-squamulose between the disc and margin; dark brownish gray with a light gray to brownish gray margin; margin incurved at first, then uplifted. Flesh: white to pale gray or brownish gray; odor variously described as farinaceous and somewhat disagreeable, sweet-fruity and somewhat spicy-peppery, or not distinctive; taste farinaceous to slightly bitter. Gills: sinuate, close to crowded; light gray fading to nearly white in age, discoloring dark gray on the edges and often staining salmon-pink in some areas; entire; lamellulae numerous. Stalk: 4–8 cm long, 5–15 mm thick, nearly equal with a rounded or slightly bulbous base, solid; surface with grayish to brownish appressed or recurved squamules, or with portions silky-fibrillose and pale gray. Microscopic features: spores 5.7–7.2 × 3.8– 4.8 µm; oblong or narrowly elliptic to elliptic.

scattered or in groups under oak or beech, or in mixed woodlands (but see comments, below); widely distributed in North America; occasional. Edibility: some European sources report it as edible and good, however we have no firsthand information on North American experience. Comments: Tricholoma michiganense A.H. Smith is thought to be a synonym and Smith’s photograph accompanying the type description certainly is consistent with that interpretation. However, Smith described his fungus as having a coal-tar odor like that of T. sulphureum (p. 157), albeit milder. Thus, it is possible that two species are involved. Tricholoma atrosquamosum (p. 43) is very similar but has a smooth stalk. In light of the minor differences between them, several mycologists consider T. squarrulosum to be merely a variety of T. atrosquamosum. Our photos all are of collections made in conifer forests, so they might not be the “real” T. squarrulosum. The epithet, squarrulosum, means having small scales.

De scr ip t ions of Speci e s

146

Bessette_4937_BK.indd 146

Occurrence:

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T. squarrulosum (A) CO

T. squarrulosum (B) ID

T. squarrulosum (C) ID 147

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Tricholoma subaureum Ovrebo 

(Not illustrated)

Cap: 2.5–7 cm wide, broadly conic, convex, or plane, usually with a rounded or somewhat pointed umbo; surface dry, glabrous on the disc, the remainder with appressed radial fibrils, occasionally with scattered squamules near the margin, sometimes rimose; dark buff-brown, light golden brown, or dark olive-brown over the disc, dull yellow to yellowish buff toward the margin. Flesh: 3–10 mm thick; white to pale buff; odor and taste strongly farinaceous. Gills: sinuate, close to subdistant, 3–10 mm broad; light buff to pale yellow, often discoloring brown or olivaceous brown on the edges; entire, often split transversely; lamellulae numerous. Stalk: 3–7 cm long, 9–14 mm thick, nearly equal with a rounded base, solid; surface silky-fibrillose with scattered projecting fibrils; light buff overall or dingy buff at the base.

Microscopic features: spores 6.2–7.6 × 4.8– 5.7 µm, narrowly elliptic to elliptic; cylindric, clavate, or balloon-like cheilocystidia and a pseudoparenchymatous hypodermium present. Occurrence: solitary or in groups under beech; described from Michigan and Ontario, but likely to occur in similar habitats elsewhere in eastern North America. Edibility: unknown, however the strong farinaceous odor and taste suggest it is inedible. Comments: Tricholoma luteomaculosum (p. 92) has a darker cap that is often tinged yellowish near the edge, drab gray flesh, and pleurocystidia. The epithet, subaureum, means almost golden.

De scr ip t ions of Speci e s

148

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Tricholoma subluteum Peck Cap: 4–11 cm wide, convex or obtusely conic when young, becoming broadly convex with a low, broad umbo; surface dry to slightly viscid when moist, bald over the disc, with long yellowish radial fibrils near the margin; color variable when young, dull orange or golden yellow to yellow, usually pale yellow at maturity; often rimose. Flesh: white; odor and taste farinaceous. Gills: emarginate to adnexed to nearly free, close; white. Stalk: 7.5–12 cm long, 1–2 cm thick, nearly equal or slightly enlarged downward, dry, solid at first, hollow in age; yellow on the upper portion and white near the base, or sometimes white or yellow overall. Microscopic features: spores 5.7–7.5 × 4.8– 6 µm, broadly elliptic to subglobose.

solitary, scattered, or in groups under conifers or in mixed woodlands; widely distributed in northeastern North America, but not common. Edibility: unknown. Comments: Tricholoma intermedium (p. 89) is very similar but lacks the long radial fibrils on the cap. Tricholoma sejunctum (p. 140) has a viscid cap with dark disc and radial fibrils and tastes bitter rather than farinaceous, although the distinction is not always obvious. Microscopically, T. sejunctum differs in having abundant clamp connections. Tricholoma fumosoluteum (p. 76) has a naked yellow cap, sometimes with a brownish disc, and occurs with conifers such as spruce, fir, and hemlock. The epithet, subluteum, means almost yellow. Occurrence:

T. subluteum (A) NY

149

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T. subluteum (B) QC

T. subluteum (C) QC

De scr ip t ions of Speci e s

Bessette_4937_BK.indd 150

150

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Tricholoma subresplendens (Murrill) Murrill Cap: 2–11 cm wide, convex when young, becoming broadly convex to nearly plane, sometimes with a low, broad umbo; margin incurved initially, sometimes wavy and lobed in age; surface viscid when moist, soon dry, smooth, often slightly silky; white to cream, becoming tinged or spotted yellow or yellowish tan to tan or pale pinkish cinnamon, especially on the disc, sometimes with bluish green or blue discolorations. Flesh: white; odor and taste farinaceous or sometimes not distinctive. Gills: sinuate to adnexed, becoming nearly free at maturity, broad, close to subdistant; white to whitish; edges uneven, sometimes finely scalloped in age; lamellulae numerous. Stalk: 5–10 cm long, 1–2.5 cm thick, nearly equal or tapering slightly downward, smooth to silkyfibrillose or slightly scurfy, dry; whitish, often stained yellow to brownish yellow or bruising bluish green or blue near the base. Microscopic features: spores 5.6–7 × 4–5 µm, elliptic; clamp connections present but only in the hymenium and not abundant there. Occurrence: solitary, scattered, or in groups under hardwoods and in mixed woodlands, often near oak and sometimes hemlock; widely distributed and fairly common in eastern North America.

unknown; in Europe the very similar Tricholoma columbetta is considered edible. Comments: Tricholoma albatum (Quélet) Maublanc and d’Astis, T. albidum (p. 34), T. columbetta (Fries) P. Kummer, and T. resplendens (Fries) P. Karsten are very similar to each other and to American collections, all being white European species whose names have sometimes been applied in America. Research is now underway to clarify species concepts in this group, determine whether or not the North American and European fungi belong to the same species, and decide how to apply names. Preliminary DNA analysis of a collection made in North Carolina by one of the authors and identified as T. subresplendens (Photo C) was a close enough match (99%) to European material assigned to T. columbetta to be considered the same species. However, much more work needs to be done before any general conclusions can be drawn. Another all-white species, T. albidulum Ayala, G. Moreno and Esteve-Raventós, was described in 1997 from a single collection made under Jeffrey pine in Baja California. It is unstaining, and has globose to broadly elliptical spores and a characteristic cap cuticle with capitate terminal cells. The epithet, subresplendens, refers to the similarity of this species to T. resplendens, whose epithet means glittering or shining. Edibility:

151

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T. subresplendens (A) WV

T. subresplendens (B) WV

T. subresplendens (C) NC De scr ip t ions of Speci e s

Bessette_4937_BK.indd 152

152

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T. subresplendens (D) FL

T. subresplendens (E) FL

153

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Tricholoma sulphurescens Bresadola 5–12+ cm wide, hemispheric to convex when young, becoming nearly plane, sometimes with an umbo; surface dry, smooth, white at first, becoming creamy white, quickly staining yellow to dull yellow or brown when handled or in age. Flesh: white; odor highly variable based on reports in the literature, described as stinking, sulfurous, slightly farinaceous, pungent, disagreeable, like coal tar (may be faint), sometimes with a coconut or other fruity component; taste mild or somewhat acrid. Gills: emarginate to adnate or adnexed, narrow, close; white to creamy white, becoming yellowish in age, especially near the edge. Stalk: 3–10+ cm long, 1–3 cm thick, clavate when young, becoming nearly equal, dry, smooth to somewhat fibrillose; white at first, becoming creamy white to yellow, staining yellow when handled or in age. Microscopic features: spores 5–7 × 4–5 µm, broadly elliptic to subglobose; Ovrebo (1980) reported that clamp connections are abundant throughout the fruitbody, however, European authors characterize them as rather uncommon. Occurrence: solitary, scattered, or in groups under hardwoods, especially beech and oak in northeastern North America, but with conifers or in mixed woodlands in the West; infrequent.

unknown in North America, but considered inedible or possibly poisonous in Europe. Comments: Tricholoma sulphurescens belongs to a small group of malodorous tricholomas, the members of which differ principally by their color. The white to creamy white caps that quickly stain yellow to dull yellow or brown, relatively close gills, and the coal tar or otherwise strong odor are the distinctive features of this mushroom. Some European mycologists consider T. resplendens (Fries) P. Karsten to be the same species. Tricholoma sulphureum (p. 157) has a sulfur-yellow cap and stalk, reddish brown streaks and spots on the cap, subdistant gills, and somewhat smaller spores. Tricholoma inamoenum (p. 86) is whitish to buff, doesn’t turn yellow, has subdistant to distant gills, and often is smaller and more slender. Tricholoma odorum (p. 114) is greenish yellow when young and then fades to yellowish buff or yellowish tan, and often has a brownish disc. The odor of all three of these species is more consistently coal-tar-like than that of T. sulphurescens. The epithet, sulphurescens, means turning sulfur-yellow.

De scr ip t ions of Speci e s

154

Cap:

Bessette_4937_BK.indd 154

Edibility:

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T. sulphurescens (A) MN

T. sulphurescens (B) FL

155

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T. sulphurescens (C) ID

T. sulphurescens (D) WA

De scr ip t ions of Speci e s

Bessette_4937_BK.indd 156

156

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Tricholoma sulphureum (Bulliard) P. Kummer  Cap: 2–7.5 cm wide, hemispheric to conic when young, becoming convex, then broadly convex; surface smooth, dull, pale to medium-dark sulfuryellow, sometimes with reddish brown streaks and spots; margin incurved initially and remaining so well into maturity. Flesh: thin; sulfur-yellow; odor strongly of coal tar or a heavy floral scent like that of hyacinths or paperwhites; taste unpleasant or mild. Gills: sinuate to adnate, broad, subdistant; yellow; edges even; lamellulae numerous. Stalk: 2.5–7.5 cm long, 7–20 mm thick, nearly equal or enlarged downward, dry, solid; usually concolorous with the cap and with a white tomentose base. Microscopic features: spores 8.5–11 × 5–7 µm, broadly elliptic to almond-shaped; clamp connections rare (at the base of basidia) or absent. Occurrence: solitary, scattered, or in groups under both hardwoods and conifers; widely distributed in North America, but generally not common.

Sulfur Trich

inedible at best, possibly poisonous. Tricholoma sulphureum belongs to a small group of malodorous tricholomas, the members of which differ principally by their color. In a recent European study, Comandini et al. (2004) concluded that T. bufonium (Persoon) Gillet, a similar species with a reddish to purplish cap, represents a color form of T. sulphureum. However, they considered only a small number of collections. Tricholoma sulphurescens (p. 154) has a more variable and often less unpleasant odor, its cap is white to creamy white and stains yellow or brown when bruised, the gills are relatively close, and it has white flesh. Tricholoma odorum (p. 114) has a cap that is greenish yellow when young and fades to yellowish buff to yellowish tan, paler yellow flesh that tastes farinaceous to disagreeable, close gills, and a paler yellow stalk. Tricholoma inamoenum (p. 86) is pale buff to whitish. The epithet, sulphureum, refers to the yellow color of the mushroom. Edibility:

Comments:

157

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T. sulphureum (A) ME

T. sulphureum (B) WA

T. sulphureum (C) WA De scr ip t ions of Speci e s

Bessette_4937_BK.indd 158

158

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Tricholoma terreum (Schaeffer) P. Kummer Earth-colored Trich, Gray Trich

Cap: 2.5–7.5 cm wide, convex when young, becoming broadly convex to plane, sometimes with an umbo; surface dry, densely tomentose to radially fibrillose-scaly; grayish brown to blackish brown or dull brownish; margin incurved at first, becoming uplifted and often wavy. Flesh: thin, fragile; white to grayish; odor weakly farinaceous or not distinctive; taste mild. Gills: adnexed to sinuate, subdistant; white and entire when young, not covered by a cortina in the button stage, becoming grayish and often eroded; with several tiers of lamellulae. Stalk: 2.5–6 cm long, 5–10 mm thick, nearly equal or slightly enlarged downward, dry, solid at first, becoming hollow; sometimes slightly fibrillose but not scaly; white to grayish white. Microscopic features: spores 5–8 × 4–5.5 µm, broadly elliptic; well-differentiated pseudoparenchymatous hypodermium usually present. Occurrence: scattered or in groups or clusters under conifers; widely reported in North America; seemingly fairly common, but many reports probably represent misidentifications. Edibility: edible, but not recommended because it can be confused with other species, some of which are poisonous.

Comments: Whether the “real” T. terreum occurs in North America (at least some mycologists feel it does not) is an open question that will not be answered easily. Even in Europe, mycologists hold differing views over how broadly or narrowly to apply this name, and so several similar species and varieties are variously reported in the literature. Certainly we have one or more fungi that generally fit the broad concept of T. terreum, so we have chosen to use the name until things get clarified. Tricholoma myomyces (p. 108), considered a synonym by some mycologists, is very similar but has a cortina (the mycologists that synonymize the two species claim that T. terreum also usually has a cortina). Tricholoma moseri (p. 101) is smaller with a more scaly cap, lacks a pseudoparenchymatous hypodermium, and occurs in the Rockies, Sierra Nevada, and other western mountain ranges in late spring and early summer. Tricholoma scalpturatum (p. 138) also is similar but has yellowish stains on the gills and farinaceous odor and taste, and lacks a pseudoparenchymatous hypodermium. Tricholoma orirubens Quélet, a European species occasionally said to occur in North America, differs by growing with hardwoods and having salmon to reddish tinges on the gills and bruised flesh. The epithet, terreum, means earthy.

159

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T. terreum (A) WV

T. terreum (B) WV

T. terreum (C) WV De scr ip t ions of Speci e s

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160

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Tricholoma transmutans Peck Cap: 2.5–11 cm wide, conic when young, becoming convex to broadly convex or nearly plane, often with a low, broad umbo; surface slightly viscid then dry, with densely matted fibrils over the disc, dark brown to reddish brown with paler, sometimes striate, margin; margin inrolled, often remaining so or becoming down-turned or straight. Flesh: white to buff; odor and taste farinaceous. Gills: adnate to emarginate or nearly free, close, thin; yellowish white to buff, faces becoming spotted or edge entirely reddish. Stalk: 4–15 cm long, 7–20 mm thick, equal or slightly clavate, often long relative to the cap diameter and slightly rooting, solid or hollow; surface dry, appressed-fibrillose, with occasional projecting fibrils, apex pruinose; yellowish buff to grayish orange, dull reddish brown toward the base at maturity. Microscopic features: spores 5–8.6 × 3.4– 6.2 µm, elliptic to broadly elliptic. Occurrence: scattered or in groups under conifers or in mixed woodlands; apparently widespread in North America (for instance it occurs on the West Coast and is said to be perhaps the most common tricholoma in Québec), but confusion over species concepts makes it difficult to determine its distribution, frequency, and abundance.

unknown, but several similar species are poisonous. Comments: Tricholoma transmutans belongs to the subset of viscid brownish tricholomas with yellowish colors, although historically there has been some uncertainty over whether and to what degree the yellow coloration was present in the material Peck studied in describing the species. The members of this group all have gone by the name T. flavobrunneum at one time or another, and they include T. fulvum (p. 73) and T. pseudonictitans Bon. The yellowish tones in the gills and stalk separate them from species such as T. pessundatum (p. 121). Whether T. transmutans merits recognition as a separate species remains to be determined, however, we have chosen to take a conservative approach and retain it pending future study. It was treated as T. nictitans (Fries) Gillet by Ovrebo (1980) and Shanks (1997). However, that name has been abandoned by most European mycologists. There, depending on which concept is accepted, either T. fulvum or T. pseudonictitans is applied, and some consider the two to be synonymous. The buff gill color and conifer forest habitat of our fungus suggest it is more similar to the latter species. The epithet, transmutans, means shifting, presumably in reference to the change in gill color as the mushroom ages. Edibility:

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T. transmutans (A) NS

T. transmutans (B) QC

T. transmutans (C) BC De scr ip t ions of Speci e s

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Tricholoma tumidum (Persoon) Ricken  Cap: 5–10 cm wide, convex when young, becoming broadly convex to nearly plane with a low, broad umbo; surface dry, appressed-fibrillose, often virgate, sometimes squamulose over the disc, cracking radially in age and exposing the underlying white flesh; color a mixture of yellow-ochre, greenish to grayish yellow, and bright yellow, often whitish on the margin, with a dark brown to yellowish brown disc; margin incurved when young, becoming uplifted and wavy. Flesh: thin; white; odor and taste faintly farinaceous. Gills: sinuate, close; whitish to pale cream or pale buff, staining yellow on the edges. Stalk: 3–11 cm long, 8–30 mm thick, ventricose, base tapering to a point, solid; surface silkyfibrillose, pruinose at the apex; pinkish or pale orange toward the base, white elsewhere, with fibrils discoloring pale brown. Microscopic features: spores 4–7.2 × 3– 5.3 µm, subglobose to broadly elliptic; broadly clavate cheilocystidia present; clamp connections present, but not abundant.

(Not illustrated)

in groups in coastal mixed evergreen forest; reported from California, where it apparently is rare and almost unknown. Edibility: unknown. Comments: We have followed the description given by Shanks (1997), which differs from European descriptions we have reviewed, especially with respect to color. The dominant European concept is of a dark-gray-capped mushroom with a whitish or pale gray stalk. However, according to Shanks (1997), the distinguishing features of this mushroom include the dry radially fibrillose cap with yellow and brown colors, and the ventricose stalk. Unfortunately, Persoon’s original description does not mention color, so it is difficult to know what his original concept of the species was. In any event, it seems quite possible that the California and European material represent different species, but critical comparative study of additional North American collections will be required before the situation can be clarified. It is interesting to note that Shanks’s description is very similar to that of T. arvernense (p. 40), which recently has been found in Washington and southern Oregon. The epithet, tumidum, means swollen, in reference to the ventricose stalk. Occurrence:

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Tricholoma ustale (Fries) P. Kummer 

(Not illustrated)

Cap:

4–15 cm wide, convex at first, becoming broadly convex, occasionally broadly umbonate; surface viscid then dry, glabrous nearly overall except for scattered appressed radial fibrils on the disc; grayish orange to brownish orange with a dark brown disc, darkening in age; margin incurved at first, becoming uplifted, lobed and wavy. Flesh: whitish; odor slightly farinaceous or not distinctive; taste mild to bitter. Gills: adnate to sinuate, sometimes with a subdecurrent tooth, close; pale buff or yellowish white, discoloring reddish brown in spots or on the edges. Stalk: 3.5–9 cm long, 1.5–3.5 cm thick, nearly equal or tapered slightly toward the base, solid or hollow; surface appressed-fibrillose to finely squamulose, pruinose at the apex; whitish to pale yellow on the upper portion, progressively darker toward the base, sometimes colored like the cap in age. Microscopic features: spores 5–7.5 × 3–5 µm, elliptic.

Scattered or in groups under hardwoods, especially oaks; reported from California, may occur more widely; apparently not common. Edibility: unknown here, but considered inedible in Europe and some similar species are poisonous. Comments: Tricholoma fulvellum (Fries) Gillet is a synonym. Tricholoma ustale is one of several very similar mushrooms with brownish viscid caps, a group that is in need of critical study in North America. Thus, as is the case for many of the other species, it is not clear whether the “real” T. ustale occurs in North America and, if it does, what its distribution, frequency, and abundance are. For instance, it is possible that the California reports actually refer to T. dryophilum (p. 58). However, until additional information is available to clarify the situation, we will use T. ustale for the mushroom that is very similar to T. pessundatum (p. 121) but grows with oaks. See the comments under the latter species for a comparison of the brownish species. The epithet, ustale, is derived from the Latin for scorched or burned and here is used in the sense of a warm russet color.

De scr ip t ions of Speci e s

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Occurrence:

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Tricholoma vaccinum (Schaeffer) P. Kummer Scaly Trich, Russet-scaly Trich

Cap: 2.5–10 cm wide, conic to convex when young, becoming broadly convex to nearly plane, usually retaining a low umbo; surface coated with rusty brown fibrillose scales on a whitish background; margin initially inrolled and cottony-fibrillose, becoming expanded but remaining cottony-fibrillose and somewhat thickened, with a thin veil. Flesh: white, occasionally developing reddish tinges when cut; odor farinaceous or not distinctive; taste farinaceous, bitter, or mild. Gills: adnexed to sinuate, close; whitish buff when young, staining reddish brown when bruised or in age. Stalk: 2.5–10 cm long, 1–2 cm thick, nearly equal or slightly enlarged downward, typically hollow at maturity; fibrillose, especially in lower part; reddish brown or paler, often whitish at apex; partial veil pale brown, cottony-fibrous, not leaving a ring. Microscopic features: spores 5.5–7.6 × 4– 5.2 µm, oval to elliptic.

scattered, in groups (which can be large), or clusters under conifers, especially spruce but occasionally pine; widely distributed in northern North America and extending southward in mountain ranges such as the Appalachians and Rockies; can be quite common and abundant, especially in western spruce forests. Edibility: considered edible by some, but of poor quality. Comments: Its dry scaly cap separates T. vaccinum from most of the other hard-to-identify brownish species such as T. pessundatum (p. 121). Tricholoma imbricatum (p. 82) is similar in having a dry cap, but differs by its darker dull brown fibrillose-scaly cap that often becomes cracked on the disc in age, solid stalk, lack of a partial veil, and preference for pines. The epithet, vaccinum, is derived from the Latin word for cow and here means dun-colored. Dun is a variable neutral gray-brown with reddish to yellowish hue, presumably the color of some Roman cows. Occurrence:

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T. vaccinum (A) CO

T. vaccinum (B) AZ

T. vaccinum (C) AZ De scr ip t ions of Speci e s

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T. vaccinum (D) WA

T. vaccinum (E) WA

T. vaccinum (F) AK 167

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Tricholoma venenatum G.F. Atkinson Cap: 2.5–13 cm wide, convex when young, becoming broadly convex-umbonate; surface dry, densely matted-fibrillose over the center; whitish with scattered tan squamules that become less distinct and lighter in color toward the whitish margin; margin incurved at first, becoming uplifted and wavy. Flesh: moderately thick; white or watery gray; odor and taste farinaceous. Gills: sinuate, close, moderately broad; white when young, becoming pale ivory buff, not discoloring; edges entire; with numerous lamellulae. Stalk: 3–10 cm long, 7–25 mm thick, equal or slightly clavate, sometimes slightly bulbous, solid; surface silky-fibrillose; light buff overall, becoming dingy buff over the lower half or overall in age. Microscopic features: spores originally reported as 5–7 × 3.5–5 µm, more recent reports indicate a larger size, 7.2–10.7 × 4.8–7.2 µm, elliptic; clamp connections usually common. Occurrence: scattered or in groups or clusters; under hardwoods and hemlock in northeastern

North America, recorded from conifer forests often containing aspen in the Rockies, Grand Canyon, Sierra Nevada, and Washington Cascades; not a well-known species, so distribution, frequency, and abundance are unclear. Edibility: poisonous, causing severe gastrointestinal upset that can last for several days. Comments: It was the occurrence of several poisonings in Rochester, Michigan, that brought this previously unnamed fungus to the attention of George F. Atkinson, mycologist at Cornell University. It is more or less a tan version of T. pardinum (p. 119), which has brownish gray to blackish cap scales and scattered to abundant cheilocystidia and grows under conifers. Tricholoma huronense (p. 81), which grows under hardwoods, is also similar, but its cap margin, gills, and stalk often have drops of pinkish fluid or pinkish streaks, and it has grayish scales on its cap and the lower portion of the stalk. Like T. pardinum, it has cheilocystidia. The epithet, venenatum, means poisonous.

T. venenatum (A) NM De scr ip t ions of Speci e s

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T. venenatum (B) NM

T. venenatum (C) NM

T. venenatum (D) AZ 169

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Tricholoma vernaticum Shanks Cap: 5–17 cm wide, convex to broadly convex when young, becoming flattened-convex to plane; surface dry to moist but not viscid, glabrous, often with a whitish bloom at first, appearing streaked or appressed-fibrillose in age; white when young or when covered with litter, rapidly becoming fuscous to brown, often with olive, grayish, or pale tan areas; margin down-turned at first, becoming uplifted and often lobed and irregular. Flesh: thick; white; odor and taste strongly farinaceous, often like cucumber or watermelon rind. Gills: emarginate or adnate to adnexed when young, often seceding and appearing free in age, close, thick; white, sometimes becoming pale pinkish. Stalk: 4–14 cm long, 1.3–3.5 cm thick, equal or slightly clavate, often with an enlarged base, solid; surface dry, glabrous to silky-fibrillose above the ring, appressed-fibrillose to fibrillose-scaly below; white, staining tan to orange-brown on the lower portion; ring median to superior, fibrillose, not always obvious. Microscopic features: spores 8–12 × 4.8– 6.2 µm, narrowly elliptic to elliptic; clamp connections present. Occurrence: solitary to gregarious under conifers at higher elevations, in California common throughout the Sierra Nevada and rare in coastal forests, extends northward in the Oregon and Washington Cascades; late spring to early summer; fairly common.

unknown, but the strong farinaceous odor and taste suggest it is not edible. Comments: This fungus previously was known as Armillaria olida Thiers and Sundberg. Like many spring montane fungi, T. vernaticum often occurs as “mushrumps,” where one or more fruitbodies barely emerge from the soil, often being visible only as cracked mounds of soil and leaf litter. When found in its typical habitat and season, it is easy to identify by its white color, stocky stature, cucumber-like odor, and (often faint) ring. The nearest look-alikes in such situations are Hygrophorus subalpinus, another stocky, white, ringed mushroom, and H. camarophyllus, a similar species with gray or brownish gray cap. However, both of these can be distinguished by their broad waxy gills and lack of a strong farinaceous odor. Somewhat similar tricholomas include T. portentosum (p. 125), with a viscid, radially fibrillose, gray cap, gills and stalk often with yellow tints, and no ring; T. mutabile (p. 107), with a pale warm gray or pale violet-gray cap that is often radially rugulose between the disc and margin, stalk flesh that is dull pink to pale orange at the base, and no ring; and T. griseoviolaceum (p. 77), with a smaller viscid cap, white sinuate gills that discolor pinkish brown to grayish orange in patches, no ring, and growth with hardwoods. However, all of these occur most commonly in the fall (and winter in California), at lower elevation, and in different habitats than T. vernaticum. The epithet, vernaticum, means of the spring.

De scr ip t ions of Speci e s

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Edibility:

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T. vernaticum (A) OR

T. vernaticum (B) CA

T. vernaticum (C) CA 171

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Tricholoma virgatum (Fries) P. Kummer 

Fibril Trich, Streaked Trich

Cap: 3.5–10 cm wide, conic or obtuse with an acute umbo when young, expanding to broadly conic, convex-umbonate or nearly plane, the umbo remaining acute; surface dry, glabrous, with long, innate, radial fibrils; silvery gray to gray, darkest on the disc, light gray, silvery buff, or nearly white toward the margin; margin incurved at first, occasionally uplifted in age. Flesh: thin; pale gray; odor not distinctive; taste very acrid or slowly bitter. Gills: sinuate, close, moderately broad; whitish to very pale gray, often dingy buff in mature specimens, occasionally discoloring fuscous on the edges; lamellulae numerous. Stalk: 6–10 cm long, 9–15 mm thick, solid or hollow, equal or slightly clavate, with a rounded or rarely bulbous base; silky fibrillose, white. Microscopic features: spores 6–8.6 × 5–7 µm, broadly elliptic; cylindric to clavate cheilocystidia present. Occurrence: usually in groups, under conifers; widely distributed in North America; fairly common.

inedible, possibly poisonous. The stalk base stains light yellow with the application of KOH and the stalk surface, stalk flesh, and gills stain bright pink with the application of PDAB. Tricholoma subacutum Peck is a synonym. Tricholoma argenteum (p. 38) is similar but has a rounded to sub-acute umbo, lighter cap color, and is, at most, slightly virgate. Tricholoma pullum (p. 130) has a less acute umbo, dark gray gill edges, and grows with hardwoods, especially beech and oak. Tricholoma portentosum (p. 125), a conifer associate with mild-tasting flesh, has a darker gray or violet-gray cap, which lacks an acute umbo, and often develops yellowish flushes on the gills and stalk. Tricholoma hordum (p. 80) has a darker virgate cap, pale pinkish or violaceous tints on young gills, and grows with hardwoods. Tricholoma virgatum var. vinaceum differs only in having a pale pinkish blush on the stalk. The epithet, virgatum, means having streaks, in reference to the cap of this fungus.

De scr ip t ions of Speci e s

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Edibility:

Comments:

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T. virgatum (A) NY

T. virgatum (B) WA

T. virgatum (C) OR 173

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Some Additional Tricholomas

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T

h e fol l ow i ng pho togr a phs illustrate some tricholomas that we have not been able to identify to species with reasonable certainty. Although at least some of them seem to represent undescribed taxa (such as NAT-12) and one appears to represent a little-known species (NAT-13, T. grave?), additional collections will be needed before their taxonomic status can be confidently assessed. To the extent they are available, supporting collection notes are provided in the hope that they will assist others in recognizing these mushrooms in the field and adding to our knowledge of them.

NAT–1 5–12 cm wide, with grayish radial fibrils and scurfy punctae over a whitish background; margin incurved when young. Flesh: white, grayish under the cuticle, weakly pinkish orange (pale salmon) near the base of the stalk, not staining on exposure; odor not distinctive; taste mild or slowly slightly bitter. Gills: adnexed, close, up to 1.2 cm broad, entire, sometimes forked; white; with several tiers of lamellulae. Stalk: 5–9 cm long, 2–3.5 cm thick, white, nearly equal, deeply rooting, solid, stuffed to somewhat hollow in age; base pinched. Macrochemicals: Ammonium hydroxide on cuticle and flesh produced no color change. Occurrence: scattered and in groups in sandy soil under oak; December 2009; Dunnellon, Marion County, Florida.

thick, firm; pale gray, not staining on exposure; odor and taste strongly farinaceous. Gills: adnexed, close, up to 1.8 cm broad; dull grayish cream to pale gray, sometimes developing brown stains on the edges; with several tiers of lamellulae. Stalk: 3–9 cm long, 1–3.5 cm thick, nearly equal or enlarged downward, solid; surface pruinose at the apex and longitudinally striate below to the base; whitish to pale grayish, sometimes with reddish brown stains. Macrochemicals: ammonium hydroxide on cuticle somewhat reddish brown; no color change when applied to the flesh. Occurrence: scattered or in groups in oak woods; December, 2009; Dunnellon, Marion County, Florida. Flesh:

Cap:

NAT–2 4–13 cm wide, scurfy tomentose to appressed tomentose or scaly; brownish gray on disc and paler near the margin, often with dull reddish brown tones; margin incurved at first, becoming wavy, uneven, and often split. Cap:

NAT–3 Cap: 4–8 cm wide, umbonate; surface viscid, shinywaxy, hygrophanous, somewhat translucent-striate, pale tan to buff, darker on the disc, sometimes with grayish or yellowish tints; margin nearly plane to slightly uplifted. Flesh: whitish to pale tan; odor resembling watermelon rind or somewhat farinaceous; taste farinaceous then bitter and unpleasant. Gills: sinuate, becoming nearly free at maturity, close to subdistant, often forked near the cap margin, up to 2 cm broad; white; with several tiers of lamellulae. Stalk: 4.5–11.5 cm long, 1–2 cm thick, nearly equal or tapered downward, often curved at the base, hollow; base sometimes pinched; surface dry, scurfyfibrillose, white on the upper portion, pale orangebrown on the lower half or more, flesh staining pale orange-brown. Macrochemicals: Ammonium hydroxide on cuticle and flesh produced no color change. Occurrence: scattered or in groups in oak woods; December, 2009; Dunnellon, Marion County, Florida.

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NAT–4

NAT–10

NAT–4 (A) Photographed in mixed woods with hemlock, birch, and maple; September 2007; Cathedral State Park, Preston County, West Virginia.

Photographed under oak and other hardwoods; August 2008; Holly River State Park, Webster County, West Virginia.

NAT–4 (B) Photographed in mixed woods with hemlock, yellow birch, and rhododendron; September 2005; Little Beaver State Park, Raleigh County, West Virginia.

NAT–11

NAT–5 Photographed under oak and pine; January 2007; Silver Lake, Apalachicola National Forest, Florida.

NAT–6 Photographed in oak woodland; October 2002; Sleep Creek Wildlife Management Area, Berkeley County, West Virginia.

NAT–7 Photographed under oak and sand pine (Pinus clausa); December 2004; Alligator Point, Franklin County, Florida.

NAT–8 Photographed under oak and sand pine (Pinus clausa); December 2004; Alligator Point, Franklin County, Florida.

NAT–9 Photographed under oak and longleaf pine (Pinus palustris); January 2004; Apalachicola Bluffs, Liberty County, Florida.

De scr ip t ions of Speci e s

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Cap: 3–7 cm wide, dark gray when young, becoming light gray with a brownish disc; surface dry, with appressed gray fibrils, smaller appressed scales on disc. Flesh: thin, fragile, flesh of stalk fibrous; white; odor faint, farinaceous; taste farinaceous. Gills: attached, notched, close to subdistant; offwhite; edges with a somewhat ragged appearance. Stalk: 2.5–5 cm long, 7–15 mm thick, off-white to light gray; partial veil cortina-like, leaving a faint ring-zone on the stipe and a few hairs on the cap margin. Microscopic features: spores average 6.4 × 3.8 µm, length/width ratio = 1.7; pileipellis a cutis; no cystidia or clamp connections observed. Occurrence: Found during January 2011, on soil under coast live oak (Quercus agrifolia) on the campus of Stanford University, Santa Clara County, California.

NAT–12 Tricholoma “quercetorum” 5–20 cm across, parabolic to almost campanulate when young, becoming convex to broadly umbonate, eventually irregular, margin wavy, splitting, uplifted in age; color highly variable, dark olive to olive-brown or yellow-olive when young, becoming olive-yellow to yellow with a darker umbo and often streaked, eventually fading to pale yellow, pinkishyellow, or pinkish; surface dry, with minute floccules that disappear as the cap expands, older caps develop concentric cracking and small upturned scales. Cap:

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fragile, fibrous in stalk; white to pale yellow, developing pinkish stains, especially towards base of stalk; odor and taste farinaceous. Gills: adnexed, close to crowded; pale pink to pinkish-yellow when young, becoming pale yellow, with pinkish stains in age. Stalk: 7–20 cm long, 1.5–5 cm thick; equal to slightly enlarged towards base, often tapering at extreme base; off-white to pale yellowish to pinkish; surface covered with fine light to dark scales when young. Occurrence: solitary, scattered, or in large groups in mixed oak-beech forest; very common in southern New Hampshire and north-central Massachusetts and also reported from Maine and Québec. Comments: This apparently common but undescribed species is recognized by its variable cap color, large size, and tendency to discolor pink all over, starting on the stalk. Tricholoma davisiae (p. 56) has similar variable cap colors and pink staining but has a sharply umbonate cap and grows with conifers. Tricholoma saponaceum (p. 134) also has variable cap color and stains pink, but usually only on the stipe base, and it has paler gills. The name T. quercetorum has been proposed for this species, but there already is a species by that name (Contu 2003), so a new epithet will have to be selected if this species is to be formally published. Flesh:

NAT–13 Tricholoma grave Peck(?) 12–18 cm wide, rounded to irregularly convex, margin inrolled; pale salmon to pinkish-brown, fading in age; surface dry, appressed fibrillose when young, cracking in age. Flesh: stalk fibrous, tough, off-white, staining pale pinkish-brown; cap white, staining faint reddishbrown; odor disagreeable; taste unpleasant, slightly bitter. Cap:

adnexed, close to subdistant, thick; salmonbuff, staining reddish-brown when rubbed. Stalk: 12–20 cm long, 3–6 cm thick, enlarged below and then tapering to the base, which is well buried in the ground; pale buff to pinkish-buff to off-white; surface dry, developing uplifted scales. Occurrence: Found in mixed woods with oak, beech, and hemlock, north-central Massachusetts and southern New Hampshire. Comments: The illustrated collection has been tentatively identified as T. grave. The gill spacing, thickness, and color, along with its robust stature, appear distinctive. The epithet grave means heavy. Peck’s (1890) original description of T. grave is given below, for comparison. Gills:

Tricholoma grave, n. sp. Pileus at first hemispherical, then broadly convex, compact, glabrous, grayish-tawny and somewhat spotted when moist, paler when dry, the margin paler, involute, often irregular, clothed with a minute appressed grayish-white tomentum or silkiness, flesh grayish-white; lamellae subdistant, rounded behind or sinuate, adnexed, at first whitish, then pale ochraceous-tawny; stem stout, compact, solid, subsquamulose or furfuraceous, abruptly attenuated at the base, penetrating the soil deeply, grayish-white; spores broadly elliptical, .0003 in. [7.6 µm] long, .0002 in. [5.1 µm] broad. Pileus 5 to 8 in. [12.7–20 cm] broad; stem 3 to 4 in. [7.6–10 cm] long, 1 to 1.5 in. [2.5–3.8 cm] thick. Mixed woods of pine and oak. Manor. September. This species is remarkable for its great size and weight. It is apparently allied to Tricholoma colossus (p. 54), from which it is separated by the absence of any viscidity of the pileus, the radicating character of the base of the stem and by the flesh not assuming a reddish color.

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NAT-14 3–7 cm wide, convex becoming broadly convex to nearly plane with a distinct umbo, slightly viscid, appressed-fibrillose, silky, somewhat shiny and satiny; grayish brown to smoky brown on the disc, pale grayish brown extending outward toward a paler whitish margin, darkening and becoming brownish gray nearly overall in age; margin even, entire or sometimes splitting. Flesh: whitish, that of cap unchanging, that of stalk sometimes staining reddish brown to dull brown when cut and around larva tunnels; firm; odor strongly farinaceous or like watermelon rind; taste mild. Gills: arcuate to free, 1–1.7 cm broad, subdistant, with several tiers of lamellulae, not forked, edges entire, sometimes split; white to cream with a pale pinkish cast, unchanging when cut or bruised. Stalk: 5–7 cm long, 1–1.8 cm thick, nearly equal with an enlarged and typically curved base, covered in sandy soil from ⅓ to ¾ of its length; solid, scurfy, longitudinally striate; whitish at the apex, dull white to pale grayish brown below, staining dull brown when handled or in age. Occurrence: scattered or in groups under water oak (Quercus nigra)and southern live oak (Quercus virginiana); 17 December 2011; Okefenokee Swamp, Folkston, Charlton County, Georgia; locally common; edibility unknown. Cap:

De scr ip t ions of Speci e s

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178

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NAT-1 FL

NAT-2 FL

NAT-3 FL 179

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NAT-4 (A) WV

NAT-4 (B) WV

NAT-5 FL De scr ip t ions of Speci e s

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180

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NAT-6 WV

NAT-7 FL

NAT-8 FL 181

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NAT-9 FL

NAT-10 WV

NAT-11 CA De scr ip t ions of Speci e s

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182

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NAT-12 MA

NAT-13 MA

NAT-14 GA 183

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Excluded Species

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T

he following species ar e not described in this book because either they are no longer considered to belong to the genus Tricholoma, or we had too little information to be confident of their status as a valid species or their occurrence in North America. There are a number of possible reasons for the latter: (1) the species is unknown to us, it has not been widely or commonly recognized, and its original description is too brief to provide a well-defined concept that would allow confident identification; (2) although the species occasionally has been reported as occurring here, those reports have been scattered, not thoroughly documented, and often involve the application of European names; (3) the species is thought to be a new taxon, but has not yet been formally described. Many of the species on this list were described by the American mycologist William Alphonso Murrill. The combination of the brevity of his original descriptions and the fact that many of his collections were made in areas that have been little studied by other mycologists, particularly Florida, makes it very difficult to apply many of his names. Many of Charles Horton Peck’s species descriptions are similarly brief and difficult to interpret and apply. Inclusion on this list does not necessarily mean the species does not occur in North America. Indeed it is quite likely that some of them do, and hopefully publication of this book will spur efforts to document them. The dates following North American epithets are not necessarily those that are part of the formal authority, rather they are intended to lead to the original description of the species, by whatever name, in conjunction with the References section. The acronym ARSB indicates a species published by Charles Peck in the Annual Report of the State Botanist (of New York). ARSB dates reflect the date of publication of the report, not the year it covers.

acerbum (Bulliard) Ventura adustum (Murrill) Murrill 1939 alabamense Murrill 1914a, b alachuanum (Murrill) Murrill 1938 albobrunneum (Persoon) P. Kummer alboflavidum Peck 1872 (ARSB) album (Schaeffer) P. Kummer altaicum Singer angustifolium (Murrill) Murrill 1914a, b anomalum (Murrill) Murrill 1913 arenicola (Murrill) Murrill 1913 argyraceum (Bulliard) Gillet (but see Comments under T. scalpturatum [p. 138]) aromaticum (Murrill) Murrill 1914a, b auratocephalum (Ellis) Singer (Ellis 1876) australe (Murrill) Murrill 1944b avellaneifolium (Murrill) Murrill 1913 avellaneum (Murrill) Murrill 1913 azalearum (Murrill) Murrill 1942 bicolor (Murrill) Murrill 1913 calceifolium (Murrill) Murrill 1945a californicum (Murrill) Murrill 1913 (= T. dryophilum?) chlorinosmum (Peck) Saccardo (Peck 1879) chrysenteroides Peck 1872 (ARSB) cinnamomeum (Murrill) Murrill 1914a, b citrinifolium (Murrill) Murrill 1938 collybiiforme (Murrill) Murrill 1913 compressipes (Murrill) Murrill 1914a, b cortinatellum Singer cortinatum Singer cystidiosum A.H. Smith (= Inocybe cystidiosa Singer) czuicum (Singer) Singer (= Humidicutis czuica [Singer] Singer) decorosum (Peck) Saccardo (= Leucopholiota decorosa [Peck] O.K. Miller, T.J. Volk, and A.E. Bessette) decorum (Fries) Quélet (= Tricholomopsis decora [Fries] Singer) diemii Singer

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earleae (Murrill) Murrill 1914a, b eduriforme (Murrill) Murrill 1914a, b edurum Banning and Peck 1891 (Peck ARSB) entoloma (Murrill) Murrill 1942 fagnani Singer ferruginescens (Murrill) Murrill 1944b ferrugineum Lamoureux nom. prov. flavescens Peck 1873 formosum (Murrill) Murrill (= Tricholomopsis formosa [Murrill] Singer) fuligineum Peck 1888 (ARSB) fulvidiscum (Murrill) Murrill 1944b fumescens (Peck) Saccardo (Peck 1879 ARSB) fumidellum (Peck) Saccardo (Peck 1873) fumosellum (Murrill) Murrill 1914a, b fumosifolium Hesler (Smith and Hesler 1944) fusisporum Singer glatfelteri (Murrill) Murrill 1914a, b glaucoalbum Singer 1948 grande Peck 1891 (ARSB) grave Peck 1890 (ARSB) (but see NAT-13 [p. 177]) harperi (Murrill) Murrill 1913 hebeloma Peck 1873 hirtellum Peck 1907 humile var. bulbosum Peck (in Ricker 1902) hygrophorus (Murrill) Murrill 1944b imbricatum var. felleum (Murrill) Murrill 1944b infantile Peck 1887 (ARSB) inocybiforme (Murrill) Murrill 1914b (based on T. hebeloma Peck) ionides var. farinaceum A.H. Smith 1944a jalapense (Murrill) Murrill 1912 jamaicense (Murrill) Murrill 1912 kauffmanii (Murrill) Murrill 1914a, b lacunosum Peck 1873 lasciviforme (Murrill) Murrill 1945a laterarium (Peck) Saccardo (Peck 1873) laticeps Kauffman (= Lyophyllum laticeps [Kauffman] Clémençon) latum Peck 1913 (ARSB) leucocephaloides Peck 1897 (ARSB) limonium Peck 1873

longipes (Murrill) Murrill 1914a, b lugubre Peck 1896 (ARSB) luteomaculans G.F. Atkinson 1909 maculatescens Peck 1891 (ARSB) maculatum (Murrill) Murrill 1940 magnum Banning and Peck 1891 (Peck ARSB) malodorum (Murrill) Murrill 1942 margarita (Murrill) Murrill 1940 marginatum (Peck) Singer (= Humidicutis marginata [Peck] Singer) melaleuciforme (Murrill) Murrill 1945a melaleucum var. caespitosum (Murrill) Murrill (= Melanoleuca melaleuca var. caespitosa Murrill) memmingeri (Murrill) Murrill 1914a, b Michiganeuse A.H. Smith 1942 microspermum (Murrill) Murrill 1945a mostnyae Singer multifolium (Murrill) Murrill 1914a, b multipunctum Peck 1873 murrillianum Singer (= Armillaria arenicola Murrill 1912a = T. magnivelare [Peck] Redhead) naucoria (Murrill) Murrill (= Tricholoma fallax Saccardo = Calocybe fallax [Saccardo] Redhead and Singer) nigrocystidium Ovrebo (1973) nom. prov. nobile Peck 1889 (ARSB) nuciolens (Murrill) Murrill 1913 odoriferum (Murrill) Murrill 1914a, b olesonii (Murrill) Murrill 1913 olivaceiflavum (Murrill) Murrill 1916 oliveum Farlow and Burt 1929 oreades (Murrill) Murrill 1913 orirubens Quélet pallidum Peck 1897 panaeoliforme (Murrill) Murrill 1943 peralbum (Murrill) Murrill 1942 pinicola (Murrill) Murrill 1913 piperatiforme (Murrill) Murrill 1946 piperatum Peck 1899 planiceps Peck 1913 (ARSB) platyphylloides (Murrill) Murrill 1944b portolense (Murrill) Murrill 1913

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praebulbosum (Murrill) Murrill 1942 praecox (Murrill) Murrill 1914a, b praemagnum (Murrill) Murrill 1914a, b praevillosum (Murrill) Murrill 1942 psammopus (Kalchbrenner) Quélet pseudolilaceum (Singer) Pilát pseudosordidum Singer 1945 pseudoterreum Lamoureux nom. prov. pulverulentipes (Murrill) Saccardo and Trotter (Murrill 1917b) quercetorum Lamoureux nom. prov. (but see T. quercetorum Contu 2003) quercicola (Murrill) Murrill 1949 radicatum Peck 1903 (ARSB) rancidulum Banning and Peck 1891 (Peck ARSB) resplendens (Fries) P. Karsten rhizoideum A.H. Smith 1944a rickenii (Singer) Pilát (= Lepista rickenii Singer) rimosum Peck 1902 (ARSB) robinsoniae (Murrill) Murrill 1914a, b roseibrunneum (Murrill) Murrill 1913 rubescentifolium (Peck) Saccardo (= Callistosporium luteo-olivaceum [Berkeley and M.A. Curtis] Singer) rubicundum Peck 1873 rudericola (Murrill) Murrill 1913 russuloides Murrill 1952 rutilans (Schaeffer) P. Kummer (= Tricholomopsis rutilans [Schaeffer] Singer) secedifolium (Murrill) Murrill 1913 semivestitum Peck (= Lentinellus semivestitus [Peck] Singer) sienna (Peck) Saccardo (Peck 1872 ARSB) silvaticum Peck 1889 (ARSB) sordidum (= Lepista sordida [Schumacher] Singer) stans (Fries) Saccardo striatellum (Murrill) Murrill 1913 striatifolium Peck 1878 (ARSB) subacidum (Murrill) Murrill 1914a, b subacre (Murrill) Murrill 1942 subacutum Peck 1889 (ARSB) subannulatum (Peck) Zeller 1922

subargillaceum (Murrill) Murrill 1914a, b subcerinum A.H. Smith 1942 subcinereiforme (Murrill) Murrill 1914a, b subcinereum Peck 1909 (ARSB) subcylindrisporum (Murrill) Murrill 1945c subdecorosum (Murrill) Murrill 1943 subdurum Banning and Peck 1891 (Peck ARSB) subfuligineum (Murrill) Murrill 1914a, b subfulvidiscum (Murrill) Murrill 1949 subisabellinum (Murrill) Murrill 1911, 1912 sublatum Murrill 1942 subluridum (Murrill) Murrill 1913 submaculatum Peck 1893 (ARSB) submulticeps (Murrill) Murrill 1913 subpessundatum (Murrill) Murrill 1913 subrimosum (Murrill) Murrill 1942 subrufescens Ellis and Everhart 1893 subsaponaceum Peck 1912 (ARSB) subsejunctum Peck 1912 (ARSB) subsilvaticum (Murrill) Murrill 1949 subterreiforme (Murrill) Murrill 1945b subterreum (Murrill) Murrill 1914a, b subtransmutans (Murrill) Murrill 1914a, b subumbrinum A.H. Smith 1944a subvelatum (Murrill) Murrill 1913 subvolkertii (Murrill) Murrill 1945c tenuipes (Murrill) Murrill 1913 terraeolens Peck 1885 (ARSB) thujinum Peck 1873 titans H.E. Bigelow and Kimbrough (= Macrocybe titans [H.E. Bigelow and Kimbrough] Pegler, Lodge, and Nakasone) totilividum (Murrill) Murrill 1943 tottenii (Murrill) Murrill 1914a, b trentonense Peck 1872 (ARSB) tricolor Peck (= Leucopaxillus tricolor [Peck] Kühner) tristiforme Kauffman 1921 umbrosum A.H. Smith and M.B. Walters 1943 unakense (Murrill) Murrill 1914a, b unifactum Peck 1906 (ARSB) ustaliforme Murrill 1944b

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ustaloides Romagnesi virescens Peck 1891 (ARSB) virgineum (Murrill) Murrill 1942 viscidipes (= Armillaria viscidipes Peck; not an armillaria in the current sense, but the transfer to Tricholoma apparently has not been made) viscosum Peck 1904 volkertii (Murrill) Murrill 1914a, b watsonii (Murrill) Murrill 1945b westiana (Murrill) Murrill 1940 westii (Murrill) Murrill 1944b yatesii (Murrill) Murrill 1914a, b

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Glossary

aborted, abortive: arrested in development and only partly formed aciculate: superficially marked as if scratched with a pin acrid (taste): sharp, biting, peppery, producing a burning sensation in the mouth acute: pointed or sharp-edged; compare with obtuse adnate (gill attachment): describes gills that are attached to the stalk along their entire height adnexed (gill attachment): narrowly attached; describes gills that are attached to the stalk along only a part of their height, or appear as though a small part of the gill had been cut away next to the stalk agaric: a gilled mushroom agglutinated: firmly attached as if glued together, often used in reference to fibrils allantoid: curved and sausage-shaped ammonium hydroxide: see NH4OH amyloid: staining blue, blue-gray, gray, or nearly black when exposed to iodine such as in Melzer’s reagent or Lugol’s solution; compare dextrinoid anastomosing: describes gills or other structures that split and re-join in network fashion, like a braided stream annular zone: remnants of the partial veil occurring as a thin band of fibrils or gluten on the stalk; less substantial than an annulus; (= ring-zone) annulus: remnants of the partial veil occurring as an attached or loose skirt-like ring of cottony or membranous tissue left on the stalk of a mushroom after expansion of the cap; (= ring) apex (–ices): the uppermost part of something; with respect to mushrooms, usually in reference to a stalk, hypha, cystidium, or spore apical: occurring at the apex of a structure apiculate: having one or more short projections

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apiculus: short projection at the base of a basidiospore by which it was attached to the sterigma appendiculate (cap margin): hung with bits of veil tissue appressed: flattened onto a surface appressed-fibrillose: composed of fibrils that lie flat on a surface appressed-pubescent: having short, soft, downy hairs that lie flat on a surface appressed-scaly: having flattened scales appressed-tomentose: having a thick covering of wooly hairs that are flattened onto a surface arachnoid: silky fibrillose, like a cobweb arcuate-decurrent (gill attachment): describes gills whose edges are turned upward near the stalk and then descend for a distance down the stalk areolate: marked out into angular block-like units by cracks or crevices, like dry mud ascomycete: informal term for fungi that produce spores within an ascus, such as morels, cup-fungi, and truffles; the group formally is referred to as the Ascomycota or Ascomycotina ascospore: a sexual spore produced within an ascus; characteristic of ascomycetes ascus: sac-like, usually elongated cell within which ascospores are produced; characteristic of ascomycetes astringent (taste): causing the mouth to pucker attached (gill attachment): describes gills that are joined to the stalk over at least a portion of their height (so not free) base: the lowest portion of the stalk basidiocarp: fruitbody of a basidiomycete basidiomycete: informal term for fungi that produce spores on a basidium, such as gilled mushrooms, boletes, coral mushrooms, jelly fungi, puffballs, and

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polypores; the group formally is referred to as the Basidiomycota or Basidiomycotina basidiospore: a sexual spore produced by a basidium; characteristic of basidiomycetes basidium (–ia): an often club-shaped cell that produces sexual basidiospores; characteristic of basidiomycetes bearded: adorned with hairs bloom: a thin surface coating with a delicate, typically whitish, powdery appearance, such as that often found on blueberries broadleaf: flowering trees with broad leaves such as alder, beech, birch, hickory, and oak; may be deciduous or evergreen; (= hardwood) buff: pale creamy yellow, brownish yellow, or grayish yellow bulb: swollen base of a stalk bulbous: having a swollen base button: young, unopened mushroom caespitose: see cespitose campanulate: bell-shaped canescence: a whitish to grayish dust-like bloom canescent: having a whitish to grayish dust-like bloom cap: the broad expanded umbrella-like portion of a typical mushroom, which bears gills, ridges, tubes, spines, or a smooth surface on its underside capitate: having a head or small knob at the tip carminophilous granules: see siderophilous granules cartilaginous: firm and tough but pliable caulocystidium (–ia): cystidium located on the surface of a stalk cellular: composed of more or less spherical cells cespitose (caespitose): growth form (habit) in which stalks of several fruitbodies arise very close together, but are not fused cheilocystidium (–ia): cystidium located on the edge of a gill or pore chlamydospore: a thick-walled spore produced asexually clamp connection: half bagel- or doughnut-like protrusion at the crosswall of a hypha in many basidiomycetes; often shortened to clamp clavate: club-shaped close (gill spacing): a relative term for gills that nearly touch each other but clearly have space between them; farther apart than crowded, but closer together than subdistant

cm: abbreviation for centimeter, unit of measurement in the metric system, equal to ¹⁄₁₀₀ meter, approximately ⅜ inch compressed (stalk shape): flattened in cross section concentric: occurring as a series of circles of different sizes, all having the same center concolorous: having the same color as something else conic (cap shape): cone-shaped conidium (–ia): an asexual spore conifer: a cone-bearing tree, usually evergreen, such as pine, spruce, or Douglas-fir connate: fused at the base and for some distance upward; in reference to mushrooms growing in a cluster context: the flesh of a mushroom, particularly that of the cap convex (cap shape): rounded upward, like an upsidedown bowl coprophilous: living on dung corrugated: coarsely ridged or wrinkled cortex (of stalk): rind, outer layer cortina: a silky fibrillose, cobwebby, partial veil, found in cortinariuses and certain other mushrooms, including some tricholomas cortinate: having a cortina costate: having ridges (costae) crenate, crenulate: scalloped, finely scalloped; usually in reference to the edge of a gill or margin of a cap cross-veined: having small ridges that connect adjoining gills crowded (gill spacing): a relative term for gills that very nearly touch each other; closer together than close cuneiform: shaped like a wedge cuticle (cutis): the outermost layer (skin) of a cap or stalk cutis: an outermost layer of the cap or stalk made up of hyphae aligned parallel to the surface, thus giving a smooth appearance to the surface cystidium (–ia): a sterile cell (actually a hyphal end), usually larger than and projecting beyond nearby cells; often with distinctive appearance and much used for identification purposes decurrent (gill attachment): gills that are attached to and extend down the stalk decurved: bent downward

Glossary

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dentate, denticulate: toothed, finely toothed depressed (cap shape): with a broadly sunken center dextrinoid: staining dark red or red-brown in Melzer’s reagent; compare amyloid dichotomous: split in two, forking, like a wishbone or tuning fork disc: the central area of the surface of a mushroom cap distant (gill spacing): a relative term for gills that are spaced very widely apart duff: plant litter making up the surface layer of forest soil; usually applied in conifer forests eccentric (or excentric): off-center, often in reference to the attachment of stalk to cap ectomycorrhiza: a mutualistic symbiosis between the hyphae of certain basidiomycetes and ascomycetes and the fine roots of certain (mostly woody) plants elliptical, ellipsoid: elongated with rounded ends and curved sides, usually in reference to spores emarginate (gill attachment): describes gills that are notched near the stalk and only narrowly attached; essentially the same as notched and, in practice, often very difficult to distinguish from sinuate and adnexed entire: even, smooth, not toothed or broken up; usually in reference to the cap margin or gill edge epithet: the second half of a species’s scientific name (the first half is the genus to which the species belongs) equal: of the same diameter throughout; usually in reference to a stalk eroded: irregularly toothed; usually in reference to the gill edge evanescent: present briefly, then vanishing; usually in reference to a cortina or annulus; (= fugacious) even: in reference to the cap cuticle, extending just to the tips of the gills, not projecting as a band of sterile tissue beyond them excentric: see eccentric farinaceous: odor or taste like that of freshly ground meal; when strong, reminiscent of cucumber or watermelon rind fascicles: bundles, like pine needles or sheaves of grain fertile surface: layer of tissue on which the spore-bearing structures are produced; (= hymenium) FeSO4: iron sulfate, usually a 10% aqueous solution; used macroscopically to test for color changes

fetid: having a heavy offensive odor, rotten, decayed, or fecal fiber: a hair-like structure fibril: a slender fiber fibrillose: composed of fibrils fibrous: tough and stringy filamentous: composed of filaments filiform: long and very slender floccose: with tufts of loose cottony material fragrant: having an odor that is more or less pleasant, but hard to describe in an unequivocal way free (gill attachment): describes gills that, from the beginning, are not at all attached to the stalk friable: easily breaking into pieces, crumbly fruitbody: macroscopic sexual reproductive structure produced by certain fungi; also known as sporocarp, carpophore, basidiocarp, basidiome, ascocarp, ascoma, or mushroom fugacious: present briefly, then vanishing; (= evanescent) fulvous: reddish cinnamon; colored like a red fox funga: all the species of fungi that inhabit a given area; (= mycoflora or mycota) furfuraceous: covered with coarsely granular or branlike particles; (= scurfy) fuscous: dark brownish gray to brownish black fusiform, fusoid: tapering from the middle toward both ends, spindle-shaped genera: plural of genus genus: a formally designated group of similar, presumably closely related, species gill trama: the sterile supporting tissue in the center of a gill, “sandwiched” between the two fertile layers gill: plate-, blade-, or sometimes fold-like structure, borne radially on the underside of a cap, and the faces of which are covered by fertile tissue; (= lamella) glabrous: smooth, naked glaucous: covered with a thin, whitish bloom that is easily rubbed off globose: more or less spherical gluten: a viscous substance resulting from the dissolution of gelatinous tissue glutinous: covered with gluten, gooey granulose: covered with small sugar-like particles gregarious (habit): fruitbodies occurring close together in a more or less dense group

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habit: morphological form, such as tricholomatoid, or manner of growth, such as solitary, scattered, gregarious, or clustered hardwood: flowering trees with broad leaves such as alder, beech, birch, hickory, and oak; may be deciduous or evergreen; (= broadleaf) herbaceous: describing plants that have no (or very little) woody tissue; most live for only one growing season herbarium: a permanent organized repository for dried or otherwise preserved fungus or plant specimens hoary: having a whitish to grayish sheen, like frost hyaline: transparent and nearly colorless; usually in reference to spores or other structures as viewed with a microscope hygrophanous: changing from translucent to opaque, and often markedly fading in color (expallent), upon drying; usually in reference to the mushroom’s cap surface hygroscopic: readily absorbing moisture and altering the tissue structure as a result hymeniform: made up of erect club-shaped to pearshaped cells arranged more or less perpendicularly to the surface hymenium: the spore-producing, or fertile, tissue of a mushroom, such as the layer of basidia and other cells on the face of a gill hypha (–ae): long tube-like element that, with many others, makes up the body (mycelium) of a filamentous fungus; may or may not be septate hypodermium: zone of tissue lying immediately beneath the cuticle (derm); usually in reference to the mushroom’s cap imbricate: overlapping, like roof shingles inferior ring: ring borne on the lower portion of a stalk infundibuliform: deeply depressed, funnel-like intervenose: having shallow cross-veins between adjacent gills involute: inrolled, especially in reference to the margin of a young mushroom cap iron sulfate: see FeSO4 isodiametric: having more or less equal diameters in all directions ixocutis: a cuticle consisting of gelatinized hyphae lying parallel to the surface ixolattice: a cuticle consisting of gelatinized, entangled,

branching hyphae arranged more or less perpendicularly to the surface ixotrichoderm: a cuticle consisting of gelatinized narrow hyphae of unequal length arranged nearly perpendicularly to the surface KOH: potassium hydroxide, usually a 3–5% aqueous solution; used, both microscopically and macroscopically, to test for color changes lageniform: flask- or gourd-shaped lamella (–ae): gill lamellula (–ae): a gill that does not extend the entire distance from cap margin to stalk large: (for purposes of this book) in reference to a mushroom, one with a cap diameter usually greater than 10 cm lateral: attached at one side, as in the short stalk of many shelf-fungi lato: broad; for instance, sensu lato, meaning “in a broad sense” lignicolous: growing on or in wood lobed: having rounded divisions lubricous: slightly greasy or slippery to the touch macrochemical: pertaining to the application of reagents to a mushroom and observing possible color changes with the unaided eye or a handlens (i.e., without the use of a microscope) macroscopic: visible with the unaided eye or at low magnification such as obtained with a 10-power handlens margin: edge; especially in reference to a cap or gill marginate: having a distinctly marked border, such as a gill with differently colored (darker) edge or bulbous stalk base with a sharp horizontal or upturned rim medium: (for purposes of this book) in reference to a mushroom, one with a cap diameter usually between 5 and 10 cm Melzer’s reagent: an iodine-containing solution used, both microscopically and macroscopically, to test for color changes; see amyloid and dextrinoid µm: abbreviation for micrometer, unit of measurement in the metric system, equal to one-millionth of a meter; informally (and decreasingly) referred to as a micron microscopic: visible only with a microscope mm: abbreviation for millimeter, unit of measurement in the metric system, equal to ¹⁄₁₀ centimeter and ¹⁄₁₀₀₀ meter, approximately 1⁄25 inch

Glossary

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mixed woodlands: wooded area or forest containing both conifer and hardwood trees morphologic(al): pertaining to the form and structure of an organism or other object mushroom: a loosely defined term for the large conspicuous fruitbodies of certain fungi, often confined to those with fleshy texture, those with cap and stalk, or those bearing gills mycelial cord: root-like structure formed by the aggregation of hyphae; similar to but structurally different from rhizomorphs mycelium (–ia): the body of a filamentous fungus, composed of a network of complexly branched and interconnected hyphae mycoflora: all the species of fungi that inhabit a given area; (= mycota or funga) mycologist: one who studies fungi mycology: the study of fungi mycoparasite: a fungus that parasitizes another fungus mycophagy, mycophagist: the eating of fungi, person or other animal that eats fungi mycorrhiza: a nearly universal mutualistic symbiosis between the hyphae of fungi and the roots of plants mycota: all the species of fungi that inhabit a given area; (= mycoflora or funga) NH4OH: ammonium hydroxide; used, both microscopically and macroscopically, to test for color changes nomen provisorium (nom. prov.): a provisionally proposed name non-amyloid: remaining colorless or yellowish in Melzer’s reagent notched (gill attachment): describes gills that have a wedge-like space where the gill turns upward before contacting the stalk; essentially the same as emarginate and, in practice, often very difficult to distinguish from sinuate and adnexed oblong: longer than wide, and with somewhat flattened ends obovate: inversely ovate (egg-shaped), with broadest portion uppermost obtuse: blunt, not pointed; compare with acute ocher: pale brownish orange-yellow; also spelled ochre ochraceous: having an ocher color olivaceous: having an olive-green color ornamentation: the surface features of a non-smooth spore, such as spines, warts, ridges, and wrinkles

ovate, ovoid: egg-shaped palisade: a side-by-side arrangement of elongated, perpendicular cells pallid: a very pale shade of any color, almost white papilla: a small, nipple-shaped projection papillate: having a papilla parasite: a fungus (or other organism) that obtains its nutrients from another living organism (its host), which suffers a negative impact as a result parasitism: a form of symbiosis, in which one organism (the parasite) obtains its nutrition from another living, and usually much larger, organism (the host), which suffers a negative impact as a result partial veil: a membranous or fibrillose tissue, occurring in some mushrooms, that extends from the cap margin to the stalk and initially covers the gills or pores pathogen: a parasitic or disease-causing organism PCR: acronym for polymerase chain reaction, a method used to produce multiple copies of selected segments of DNA molecules PDAB: acronym for paradimethylaminobenzaldehyde, used macroscopically to test for color changes pellicle: a viscid, peelable outermost layer of a mushroom cap pendant: hanging downward like an icicle or stalactite persistent: retaining its shape or position for a long time pileate: having a pileus (cap) pileipellis: the outermost layer of the pileus; (= cap cuticle) pileocystidium (–ia): cystidium located on the pileus (cap) surface pileus: cap plage: a smooth area near the point of attachment of a spore pleurocystidium (–ia): cystidium located on the face of a gill or tube pliant: flexible, can be bent without breaking plicate: folded like a fan or pleated skirt pore: small opening, or the mouth of the tubes of boletes and polypores potassium hydroxide: see KOH primordium (–ia): very young undeveloped mushroom pruinose: appearing to be covered with a fine powder

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pseudoparenchymatous: having the form of plant parenchyma cells, which are nearly isodiametric and look like crowded soap bubbles when in mass pubescent: finely hairy punctae: tiny scales or spots punctate: covered with tiny scales or spots pungent: sharp and piercing taste or smell Q (value): the ratio between a spore’s length and its width radial: pointed away from a common central point, like the spokes of a wheel radicating: with a root-like projection recurved: scale or fibril that is curved backward; also applies to a cap margin repent: prostrate reticulate: having a net-like pattern reticulum: net-like system of ridges; often in reference to a stalk or spore rhizomorph: root-like structure formed by the aggregation of hyphae and often found at the base of a stalk; similar to, but structurally different from, mycelial cords rimose: having distinct cracks or crevices ring: membranous skirt-like remnant of a partial veil borne on a stalk; (= annulus) ring-zone: band of fibrillose remnants of a partial veil borne on a stalk; often colored by the accumulation of falling spores; (= annular zone) rugose, rugulose: wrinkled, finely wrinkled russet: reddish brown saccate: shaped like a sack saprobe, saprotroph: an organism that obtains its nutrition by decomposing dead organic material scale: an erect, flattened, or recurved projection or torn portion of the cap or stalk surface scattered (habit): fruitbodies occurring closely enough together to be considered a group, but farther apart than for gregarious sclerotium (–ia): a hard, compact mass of hyphae, sometimes including soil, that is a resting stage for some fungi and from which a fruitbody may arise scrobiculate: with shallow, shiny pits or depressions scurfy: covered with coarsely granular or bran-like particles; (= furfuraceous) seceding: refers to gills that are initially attached, then tear away from the stalk as the fruitbody develops, usually leaving lines on the stalk surface

sensu: in the sense of, or according to septate: divided by crosswalls (septa) septum (–a): cross-wall; usually in reference to a hypha sericeous: silky serrate, serrulate: saw-toothed, finely toothed; usually in reference to a gill edge siderophilous granules: small dark-purple to nearly black particle observed in the basidia of some mushrooms when heated in acetocarmine sinuate (gill attachment): describes gills that are attached to the stalk along more or less their entire height, but which have a concave indentation a short distance from the stalk; in practice, often very hard to distinguish from emarginate, notched, and adnexed small: (for purposes of this book) in reference to a mushroom, one with a cap diameter usually less than 5 cm solitary (habit): fruitbodies occurring singly without others nearby sordid: dirty or dingy spathulate: shaped like a spoon or spatula, oval with a narrowed base species: the lowest category in the classification of organisms (although some species are further divided into subspecies, varieties, or forms); variously defined, but can be considered to represent a single kind or type of organism spermatic: resembling the odor of human seminal fluid spore: single- or sometimes multi-celled sexual or asexual reproductive propagule formed by fungi and many of the “lower” plants such as ferns and mosses; functionally similar to the seeds of gymnosperms and flowering plants spore-print (or spore-deposit): an accumulation of spores beneath, for instance, a mushroom’s cap; the color of the spore-print can be important in mushroom identification sporocarp: a fungal fruitbody; also known as carpophore, basidiocarp, basidiome, ascocarp, ascoma, or mushroom squamules: small scales squamulose: covered with small scales (squamules) squarrose: covered with stiff upright scales squarrulose: covered with small stiff upright scales stalk: the structure that arises from the substrate and

Glossary

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supports the cap of a mushroom; also referred to as stipe or stem stature: overall form, such as tall and slender or short and stocky stem: the stalk or stipe of a mushroom sterigma (–ata): branch-like extension of a basidium, on which a spore is formed sterile: non-fertile stipe: the stalk or stem of a mushroom stipitate: having a stipe (stalk) striate: having fine parallel or radiating lines or grooves; often in reference to a cap surface (especially the margin) or spore striatulate: having very fine, more or less parallel lines or furrows stricto: strict or narrow; for instance, sensu stricto, meaning “in a narrow or restricted sense” strigose: coated with long, coarse, stiff, usually appressed hairs stuffed: containing a soft tissue that usually disappears in age, leaving a hollow space; usually in reference to a stalk subdecurrent (gill attachment): extending slightly down the stalk subdistant (gill spacing): a relative term for gills that are farther apart than close, but closer together than distant substrate: substance on or within which the vegetative body of the fungus is growing and from which the mushroom arises; usually soil, woody material, leaf litter, or dung, but can be such things as insects or other mushrooms subulate: awl-shaped sulcate: having parallel or radial grooves, the depth of which is greater than in striate and less than in plicate superior ring: ring borne on the upper portion of a stalk synonym: an alternate name for the same organism tacky: slightly sticky tawny: dull yellowish brown, like a lion’s coat taxonomy: the naming and classifying of organisms terrestrial: growing on the ground toadstool: informal term for mushroom; sometimes used to refer to poisonous species tomentose: covered with long, soft, densely matted hairs; wooly

tomentum: covering of long, soft, matted hairs trama: see gill trama translucent-striate: appearing striate as a result of the gills being visible through the very thin flesh of the cap trichoderm: a cap cuticle with slender upright hyphae of unequal length, arranged perpendicularly to the surface troop: large numbers of fruitbodies in a dense group truncate: ending abruptly, appearing chopped off umbilicate: having a small central depression, like a navel; usually in reference to a cap umbo: knob, protrusion, or raised area at the center of a cap; may be narrow or broad, rounded or pointed umbonate: having an umbo universal veil: a membranous, fibrillose, or friable tissue that entirely covers an unopened mushroom; often leaves remnants in the form of a volva, bands on the stalk, or a patch or warts on the cap veil: a membranous, fibrillose, or friable tissue that covers all or part of a young mushroom; see partial veil and universal veil velutinous: coated with short, fine, soft hairs ventricose: fattest in the middle and tapering toward both ends; usually in reference to a stalk or a cystidium verrucose, verruculose: warty, finely warty vesiculose: bladder-like or somewhat spherical vinaceous: said to be the color of red wine; in practice, not so deeply colored and with duller gray to brown tones yielding more of a dark salmon-pink violaceous: having purplish tones virgate: streaked viscid: sticky volva: the remains of a universal veil, situated at the stalk base; may take several forms such as membranous and cup-like or friable bands of tissue zonate, zoned: with concentric bands of different color or different shades of one basic color; usually in reference to the cap surface

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References

The following list, although not exhaustive (for instance, field guides are not included), includes most of the literature on North American tricholomas, several European treatments of the genus, and a variety of other relevant publications. Ammirati, J. F., and C. L. Ovrebo. 1979. Type studies: some Cortinarius and Tricholoma species described by Charles Horton Peck. Mycotaxon 8(2): 353–371. Anand, J. Sein, P. Kabata, M. Barwina, and W. Waldman. 2011. Some aspects of intoxication with Tricholoma equestre. Clinical Toxicology 49: 230. Arde Jr., W. R. 1942. Studies in the genus Tricholoma I. Mycologia 34(4): 416–423. Atkinson, G. F. 1908. A new poisonous mushroom. Botanical Gazette 46: 461–463. Atkinson, G. F. 1909. Preliminary notes on some new species of Agaricaceae and Clavaria. Annales Mycologici 7(4): 365–376. Baroni, T. J., and C. L. Ovrebo. 1983. Tricholoma manzanitae—a new species from California. Mycotaxon 18(2): 299–302. Bedry, R., I. Baudrimont, G. Deffieux, E. E. Creppy, J. P. Pomies, J. M. Ragnaud, M. Dupon, D. Neau, C. Gabinski, S. De Witte, J. C. Chapalain, J. Beylot, and P. Godeau. 2001. Wild-mushroom intoxication as a cause of rhabdomyolysis. New England Journal of Medicine 345(11): 798–802. Bigelow, H. E. 1979. A contribution to Tricholoma. Beihefte zur Sydowia, Annales Mycologici Series II 8: 54–62. Bon, M. 1984. Les tricholomes de France et d’Europe occidentale. Paris, France: Editions Lechevalier S.A.R.L.

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Bon, M. 1991. Flore mycologique d’Europe 2: les tricholomes et ressemblants. Documents Mycologiques Mémoire Hors Série no. 2. Carriconde, F., M. Gardes, P. Jargeat, J. HeilmannClausen, B. Mouhamadou, and H. Gryta. 2008. Population evidence of cryptic species and geographical structure in the cosmopolitan ectomycorrhizal fungus, Tricholoma scalpturatum. Microbial Ecology 56: 513–524. Carriconde, F., H. Gryta, P. Jargeat, B. Mouhamadou, and M. Gardes. 2008. High sexual reproduction and limited contemporary dispersal in the ectomycorrhizal fungus, Tricholoma scalpturatum: New insights from population genetics and spatial autocorrelation analysis. Molecular Ecology 17: 4433–4445. Chapela, I. H., and M. Garbelotto. 2004. Phylogeography and evolution in matsutake and close allies inferred by analyses of ITS sequences and AFLPs. Mycologia 96(4): 730–741. Christensen, M., and J. Heilmann-Clausen. 2008. “Tricholoma (Fr.) P. Kumm.,” in Funga Nordica: Agaricoid, Boletoid, and Cyphelloid Genera, ed. H. Knudsen and J. Vesterholt, 413–429. Copenhagen, Denmark: Nordsvamp. Christensen, M., and J. Heilmann-Clausen. 2009. Two new boreal species of Tricholoma from Fennoscandia. Mycotaxon 107: 431–440. Comandini, O., I. Haug, A. C. Rinaldi, and T. W. Kuyper. 2004. Uniting Tricholoma sulphureum and T. bufonium. Mycological Research 108(10): 1162–1171. Contu, M. 2003. “Rare, critical, and new species of the genus Tricholoma collected in Sardinia, with two new records for its mycoflora.” [In Italian.] Micologia e Vegetazione Mediterranea 18(2): 91–100.

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Deng, H., and Y.-J. Yao. 2005. Tricholoma equestre, the correct name for T. flavovirens. Mycotaxon 94: 325–329. Ellis J. B. 1876. New fungi found at New Field, New Jersey (continued). Bulletin of the Torrey Botanical Club 6(14): 75-77. Ellis, J. B., and B. M. Everhart. 1893. New species of fungi from various localities. Proceedings of the Academy of Natural Sciences of Philadelphia 45(3): 440–466. Farlow, W. G., and E. A. Burt. 1929. Icones Farlowianae: Illustrations of the larger fungi of eastern North America. Cambridge, Massachusetts: The Farlow Library and Herbarium of Harvard University. Fries, E. M. 1821. Systema mycologicum, Vol. 1. Greifswald, Germany: Sumtibus Ernesti Mauritii. Fries, E. M. 1836–1838. Epicrisis systematis mycologici. Uppsala, Sweden: Typographia Academica. Galli, R. 1999. I tricolomi. Milan, Italy: Edinatura. Gryta, H., F. Carriconde, J. Y. Charcosset, P. Jargeat, and M. Gardes. 2006. Population dynamics of the ectomycorrhizal fungal species Tricholoma populinum and Tricholoma scalpturatum associated with black poplar under differing environmental conditions. Environmental Microbiology 8(5): 773–786. Hesler, L. R. 1958. Southeastern Agaricales II. Studies of Tricholoma types. Journal of the Tennessee Academy of Science 33: 186–191. Jargeat, P., F. Martos, F. Carriconde, H. Gryta, P.-A. Moreau, and M. Gardes. 2010. Phylogenetic species delimitation in ectomycorrhizal fungi and implications for barcoding: the case of the Tricholoma scalpturatum complex (Basidiomycota). Molecular Ecology 19: 5216–5230. Kauffman, C. H. 1918. The Agaricaceae of Michigan. Biological Series 5, Publication 26, Michigan Geological and Biological Survey. Kauffman, C. H. 1921. Mycological flora of the Rockies. Papers of the Michigan Academy of Science, Arts, and Letters 1: 101–150. Kibby, G. 2010. The genus Tricholoma in Britain. Field Mycology 11(4): 1131–40. Kummer, P. 1871. Der führer in die pilzkunde. Verlag von E. Luppe’s Buchhandlung. Leonard, L. M. 2000. Is it sinuate or emarginate? McIlvainea 14(2): 152–6.

Mankel, A., G. Kost, and E. Kothe. 1998. Re-evaluation of the phylogenetic relationship among species of the genus Tricholoma. Microbiological Research 153: 3773–3788. Marchand, A. 1986. Champignons du nord et du midi, Vol. 9: les tricholomes. Perpignan, France: Société Mycologique des Pyrénées Méditerranéennes. Mitchel, D. H., and A. H. Smith. 1978. Notes on Colorado fungi III: New and interesting mushrooms from the aspen zone. Mycologia 70(5): 1040–1063. Moreno, G., N. Ayala, and F. Esteve Raventós. 1997. Tricholoma albidulum sp. nov. (Tricholomataceae, Agaricales) from Baja California, Mexico. Mycotaxon 63: 383–387. Moser, M. 1978. Fungorum rariorum icones coloratae, Part VII. Vaduz, Germany: J. Cramer. Mouhamadou, B., F. Carriconde, H. Gryta, P. Jargeat, S. Manzi, and M. Gardes. 2008. Molecular evolution of mitochondrial ribosomal DNA in the fungal genus Tricholoma: Barcoding implications. Fungal Genetics and Biology 45: 1219–1226. Murrill, W. A. 1911. The Agaricaceae of tropical North America III. Mycologia 3(4): 189–199. Murrill, W. A. 1912a. New combinations. Mycologia 4(6): 327–332. Murrill, W. A. 1912b. The Agaricaceae of the Pacific Coast—I. Mycologia 4(4): 205–217. Murrill, W. A. 1913. The Agaricaceae of the Pacific Coast IV: new species of Clitocybe and Melanoleuca. Mycologia 5(4): 206–223. Murrill, W. A. 1914a. North American Flora, Volume 10, Part 1: Agaricales, Agaricaceae (pars). The New York Botanical Garden. Murrill, W. A. 1914b. A new fungous part of North American Flora. Mycologia 6(5): 264–269. Murrill, W. A. 1916. An attractive species of Melanoleuca from Oregon. Mycologia 8(2): 113. Murrill, W. A. 1917a. New combinations. Mycologia 9(1): 37–46. Murrill, W. A. 1917b. Notes: Melanoleuca pulverulentipes. Mycologia 9(3): 173–180. Murrill, W. A. 1938. New Florida agarics. Mycologia 30(4): 359–371. Murrill, W. A. 1939. Some Florida gill-fungi. Journal of the Elisha Mitchell Scientific Society 55: 361–372. Murrill, W. A. 1940. Additions to Florida fungi III.

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from North America associated with cottonwood. Brittonia 38(3): 260–263. Ovrebo, C. L., R. E. Halling, and T. J. Baroni. 1984. A note on Tricholoma niveipes. Mycotaxon 20(2): 473–478. Ovrebo, C. L., K. W. Hughes, and R. E. Halling. 2009. A preliminary phylogeny of Tricholoma based on the rRNA ITS region. Mycological Society of America annual meeting abstracts, Inoculum, supplement to Mycologia 60(3): 33. Ovrebo, C. L., and A. H. Smith. 1979. Studies on Tricholoma vaccinum and Tricholoma imbricatum in North America. The Michigan Botanist 18: 99–106. Ovrebo, C., and E. E. Tylutki. 1975. New species and a new combination of Tricholoma (Agaricales) from the Pacific Northwest. Mycologia 67(1): 75–82. Peck, C. H. (ARSB). 1872–1914. Report of State Botanist. From the Annual Report of the New York State Museum. Peck, C. H. 1873. Descriptions of new species of fungi. Bulletin of the Buffalo Society of Natural Science 1: 41–72. Peck, C. H. 1879. New species of fungi. Botanical Gazette 4(3): 137–139. Peck, C. H. 1895. New species of fungi. Bulletin of the Torrey Botanical Club 22(12): 485–493. Peck, C. H. 1897. New species of fungi. Bulletin of the Torrey Botanical Club 24(3): 137–147. Peck, C. H. 1898. New species of fungi. Bulletin of the Torrey Botanical Club 25(6): 321–328. Peck, C. H. 1899. New species of fungi. Bulletin of the Torrey Botanical Club 26(2): 63–71. Peck, C. H. 1900. New species of fungi. Bulletin of the Torrey Botanical Club 27(12): 609–613. Peck, C. H. 1902. New species of fungi. Bulletin of the Torrey Botanical Club 29(2): 69–74. Peck, C. H. 1904. New species of fungi. Bulletin of the Torrey Botanical Club 31(4): 177–182. Peck, C. H. 1909. New species of fungi. Bulletin of the Torrey Botanical Club 36(6): 329–339. Quélet, L. 1872–1875. Les champignons du Jura et des Vosges. Redhead, S. A. 1984. Mycological observations 13–14: on Hypsizygus and Tricholoma. Transactions of the Mycological Society of Japan 25: 1–9.

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Photo Credits

Most of the photographs in this book were taken by the authors. Those have been augmented by many excellent images from a number of other accomplished mushroom photographers, as listed below. Species names are listed by epithet only. Harley Barnhart: atroviolaceum(B), fulvum(E), nigrum(C), squarrulosum(C) sulphurescens(C) Kit Scates Barnhart: farinaceum, palustre(B) Dr. Michael Beug: intermedium(E), squarrulosum(B) Dr. Bruce Horn: odorum(B) Sava Krstic: arvernense(B), arvernense(C) Renée Lebeuf: argenteum(B), atrodiscum, cingulatum(A), fulvum(B–C–D), myomyces(B), pudorinum(A–B), roseoacerbum(A–B), scalpturatum(B), subluteum(B), transmutans(B) David Lewis: niveipes(C) Dr. Clark Ovrebo: luteomaculosum, palustre(A), venenatum(C) Andrew Parker: arvernense(A), atroviolaceum(A, C), focale(C–D), imbricatum(E), inamoenum(B), moseri(C), pardinum(B–C), populinum(B), portentosum(D), sulphurescens(D), sulphureum(C), vaccinum(E), virgatum(B) John Plischke III: roseoacerbum(C-1, C-2) Christian Schwarz: cingulatum(C) Noah Siegel: apium(A), aurantio-olivaceum, cingulatum(B), colossus(B), dryophilum(A), griseoviolaceum(A, C), manzanitae, moseri(D), NAT11, NAT-12, NAT-13 Fred Stevens: moseri(E) Walt Sturgeon: NAT-4(B) Michael Wood: dryophilum(B), griseoviolaceum(D), muricatum(B), myomyces(C), vernaticum(C)

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Index to Common Names

American matsutake, 4, 94 belted trich, 52 black-scaled trich, 43 brown birch trich, 73 brown matsutake, 50 canary trich, 60 dirty trich, 119 dusky trich, 130 earth-colored trich, 159 eastern brown-caps (key), 17 eastern gray-caps (key), 15 eastern white-caps (key), 12 eastern yellow-caps (key), 13 fibril trich, 172 giant trich, 54 girdled trich, 52 gray trich, 159 hot gray trich, 30 man-on-horseback, 60 matsutake, 5, 6, 94 orange-sheathed trich, 48

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pine mushroom, 94 poisonous trich, 119 poplar trich, 123 red-brown trich, 121 russet-scaly trich, 165 sand mushroom, 123 saw-tooth trich, 143 scaly trich, 165 shingled trich, 82 soap-scented trich, 134 sticky gray trich, 125 streaked trich, 172 sulfur trich, 157 the sandy, 123 western brown-caps (key), 25 western gray-caps (key), 23 western white-caps (key), 19 western yellow-caps (key), 21 white matsutake, 94 yellow trich, 60

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Index to Scientific Names

Pages on which species descriptions and illustrations appear are indicated by boldface.

Agaricus, 1 magnivelaris, 7 Amanita, 2 smithiana, 94 Armillaria, 1, 2, 5, 50 arenicola, 186 caligatum var. glaucescens, 50 caligatum var. occidentalis, 50 olida, 170 ponderosa, 94 viscidipes, 188 Armillariella, 2 Callistosporium luteo-olivaceum, 187 Calocybe, 2 fallax, 186 Catathelasma, 2 imperiale, 94 ventricosum, 94 Clitocybe, 2 Collybia, 2 Cortinellus, 4 Dermoloma, 2 Entoloma, 3 Floccularia, 2

Hygrophorus, 2 camarophyllus, 170 subalpinus, 170 Inocybe, 3 cystidiosa, 185 Laccaria, 2 Lactarius camphoratus, 36 helvus, 36 Lentinellus semivestitus, 187 Lepista, 2 rickenii, 187 sordida, 187 Leucopaxillus, 2 tricolor, 187 Leucopholiota decorosa, 185 Lyophyllum, 2 laticeps, 186 Macrocybe titans, 187 Megacollybia, 2 Megatricholoma colossus, 54 Melanoleuca, 2, 4, 34 melaleuca var. caespitosa, 186

Gymnopus, 2 Hebeloma, 3 Humidicutis czuica, 185 marginata, 186

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Penicillium janthinellum, 50 simplicissimum, 50 Porpoloma, 2, 3

Rhodocollybia, 2 Rhodocybe, 2 Rhodopaxillus, 2 Russula brevipes, 94 Tephrocybe, 2 Tricholoma acerbum, 132, 185 acre, 15, 23, 28, 30–31, 117, 130, 143 adustum, 185 aestuans, 14, 22, 28, 32–33, 64, 117 alabamense, 185 alachuanum, 185 albatum, 34, 151 albidulum, 151 albidum, 13, 34–35, 63, 151 albobrunneum, 185 alboconicum, 138 alboflavidum, 185 album, 185 altaicum, 185 angustifolium, 185 anomalum, 185 apium, 13, 17, 21, 25, 36–37, 94, 132 arenicola, 185 argenteum, 13, 15, 21, 23, 28, 38–39, 80, 100, 172 argyraceum, 52, 138, 185 aromaticum, 185 arvernense, 22, 23, 26, 27, 28, 40–41, 163 atrodiscum, 15, 28, 42, 80

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atrosquamosum, 5, 13, 16, 21, 24, 43, 44, 92, 110, 146 atroviolaceum, 24, 28, 43, 44–45, 92, 110 aurantio-olivaceum, 22, 26, 46–47, 48 aurantium, 17, 25, 48–49, 54, 66, 70 aurantium var. olivascens, 48 auratocephalum, 185 australe, 185 avellaneifolium, 125, 185 avellaneum, 185 azalearum, 185 batschii, 70 bicolor, 185 bresadolanum, 30 bufonium, 157 calceifolium, 185 californicum, 185 caligatum, 5, 12, 17, 20, 25, 50–51, 66, 94 cheilolaminum, 56 chlorinosmum, 185 chrysenteroides, 185 cingulatum, 12, 15, 20, 23, 52–53, 138 cinnamomeum, 185 citrinifolium, 185 collybiiforme, 185 colossus, 17, 28, 54–55, 70, 177 columbetta, 5, 34, 151 compressipes, 185 cortinatellum, 185 cortinatum, 185 cystidiosum, 185 czuicum, 185 davisiae, 15, 18, 22, 26, 28, 56–57, 177 decorosum, 185 decorum, 185 diemii, 185 dryophilum, 21, 27, 28, 58–59, 98, 164, 185 earleae, 186 eduriforme, 186 edurum, 186 entoloma, 186

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equestre, 1, 5, 14, 21, 32, 40, 60–62, 89, 140 fagnani, 186 fallax, 186 farinaceum, 20, 63 ferruginescens, 186 ferrugineum, 186 flavescens, 186 flavobrunneum, 73, 161 flavovirens, 1, 5, 60 floridanum, 14, 64–65 focale, 17, 25, 48, 54, 66–69 formosum, 186 fracticum, 25, 0–71, 121 fucatum, 116 fuligineum, 186 fulvellum, 164 fulvidiscum, 186 fulvimarginatum, 17, 72, 121, 123 fulvum, 18, 73–75, 161 fumescens, 186 fumidellum, 186 fumosellum, 186 fumosifolium, 186 fumosoluteum, 14, 76, 149 fusisporum, 186 glatfelteri, 186 glaucoalbum, 186 grande, 186 grave, 177, 186 griseoviolaceum, 20, 23, 28, 77–79, 107, 125, 170 harperi, 186 hebeloma, 186 hirtellum, 186 hordum, 15, 28, 42, 80, 172 humile var. bulbosum, 186 huronense, 16, 28, 81, 119, 168 hygrophorus, 186 imbricatum, 19, 27, 36, 46, 82–85, 132, 165 imbricatum var. felleum, 186 impolitum, 30 inamoenum, 12, 20, 28, 63, 86–87, 114, 154, 157 infantile, 186 inocybeoides, 52, 138 inocybiforme, 186



insigne, 16, 19, 28, 88 intermedium, 14, 22, 28, 40, 89–91, 149 ionides var. farinaceum, 186 jalapense, 186 jamaicense, 186 kauffmanii, 186 lacunosum, 186 lasciviforme, 186 laterarium, 186 laticeps, 186 latum, 186 leucocephaloides, 186 leucophyllum, 89 limonium, 186 longipes, 186 lugubre, 186 luteomaculans, 186 luteomaculosum, 14, 16, 22, 24, 28, 43, 44, 92–93, 116, 148 maculatescens, 186 maculatum, 186 magnivelare, 4, 5, 7, 12, 17, 20, 25, 50, 66, 94–97, 186 magnum, 186 malodorum, 186 manzanitae, 21, 27, 98–99, 121 margarita, 186 marginatum, 186 marquettense, 13, 15, 28, 100, 110, 116 matsutake, 4, 94 melaleuciforme, 186 melaleucum var. caespitosum, 186 memmingeri, 186 michiganense, 146, 186 microspermum, 186 moseri, 24, 26, 28, 101–103, 108, 138, 159 mostnyae, 186 multifolium, 186 multipunctum, 186 muricatum, 7, 27, 70, 104–105, 121 murinaceum, 30 murrillianum, 186 muskokense, 18, 106 mutabile, 23, 28, 77, 107, 110, 170

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myomyces, 17, 24, 28, 101, 108–109, 112, 138, 159 myomyces var. cystidiotum, 101, 108 naucoria, 186 nictitans, 161 nigrocystidium, 186 nigromarginatum, 43 nigrum, 25, 28, 100, 107, 110–111, 119 niveipes, 15, 18, 19, 28, 100, 112–113, 125 nobile, 186 nuciolens, 186 odoriferum, 186 odorum, 14, 21, 28, 86, 114–115, 154, 157 olesonii, 186 olivaceiflavum, 186 olivaceobrunneum, 15, 18, 28, 100, 116 oliveum, 186 oreades, 186 orirubens, 5, 43, 88, 159, 186 pallidum, 186 palustre, 14, 28, 32, 117–118 panaeoliforme, 186 pardinum, 13, 17, 21, 24, 28, 81, 110, 119–120, 168 peralbum, 186 pessundatum, 18, 27, 58, 70, 72, 82, 98, 104, 106, 121–122, 123, 161, 164, 165 pinicola, 186 piperatiforme, 186 piperatum, 186 planiceps, 186 platyphylloides, 186 platyphyllum, 86 populinum, 5, 18, 27, 70, 72, 121, 123–124 portentosum, 5, 15, 23, 77, 112, 116, 125–127, 140, 170, 172 portolense, 186 praebulbosum, 186 praecox, 187 praemagnum, 187 praevillosum, 187



psammopus, 46, 187 pseudolilaceum, 187 pseudonictitans, 73, 161 pseudosordidum, 187 pseudoterreum, 187 pudorinum, 18, 128–129 pullum, 16, 28, 130–131, 172 pulverulentipes, 187 quercetorum, 176, 177, 187 quercicola, 187 radicatum, 187 ramentaceum, 138 rancidulum, 187 resplendens, 151, 154, 187 rhizoideum, 187 rickenii, 187 rimosum, 187 robinsoniae, 187 robustum, 66 roseibrunneum, 187 roseoacerbum, 18, 28, 132–133 rubescentifolium, 187 rubicundum, 187 rudericola, 187 russuloides, 187 rutilans, 187 saponaceum, 14, 16, 17, 22, 24, 25, 28, 64, 76, 134–137, 177 saponaceum var. ardosiacum, 134 saponaceum var. squamosum, 134 scalpturatum, 5, 16, 19, 24, 26, 52, 92, 101, 108, 138–139, 159 sciodes, 130 secedifolium, 187 sejunctum, 14, 22, 28, 32, 40, 60, 89, 117, 125, 140–142, 149 semivestitum, 187 serratifolium, 13, 19, 28, 30, 143–144 sienna, 187 silvaticoides, 13, 145 silvaticum, 145, 187 sordidum, 187 squarrulosum, 16, 24, 43, 146–147 stans, 121, 187 striatellum, 187 striatifolium, 187 subacidum, 187



subacre, 187 subacutum, 172, 187 subannulatum, 70, 187 subargillaceum, 187 subaureum, 15, 19, 28, 148 subcerinum, 187 subcinereiforme, 187 subcinereum, 187 subcylindrisporum, 187 subdecorosum, 187 subdurum, 187 subfuligineum, 187 subfulvidiscum, 187 subisabellinum, 187 sublatum, 187 subluridum, 187 subluteum, 14, 76, 89, 140, 149–150 submaculatum, 187 submulticeps, 187 subpessundatum, 187 subresplendens, 5, 13, 28, 34, 40, 63, 143, 151–153 subrimosum, 187 subrufescens, 187 subsaponaceum, 187 subsejunctum, 140, 187 subsilvaticum, 187 subterreiforme, 187 subterreum, 187 subtransmutans, 187 subumbrinum, 187 subvelatum, 187 subvolkertii, 187 sulphurescens, 12, 14, 20, 21, 28, 36, 86, 114, 154–156, 157 sulphureum, 14, 21, 28, 86, 114, 146, 154, 157–158 tenuipes, 187 terraeolens, 187 terreum, 5, 17, 19, 25, 27, 28, 30, 43, 108, 119, 138, 159–160 terriferum, 121 thujinum, 187 titans, 187 totilividum, 187 tottenii, 187 transmutans, 18, 27, 121, 161–162

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trentonense, 187 tricolor, 187 tridentinum, 121 tristiforme, 187 tumidum, 22, 28, 40, 163 umbrosum, 187 unakense, 187 unifactum, 187 ustale, 18, 27, 58, 121, 128, 164 ustaliforme, 188 ustaloides, 70, 188 vaccinum, 19, 26, 46, 82, 165–167 venenatum, 13, 19, 21, 26, 28, 81, 119, 143, 168–169 vernaticum, 20, 26, 28, 77, 170–171 virescens, 188 virgatum, 13, 15, 21, 23, 28, 30, 38, 80, 100, 119, 125, 130, 172–173 virgatum var. vinaceum, 172 virgineum, 188 viridilutescens, 140 viscidipes, 188 viscosum, 188 volkertii, 188 watsonii, 188 westiana, 188 westii, 188 yatesii, 188 zelleri, 66 Tricholomopsis, 2, 3 decora, 185 formosa, 186 rutilans, 187 Tricholosporum, 2, 3

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